{"entryType":"UniProtKB reviewed (Swiss-Prot)","primaryAccession":"O76365","uniProtkbId":"CTU1_CAEEL","entryAudit":{"firstPublicDate":"2009-04-14","lastAnnotationUpdateDate":"2026-01-28","lastSequenceUpdateDate":"1998-11-01","entryVersion":135,"sequenceVersion":1},"annotationScore":5.0,"organism":{"scientificName":"Caenorhabditis elegans","taxonId":6239,"lineage":["Eukaryota","Metazoa","Ecdysozoa","Nematoda","Chromadorea","Rhabditida","Rhabditina","Rhabditomorpha","Rhabditoidea","Rhabditidae","Peloderinae","Caenorhabditis"]},"proteinExistence":"1: Evidence at protein level","proteinDescription":{"recommendedName":{"fullName":{"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_03053"}],"value":"Cytoplasmic tRNA 2-thiolation protein 1"},"ecNumbers":[{"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_03053"}],"value":"2.7.7.-"}]},"alternativeNames":[{"fullName":{"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_03053"}],"value":"Cytoplasmic tRNA adenylyltransferase 1"}},{"fullName":{"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_03053"}],"value":"Thiolation of uridine in tRNA protein 1"}}]},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_03053"}],"value":"tut-1"},"synonyms":[{"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_03053"}],"value":"ctu-1"}],"orfNames":[{"value":"F29C4.6"}]}],"comments":[{"texts":[{"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_03053"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18391219"}],"value":"Plays a central role in 2-thiolation of mcm(5)S(2)U at tRNA wobble positions of tRNA(Lys), tRNA(Glu) and tRNA(Gln). Directly binds tRNAs and probably acts by catalyzing adenylation of tRNAs, an intermediate required for 2-thiolation. It is unclear whether it acts as a sulfurtransferase that transfers sulfur from thiocarboxylated URM1 onto the uridine of tRNAs at wobble position"}],"commentType":"FUNCTION"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_03053"}],"value":"tRNA modification; 5-methoxycarbonylmethyl-2-thiouridine-tRNA biosynthesis"}],"commentType":"PATHWAY"},{"commentType":"SUBCELLULAR LOCATION","subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_03053"}],"value":"Cytoplasm","id":"SL-0086"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18391219"}],"value":"Required for normal germline maturation and for viability. Viable at 20 degrees Celsius. At a higher temperature (25 degrees Celsius), the hatched eggs progress through the L1-L4 larval stages with kinetics similar to those of wild type but present an important delay in germline maturation. Whereas gravid wild-type worms are present 53 hours after hatching, germline maturation are still ongoing in the mutant after 66 hours, and the first adults containing eggs are observed 75 hours after hatching, when a large number of new L1 are already present in the wild type. Strikingly, the eggs that finally appeared in the mutant display aberrant morphology and size resulting in both low progeny and high mortality"}],"commentType":"DISRUPTION PHENOTYPE"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_03053"}],"value":"Belongs to the TtcA family. CTU1/NCS6/ATPBD3 subfamily"}],"commentType":"SIMILARITY"}],"features":[{"type":"Chain","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":373,"modifier":"EXACT"}},"description":"Cytoplasmic tRNA 2-thiolation protein 1","featureId":"PRO_0000368239"}],"keywords":[{"id":"KW-0963","category":"Cellular component","name":"Cytoplasm"},{"id":"KW-1185","category":"Technical term","name":"Reference proteome"},{"id":"KW-0694","category":"Molecular function","name":"RNA-binding"},{"id":"KW-0808","category":"Molecular function","name":"Transferase"},{"id":"KW-0819","category":"Biological process","name":"tRNA processing"},{"id":"KW-0820","category":"Molecular function","name":"tRNA-binding"}],"references":[{"referenceNumber":1,"citation":{"id":"9851916","citationType":"journal article","authoringGroup":["The C. elegans sequencing consortium"],"citationCrossReferences":[{"database":"PubMed","id":"9851916"},{"database":"DOI","id":"10.1126/science.282.5396.2012"}],"title":"Genome sequence of the nematode C. elegans: a platform for investigating biology.","publicationDate":"1998","journal":"Science","firstPage":"2012","lastPage":"2018","volume":"282"},"referencePositions":["NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]"],"referenceComments":[{"value":"Bristol N2","type":"STRAIN"}]},{"referenceNumber":2,"citation":{"id":"18391219","citationType":"journal article","authors":["Dewez M.","Bauer F.","Dieu M.","Raes M.","Vandenhaute J.","Hermand D."],"citationCrossReferences":[{"database":"PubMed","id":"18391219"},{"database":"DOI","id":"10.1073/pnas.0709404105"}],"title":"The conserved wobble uridine tRNA thiolase Ctu1-Ctu2 is required to maintain genome integrity.","publicationDate":"2008","journal":"Proc. Natl. Acad. Sci. U.S.A.","firstPage":"5459","lastPage":"5464","volume":"105"},"referencePositions":["FUNCTION IN 2-THIOLATION OF TRNA","DISRUPTION PHENOTYPE"]},{"referenceNumber":3,"citation":{"id":"19145231","citationType":"journal article","authors":["Leidel S.","Pedrioli P.G.A.","Bucher T.","Brost R.","Costanzo M.","Schmidt A.","Aebersold R.","Boone C.","Hofmann K.","Peter M."],"citationCrossReferences":[{"database":"PubMed","id":"19145231"},{"database":"DOI","id":"10.1038/nature07643"}],"title":"Ubiquitin-related modifier Urm1 acts as a sulphur carrier in thiolation of eukaryotic transfer RNA.","publicationDate":"2009","journal":"Nature","firstPage":"228","lastPage":"233","volume":"458"},"referencePositions":["TRNA-BINDING"]}],"uniProtKBCrossReferences":[{"database":"EMBL","id":"FO080227","properties":[{"key":"ProteinId","value":"CCD62179.1"},{"key":"Status","value":"-"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"PIR","id":"T33145","properties":[{"key":"EntryName","value":"T33145"}]},{"database":"RefSeq","id":"NP_499865.1","properties":[{"key":"NucleotideSequenceId","value":"NM_067464.5"}]},{"database":"AlphaFoldDB","id":"O76365","properties":[{"key":"Description","value":"-"}]},{"database":"SMR","id":"O76365","properties":[{"key":"Description","value":"-"}]},{"database":"BioGRID","id":"41989","properties":[{"key":"Interactions","value":"2"}]},{"database":"FunCoup","id":"O76365","properties":[{"key":"Number of interactors","value":"1404"}]},{"database":"STRING","id":"6239.F29C4.6.1","properties":[{"key":"Description","value":"-"}]},{"database":"PaxDb","id":"6239-F29C4.6.1","properties":[{"key":"Description","value":"-"}]},{"database":"PeptideAtlas","id":"O76365","properties":[{"key":"Description","value":"-"}]},{"database":"EnsemblMetazoa","id":"F29C4.6.1","properties":[{"key":"ProteinId","value":"F29C4.6.1"},{"key":"GeneId","value":"WBGene00017928"}]},{"database":"EnsemblMetazoa","id":"F29C4.6.2","properties":[{"key":"ProteinId","value":"F29C4.6.2"},{"key":"GeneId","value":"WBGene00017928"}]},{"database":"GeneID","id":"176826","properties":[{"key":"Description","value":"-"}]},{"database":"KEGG","id":"cel:CELE_F29C4.6","properties":[{"key":"Description","value":"-"}]},{"database":"UCSC","id":"F29C4.6.1","properties":[{"key":"OrganismName","value":"c. elegans"}]},{"database":"AGR","id":"WB:WBGene00017928","properties":[{"key":"Description","value":"-"}]},{"database":"CTD","id":"176826","properties":[{"key":"Description","value":"-"}]},{"database":"WormBase","id":"F29C4.6","properties":[{"key":"ProteinId","value":"CE17723"},{"key":"GeneId","value":"WBGene00017928"},{"key":"GeneName","value":"tut-1"}]},{"database":"eggNOG","id":"KOG2840","properties":[{"key":"ToxonomicScope","value":"Eukaryota"}]},{"database":"HOGENOM","id":"CLU_026481_1_2_1","properties":[{"key":"Description","value":"-"}]},{"database":"InParanoid","id":"O76365","properties":[{"key":"Description","value":"-"}]},{"database":"OMA","id":"KPVRGIC","properties":[{"key":"Fingerprint","value":"-"}]},{"database":"OrthoDB","id":"198857at2759","properties":[{"key":"Description","value":"-"}]},{"database":"PhylomeDB","id":"O76365","properties":[{"key":"Description","value":"-"}]},{"database":"UniPathway","id":"UPA00988","properties":[{"key":"RectionId","value":"-"}]},{"database":"PRO","id":"PR:O76365","properties":[{"key":"Description","value":"-"}]},{"database":"Proteomes","id":"UP000001940","properties":[{"key":"Component","value":"Chromosome IV"}]},{"database":"Bgee","id":"WBGene00017928","properties":[{"key":"ExpressionPatterns","value":"Expressed in germ line (C elegans) and 4 other cell types or tissues"}]},{"database":"GO","id":"GO:0005829","properties":[{"key":"GoTerm","value":"C:cytosol"},{"key":"GoEvidenceType","value":"ISS:UniProtKB"}]},{"database":"GO","id":"GO:0002144","properties":[{"key":"GoTerm","value":"C:cytosolic tRNA wobble base thiouridylase complex"},{"key":"GoEvidenceType","value":"IBA:GO_Central"}]},{"database":"GO","id":"GO:0016779","properties":[{"key":"GoTerm","value":"F:nucleotidyltransferase activity"},{"key":"GoEvidenceType","value":"IEA:UniProtKB-UniRule"}]},{"database":"GO","id":"GO:0016783","properties":[{"key":"GoTerm","value":"F:sulfurtransferase activity"},{"key":"GoEvidenceType","value":"IMP:WormBase"}],"evidences":[{"evidenceCode":"ECO:0000315","source":"PubMed","id":"19145231"}]},{"database":"GO","id":"GO:0000049","properties":[{"key":"GoTerm","value":"F:tRNA binding"},{"key":"GoEvidenceType","value":"IDA:WormBase"}],"evidences":[{"evidenceCode":"ECO:0000314","source":"PubMed","id":"19145231"}]},{"database":"GO","id":"GO:0048598","properties":[{"key":"GoTerm","value":"P:embryonic morphogenesis"},{"key":"GoEvidenceType","value":"IGI:WormBase"}],"evidences":[{"evidenceCode":"ECO:0000316","source":"PubMed","id":"19593383"}]},{"database":"GO","id":"GO:0048599","properties":[{"key":"GoTerm","value":"P:oocyte development"},{"key":"GoEvidenceType","value":"IGI:WormBase"}],"evidences":[{"evidenceCode":"ECO:0000316","source":"PubMed","id":"19593383"}]},{"database":"GO","id":"GO:0032447","properties":[{"key":"GoTerm","value":"P:protein urmylation"},{"key":"GoEvidenceType","value":"IEA:UniProtKB-UniRule"}]},{"database":"GO","id":"GO:0007283","properties":[{"key":"GoTerm","value":"P:spermatogenesis"},{"key":"GoEvidenceType","value":"IGI:WormBase"}],"evidences":[{"evidenceCode":"ECO:0000316","source":"PubMed","id":"19593383"}]},{"database":"GO","id":"GO:0006412","properties":[{"key":"GoTerm","value":"P:translation"},{"key":"GoEvidenceType","value":"IMP:WormBase"}],"evidences":[{"evidenceCode":"ECO:0000315","source":"PubMed","id":"19593383"}]},{"database":"GO","id":"GO:0034227","properties":[{"key":"GoTerm","value":"P:tRNA thio-modification"},{"key":"GoEvidenceType","value":"IMP:WormBase"}],"evidences":[{"evidenceCode":"ECO:0000315","source":"PubMed","id":"19145231"}]},{"database":"GO","id":"GO:0002143","properties":[{"key":"GoTerm","value":"P:tRNA wobble position uridine thiolation"},{"key":"GoEvidenceType","value":"IBA:GO_Central"}]},{"database":"GO","id":"GO:0002098","properties":[{"key":"GoTerm","value":"P:tRNA wobble uridine modification"},{"key":"GoEvidenceType","value":"IMP:WormBase"}],"evidences":[{"evidenceCode":"ECO:0000315","source":"PubMed","id":"19593383"}]},{"database":"GO","id":"GO:0040025","properties":[{"key":"GoTerm","value":"P:vulval development"},{"key":"GoEvidenceType","value":"IGI:WormBase"}],"evidences":[{"evidenceCode":"ECO:0000316","source":"PubMed","id":"19593383"}]},{"database":"CDD","id":"cd01713","properties":[{"key":"EntryName","value":"CTU1-like"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"3.40.50.620:FF:000132","properties":[{"key":"EntryName","value":"Cytoplasmic tRNA 2-thiolation protein 1"},{"key":"MatchStatus","value":"1"}]},{"database":"Gene3D","id":"3.40.50.620","properties":[{"key":"EntryName","value":"HUPs"},{"key":"MatchStatus","value":"1"}]},{"database":"HAMAP","id":"MF_03053","properties":[{"key":"EntryName","value":"CTU1"},{"key":"MatchStatus","value":"1"}]},{"database":"InterPro","id":"IPR056369","properties":[{"key":"EntryName","value":"CTU1-like_ATP-bd"}]},{"database":"InterPro","id":"IPR032442","properties":[{"key":"EntryName","value":"CTU1_C"}]},{"database":"InterPro","id":"IPR000541","properties":[{"key":"EntryName","value":"Ncs6/Tuc1/Ctu1"}]},{"database":"InterPro","id":"IPR014729","properties":[{"key":"EntryName","value":"Rossmann-like_a/b/a_fold"}]},{"database":"InterPro","id":"IPR011063","properties":[{"key":"EntryName","value":"TilS/TtcA_N"}]},{"database":"InterPro","id":"IPR035107","properties":[{"key":"EntryName","value":"tRNA_thiolation_TtcA_Ctu1"}]},{"database":"NCBIfam","id":"TIGR00269","properties":[{"key":"EntryName","value":"TIGR00269 family protein"},{"key":"MatchStatus","value":"1"}]},{"database":"PANTHER","id":"PTHR11807","properties":[{"key":"EntryName","value":"ATPASES OF THE PP SUPERFAMILY-RELATED"},{"key":"MatchStatus","value":"1"}]},{"database":"PANTHER","id":"PTHR11807:SF12","properties":[{"key":"EntryName","value":"CYTOPLASMIC TRNA 2-THIOLATION PROTEIN 1"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF01171","properties":[{"key":"EntryName","value":"ATP_bind_3"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF16503","properties":[{"key":"EntryName","value":"zn-ribbon_14"},{"key":"MatchStatus","value":"1"}]},{"database":"PIRSF","id":"PIRSF004976","properties":[{"key":"EntryName","value":"ATPase_YdaO"},{"key":"MatchStatus","value":"1"}]},{"database":"SUPFAM","id":"SSF52402","properties":[{"key":"EntryName","value":"Adenine nucleotide alpha hydrolases-like"},{"key":"MatchStatus","value":"1"}]}],"sequence":{"value":"MEKRRGPPPCQSGSGCSNPAKIRKAKDGAQLCGPCFSRNFEDDVHEAIVNNKLFKRGERVAIGASGGKDSTVLAYVMKTLNDRHDYGLDLQLLSIDEGIKGYRDDSLLAVEKNRVEYGLPLTILSYRDLYGWTMDDIVAKIGKKNNCTFCGVFRRQALDRGAFKIGATKLVTGHNADDMAETLLMNVLRGDIARLERCTNIVTGEEGDLPRAKPLKYCFERDIVMYARTNQLEYFYTECIYAPNAYRGYARKYVRDLEKVHPRAILDLIRSGEKVSVKKEVEMPTLKICERCGYMTSQKLCKACLLIEGLNTGNTDLGVRKSKKSKKVTVEADELNKEGGCGSGGGGGGCGCAGAEDAAENEETRQRLKDLQF","length":373,"molWeight":41332,"crc64":"B5E4A95EE348B841","md5":"EC29CC0023416E35EAFC153CF9A055C0"},"extraAttributes":{"countByCommentType":{"FUNCTION":1,"PATHWAY":1,"SUBCELLULAR LOCATION":1,"DISRUPTION PHENOTYPE":1,"SIMILARITY":1},"countByFeatureType":{"Chain":1},"uniParcId":"UPI00000801A2"}}