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Entry version 196 (05 Jun 2019)
Sequence version 3 (11 Jan 2011)
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Protein

Probable ubiquitin carboxyl-terminal hydrolase FAF-X

Gene

USP9X

Organism
Homo sapiens (Human)
Status
Reviewed-Annotation score:

Annotation score:5 out of 5

<p>The annotation score provides a heuristic measure of the annotation content of a UniProtKB entry or proteome. This score <strong>cannot</strong> be used as a measure of the accuracy of the annotation as we cannot define the ‘correct annotation’ for any given protein.<p><a href='/help/annotation_score' target='_top'>More...</a></p>
-Experimental evidence at protein leveli <p>This indicates the type of evidence that supports the existence of the protein. Note that the ‘protein existence’ evidence does not give information on the accuracy or correctness of the sequence(s) displayed.<p><a href='/help/protein_existence' target='_top'>More...</a></p>

<p>This section provides any useful information about the protein, mostly biological knowledge.<p><a href='/help/function_section' target='_top'>More...</a></p>Functioni

Deubiquitinase involved both in the processing of ubiquitin precursors and of ubiquitinated proteins. May therefore play an important regulatory role at the level of protein turnover by preventing degradation of proteins through the removal of conjugated ubiquitin. Specifically hydrolyzes 'Lys-48'-, 'Lys-29'- and 'Lys-33'-linked polyubiquitins chains. Essential component of TGF-beta/BMP signaling cascade. Specifically deubiquitinates monoubiquitinated SMAD4, opposing the activity of E3 ubiquitin-protein ligase TRIM33. Deubiquitinates alkylation repair enzyme ALKBH3. OTUD4 recruits USP7 and USP9X to stabilize ALKBH3, thereby promoting the repair of alkylated DNA lesions (PubMed:25944111). Regulates chromosome alignment and segregation in mitosis by regulating the localization of BIRC5/survivin to mitotic centromeres. Involved in axonal growth and neuronal cell migration (PubMed:16322459, PubMed:18254724, PubMed:19135894, PubMed:24607389). Regulates cellular clock function by enhancing the protein stability and transcriptional activity of the core circadian protein ARNTL/BMAL1 via its deubiquitinating activity (PubMed:29626158).6 Publications

Miscellaneous

Escapes X-inactivation.

<p>This subsection of the <a href="http://www.uniprot.org/help/function_section">Function</a> section describes the catalytic activity of an enzyme, i.e. a chemical reaction that the enzyme catalyzes.<p><a href='/help/catalytic_activity' target='_top'>More...</a></p>Catalytic activityi

  • 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). EC:3.4.19.12

Sites

Feature keyPosition(s)DescriptionActionsGraphical viewLength
<p>This subsection of the <a href="http://www.uniprot.org/help/function_section">Function</a> section is used for enzymes and indicates the residues directly involved in catalysis.<p><a href='/help/act_site' target='_top'>More...</a></p>Active sitei1566NucleophilePROSITE-ProRule annotation1
Active sitei1879Proton acceptorPROSITE-ProRule annotation1

<p>The <a href="http://www.geneontology.org/">Gene Ontology (GO)</a> project provides a set of hierarchical controlled vocabulary split into 3 categories:<p><a href='/help/gene_ontology' target='_top'>More...</a></p>GO - Molecular functioni

GO - Biological processi

<p>UniProtKB Keywords constitute a <a href="http://www.uniprot.org/keywords">controlled vocabulary</a> with a hierarchical structure. Keywords summarise the content of a UniProtKB entry and facilitate the search for proteins of interest.<p><a href='/help/keywords' target='_top'>More...</a></p>Keywordsi

Molecular functionHydrolase, Protease, Thiol protease
Biological processBiological rhythms, Cell cycle, Cell division, Chromosome partition, Mitosis, Ubl conjugation pathway

Enzyme and pathway databases

Reactome - a knowledgebase of biological pathways and processes

More...
Reactomei
R-HSA-2173795 Downregulation of SMAD2/3:SMAD4 transcriptional activity
R-HSA-5689880 Ub-specific processing proteases
R-HSA-8866652 Synthesis of active ubiquitin: roles of E1 and E2 enzymes
R-HSA-9033241 Peroxisomal protein import
R-HSA-977225 Amyloid fiber formation

SIGNOR Signaling Network Open Resource

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SIGNORi
Q93008

Protein family/group databases

MEROPS protease database

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MEROPSi
C19.017

<p>This section provides information about the protein and gene name(s) and synonym(s) and about the organism that is the source of the protein sequence.<p><a href='/help/names_and_taxonomy_section' target='_top'>More...</a></p>Names & Taxonomyi

<p>This subsection of the <a href="http://www.uniprot.org/help/names_and_taxonomy_section">Names and taxonomy</a> section provides an exhaustive list of all names of the protein, from commonly used to obsolete, to allow unambiguous identification of a protein.<p><a href='/help/protein_names' target='_top'>More...</a></p>Protein namesi
Recommended name:
Probable ubiquitin carboxyl-terminal hydrolase FAF-X (EC:3.4.19.12)
Alternative name(s):
Deubiquitinating enzyme FAF-X
Fat facets in mammals
Short name:
hFAM
Fat facets protein-related, X-linked
Ubiquitin thioesterase FAF-X
Ubiquitin-specific protease 9, X chromosome
Ubiquitin-specific-processing protease FAF-X
<p>This subsection of the <a href="http://www.uniprot.org/help/names_and_taxonomy_section">Names and taxonomy</a> section indicates the name(s) of the gene(s) that code for the protein sequence(s) described in the entry. Four distinct tokens exist: ‘Name’, ‘Synonyms’, ‘Ordered locus names’ and ‘ORF names’.<p><a href='/help/gene_name' target='_top'>More...</a></p>Gene namesi
Name:USP9XImported
Synonyms:DFFRX, FAM, USP9
<p>This subsection of the <a href="http://www.uniprot.org/help/names_and_taxonomy_section">Names and taxonomy</a> section provides information on the name(s) of the organism that is the source of the protein sequence.<p><a href='/help/organism-name' target='_top'>More...</a></p>OrganismiHomo sapiens (Human)
<p>This subsection of the <a href="http://www.uniprot.org/help/names_and_taxonomy_section">Names and taxonomy</a> section shows the unique identifier assigned by the NCBI to the source organism of the protein. This is known as the ‘taxonomic identifier’ or ‘taxid’.<p><a href='/help/taxonomic_identifier' target='_top'>More...</a></p>Taxonomic identifieri9606 [NCBI]
<p>This subsection of the <a href="http://www.uniprot.org/help/names_and_taxonomy_section">Names and taxonomy</a> section contains the taxonomic hierarchical classification lineage of the source organism. It lists the nodes as they appear top-down in the taxonomic tree, with the more general grouping listed first.<p><a href='/help/taxonomic_lineage' target='_top'>More...</a></p>Taxonomic lineageiEukaryotaMetazoaChordataCraniataVertebrataEuteleostomiMammaliaEutheriaEuarchontogliresPrimatesHaplorrhiniCatarrhiniHominidaeHomo
<p>This subsection of the <a href="http://www.uniprot.org/help/names_and_taxonomy_section">Names and taxonomy</a> section is present for entries that are part of a <a href="http://www.uniprot.org/proteomes">proteome</a>, i.e. of a set of proteins thought to be expressed by organisms whose genomes have been completely sequenced.<p><a href='/help/proteomes_manual' target='_top'>More...</a></p>Proteomesi
  • UP000005640 <p>A UniProt <a href="http://www.uniprot.org/manual/proteomes_manual">proteome</a> can consist of several components. <br></br>The component name refers to the genomic component encoding a set of proteins.<p><a href='/help/proteome_component' target='_top'>More...</a></p> Componenti: Chromosome X

Organism-specific databases

Human Gene Nomenclature Database

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HGNCi
HGNC:12632 USP9X

Online Mendelian Inheritance in Man (OMIM)

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MIMi
300072 gene

neXtProt; the human protein knowledge platform

More...
neXtProti
NX_Q93008

<p>This section provides information on the location and the topology of the mature protein in the cell.<p><a href='/help/subcellular_location_section' target='_top'>More...</a></p>Subcellular locationi

Extracellular region or secreted Cytosol Plasma membrane Cytoskeleton Lysosome Endosome Peroxisome ER Golgi apparatus Nucleus Mitochondrion Manual annotation Automatic computational assertionGraphics by Christian Stolte & Seán O’Donoghue; Source: COMPARTMENTS

Keywords - Cellular componenti

Cell projection, Cytoplasm

<p>This section provides information on the disease(s) and phenotype(s) associated with a protein.<p><a href='/help/pathology_and_biotech_section' target='_top'>More...</a></p>Pathology & Biotechi

<p>This subsection of the ‘Pathology and Biotech’ section provides information on the disease(s) associated with genetic variations in a given protein. The information is extracted from the scientific literature and diseases that are also described in the <a href="http://www.ncbi.nlm.nih.gov/sites/entrez?db=omim">OMIM</a> database are represented with a <a href="http://www.uniprot.org/diseases">controlled vocabulary</a> in the following way:<p><a href='/help/involvement_in_disease' target='_top'>More...</a></p>Involvement in diseasei

Mental retardation, X-linked 99 (MRX99)1 Publication
The disease is caused by mutations affecting the gene represented in this entry.
Disease descriptionA disorder characterized by significantly below average general intellectual functioning associated with impairments in adaptive behavior and manifested during the developmental period. Intellectual deficiency is the only primary symptom of non-syndromic X-linked mental retardation, while syndromic mental retardation presents with associated physical, neurological and/or psychiatric manifestations.
Related information in OMIM
Feature keyPosition(s)DescriptionActionsGraphical viewLength
<p>This subsection of the ‘Sequence’ section describes natural variant(s) of the protein sequence.<p><a href='/help/variant' target='_top'>More...</a></p>Natural variantiVAR_0711312093L → H in MRX99; does not affect interaction with DCX; reduced subcellular localization in the axonal growth cones. 1 PublicationCorresponds to variant dbSNP:rs587777317EnsemblClinVar.1
Natural variantiVAR_0711322157L → I in MRX99; unknown pathological significance; does not affect interaction with DCX; reduced subcellular localization in the axonal growth cones. 1 PublicationCorresponds to variant dbSNP:rs587777319EnsemblClinVar.1

Keywords - Diseasei

Disease mutation, Mental retardation

Organism-specific databases

DisGeNET

More...
DisGeNETi
8239

MalaCards human disease database

More...
MalaCardsi
USP9X
MIMi300919 phenotype

Open Targets

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OpenTargetsi
ENSG00000124486

Orphanet; a database dedicated to information on rare diseases and orphan drugs

More...
Orphaneti
480880 X-linked female restricted facial dysmorphism-short stature-choanal atresia-intellectual disability
777 X-linked non-syndromic intellectual disability

The Pharmacogenetics and Pharmacogenomics Knowledge Base

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PharmGKBi
PA37257

Chemistry databases

ChEMBL database of bioactive drug-like small molecules

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ChEMBLi
CHEMBL2406899

Polymorphism and mutation databases

BioMuta curated single-nucleotide variation and disease association database

More...
BioMutai
USP9X

Domain mapping of disease mutations (DMDM)

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DMDMi
317373496

<p>This section describes post-translational modifications (PTMs) and/or processing events.<p><a href='/help/ptm_processing_section' target='_top'>More...</a></p>PTM / Processingi

Molecule processing

Feature keyPosition(s)DescriptionActionsGraphical viewLength
<p>This subsection of the ‘PTM / Processing’ section describes the extent of a polypeptide chain in the mature protein following processing.<p><a href='/help/chain' target='_top'>More...</a></p>ChainiPRO_00000806891 – 2570Probable ubiquitin carboxyl-terminal hydrolase FAF-XAdd BLAST2570

Amino acid modifications

Feature keyPosition(s)DescriptionActionsGraphical viewLength
<p>This subsection of the ‘PTM / Processing’ section specifies the position and type of each modified residue excluding <a href="http://www.uniprot.org/manual/lipid">lipids</a>, <a href="http://www.uniprot.org/manual/carbohyd">glycans</a> and <a href="http://www.uniprot.org/manual/crosslnk">protein cross-links</a>.<p><a href='/help/mod_res' target='_top'>More...</a></p>Modified residuei588PhosphoserineCombined sources1
Modified residuei590PhosphothreonineCombined sources1
Modified residuei1600PhosphoserineCombined sources1
Modified residuei2443PhosphoserineCombined sources1
Modified residuei2556PhosphotyrosineCombined sources1
Modified residuei2563PhosphoserineCombined sources1
Modified residuei2567PhosphothreonineCombined sources1

Keywords - PTMi

Phosphoprotein

Proteomic databases

Encyclopedia of Proteome Dynamics

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EPDi
Q93008

jPOST - Japan Proteome Standard Repository/Database

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jPOSTi
Q93008

MaxQB - The MaxQuant DataBase

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MaxQBi
Q93008

PaxDb, a database of protein abundance averages across all three domains of life

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PaxDbi
Q93008

PeptideAtlas

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PeptideAtlasi
Q93008

PRoteomics IDEntifications database

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PRIDEi
Q93008

ProteomicsDB human proteome resource

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ProteomicsDBi
75667
75668 [Q93008-1]

PTM databases

iPTMnet integrated resource for PTMs in systems biology context

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iPTMneti
Q93008

Comprehensive resource for the study of protein post-translational modifications (PTMs) in human, mouse and rat.

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PhosphoSitePlusi
Q93008

SwissPalm database of S-palmitoylation events

More...
SwissPalmi
Q93008

<p>This section provides information on the expression of a gene at the mRNA or protein level in cells or in tissues of multicellular organisms.<p><a href='/help/expression_section' target='_top'>More...</a></p>Expressioni

<p>This subsection of the ‘Expression’ section provides information on the expression of a gene at the mRNA or protein level in cells or in tissues of multicellular organisms. By default, the information is derived from experiments at the mRNA level, unless specified ‘at protein level’. <br></br>Examples: <a href="http://www.uniprot.org/uniprot/P92958#expression">P92958</a>, <a href="http://www.uniprot.org/uniprot/Q8TDN4#expression">Q8TDN4</a>, <a href="http://www.uniprot.org/uniprot/O14734#expression">O14734</a><p><a href='/help/tissue_specificity' target='_top'>More...</a></p>Tissue specificityi

Widely expressed in embryonic and adult tissues.

Gene expression databases

Bgee dataBase for Gene Expression Evolution

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Bgeei
ENSG00000124486 Expressed in 246 organ(s), highest expression level in placenta

Genevisible search portal to normalized and curated expression data from Genevestigator

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Genevisiblei
Q93008 HS

Organism-specific databases

Human Protein Atlas

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HPAi
CAB011618
CAB070164
HPA047417

<p>This section provides information on the quaternary structure of a protein and on interaction(s) with other proteins or protein complexes.<p><a href='/help/interaction_section' target='_top'>More...</a></p>Interactioni

<p>This subsection of the <a href="http://www.uniprot.org/help/interaction_section">'Interaction'</a> section provides information about the protein quaternary structure and interaction(s) with other proteins or protein complexes (with the exception of physiological receptor-ligand interactions which are annotated in the <a href="http://www.uniprot.org/help/function_section">'Function'</a> section).<p><a href='/help/subunit_structure' target='_top'>More...</a></p>Subunit structurei

Interacts with SMAD4, MARK4, NUAK1 and BIRC5/survivin. Interacts with DCX. Interacts with OTUD4 and USP7; the interaction is direct (PubMed:25944111). Interacts with ARNTL/BMAL1 (PubMed:29626158).6 Publications

<p>This subsection of the '<a href="http://www.uniprot.org/help/interaction_section%27">Interaction</a> section provides information about binary protein-protein interactions. The data presented in this section are a quality-filtered subset of binary interactions automatically derived from the <a href="http://www.ebi.ac.uk/intact/">IntAct database</a>. It is updated on a monthly basis. Each binary interaction is displayed on a separate line.<p><a href='/help/binary_interactions' target='_top'>More...</a></p>Binary interactionsi

GO - Molecular functioni

Protein-protein interaction databases

The Biological General Repository for Interaction Datasets (BioGrid)

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BioGridi
113867, 180 interactors

CORUM comprehensive resource of mammalian protein complexes

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CORUMi
Q93008

Database of interacting proteins

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DIPi
DIP-27562N

Protein interaction database and analysis system

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IntActi
Q93008, 49 interactors

Molecular INTeraction database

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MINTi
Q93008

STRING: functional protein association networks

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STRINGi
9606.ENSP00000316357

Chemistry databases

BindingDB database of measured binding affinities

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BindingDBi
Q93008

<p>This section provides information on the tertiary and secondary structure of a protein.<p><a href='/help/structure_section' target='_top'>More...</a></p>Structurei

Secondary structure

12570
Legend: HelixTurnBeta strandPDB Structure known for this area
Show more details

3D structure databases

SWISS-MODEL Repository - a database of annotated 3D protein structure models

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SMRi
Q93008

Database of comparative protein structure models

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ModBasei
Search...

MobiDB: a database of protein disorder and mobility annotations

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MobiDBi
Search...

<p>This section provides information on sequence similarities with other proteins and the domain(s) present in a protein.<p><a href='/help/family_and_domains_section' target='_top'>More...</a></p>Family & Domainsi

Domains and Repeats

Feature keyPosition(s)DescriptionActionsGraphical viewLength
<p>This subsection of the <a href="http://www.uniprot.org/help/family_and_domains_section">Family and Domains</a> section describes the position and type of a domain, which is defined as a specific combination of secondary structures organized into a characteristic three-dimensional structure or fold.<p><a href='/help/domain' target='_top'>More...</a></p>Domaini1557 – 1956USPAdd BLAST400

<p>This subsection of the ‘Family and domains’ section provides information about the sequence similarity with other proteins.<p><a href='/help/sequence_similarities' target='_top'>More...</a></p>Sequence similaritiesi

Belongs to the peptidase C19 family.Curated

Phylogenomic databases

evolutionary genealogy of genes: Non-supervised Orthologous Groups

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eggNOGi
KOG1866 Eukaryota
COG5077 LUCA

Ensembl GeneTree

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GeneTreei
ENSGT00940000155375

The HOGENOM Database of Homologous Genes from Fully Sequenced Organisms

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HOGENOMi
HOG000231283

InParanoid: Eukaryotic Ortholog Groups

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InParanoidi
Q93008

KEGG Orthology (KO)

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KOi
K11840

Identification of Orthologs from Complete Genome Data

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OMAi
DYMGEIY

Database of Orthologous Groups

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OrthoDBi
625455at2759

Database for complete collections of gene phylogenies

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PhylomeDBi
Q93008

TreeFam database of animal gene trees

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TreeFami
TF323966

Family and domain databases

Integrated resource of protein families, domains and functional sites

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InterProi
View protein in InterPro
IPR016024 ARM-type_fold
IPR038765 Papain-like_cys_pep_sf
IPR001394 Peptidase_C19_UCH
IPR018200 USP_CS
IPR028889 USP_dom

Pfam protein domain database

More...
Pfami
View protein in Pfam
PF00443 UCH, 1 hit

Superfamily database of structural and functional annotation

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SUPFAMi
SSF48371 SSF48371, 1 hit
SSF54001 SSF54001, 1 hit

PROSITE; a protein domain and family database

More...
PROSITEi
View protein in PROSITE
PS00972 USP_1, 1 hit
PS00973 USP_2, 1 hit
PS50235 USP_3, 1 hit

<p>This section displays by default the canonical protein sequence and upon request all isoforms described in the entry. It also includes information pertinent to the sequence(s), including <a href="http://www.uniprot.org/help/sequence_length">length</a> and <a href="http://www.uniprot.org/help/sequences">molecular weight</a>. The information is filed in different subsections. The current subsections and their content are listed below:<p><a href='/help/sequences_section' target='_top'>More...</a></p>Sequences (2)i

<p>This subsection of the <a href="http://www.uniprot.org/help/sequences_section">Sequence</a> section indicates if the <a href="http://www.uniprot.org/help/canonical_and_isoforms">canonical sequence</a> displayed by default in the entry is complete or not.<p><a href='/help/sequence_status' target='_top'>More...</a></p>Sequence statusi: Complete.

This entry describes 2 <p>This subsection of the ‘Sequence’ section lists the alternative protein sequences (isoforms) that can be generated from the same gene by a single or by the combination of up to four biological events (alternative promoter usage, alternative splicing, alternative initiation and ribosomal frameshifting). Additionally, this section gives relevant information on each alternative protein isoform.<p><a href='/help/alternative_products' target='_top'>More...</a></p> isoformsi produced by alternative splicing. AlignAdd to basket
Isoform 1 (identifier: Q93008-3) [UniParc]FASTAAdd to basket
Also known as: Long

This isoform has been chosen as the 'canonical' sequence. All positional information in this entry refers to it. This is also the sequence that appears in the downloadable versions of the entry.

« Hide
        10         20         30         40         50
MTATTRGSPV GGNDNQGQAP DGQSQPPLQQ NQTSSPDSSN ENSPATPPDE
60 70 80 90 100
QGQGDAPPQL EDEEPAFPHT DLAKLDDMIN RPRWVVPVLP KGELEVLLEA
110 120 130 140 150
AIDLSKKGLD VKSEACQRFF RDGLTISFTK ILTDEAVSGW KFEIHRCIIN
160 170 180 190 200
NTHRLVELCV AKLSQDWFPL LELLAMALNP HCKFHIYNGT RPCESVSSSV
210 220 230 240 250
QLPEDELFAR SPDPRSPKGW LVDLLNKFGT LNGFQILHDR FINGSALNVQ
260 270 280 290 300
IIAALIKPFG QCYEFLTLHT VKKYFLPIIE MVPQFLENLT DEELKKEAKN
310 320 330 340 350
EAKNDALSMI IKSLKNLASR VPGQEETVKN LEIFRLKMIL RLLQISSFNG
360 370 380 390 400
KMNALNEVNK VISSVSYYTH RHGNPEEEEW LTAERMAEWI QQNNILSIVL
410 420 430 440 450
RDSLHQPQYV EKLEKILRFV IKEKALTLQD LDNIWAAQAG KHEAIVKNVH
460 470 480 490 500
DLLAKLAWDF SPEQLDHLFD CFKASWTNAS KKQREKLLEL IRRLAEDDKD
510 520 530 540 550
GVMAHKVLNL LWNLAHSDDV PVDIMDLALS AHIKILDYSC SQDRDTQKIQ
560 570 580 590 600
WIDRFIEELR TNDKWVIPAL KQIREICSLF GEAPQNLSQT QRSPHVFYRH
610 620 630 640 650
DLINQLQHNH ALVTLVAENL ATYMESMRLY ARDHEDYDPQ TVRLGSRYSH
660 670 680 690 700
VQEVQERLNF LRFLLKDGQL WLCAPQAKQI WKCLAENAVY LCDREACFKW
710 720 730 740 750
YSKLMGDEPD LDPDINKDFF ESNVLQLDPS LLTENGMKCF ERFFKAVNCR
760 770 780 790 800
EGKLVAKRRA YMMDDLELIG LDYLWRVVIQ SNDDIASRAI DLLKEIYTNL
810 820 830 840 850
GPRLQVNQVV IHEDFIQSCF DRLKASYDTL CVLDGDKDSV NCARQEAVRM
860 870 880 890 900
VRVLTVLREY INECDSDYHE ERTILPMSRA FRGKHLSFVV RFPNQGRQVD
910 920 930 940 950
DLEVWSHTND TIGSVRRCIL NRIKANVAHT KIELFVGGEL IDPADDRKLI
960 970 980 990 1000
GQLNLKDKSL ITAKLTQISS NMPSSPDSSS DSSTGSPGNH GNHYSDGPNP
1010 1020 1030 1040 1050
EVESCLPGVI MSLHPRYISF LWQVADLGSS LNMPPLRDGA RVLMKLMPPD
1060 1070 1080 1090 1100
STTIEKLRAI CLDHAKLGES SLSPSLDSLF FGPSASQVLY LTEVVYALLM
1110 1120 1130 1140 1150
PAGAPLADDS SDFQFHFLKS GGLPLVLSML TRNNFLPNAD METRRGAYLN
1160 1170 1180 1190 1200
ALKIAKLLLT AIGYGHVRAV AEACQPGVEG VNPMTQINQV THDQAVVLQS
1210 1220 1230 1240 1250
ALQSIPNPSS ECMLRNVSVR LAQQISDEAS RYMPDICVIR AIQKIIWASG
1260 1270 1280 1290 1300
CGSLQLVFSP NEEITKIYEK TNAGNEPDLE DEQVCCEALE VMTLCFALIP
1310 1320 1330 1340 1350
TALDALSKEK AWQTFIIDLL LHCHSKTVRQ VAQEQFFLMC TRCCMGHRPL
1360 1370 1380 1390 1400
LFFITLLFTV LGSTARERAK HSGDYFTLLR HLLNYAYNSN INVPNAEVLL
1410 1420 1430 1440 1450
NNEIDWLKRI RDDVKRTGET GIEETILEGH LGVTKELLAF QTSEKKFHIG
1460 1470 1480 1490 1500
CEKGGANLIK ELIDDFIFPA SNVYLQYMRN GELPAEQAIP VCGSPPTINA
1510 1520 1530 1540 1550
GFELLVALAV GCVRNLKQIV DSLTEMYYIG TAITTCEALT EWEYLPPVGP
1560 1570 1580 1590 1600
RPPKGFVGLK NAGATCYMNS VIQQLYMIPS IRNGILAIEG TGSDVDDDMS
1610 1620 1630 1640 1650
GDEKQDNESN VDPRDDVFGY PQQFEDKPAL SKTEDRKEYN IGVLRHLQVI
1660 1670 1680 1690 1700
FGHLAASRLQ YYVPRGFWKQ FRLWGEPVNL REQHDALEFF NSLVDSLDEA
1710 1720 1730 1740 1750
LKALGHPAML SKVLGGSFAD QKICQGCPHR YECEESFTTL NVDIRNHQNL
1760 1770 1780 1790 1800
LDSLEQYVKG DLLEGANAYH CEKCNKKVDT VKRLLIKKLP PVLAIQLKRF
1810 1820 1830 1840 1850
DYDWERECAI KFNDYFEFPR ELDMEPYTVA GVAKLEGDNV NPESQLIQQS
1860 1870 1880 1890 1900
EQSESETAGS TKYRLVGVLV HSGQASGGHY YSYIIQRNGG DGERNRWYKF
1910 1920 1930 1940 1950
DDGDVTECKM DDDEEMKNQC FGGEYMGEVF DHMMKRMSYR RQKRWWNAYI
1960 1970 1980 1990 2000
LFYERMDTID QDDELIRYIS ELAITTRPHQ IIMPSAIERS VRKQNVQFMH
2010 2020 2030 2040 2050
NRMQYSMEYF QFMKKLLTCN GVYLNPPPGQ DHLLPEAEEI TMISIQLAAR
2060 2070 2080 2090 2100
FLFTTGFHTK KVVRGSASDW YDALCILLRH SKNVRFWFAH NVLFNVSNRF
2110 2120 2130 2140 2150
SEYLLECPSA EVRGAFAKLI VFIAHFSLQD GPCPSPFASP GPSSQAYDNL
2160 2170 2180 2190 2200
SLSDHLLRAV LNLLRREVSE HGRHLQQYFN LFVMYANLGV AEKTQLLKLS
2210 2220 2230 2240 2250
VPATFMLVSL DEGPGPPIKY QYAELGKLYS VVSQLIRCCN VSSRMQSSIN
2260 2270 2280 2290 2300
GNPPLPNPFG DPNLSQPIMP IQQNVADILF VRTSYVKKII EDCSNSEETV
2310 2320 2330 2340 2350
KLLRFCCWEN PQFSSTVLSE LLWQVAYSYT YELRPYLDLL LQILLIEDSW
2360 2370 2380 2390 2400
QTHRIHNALK GIPDDRDGLF DTIQRSKNHY QKRAYQCIKC MVALFSNCPV
2410 2420 2430 2440 2450
AYQILQGNGD LKRKWTWAVE WLGDELERRP YTGNPQYTYN NWSPPVQSNE
2460 2470 2480 2490 2500
TSNGYFLERS HSARMTLAKA CELCPEEVKK ATSVQQIEME ESKEPDDQDA
2510 2520 2530 2540 2550
PDEHESPPPE DAPLYPHSPG SQYQQNNHVH GQPYTGPAAH HMNNPQRTGQ
2560 2570
RAQENYEGSE EVSPPQTKDQ
Length:2,570
Mass (Da):292,280
Last modified:January 11, 2011 - v3
<p>The checksum is a form of redundancy check that is calculated from the sequence. It is useful for tracking sequence updates.</p> <p>It should be noted that while, in theory, two different sequences could have the same checksum value, the likelihood that this would happen is extremely low.</p> <p>However UniProtKB may contain entries with identical sequences in case of multiple genes (paralogs).</p> <p>The checksum is computed as the sequence 64-bit Cyclic Redundancy Check value (CRC64) using the generator polynomial: x<sup>64</sup> + x<sup>4</sup> + x<sup>3</sup> + x + 1. The algorithm is described in the ISO 3309 standard. </p> <p class="publication">Press W.H., Flannery B.P., Teukolsky S.A. and Vetterling W.T.<br /> <strong>Cyclic redundancy and other checksums</strong><br /> <a href="http://www.nrbook.com/b/bookcpdf.php">Numerical recipes in C 2nd ed., pp896-902, Cambridge University Press (1993)</a>)</p> Checksum:i84CB979A405AA56F
GO
Isoform 2 (identifier: Q93008-1) [UniParc]FASTAAdd to basket
Also known as: Short

The sequence of this isoform differs from the canonical sequence as follows:
     2478-2493: Missing.

Show »
Length:2,554
Mass (Da):290,463
Checksum:i16B87B7FCC1428AF
GO

Experimental Info

Feature keyPosition(s)DescriptionActionsGraphical viewLength
<p>This subsection of the ‘Sequence’ section reports difference(s) between the canonical sequence (displayed by default in the entry) and the different sequence submissions merged in the entry. These various submissions may originate from different sequencing projects, different types of experiments, or different biological samples. Sequence conflicts are usually of unknown origin.<p><a href='/help/conflict' target='_top'>More...</a></p>Sequence conflicti25Q → L in CAA66942 (PubMed:8922996).Curated1
Sequence conflicti148 – 154Missing in CAA66942 (PubMed:8922996).Curated7
Sequence conflicti468L → P in CAA66942 (PubMed:8922996).Curated1
Sequence conflicti476W → R in CAA66942 (PubMed:8922996).Curated1
Sequence conflicti506K → R in CAA66942 (PubMed:8922996).Curated1
Sequence conflicti621A → V in CAA66942 (PubMed:8922996).Curated1
Sequence conflicti1400L → F in CAA66942 (PubMed:8922996).Curated1
Sequence conflicti1951L → P in CAA66942 (PubMed:8922996).Curated1
Sequence conflicti2330T → P in CAA66942 (PubMed:8922996).Curated1

Natural variant

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Natural variantiVAR_0711312093L → H in MRX99; does not affect interaction with DCX; reduced subcellular localization in the axonal growth cones. 1 PublicationCorresponds to variant dbSNP:rs587777317EnsemblClinVar.1
Natural variantiVAR_0711322157L → I in MRX99; unknown pathological significance; does not affect interaction with DCX; reduced subcellular localization in the axonal growth cones. 1 PublicationCorresponds to variant dbSNP:rs587777319EnsemblClinVar.1

Alternative sequence

Feature keyPosition(s)DescriptionActionsGraphical viewLength
<p>This subsection of the ‘Sequence’ section describes the sequence of naturally occurring alternative protein isoform(s). The changes in the amino acid sequence may be due to alternative splicing, alternative promoter usage, alternative initiation, or ribosomal frameshifting.<p><a href='/help/var_seq' target='_top'>More...</a></p>Alternative sequenceiVSP_0404782478 – 2493Missing in isoform 2. 1 PublicationAdd BLAST16

Sequence databases

Select the link destinations:

EMBL nucleotide sequence database

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EMBLi

GenBank nucleotide sequence database

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GenBanki

DNA Data Bank of Japan; a nucleotide sequence database

More...
DDBJi
Links Updated
X98296 mRNA Translation: CAA66942.1
AL109797 Genomic DNA No translation available.
AL391259 Genomic DNA No translation available.
AF070645 mRNA Translation: AAC25395.1

The Consensus CDS (CCDS) project

More...
CCDSi
CCDS43930.1 [Q93008-3]
CCDS55403.1 [Q93008-1]

NCBI Reference Sequences

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RefSeqi
NP_001034679.2, NM_001039590.2 [Q93008-3]
NP_001034680.2, NM_001039591.2 [Q93008-1]

Genome annotation databases

Ensembl eukaryotic genome annotation project

More...
Ensembli
ENST00000324545; ENSP00000316357; ENSG00000124486 [Q93008-3]
ENST00000378308; ENSP00000367558; ENSG00000124486 [Q93008-1]

Database of genes from NCBI RefSeq genomes

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GeneIDi
8239

KEGG: Kyoto Encyclopedia of Genes and Genomes

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KEGGi
hsa:8239

UCSC genome browser

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UCSCi
uc004dfb.3 human [Q93008-3]

Keywords - Coding sequence diversityi

Alternative splicing

<p>This section provides links to proteins that are similar to the protein sequence(s) described in this entry at different levels of sequence identity thresholds (100%, 90% and 50%) based on their membership in UniProt Reference Clusters (<a href="http://www.uniprot.org/help/uniref">UniRef</a>).<p><a href='/help/similar_proteins_section' target='_top'>More...</a></p>Similar proteinsi

<p>This section is used to point to information related to entries and found in data collections other than UniProtKB.<p><a href='/help/cross_references_section' target='_top'>More...</a></p>Cross-referencesi

Sequence databases

Select the link destinations:
EMBLi
GenBanki
DDBJi
Links Updated
X98296 mRNA Translation: CAA66942.1
AL109797 Genomic DNA No translation available.
AL391259 Genomic DNA No translation available.
AF070645 mRNA Translation: AAC25395.1
CCDSiCCDS43930.1 [Q93008-3]
CCDS55403.1 [Q93008-1]
RefSeqiNP_001034679.2, NM_001039590.2 [Q93008-3]
NP_001034680.2, NM_001039591.2 [Q93008-1]

3D structure databases

Select the link destinations:

Protein Data Bank Europe

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PDBei

Protein Data Bank RCSB

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RCSB PDBi

Protein Data Bank Japan

More...
PDBji
Links Updated
PDB entryMethodResolution (Å)ChainPositionsPDBsum
5VBDX-ray1.50A880-970[»]
5WCHX-ray2.50A/B/C/D1551-1970[»]
SMRiQ93008
ModBaseiSearch...
MobiDBiSearch...

Protein-protein interaction databases

BioGridi113867, 180 interactors
CORUMiQ93008
DIPiDIP-27562N
IntActiQ93008, 49 interactors
MINTiQ93008
STRINGi9606.ENSP00000316357

Chemistry databases

BindingDBiQ93008
ChEMBLiCHEMBL2406899

Protein family/group databases

MEROPSiC19.017

PTM databases

iPTMnetiQ93008
PhosphoSitePlusiQ93008
SwissPalmiQ93008

Polymorphism and mutation databases

BioMutaiUSP9X
DMDMi317373496

Proteomic databases

EPDiQ93008
jPOSTiQ93008
MaxQBiQ93008
PaxDbiQ93008
PeptideAtlasiQ93008
PRIDEiQ93008
ProteomicsDBi75667
75668 [Q93008-1]

Protocols and materials databases

Structural Biology KnowledgebaseSearch...

Genome annotation databases

EnsembliENST00000324545; ENSP00000316357; ENSG00000124486 [Q93008-3]
ENST00000378308; ENSP00000367558; ENSG00000124486 [Q93008-1]
GeneIDi8239
KEGGihsa:8239
UCSCiuc004dfb.3 human [Q93008-3]

Organism-specific databases

Comparative Toxicogenomics Database

More...
CTDi
8239
DisGeNETi8239

GeneCards: human genes, protein and diseases

More...
GeneCardsi
USP9X
HGNCiHGNC:12632 USP9X
HPAiCAB011618
CAB070164
HPA047417
MalaCardsiUSP9X
MIMi300072 gene
300919 phenotype
neXtProtiNX_Q93008
OpenTargetsiENSG00000124486
Orphaneti480880 X-linked female restricted facial dysmorphism-short stature-choanal atresia-intellectual disability
777 X-linked non-syndromic intellectual disability
PharmGKBiPA37257

GenAtlas: human gene database

More...
GenAtlasi
Search...

Phylogenomic databases

eggNOGiKOG1866 Eukaryota
COG5077 LUCA
GeneTreeiENSGT00940000155375
HOGENOMiHOG000231283
InParanoidiQ93008
KOiK11840
OMAiDYMGEIY
OrthoDBi625455at2759
PhylomeDBiQ93008
TreeFamiTF323966

Enzyme and pathway databases

ReactomeiR-HSA-2173795 Downregulation of SMAD2/3:SMAD4 transcriptional activity
R-HSA-5689880 Ub-specific processing proteases
R-HSA-8866652 Synthesis of active ubiquitin: roles of E1 and E2 enzymes
R-HSA-9033241 Peroxisomal protein import
R-HSA-977225 Amyloid fiber formation
SIGNORiQ93008

Miscellaneous databases

ChiTaRS: a database of human, mouse and fruit fly chimeric transcripts and RNA-sequencing data

More...
ChiTaRSi
USP9X human

The Gene Wiki collection of pages on human genes and proteins

More...
GeneWikii
USP9X

Database of phenotypes from RNA interference screens in Drosophila and Homo sapiens

More...
GenomeRNAii
8239

Protein Ontology

More...
PROi
PR:Q93008

The Stanford Online Universal Resource for Clones and ESTs

More...
SOURCEi
Search...

Gene expression databases

BgeeiENSG00000124486 Expressed in 246 organ(s), highest expression level in placenta
GenevisibleiQ93008 HS

Family and domain databases

InterProiView protein in InterPro
IPR016024 ARM-type_fold
IPR038765 Papain-like_cys_pep_sf
IPR001394 Peptidase_C19_UCH
IPR018200 USP_CS
IPR028889 USP_dom
PfamiView protein in Pfam
PF00443 UCH, 1 hit
SUPFAMiSSF48371 SSF48371, 1 hit
SSF54001 SSF54001, 1 hit
PROSITEiView protein in PROSITE
PS00972 USP_1, 1 hit
PS00973 USP_2, 1 hit
PS50235 USP_3, 1 hit

ProtoNet; Automatic hierarchical classification of proteins

More...
ProtoNeti
Search...

<p>This section provides general information on the entry.<p><a href='/help/entry_information_section' target='_top'>More...</a></p>Entry informationi

<p>This subsection of the ‘Entry information’ section provides a mnemonic identifier for a UniProtKB entry, but it is not a stable identifier. Each reviewed entry is assigned a unique entry name upon integration into UniProtKB/Swiss-Prot.<p><a href='/help/entry_name' target='_top'>More...</a></p>Entry nameiUSP9X_HUMAN
<p>This subsection of the ‘Entry information’ section provides one or more accession number(s). These are stable identifiers and should be used to cite UniProtKB entries. Upon integration into UniProtKB, each entry is assigned a unique accession number, which is called ‘Primary (citable) accession number’.<p><a href='/help/accession_numbers' target='_top'>More...</a></p>AccessioniPrimary (citable) accession number: Q93008
Secondary accession number(s): O75550, Q8WWT3, Q8WX12
<p>This subsection of the ‘Entry information’ section shows the date of integration of the entry into UniProtKB, the date of the last sequence update and the date of the last annotation modification (‘Last modified’). The version number for both the entry and the <a href="http://www.uniprot.org/help/canonical_and_isoforms">canonical sequence</a> are also displayed.<p><a href='/help/entry_history' target='_top'>More...</a></p>Entry historyiIntegrated into UniProtKB/Swiss-Prot: November 1, 1997
Last sequence update: January 11, 2011
Last modified: June 5, 2019
This is version 196 of the entry and version 3 of the sequence. See complete history.
<p>This subsection of the ‘Entry information’ section indicates whether the entry has been manually annotated and reviewed by UniProtKB curators or not, in other words, if the entry belongs to the Swiss-Prot section of UniProtKB (<strong>reviewed</strong>) or to the computer-annotated TrEMBL section (<strong>unreviewed</strong>).<p><a href='/help/entry_status' target='_top'>More...</a></p>Entry statusiReviewed (UniProtKB/Swiss-Prot)
Annotation programChordata Protein Annotation Program
DisclaimerAny medical or genetic information present in this entry is provided for research, educational and informational purposes only. It is not in any way intended to be used as a substitute for professional medical advice, diagnosis, treatment or care.

<p>This section contains any relevant information that doesn’t fit in any other defined sections<p><a href='/help/miscellaneous_section' target='_top'>More...</a></p>Miscellaneousi

Keywords - Technical termi

3D-structure, Complete proteome, Reference proteome

Documents

  1. PDB cross-references
    Index of Protein Data Bank (PDB) cross-references
  2. SIMILARITY comments
    Index of protein domains and families
  3. Human chromosome X
    Human chromosome X: entries, gene names and cross-references to MIM
  4. Human entries with polymorphisms or disease mutations
    List of human entries with polymorphisms or disease mutations
  5. Human polymorphisms and disease mutations
    Index of human polymorphisms and disease mutations
  6. Peptidase families
    Classification of peptidase families and list of entries
  7. MIM cross-references
    Online Mendelian Inheritance in Man (MIM) cross-references in UniProtKB/Swiss-Prot
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