{"entryType":"UniProtKB reviewed (Swiss-Prot)","primaryAccession":"Q93008","secondaryAccessions":["O75550","Q8WWT3","Q8WX12"],"uniProtkbId":"USP9X_HUMAN","entryAudit":{"firstPublicDate":"1997-11-01","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"2020-10-07","entryVersion":234,"sequenceVersion":4},"annotationScore":5.0,"organism":{"scientificName":"Homo sapiens","commonName":"Human","taxonId":9606,"lineage":["Eukaryota","Metazoa","Chordata","Craniata","Vertebrata","Euteleostomi","Mammalia","Eutheria","Euarchontoglires","Primates","Haplorrhini","Catarrhini","Hominidae","Homo"]},"proteinExistence":"1: Evidence at protein level","proteinDescription":{"recommendedName":{"fullName":{"value":"Ubiquitin carboxyl-terminal hydrolase 9X"},"ecNumbers":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18254724"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19135894"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25944111"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"26598551"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"29626158"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"30914461"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"37454738"}],"value":"3.4.19.12"}]},"alternativeNames":[{"fullName":{"value":"Deubiquitinating enzyme FAF-X"}},{"fullName":{"value":"Fat facets in mammals"},"shortNames":[{"value":"hFAM"}]},{"fullName":{"value":"Fat facets protein-related, X-linked"}},{"fullName":{"value":"Ubiquitin thioesterase FAF-X"}},{"fullName":{"value":"Ubiquitin-specific protease 9, X chromosome"}},{"fullName":{"value":"Ubiquitin-specific-processing protease FAF-X"}}]},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000303","source":"PubMed","id":"18254724"},{"evidenceCode":"ECO:0000312","source":"HGNC","id":"HGNC:12632"}],"value":"USP9X"},"synonyms":[{"value":"DFFRX"},{"value":"FAM"},{"value":"USP9"}]}],"comments":[{"texts":[{"evidences":[{"evidenceCode":"ECO:0000250","source":"UniProtKB","id":"P70398"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16322459"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18254724"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19135894"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22371489"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"24607389"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25944111"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"26598551"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"29626158"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"30914461"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"33378666"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"34404733"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"37454738"}],"value":"Deubiquitinase involved both in the processing of ubiquitin precursors and of ubiquitinated proteins (PubMed:18254724, PubMed:19135894, PubMed:22371489, PubMed:25944111, PubMed:29626158, PubMed:30914461, PubMed:37454738). May therefore play an important regulatory role at the level of protein turnover by preventing degradation of proteins through the removal of conjugated ubiquitin (PubMed:18254724, PubMed:19135894, PubMed:22371489, PubMed:25944111, PubMed:29626158, PubMed:30914461, PubMed:37454738). Specifically hydrolyzes 'Lys-11'-, followed by 'Lys-63'-, 'Lys-48'- and 'Lys-6'-linked polyubiquitins chains (PubMed:30914461). Essential component of TGF-beta/BMP signaling cascade (PubMed:19135894). Specifically deubiquitinates monoubiquitinated SMAD4, opposing the activity of E3 ubiquitin-protein ligase TRIM33 (PubMed:19135894). Deubiquitinates alkylation repair enzyme ALKBH3 (PubMed:25944111). OTUD4 recruits USP7 and USP9X to stabilize ALKBH3, thereby promoting the repair of alkylated DNA lesions (PubMed:25944111). Deubiquitinates RNA demethylase enzyme ALKBH5, promoting its stability (PubMed:37454738). Deubiquitinates mTORC2 complex component RICTOR at 'Lys-294' by removing 'Lys-63'-linked polyubiquitin chains, stabilizing RICTOR and enhancing its binding to MTOR, thus promoting mTORC2 complex assembly (PubMed:33378666). Regulates chromosome alignment and segregation in mitosis by regulating the localization of BIRC5/survivin to mitotic centromeres (PubMed:16322459). Involved in axonal growth and neuronal cell migration (PubMed:24607389). Regulates cellular clock function by enhancing the protein stability and transcriptional activity of the core circadian protein BMAL1 via its deubiquitinating activity (PubMed:29626158). Acts as a regulator of peroxisome import by mediating deubiquitination of PEX5: specifically deubiquitinates PEX5 monoubiquitinated at 'Cys-11' following its retrotranslocation into the cytosol, resetting PEX5 for a subsequent import cycle (PubMed:22371489). Deubiquitinates PEG10 (By similarity). Inhibits the activation of the Hippo signaling pathway via deubiquitination of AMOTL2 at 'Lys-347' and 'Lys-408' which prohibits its interaction with and activation of LATS2. Loss of LATS2 activation and subsequent loss of YAP1 phosphorylation results in an increase in YAP1-driven transcription of target genes (PubMed:26598551, PubMed:34404733)"}],"commentType":"FUNCTION"},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"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).","ecNumber":"3.4.19.12","evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18254724"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19135894"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"22371489"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25944111"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"26598551"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"29626158"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"30914461"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"34404733"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"37454738"}]}},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"16322459"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"18254724"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19135894"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"24607389"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"25944111"}],"value":"Interacts with SMAD4, MARK4, NUAK1 and BIRC5/survivin (PubMed:16322459, PubMed:18254724, PubMed:19135894). Interacts with DCX (PubMed:24607389). Interacts with OTUD4 and USP7; the interaction is direct (PubMed:25944111)"}],"commentType":"SUBUNIT"},{"commentType":"INTERACTION","interactions":[{"interactantOne":{"uniProtKBAccession":"Q93008","intActId":"EBI-302524"},"interactantTwo":{"uniProtKBAccession":"P42858","geneName":"HTT","intActId":"EBI-466029"},"numberOfExperiments":8,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"Q93008","intActId":"EBI-302524"},"interactantTwo":{"uniProtKBAccession":"Q07820","geneName":"MCL1","intActId":"EBI-1003422"},"numberOfExperiments":10,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"Q93008","intActId":"EBI-302524"},"interactantTwo":{"uniProtKBAccession":"O60285","geneName":"NUAK1","intActId":"EBI-1046789"},"numberOfExperiments":2,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"Q93008","intActId":"EBI-302524"},"interactantTwo":{"uniProtKBAccession":"Q13485","geneName":"SMAD4","intActId":"EBI-347263"},"numberOfExperiments":2,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"Q93008","intActId":"EBI-302524"},"interactantTwo":{"uniProtKBAccession":"P40337","geneName":"VHL","intActId":"EBI-301246"},"numberOfExperiments":5,"organismDiffer":false},{"interactantOne":{"uniProtKBAccession":"Q93008","intActId":"EBI-302524"},"interactantTwo":{"uniProtKBAccession":"P08393","geneName":"ICP0","intActId":"EBI-6148881"},"numberOfExperiments":3,"organismDiffer":true},{"interactantOne":{"uniProtKBAccession":"Q93008","intActId":"EBI-302524"},"interactantTwo":{"uniProtKBAccession":"P0DTC9","geneName":"N","intActId":"EBI-25475856"},"numberOfExperiments":3,"organismDiffer":true}]},{"commentType":"SUBCELLULAR LOCATION","subcellularLocations":[{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"19135894"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"26833328"}],"value":"Cytoplasm, cytosol","id":"SL-0091"}},{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"24607389"}],"value":"Cell projection, growth cone","id":"SL-0288"}},{"location":{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"26833328"}],"value":"Cytoplasm, cytoskeleton, cilium axoneme","id":"SL-0304"}}]},{"commentType":"ALTERNATIVE PRODUCTS","events":["Alternative splicing"],"isoforms":[{"name":{"value":"1"},"synonyms":[{"value":"Short"}],"isoformIds":["Q93008-1"],"isoformSequenceStatus":"Displayed"},{"name":{"value":"2"},"synonyms":[{"value":"Long"}],"isoformIds":["Q93008-3"],"sequenceIds":["VSP_060711"],"isoformSequenceStatus":"Described"}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"8922996"}],"value":"Widely expressed in embryonic and adult tissues"}],"commentType":"TISSUE SPECIFICITY"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"33378666"}],"value":"By growth factors"}],"commentType":"INDUCTION"},{"commentType":"DISEASE","disease":{"diseaseId":"Intellectual developmental disorder, X-linked 99","diseaseAccession":"DI-04101","acronym":"XLID99","description":"A disorder characterized by significantly below average general intellectual functioning associated with impairments in adaptive behavior and manifested during the developmental period. 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MRXS99F affected females manifest intellectual disability, developmental delay, facial dysmorphism, short stature, and distinct congenital malformations comprising choanal atresia, anal abnormalities, post-axial polydactyly, heart defects, hypomastia, cleft palate/bifid uvula, progressive scoliosis, and structural brain abnormalities. 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