ITCH itchy E3 ubiquitin protein ligase [Homo sapiens (human)] - Gene

Summary

Official Symbol: ITCHprovided by HGNC
Official Full Name: itchy E3 ubiquitin protein ligaseprovided by HGNC
Primary source: HGNC:HGNC:13890
See related: Ensembl:ENSG00000078747 MIM:606409; AllianceGenome:HGNC:13890
Gene type: protein coding
RefSeq status: REVIEWED
Organism: Homo sapiens
Lineage: Eukaryota; Metazoa; Chordata; Craniata; Vertebrata; Euteleostomi; Mammalia; Eutheria; Euarchontoglires; Primates; Haplorrhini; Catarrhini; Hominidae; Homo
Also known as: AIF4; AIP4; ADMFD; NAPP1
Summary: This gene encodes a member of the Nedd4 family of HECT domain E3 ubiquitin ligases. HECT domain E3 ubiquitin ligases transfer ubiquitin from E2 ubiquitin-conjugating enzymes to protein substrates, thus targeting specific proteins for lysosomal degradation. The encoded protein plays a role in multiple cellular processes including erythroid and lymphoid cell differentiation and the regulation of immune responses. Mutations in this gene are a cause of syndromic multisystem autoimmune disease. Alternatively spliced transcript variants encoding multiple isoforms have been observed for this gene. [provided by RefSeq, Mar 2012]
Expression: Ubiquitous expression in testis (RPKM 12.7), esophagus (RPKM 11.6) and 25 other tissues See more
Orthologs: mouse all
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Genomic context

Location:: 20q11.22

Exon count:: 31

Annotation release	Status	Assembly	Chr	Location
RS_2023_10	current	GRCh38.p14 (GCF_000001405.40)	20	NC_000020.11 (34363273..34511773)
RS_2023_10	current	T2T-CHM13v2.0 (GCF_009914755.1)	20	NC_060944.1 (36086401..36234950)
105.20220307	previous assembly	GRCh37.p13 (GCF_000001405.25)	20	NC_000020.10 (32951079..33099578)

Chromosome 20 - NC_000020.11 Genomic Context describing neighboring genes

Genomic regions, transcripts, and products

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Go to reference sequence details

Genomic Sequence:

Go to nucleotide: Graphics FASTA GenBank

Expression

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See details

Project title: HPA RNA-seq normal tissues
Description: RNA-seq was performed of tissue samples from 95 human individuals representing 27 different tissues in order to determine tissue-specificity of all protein-coding genes
BioProject: PRJEB4337
Publication: PMID 24309898
Analysis date: Wed Apr 4 07:08:55 2018

Bibliography

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GeneRIFs: Gene References Into Functions

What's a GeneRIF?

Host E3 ubiquitin ligase ITCH mediates Toxoplasma gondii effector GRA35-triggered NLRP1 inflammasome activation and cell-autonomous immunity.
Title: Host E3 ubiquitin ligase ITCH mediates Toxoplasma gondii effector GRA35-triggered NLRP1 inflammasome activation and cell-autonomous immunity.
The E3 ubiquitin ligase Itch regulates death receptor and cholesterol trafficking to affect TRAIL-mediated apoptosis.
Title: The E3 ubiquitin ligase Itch regulates death receptor and cholesterol trafficking to affect TRAIL-mediated apoptosis.
UBE4B interacts with the ITCH E3 ubiquitin ligase to induce Ku70 and c-FLIPL polyubiquitination and enhanced neuroblastoma apoptosis.
Title: UBE4B interacts with the ITCH E3 ubiquitin ligase to induce Ku70 and c-FLIPL polyubiquitination and enhanced neuroblastoma apoptosis.
The Ubiquitin Ligase Itch Skews Light Zone Selection in Germinal Centers.
Title: The Ubiquitin Ligase Itch Skews Light Zone Selection in Germinal Centers.
Circ-ITCH overexpression promoted cell proliferation and migration in Hirschsprung disease through miR-146b-5p/RET axis.
Title: Circ-ITCH overexpression promoted cell proliferation and migration in Hirschsprung disease through miR-146b-5p/RET axis.
Enhancing PD-L1 Degradation by ITCH during MAPK Inhibitor Therapy Suppresses Acquired Resistance.
Title: Enhancing PD-L1 Degradation by ITCH during MAPK Inhibitor Therapy Suppresses Acquired Resistance.
H5N1 infection impairs the alveolar epithelial barrier through intercellular junction proteins via Itch-mediated proteasomal degradation.
Title: H5N1 infection impairs the alveolar epithelial barrier through intercellular junction proteins via Itch-mediated proteasomal degradation.
Circular RNA ITCH suppresses metastasis of gastric cancer via regulating miR-199a-5p/Klotho axis.
Title: Circular RNA ITCH suppresses metastasis of gastric cancer via regulating miR-199a-5p/Klotho axis.
Pirh2, an E3 ligase, regulates the AIP4-p73 regulatory pathway by modulating AIP4 expression and ubiquitination.
Title: Pirh2, an E3 ligase, regulates the AIP4-p73 regulatory pathway by modulating AIP4 expression and ubiquitination.
ITCH as a potential therapeutic target in human cancers.
Title: ITCH as a potential therapeutic target in human cancers.

Submit: New GeneRIF Correction See all GeneRIFs (118)

Phenotypes

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Find tests for this gene in the NIH Genetic Testing Registry (GTR)

Review eQTL and phenotype association data in this region using PheGenI

Associated conditions

Description	Tests
Syndromic multisystem autoimmune disease due to ITCH deficiency MedGen: C3150649 OMIM: 613385 GeneReviews: Not available	Compare labs

Variation

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See variants in ClinVar

See studies and variants in dbVar

See Variation Viewer (GRCh37.p13)

See Variation Viewer (GRCh38)

HIV-1 interactions

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Protein interactions

Protein	Gene	Interaction	Pubs
Nef	nef	HIV-1 Nef co-localizes with CXCR4, AIP4, NEDD4, CD63/LAMP-1-positive vesicles, and HRS/VPS27-positive ESCRT-0 structures in HeLa cells	PubMed
	nef	The Lys residues (K327/331/333) in the C-terminal tail of CXCR4 and the catalytic Cys830 in the HECT domain of AIP4 are critically required for HIV-1 Nef-mediated downregulation of CXCR4	PubMed
	nef	HIV-1 Nef dramatically enhances the binding of AIP4 with CXCR4 and alanine substitutions at positions Q297 and N329 of AIP4 eliminate this binding	PubMed
	nef	HIV-1 Nef-mediated CXCR4 downregulation is dependent on ubiquitinylation, which is mediated by the E3 ubiquitin ligase AIP4	PubMed
	nef	HIV-1 Nef co-localizes with E3 ubiquitin ligases AIP4 and NEDD4. HIV-1 Nef induced degradation of GNAI2 involves AIP4 and NEDD4 through recruitment of AIP4 and NEDD4 by Nef	PubMed
Vif	vif	HIV-1 Vif, through amino acids 20-128, binds to the WW domains of hNedd4 and AIP4, two HECT E3 ubiquitin ligases	PubMed

Go to the HIV-1, Human Interaction Database

Interactions

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Products	Interactant	Other Gene	Complex	Source	Pubs	Description

General gene information

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Markers

A007H06 (e-PCR)
- UniSTS:32777

RH67935 (e-PCR)
- UniSTS:92464

AL034523 (e-PCR)
- UniSTS:94275

ITCH_3687 (e-PCR)
- UniSTS:462281

Z94419 (e-PCR)
- UniSTS:32478

D20S498 (e-PCR)
- UniSTS:6370

Gene Ontology Provided by GOA

Function	Evidence Code	Pubs
enables CXCR chemokine receptor binding	IPI Inferred from Physical Interaction more info	PubMed
enables arrestin family protein binding	IPI Inferred from Physical Interaction more info	PubMed
enables ligase activity	IEA Inferred from Electronic Annotation more info
enables protein binding	IPI Inferred from Physical Interaction more info	PubMed
enables ribonucleoprotein complex binding	IPI Inferred from Physical Interaction more info	PubMed
enables ubiquitin protein ligase activity	IBA Inferred from Biological aspect of Ancestor more info
enables ubiquitin protein ligase activity	IDA Inferred from Direct Assay more info	PubMed
enables ubiquitin protein ligase activity	IMP Inferred from Mutant Phenotype more info	PubMed
enables ubiquitin protein ligase activity	TAS Traceable Author Statement more info
enables ubiquitin-like protein ligase binding	IPI Inferred from Physical Interaction more info	PubMed
enables ubiquitin-like protein transferase activity	TAS Traceable Author Statement more info
enables ubiquitin-protein transferase activity	EXP Inferred from Experiment more info	PubMed
enables ubiquitin-protein transferase activity	IDA Inferred from Direct Assay more info	PubMed
enables ubiquitin-protein transferase activity	IMP Inferred from Mutant Phenotype more info	PubMed
enables ubiquitin-protein transferase activity	NAS Non-traceable Author Statement more info	PubMed

Process	Evidence Code	Pubs
involved_in CD4-positive, alpha-beta T cell proliferation	IEA Inferred from Electronic Annotation more info
involved_in T cell anergy	IEA Inferred from Electronic Annotation more info
involved_in apoptotic process	IEA Inferred from Electronic Annotation more info
involved_in defense response to virus	IEA Inferred from Electronic Annotation more info
involved_in inflammatory response	NAS Non-traceable Author Statement more info	PubMed
involved_in innate immune response	IEA Inferred from Electronic Annotation more info
involved_in negative regulation of CD4-positive, alpha-beta T cell proliferation	IEA Inferred from Electronic Annotation more info
involved_in negative regulation of JNK cascade	ISS Inferred from Sequence or Structural Similarity more info
involved_in negative regulation of NF-kappaB transcription factor activity	ISS Inferred from Sequence or Structural Similarity more info
involved_in negative regulation of apoptotic process	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in negative regulation of cytoplasmic pattern recognition receptor signaling pathway	IDA Inferred from Direct Assay more info	PubMed
involved_in negative regulation of defense response to virus	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in negative regulation of type I interferon production	TAS Traceable Author Statement more info
involved_in nucleotide-binding domain, leucine rich repeat containing receptor signaling pathway	TAS Traceable Author Statement more info
involved_in positive regulation of T cell anergy	IEA Inferred from Electronic Annotation more info
involved_in positive regulation of protein catabolic process	IEA Inferred from Electronic Annotation more info
involved_in positive regulation of receptor catabolic process	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in proteasome-mediated ubiquitin-dependent protein catabolic process	IBA Inferred from Biological aspect of Ancestor more info
involved_in proteasome-mediated ubiquitin-dependent protein catabolic process	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in protein K29-linked ubiquitination	IBA Inferred from Biological aspect of Ancestor more info
involved_in protein K29-linked ubiquitination	IDA Inferred from Direct Assay more info	PubMed
involved_in protein K48-linked ubiquitination	IBA Inferred from Biological aspect of Ancestor more info
involved_in protein K48-linked ubiquitination	IDA Inferred from Direct Assay more info	PubMed
involved_in protein K63-linked ubiquitination	IBA Inferred from Biological aspect of Ancestor more info
involved_in protein K63-linked ubiquitination	IDA Inferred from Direct Assay more info	PubMed
involved_in protein autoubiquitination	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in protein monoubiquitination	IDA Inferred from Direct Assay more info	PubMed
involved_in protein ubiquitination	IDA Inferred from Direct Assay more info	PubMed
involved_in protein ubiquitination	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in receptor internalization	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in regulation of cell growth	NAS Non-traceable Author Statement more info	PubMed
involved_in regulation of hematopoietic stem cell differentiation	TAS Traceable Author Statement more info
involved_in regulation of necroptotic process	TAS Traceable Author Statement more info
involved_in regulation of protein deubiquitination	ISS Inferred from Sequence or Structural Similarity more info
involved_in symbiont entry into host cell	TAS Traceable Author Statement more info	PubMed
acts_upstream_of_or_within ubiquitin-dependent protein catabolic process	IDA Inferred from Direct Assay more info	PubMed
involved_in ubiquitin-dependent protein catabolic process	IMP Inferred from Mutant Phenotype more info	PubMed
involved_in ubiquitin-dependent protein catabolic process	NAS Non-traceable Author Statement more info	PubMed

Component	Evidence Code	Pubs
located_in cell cortex	IEA Inferred from Electronic Annotation more info
is_active_in cytoplasm	IBA Inferred from Biological aspect of Ancestor more info
located_in cytoplasmic vesicle	IDA Inferred from Direct Assay more info	PubMed
located_in cytosol	TAS Traceable Author Statement more info
colocalizes_with early endosome	IDA Inferred from Direct Assay more info	PubMed
located_in early endosome membrane	IEA Inferred from Electronic Annotation more info
located_in extracellular exosome	HDA more info	PubMed
located_in intracellular membrane-bounded organelle	IDA Inferred from Direct Assay more info
located_in membrane	HDA more info	PubMed
is_active_in mitochondrion	IC Inferred by Curator more info	PubMed
located_in nucleoplasm	IDA Inferred from Direct Assay more info
is_active_in plasma membrane	IDA Inferred from Direct Assay more info	PubMed
located_in plasma membrane	IDA Inferred from Direct Assay more info	PubMed
part_of protein-containing complex	IDA Inferred from Direct Assay more info	PubMed

General protein information

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Preferred Names: E3 ubiquitin-protein ligase Itchy homolog

Names: HECT-type E3 ubiquitin transferase Itchy homolog; NFE2-associated polypeptide 1; atrophin-1 interacting protein 4; itchy E3 ubiquitin protein ligase homolog

NP_001244066.1

EC 2.3.2.26

NP_001244067.1

EC 2.3.2.26

NP_001311126.1

EC 2.3.2.26

NP_001311127.1

EC 2.3.2.26

NP_113671.3

EC 2.3.2.26

XP_016883578.1

EC 2.3.2.26

XP_024307773.1

EC 2.3.2.26

XP_047296488.1

EC 2.3.2.26

XP_047296489.1

EC 2.3.2.26

XP_047296490.1

EC 2.3.2.26

XP_047296491.1

EC 2.3.2.26

XP_047296492.1

EC 2.3.2.26

XP_047296493.1

EC 2.3.2.26

XP_047296494.1

EC 2.3.2.26

XP_047296495.1

EC 2.3.2.26

XP_054180074.1

EC 2.3.2.26

XP_054180075.1

EC 2.3.2.26

XP_054180076.1

EC 2.3.2.26

XP_054180077.1

EC 2.3.2.26

XP_054180078.1

EC 2.3.2.26

XP_054180079.1

EC 2.3.2.26

XP_054180080.1

EC 2.3.2.26

XP_054180081.1

EC 2.3.2.26

XP_054180082.1

EC 2.3.2.26

XP_054180083.1

EC 2.3.2.26

NCBI Reference Sequences (RefSeq)

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RefSeqs maintained independently of Annotated Genomes

These reference sequences exist independently of genome builds. Explain

These reference sequences are curated independently of the genome annotation cycle, so their versions may not match the RefSeq versions in the current genome build. Identify version mismatches by comparing the version of the RefSeq in this section to the one reported in Genomic regions, transcripts, and products above.

Genomic

NG_023206.1 RefSeqGene

Range

4980..153137

Download

GenBank, FASTA, Sequence Viewer (Graphics), LRG_354

mRNA and Protein(s)

NM_001257137.3 → NP_001244066.1 E3 ubiquitin-protein ligase Itchy homolog isoform 1

See identical proteins and their annotated locations for NP_001244066.1

Status: REVIEWED

Description

Transcript Variant: This variant (1) encodes the longest isoform (1). Both variants 1 and 4 encode the same isoform (1).

Source sequence(s)

AA730741, AL109923, AL356299, AL705688, BC011571, BG928040

Consensus CDS

CCDS58768.1

UniProtKB/Swiss-Prot

A6NEW4, B4E234, E1P5P3, F5H217, O43584, Q5QP37, Q5TEL0, Q96F66, Q96J02, Q9BY75, Q9H451, Q9H4U5

UniProtKB/TrEMBL

A0A590UJQ1

Related

ENSP00000262650.5, ENST00000262650.11

Conserved Domains (3) summary

cd04021
Location:17 → 140

C2_E3_ubiquitin_ligase; C2 domain present in E3 ubiquitin ligase

COG5021
Location:359 → 903

HUL4; Ubiquitin-protein ligase [Posttranslational modification, protein turnover, chaperones]

pfam00397
Location:328 → 357

WW; WW domain
NM_001257138.3 → NP_001244067.1 E3 ubiquitin-protein ligase Itchy homolog isoform 3

See identical proteins and their annotated locations for NP_001244067.1

Status: REVIEWED

Description

Transcript Variant: This variant (3) lacks two internal exons and initiates translation at a downstream, in-frame start codon, compared to variant 1. The encoded isoform (3) is shorter than isoform 1.

Source sequence(s)

AA730741, AF038564, AK304090, AL109923, AL356299, AL705688, BG928040

Consensus CDS

CCDS58769.1

UniProtKB/TrEMBL

B4DN85

Related

ENSP00000445608.1, ENST00000535650.8

Conserved Domains (3) summary

COG5021
Location:208 → 752

HUL4; Ubiquitin-protein ligase [Posttranslational modification, protein turnover, chaperones]

pfam00397
Location:177 → 206

WW; WW domain

cl14603
Location:1 → 30

C2; C2 domain
NM_001324197.2 → NP_001311126.1 E3 ubiquitin-protein ligase Itchy homolog isoform 1

Status: REVIEWED

Source sequence(s)

AL109923, AL356299, BG928040

Consensus CDS

CCDS58768.1

UniProtKB/Swiss-Prot

A6NEW4, B4E234, E1P5P3, F5H217, O43584, Q5QP37, Q5TEL0, Q96F66, Q96J02, Q9BY75, Q9H451, Q9H4U5

UniProtKB/TrEMBL

A0A590UJQ1

Related

ENSP00000499786.1, ENST00000665346.1

Conserved Domains (3) summary

cd04021
Location:17 → 140

C2_E3_ubiquitin_ligase; C2 domain present in E3 ubiquitin ligase

COG5021
Location:359 → 903

HUL4; Ubiquitin-protein ligase [Posttranslational modification, protein turnover, chaperones]

pfam00397
Location:328 → 357

WW; WW domain
NM_001324198.2 → NP_001311127.1 E3 ubiquitin-protein ligase Itchy homolog isoform 2

Status: REVIEWED

Source sequence(s)

AL109923, AL356299, BG928040

Consensus CDS

CCDS13234.1

UniProtKB/TrEMBL

A0A590UJQ1

Related

ENSP00000499685.1, ENST00000660337.1

Conserved Domains (3) summary

cd04021
Location:17 → 140

C2_E3_ubiquitin_ligase; C2 domain present in E3 ubiquitin ligase

COG5021
Location:318 → 862

HUL4; Ubiquitin-protein ligase [Posttranslational modification, protein turnover, chaperones]

pfam00397
Location:287 → 316

WW; WW domain
NM_031483.7 → NP_113671.3 E3 ubiquitin-protein ligase Itchy homolog isoform 2

See identical proteins and their annotated locations for NP_113671.3

Status: REVIEWED

Description

Transcript Variant: This variant (2) lacks an exon in the 5' coding region, but maintains the reading frame, compared to variant 1. The encoded isoform (2) is shorter than isoform 1. Both variants 2 and 5 encode the same isoform (2).

Source sequence(s)

AA730741, AK315212, AL109923, AL356299, AL705688, BG928040

Consensus CDS

CCDS13234.1

UniProtKB/TrEMBL

A0A590UJQ1

Related

ENSP00000363998.4, ENST00000374864.10

Conserved Domains (3) summary

cd04021
Location:17 → 140

C2_E3_ubiquitin_ligase; C2 domain present in E3 ubiquitin ligase

COG5021
Location:318 → 862

HUL4; Ubiquitin-protein ligase [Posttranslational modification, protein turnover, chaperones]

pfam00397
Location:287 → 316

WW; WW domain

RefSeqs of Annotated Genomes: GCF_000001405.40-RS_2023_10

The following sections contain reference sequences that belong to a specific genome build. Explain

Reference GRCh38.p14 Primary Assembly

Genomic

NC_000020.11 Reference GRCh38.p14 Primary Assembly

Range

34363273..34511773

Download

GenBank, FASTA, Sequence Viewer (Graphics)

mRNA and Protein(s)

XM_047440533.1 → XP_047296489.1 E3 ubiquitin-protein ligase Itchy homolog isoform X3
XM_047440538.1 → XP_047296494.1 E3 ubiquitin-protein ligase Itchy homolog isoform X7
XM_047440539.1 → XP_047296495.1 E3 ubiquitin-protein ligase Itchy homolog isoform X8
XM_024452005.2 → XP_024307773.1 E3 ubiquitin-protein ligase Itchy homolog isoform X1

UniProtKB/Swiss-Prot

A6NEW4, B4E234, E1P5P3, F5H217, O43584, Q5QP37, Q5TEL0, Q96F66, Q96J02, Q9BY75, Q9H451, Q9H4U5

UniProtKB/TrEMBL

A0A590UJQ1

Conserved Domains (3) summary

cd04021
Location:17 → 140

C2_E3_ubiquitin_ligase; C2 domain present in E3 ubiquitin ligase

COG5021
Location:359 → 903

HUL4; Ubiquitin-protein ligase [Posttranslational modification, protein turnover, chaperones]

pfam00397
Location:328 → 357

WW; WW domain
XM_017028089.2 → XP_016883578.1 E3 ubiquitin-protein ligase Itchy homolog isoform X1

UniProtKB/Swiss-Prot

A6NEW4, B4E234, E1P5P3, F5H217, O43584, Q5QP37, Q5TEL0, Q96F66, Q96J02, Q9BY75, Q9H451, Q9H4U5

UniProtKB/TrEMBL

A0A590UJQ1

Conserved Domains (3) summary

cd04021
Location:17 → 140

C2_E3_ubiquitin_ligase; C2 domain present in E3 ubiquitin ligase

COG5021
Location:359 → 903

HUL4; Ubiquitin-protein ligase [Posttranslational modification, protein turnover, chaperones]

pfam00397
Location:328 → 357

WW; WW domain
XM_047440532.1 → XP_047296488.1 E3 ubiquitin-protein ligase Itchy homolog isoform X2
XM_047440534.1 → XP_047296490.1 E3 ubiquitin-protein ligase Itchy homolog isoform X3
XM_047440535.1 → XP_047296491.1 E3 ubiquitin-protein ligase Itchy homolog isoform X4
XM_047440537.1 → XP_047296493.1 E3 ubiquitin-protein ligase Itchy homolog isoform X6
XM_047440536.1 → XP_047296492.1 E3 ubiquitin-protein ligase Itchy homolog isoform X5

Alternate T2T-CHM13v2.0

Genomic

NC_060944.1 Alternate T2T-CHM13v2.0

Range

36086401..36234950

Download

GenBank, FASTA, Sequence Viewer (Graphics)

mRNA and Protein(s)

XM_054324102.1 → XP_054180077.1 E3 ubiquitin-protein ligase Itchy homolog isoform X3
XM_054324107.1 → XP_054180082.1 E3 ubiquitin-protein ligase Itchy homolog isoform X7
XM_054324108.1 → XP_054180083.1 E3 ubiquitin-protein ligase Itchy homolog isoform X8
XM_054324099.1 → XP_054180074.1 E3 ubiquitin-protein ligase Itchy homolog isoform X1

UniProtKB/Swiss-Prot

A6NEW4, B4E234, E1P5P3, F5H217, O43584, Q5QP37, Q5TEL0, Q96F66, Q96J02, Q9BY75, Q9H451, Q9H4U5
XM_054324100.1 → XP_054180075.1 E3 ubiquitin-protein ligase Itchy homolog isoform X1

UniProtKB/Swiss-Prot

A6NEW4, B4E234, E1P5P3, F5H217, O43584, Q5QP37, Q5TEL0, Q96F66, Q96J02, Q9BY75, Q9H451, Q9H4U5
XM_054324101.1 → XP_054180076.1 E3 ubiquitin-protein ligase Itchy homolog isoform X2
XM_054324103.1 → XP_054180078.1 E3 ubiquitin-protein ligase Itchy homolog isoform X3
XM_054324104.1 → XP_054180079.1 E3 ubiquitin-protein ligase Itchy homolog isoform X4
XM_054324106.1 → XP_054180081.1 E3 ubiquitin-protein ligase Itchy homolog isoform X6
XM_054324105.1 → XP_054180080.1 E3 ubiquitin-protein ligase Itchy homolog isoform X5