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Entry version 174 (12 Aug 2020)
Sequence version 1 (01 Dec 2001)
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Protein

Ubiquitin carboxyl-terminal hydrolase 28

Gene

USP28

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 in DNA damage response checkpoint and MYC proto-oncogene stability. Involved in DNA damage induced apoptosis by specifically deubiquitinating proteins of the DNA damage pathway such as CLSPN. Also involved in G2 DNA damage checkpoint, by deubiquitinating CLSPN, and preventing its degradation by the anaphase promoting complex/cyclosome (APC/C). In contrast, it does not deubiquitinate PLK1. Specifically deubiquitinates MYC in the nucleoplasm, leading to prevent MYC degradation by the proteasome: acts by specifically interacting with isoform 1 of FBXW7 (FBW7alpha) in the nucleoplasm and counteracting ubiquitination of MYC by the SCF(FBW7) complex. In contrast, it does not interact with isoform 4 of FBXW7 (FBW7gamma) in the nucleolus, allowing MYC degradation and explaining the selective MYC degradation in the nucleolus. Deubiquitinates ZNF304, hence preventing ZNF304 degradation by the proteasome and leading to the activated KRAS-mediated promoter hypermethylation and transcriptional silencing of tumor suppressor genes (TSGs) in a subset of colorectal cancers (CRC) cells (PubMed:24623306).5 Publications

<p>This subsection of the <a href="http://www.uniprot.org/help/function%5Fsection">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%5Fsection">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 sitei171Nucleophile1
Active sitei600Proton 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 processDNA damage, DNA repair, Ubl conjugation pathway

Enzyme and pathway databases

Pathway Commons web resource for biological pathway data

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PathwayCommonsi
Q96RU2

Reactome - a knowledgebase of biological pathways and processes

More...
Reactomei
R-HSA-5689880, Ub-specific processing proteases

SIGNOR Signaling Network Open Resource

More...
SIGNORi
Q96RU2

Protein family/group databases

MEROPS protease database

More...
MEROPSi
C19.054

<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%5Fand%5Ftaxonomy%5Fsection">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:
Ubiquitin carboxyl-terminal hydrolase 28 (EC:3.4.19.12)
Alternative name(s):
Deubiquitinating enzyme 28
Ubiquitin thioesterase 28
Ubiquitin-specific-processing protease 28
<p>This subsection of the <a href="http://www.uniprot.org/help/names%5Fand%5Ftaxonomy%5Fsection">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:USP28
Synonyms:KIAA1515
<p>This subsection of the <a href="http://www.uniprot.org/help/names%5Fand%5Ftaxonomy%5Fsection">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%5Fand%5Ftaxonomy%5Fsection">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%5Fand%5Ftaxonomy%5Fsection">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%5Fand%5Ftaxonomy%5Fsection">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%5Fmanual">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 11

Organism-specific databases

Eukaryotic Pathogen Database Resources

More...
EuPathDBi
HostDB:ENSG00000048028.11

Human Gene Nomenclature Database

More...
HGNCi
HGNC:12625, USP28

Online Mendelian Inheritance in Man (OMIM)

More...
MIMi
610748, gene

neXtProt; the human protein knowledge platform

More...
neXtProti
NX_Q96RU2

<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

Nucleus

<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

Mutagenesis

Feature keyPosition(s)DescriptionActionsGraphical viewLength
<p>This subsection of the <a href="http://www.uniprot.org/manual/pathology%5Fand%5Fbiotech%5Fsection">'Pathology and Biotech'</a> section describes the effect of the experimental mutation of one or more amino acid(s) on the biological properties of the protein.<p><a href='/help/mutagen' target='_top'>More...</a></p>Mutagenesisi171C → A: Abolishes deubiquitinase activity. 3 Publications1
Mutagenesisi899S → A: Prevents ZNF304 ubiquitination reduction. 1 Publication1

Organism-specific databases

DisGeNET

More...
DisGeNETi
57646

Open Targets

More...
OpenTargetsi
ENSG00000048028

The Pharmacogenetics and Pharmacogenomics Knowledge Base

More...
PharmGKBi
PA37250

Miscellaneous databases

Pharos NIH Druggable Genome Knowledgebase

More...
Pharosi
Q96RU2, Tbio

Chemistry databases

ChEMBL database of bioactive drug-like small molecules

More...
ChEMBLi
CHEMBL2157853

Polymorphism and mutation databases

BioMuta curated single-nucleotide variation and disease association database

More...
BioMutai
USP28

Domain mapping of disease mutations (DMDM)

More...
DMDMi
20140700

<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 or proteolytic cleavage.<p><a href='/help/chain' target='_top'>More...</a></p>ChainiPRO_00000806571 – 1077Ubiquitin carboxyl-terminal hydrolase 28Add BLAST1077

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 residuei67Phosphoserine1 Publication1
<p>This subsection of the <a href="http://www.uniprot.org/help/ptm%5Fprocessing%5Fsection">PTM / Processing</a> section describes <strong>covalent linkages</strong> of various types formed <strong>between two proteins (interchain cross-links)</strong> or <strong>between two parts of the same protein (intrachain cross-links)</strong>, except the disulfide bonds that are annotated in the <a href="http://www.uniprot.org/manual/disulfid">'Disulfide bond'</a> subsection.<p><a href='/help/crosslnk' target='_top'>More...</a></p>Cross-linki99Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in SUMO2)Combined sources
Modified residuei375PhosphoserineCombined sources1
Modified residuei550PhosphoserineBy similarity1
Modified residuei714Phosphoserine1 Publication1
Modified residuei1048PhosphothreonineCombined sources1
Isoform 2 (identifier: Q96RU2-2)
Cross-linki759Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in SUMO2)Combined sources

<p>This subsection of the <a href="http://www.uniprot.org/help/ptm%5Fprocessing%5Fsection">PTM/processing</a> section describes post-translational modifications (PTMs). This subsection <strong>complements</strong> the information provided at the sequence level or describes modifications for which <strong>position-specific data is not yet available</strong>.<p><a href='/help/post-translational_modification' target='_top'>More...</a></p>Post-translational modificationi

Degraded upon nickel ion level or hypoxia exposure.
Phosphorylated upon DNA damage at Ser-67 and Ser-714, by ATM or ATR (PubMed:16901786). Phosphorylated by PRKD1 (PubMed:24623306).2 Publications

Keywords - PTMi

Isopeptide bond, Phosphoprotein, Ubl conjugation

Proteomic databases

Encyclopedia of Proteome Dynamics

More...
EPDi
Q96RU2

jPOST - Japan Proteome Standard Repository/Database

More...
jPOSTi
Q96RU2

MassIVE - Mass Spectrometry Interactive Virtual Environment

More...
MassIVEi
Q96RU2

MaxQB - The MaxQuant DataBase

More...
MaxQBi
Q96RU2

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

More...
PaxDbi
Q96RU2

PeptideAtlas

More...
PeptideAtlasi
Q96RU2

PRoteomics IDEntifications database

More...
PRIDEi
Q96RU2

ProteomicsDB: a multi-organism proteome resource

More...
ProteomicsDBi
66797
78033 [Q96RU2-1]
78034 [Q96RU2-2]

PTM databases

iPTMnet integrated resource for PTMs in systems biology context

More...
iPTMneti
Q96RU2

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

More...
PhosphoSitePlusi
Q96RU2

<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 reports the experimentally proven effects of inducers and repressors (usually chemical compounds or environmental factors) on the level of protein (or mRNA) expression (up-regulation, down-regulation, constitutive expression).<p><a href='/help/induction' target='_top'>More...</a></p>Inductioni

Down-regulated upon hypoxia (PubMed:20046830). Up-regulated by the transcription factor c-Jun/JUN in a KRAS-dependent manner in colorectal cancer (CRC) cells (PubMed:24623306).2 Publications

Gene expression databases

Bgee dataBase for Gene Expression Evolution

More...
Bgeei
ENSG00000048028, Expressed in heart and 208 other tissues

ExpressionAtlas, Differential and Baseline Expression

More...
ExpressionAtlasi
Q96RU2, baseline and differential

Genevisible search portal to normalized and curated expression data from Genevestigator

More...
Genevisiblei
Q96RU2, HS

Organism-specific databases

Human Protein Atlas

More...
HPAi
ENSG00000048028, Tissue enhanced (heart muscle, skeletal muscle)

<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%5Fsection">'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%5Fsection">'Function'</a> section).<p><a href='/help/subunit_structure' target='_top'>More...</a></p>Subunit structurei

Interacts with ZNF304 (PubMed:24623306).

Interacts with PRKD1 (PubMed:24623306).

Interacts with TP53BP1 (PubMed:16901786).

Interacts with isoform 1 of FBXW7; following DNA damage, dissociates from FBXW7 leading to degradation of MYC (PubMed:17873522, PubMed:17558397).

4 Publications

Protein-protein interaction databases

The Biological General Repository for Interaction Datasets (BioGRID)

More...
BioGRIDi
121683, 80 interactors

Protein interaction database and analysis system

More...
IntActi
Q96RU2, 19 interactors

Molecular INTeraction database

More...
MINTi
Q96RU2

STRING: functional protein association networks

More...
STRINGi
9606.ENSP00000003302

Miscellaneous databases

RNAct, Protein-RNA interaction predictions for model organisms.

More...
RNActi
Q96RU2, protein

<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

11077
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

More...
SMRi
Q96RU2

Database of comparative protein structure models

More...
ModBasei
Search...

Protein Data Bank in Europe - Knowledge Base

More...
PDBe-KBi
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%5Fand%5Fdomains%5Fsection">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>Domaini97 – 116UIMCuratedAdd BLAST20
Domaini162 – 650USPAdd BLAST489

<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. USP28 subfamily.Curated

Phylogenomic databases

evolutionary genealogy of genes: Non-supervised Orthologous Groups

More...
eggNOGi
KOG1863, Eukaryota

Ensembl GeneTree

More...
GeneTreei
ENSGT00940000157670

The HOGENOM Database of Homologous Genes from Fully Sequenced Organisms

More...
HOGENOMi
CLU_467647_0_0_1

InParanoid: Eukaryotic Ortholog Groups

More...
InParanoidi
Q96RU2

KEGG Orthology (KO)

More...
KOi
K21122

Identification of Orthologs from Complete Genome Data

More...
OMAi
GDWPVGM

Database of Orthologous Groups

More...
OrthoDBi
194025at2759

Database for complete collections of gene phylogenies

More...
PhylomeDBi
Q96RU2

TreeFam database of animal gene trees

More...
TreeFami
TF329035

Family and domain databases

Integrated resource of protein families, domains and functional sites

More...
InterProi
View protein in InterPro
IPR038765, Papain-like_cys_pep_sf
IPR001394, Peptidase_C19_UCH
IPR009060, UBA-like_sf
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

More...
SUPFAMi
SSF46934, SSF46934, 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%5Flength">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 (3+)i

<p>This subsection of the <a href="http://www.uniprot.org/help/sequences%5Fsection">Sequence</a> section indicates if the <a href="http://www.uniprot.org/help/canonical%5Fand%5Fisoforms">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 3 <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. This section is only present in reviewed entries, i.e. in UniProtKB/Swiss-Prot.<p><a href='/help/alternative_products' target='_top'>More...</a></p> isoformsi produced by alternative splicing. AlignAdd to basket

This entry has 3 described isoforms and 11 potential isoforms that are computationally mapped.Show allAlign All

Isoform 1 (identifier: Q96RU2-1) [UniParc]FASTAAdd to basket

This isoform has been chosen as the <div> <p><b>What is the canonical sequence?</b><p><a href='/help/canonical_and_isoforms' target='_top'>More...</a></p>canonicali 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
MTAELQQDDA AGAADGHGSS CQMLLNQLRE ITGIQDPSFL HEALKASNGD
60 70 80 90 100
ITQAVSLLTD ERVKEPSQDT VATEPSEVEG SAANKEVLAK VIDLTHDNKD
110 120 130 140 150
DLQAAIALSL LESPKIQADG RDLNRMHEAT SAETKRSKRK RCEVWGENPN
160 170 180 190 200
PNDWRRVDGW PVGLKNVGNT CWFSAVIQSL FQLPEFRRLV LSYSLPQNVL
210 220 230 240 250
ENCRSHTEKR NIMFMQELQY LFALMMGSNR KFVDPSAALD LLKGAFRSSE
260 270 280 290 300
EQQQDVSEFT HKLLDWLEDA FQLAVNVNSP RNKSENPMVQ LFYGTFLTEG
310 320 330 340 350
VREGKPFCNN ETFGQYPLQV NGYRNLDECL EGAMVEGDVE LLPSDHSVKY
360 370 380 390 400
GQERWFTKLP PVLTFELSRF EFNQSLGQPE KIHNKLEFPQ IIYMDRYMYR
410 420 430 440 450
SKELIRNKRE CIRKLKEEIK ILQQKLERYV KYGSGPARFP LPDMLKYVIE
460 470 480 490 500
FASTKPASES CPPESDTHMT LPLSSVHCSV SDQTSKESTS TESSSQDVES
510 520 530 540 550
TFSSPEDSLP KSKPLTSSRS SMEMPSQPAP RTVTDEEINF VKTCLQRWRS
560 570 580 590 600
EIEQDIQDLK TCIASTTQTI EQMYCDPLLR QVPYRLHAVL VHEGQANAGH
610 620 630 640 650
YWAYIYNQPR QSWLKYNDIS VTESSWEEVE RDSYGGLRNV SAYCLMYIND
660 670 680 690 700
KLPYFNAEAA PTESDQMSEV EALSVELKHY IQEDNWRFEQ EVEEWEEEQS
710 720 730 740 750
CKIPQMESST NSSSQDYSTS QEPSVASSHG VRCLSSEHAV IVKEQTAQAI
760 770 780 790 800
ANTARAYEKS GVEAALSEVM LSPAMQGVIL AIAKARQTFD RDGSEAGLIK
810 820 830 840 850
AFHEEYSRLY QLAKETPTSH SDPRLQHVLV YFFQNEAPKR VVERTLLEQF
860 870 880 890 900
ADKNLSYDER SISIMKVAQA KLKEIGPDDM NMEEYKKWHE DYSLFRKVSV
910 920 930 940 950
YLLTGLELYQ KGKYQEALSY LVYAYQSNAA LLMKGPRRGV KESVIALYRR
960 970 980 990 1000
KCLLELNAKA ASLFETNDDH SVTEGINVMN ELIIPCIHLI INNDISKDDL
1010 1020 1030 1040 1050
DAIEVMRNHW CSYLGQDIAE NLQLCLGEFL PRLLDPSAEI IVLKEPPTIR
1060 1070
PNSPYDLCSR FAAVMESIQG VSTVTVK
Length:1,077
Mass (Da):122,491
Last modified:December 1, 2001 - v1
<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:i799C758888C67B82
GO
Isoform 2 (identifier: Q96RU2-2) [UniParc]FASTAAdd to basket

The sequence of this isoform differs from the canonical sequence as follows:
     769-800: Missing.

Show »
Length:1,045
Mass (Da):119,121
Checksum:i1A10819BCF8CCBC6
GO
Isoform 3 (identifier: Q96RU2-3) [UniParc]FASTAAdd to basket

The sequence of this isoform differs from the canonical sequence as follows:
     583-583: P → E
     584-1077: Missing.

Show »
Length:583
Mass (Da):66,243
Checksum:iA932AD8856535EBD
GO

<p>In eukaryotic reference proteomes, unreviewed entries that are likely to belong to the same gene are computationally mapped, based on gene identifiers from Ensembl, EnsemblGenomes and model organism databases.<p><a href='/help/gene_centric_isoform_mapping' target='_top'>More...</a></p>Computationally mapped potential isoform sequencesi

There are 11 potential isoforms mapped to this entry.BLASTAlignShow allAdd to basket
EntryEntry nameProtein names
Gene namesLengthAnnotation
B4E3L3B4E3L3_HUMAN
Ubiquitin carboxyl-terminal hydrola...
USP28
920Annotation score:

Annotation score:2 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>
G3V1N5G3V1N5_HUMAN
Ubiquitin carboxyl-terminal hydrola...
USP28 hCG_39594
753Annotation score:

Annotation score:1 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>
H0YG96H0YG96_HUMAN
Ubiquitin carboxyl-terminal hydrola...
USP28
360Annotation score:

Annotation score:1 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>
H0YFX3H0YFX3_HUMAN
Ubiquitin carboxyl-terminal hydrola...
USP28
244Annotation score:

Annotation score:1 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>
F5GZ74F5GZ74_HUMAN
Ubiquitin carboxyl-terminal hydrola...
USP28
194Annotation score:

Annotation score:1 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>
H0YFT9H0YFT9_HUMAN
Ubiquitin carboxyl-terminal hydrola...
USP28
194Annotation score:

Annotation score:1 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>
F5H2G4F5H2G4_HUMAN
Ubiquitin carboxyl-terminal hydrola...
USP28
77Annotation score:

Annotation score:1 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>
H0YH19H0YH19_HUMAN
Ubiquitin carboxyl-terminal hydrola...
USP28
91Annotation score:

Annotation score:1 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>
F5H6Q3F5H6Q3_HUMAN
Ubiquitin carboxyl-terminal hydrola...
USP28
46Annotation score:

Annotation score:1 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>
H0YFD7H0YFD7_HUMAN
Ubiquitin carboxyl-terminal hydrola...
USP28
51Annotation score:

Annotation score:1 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>
There is more potential isoformShow all

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 conflicti289 – 303VQLFY…EGVRE → IVIVMSFLKSLSLCL in BAA96039 (Ref. 4) CuratedAdd BLAST15

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_057359583P → E in isoform 3. 1 Publication1
Alternative sequenceiVSP_057360584 – 1077Missing in isoform 3. 1 PublicationAdd BLAST494
Alternative sequenceiVSP_015580769 – 800Missing in isoform 2. 1 PublicationAdd BLAST32

Sequence databases

Select the link destinations:

EMBL nucleotide sequence database

More...
EMBLi

GenBank nucleotide sequence database

More...
GenBanki

DNA Data Bank of Japan; a nucleotide sequence database

More...
DDBJi
Links Updated
AF266283 mRNA Translation: AAK58565.1
EF445045 Genomic DNA Translation: ACA06098.1
EF445045 Genomic DNA Translation: ACA06099.1
AP001874 Genomic DNA No translation available.
AP003170 Genomic DNA No translation available.
KF455523 Genomic DNA No translation available.
KF459544 Genomic DNA No translation available.
CH471065 Genomic DNA Translation: EAW67232.1
BC065928 mRNA Translation: AAH65928.1
AB040948 mRNA Translation: BAA96039.1

The Consensus CDS (CCDS) project

More...
CCDSi
CCDS31680.1 [Q96RU2-1]
CCDS86248.1 [Q96RU2-3]

NCBI Reference Sequences

More...
RefSeqi
NP_001333187.1, NM_001346258.1 [Q96RU2-2]
NP_001333202.1, NM_001346273.1 [Q96RU2-3]
NP_065937.1, NM_020886.3 [Q96RU2-1]

Genome annotation databases

Ensembl eukaryotic genome annotation project

More...
Ensembli
ENST00000003302; ENSP00000003302; ENSG00000048028 [Q96RU2-1]
ENST00000537706; ENSP00000445743; ENSG00000048028 [Q96RU2-3]

Database of genes from NCBI RefSeq genomes

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GeneIDi
57646

KEGG: Kyoto Encyclopedia of Genes and Genomes

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KEGGi
hsa:57646

UCSC genome browser

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UCSCi
uc001poh.4, human [Q96RU2-1]

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
AF266283 mRNA Translation: AAK58565.1
EF445045 Genomic DNA Translation: ACA06098.1
EF445045 Genomic DNA Translation: ACA06099.1
AP001874 Genomic DNA No translation available.
AP003170 Genomic DNA No translation available.
KF455523 Genomic DNA No translation available.
KF459544 Genomic DNA No translation available.
CH471065 Genomic DNA Translation: EAW67232.1
BC065928 mRNA Translation: AAH65928.1
AB040948 mRNA Translation: BAA96039.1
CCDSiCCDS31680.1 [Q96RU2-1]
CCDS86248.1 [Q96RU2-3]
RefSeqiNP_001333187.1, NM_001346258.1 [Q96RU2-2]
NP_001333202.1, NM_001346273.1 [Q96RU2-3]
NP_065937.1, NM_020886.3 [Q96RU2-1]

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
2LVANMR-A22-132[»]
2MUUNMR-A1-120[»]
6H4HX-ray3.50A/B149-707[»]
6H4IX-ray3.22A/C148-707[»]
6HEHX-ray2.26A149-399[»]
A580-703[»]
6HEIX-ray1.64A149-399[»]
A580-703[»]
6HEJX-ray2.79A/B149-703[»]
6HEKX-ray3.03A/C149-703[»]
SMRiQ96RU2
ModBaseiSearch...
PDBe-KBiSearch...

Protein-protein interaction databases

BioGRIDi121683, 80 interactors
IntActiQ96RU2, 19 interactors
MINTiQ96RU2
STRINGi9606.ENSP00000003302

Chemistry databases

ChEMBLiCHEMBL2157853

Protein family/group databases

MEROPSiC19.054

PTM databases

iPTMnetiQ96RU2
PhosphoSitePlusiQ96RU2

Polymorphism and mutation databases

BioMutaiUSP28
DMDMi20140700

Proteomic databases

EPDiQ96RU2
jPOSTiQ96RU2
MassIVEiQ96RU2
MaxQBiQ96RU2
PaxDbiQ96RU2
PeptideAtlasiQ96RU2
PRIDEiQ96RU2
ProteomicsDBi66797
78033 [Q96RU2-1]
78034 [Q96RU2-2]

Protocols and materials databases

Antibodypedia a portal for validated antibodies

More...
Antibodypediai
1708, 274 antibodies

Genome annotation databases

EnsembliENST00000003302; ENSP00000003302; ENSG00000048028 [Q96RU2-1]
ENST00000537706; ENSP00000445743; ENSG00000048028 [Q96RU2-3]
GeneIDi57646
KEGGihsa:57646
UCSCiuc001poh.4, human [Q96RU2-1]

Organism-specific databases

Comparative Toxicogenomics Database

More...
CTDi
57646
DisGeNETi57646
EuPathDBiHostDB:ENSG00000048028.11

GeneCards: human genes, protein and diseases

More...
GeneCardsi
USP28
HGNCiHGNC:12625, USP28
HPAiENSG00000048028, Tissue enhanced (heart muscle, skeletal muscle)
MIMi610748, gene
neXtProtiNX_Q96RU2
OpenTargetsiENSG00000048028
PharmGKBiPA37250

Human Unidentified Gene-Encoded large proteins database

More...
HUGEi
Search...

GenAtlas: human gene database

More...
GenAtlasi
Search...

Phylogenomic databases

eggNOGiKOG1863, Eukaryota
GeneTreeiENSGT00940000157670
HOGENOMiCLU_467647_0_0_1
InParanoidiQ96RU2
KOiK21122
OMAiGDWPVGM
OrthoDBi194025at2759
PhylomeDBiQ96RU2
TreeFamiTF329035

Enzyme and pathway databases

PathwayCommonsiQ96RU2
ReactomeiR-HSA-5689880, Ub-specific processing proteases
SIGNORiQ96RU2

Miscellaneous databases

BioGRID ORCS database of CRISPR phenotype screens

More...
BioGRID-ORCSi
57646, 5 hits in 878 CRISPR screens

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

More...
ChiTaRSi
USP28, human

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

More...
GenomeRNAii
57646
PharosiQ96RU2, Tbio

Protein Ontology

More...
PROi
PR:Q96RU2
RNActiQ96RU2, protein

The Stanford Online Universal Resource for Clones and ESTs

More...
SOURCEi
Search...

Gene expression databases

BgeeiENSG00000048028, Expressed in heart and 208 other tissues
ExpressionAtlasiQ96RU2, baseline and differential
GenevisibleiQ96RU2, HS

Family and domain databases

InterProiView protein in InterPro
IPR038765, Papain-like_cys_pep_sf
IPR001394, Peptidase_C19_UCH
IPR009060, UBA-like_sf
IPR018200, USP_CS
IPR028889, USP_dom
PfamiView protein in Pfam
PF00443, UCH, 1 hit
SUPFAMiSSF46934, SSF46934, 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...

MobiDB: a database of protein disorder and mobility annotations

More...
MobiDBi
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 nameiUBP28_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: Q96RU2
Secondary accession number(s): B0YJC0
, B0YJC1, Q6NZX9, Q9P213
<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%5Fand%5Fisoforms">canonical sequence</a> are also displayed.<p><a href='/help/entry_history' target='_top'>More...</a></p>Entry historyiIntegrated into UniProtKB/Swiss-Prot: March 27, 2002
Last sequence update: December 1, 2001
Last modified: August 12, 2020
This is version 174 of the entry and version 1 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, Reference proteome

Documents

  1. Human chromosome 11
    Human chromosome 11: entries, gene names and cross-references to MIM
  2. MIM cross-references
    Online Mendelian Inheritance in Man (MIM) cross-references in UniProtKB/Swiss-Prot
  3. PDB cross-references
    Index of Protein Data Bank (PDB) cross-references
  4. SIMILARITY comments
    Index of protein domains and families
  5. Peptidase families
    Classification of peptidase families and list of entries
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