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Protein

Muscle, skeletal receptor tyrosine-protein kinase

Gene

MUSK

Organism
Homo sapiens (Human)
Status
Reviewed-Annotation score: -Experimental evidence at protein leveli

Functioni

Receptor tyrosine kinase which plays a central role in the formation and the maintenance of the neuromuscular junction (NMJ), the synapse between the motor neuron and the skeletal muscle (PubMed:25537362). Recruitment of AGRIN by LRP4 to the MUSK signaling complex induces phosphorylation and activation of MUSK, the kinase of the complex. The activation of MUSK in myotubes regulates the formation of NMJs through the regulation of different processes including the specific expression of genes in subsynaptic nuclei, the reorganization of the actin cytoskeleton and the clustering of the acetylcholine receptors (AChR) in the postsynaptic membrane. May regulate AChR phosphorylation and clustering through activation of ABL1 and Src family kinases which in turn regulate MUSK. DVL1 and PAK1 that form a ternary complex with MUSK are also important for MUSK-dependent regulation of AChR clustering. May positively regulate Rho family GTPases through FNTA. Mediates the phosphorylation of FNTA which promotes prenylation, recruitment to membranes and activation of RAC1 a regulator of the actin cytoskeleton and of gene expression. Other effectors of the MUSK signaling include DNAJA3 which functions downstream of MUSK. May also play a role within the central nervous system by mediating cholinergic responses, synaptic plasticity and memory formation (By similarity).By similarity1 Publication

Catalytic activityi

ATP + a [protein]-L-tyrosine = ADP + a [protein]-L-tyrosine phosphate.PROSITE-ProRule annotation1 Publication

Cofactori

Mg2+1 Publication

Enzyme regulationi

Positively regulated by CK2.By similarity

Sites

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Binding sitei609ATPPROSITE-ProRule annotation1
Active sitei725Proton acceptorPROSITE-ProRule annotation1

Regions

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Nucleotide bindingi581 – 589ATPPROSITE-ProRule annotation9

GO - Molecular functioni

GO - Biological processi

Keywordsi

Molecular functionDevelopmental protein, Kinase, Muscle protein, Receptor, Transferase, Tyrosine-protein kinase
Biological processDifferentiation
LigandATP-binding, Magnesium, Metal-binding, Nucleotide-binding

Enzyme and pathway databases

ReactomeiR-HSA-3000178 ECM proteoglycans
SignaLinkiO15146
SIGNORiO15146

Names & Taxonomyi

Protein namesi
Recommended name:
Muscle, skeletal receptor tyrosine-protein kinase (EC:2.7.10.11 Publication)
Alternative name(s):
Muscle-specific tyrosine-protein kinase receptor
Short name:
MuSK
Short name:
Muscle-specific kinase receptor
Gene namesi
Name:MUSK
OrganismiHomo sapiens (Human)
Taxonomic identifieri9606 [NCBI]
Taxonomic lineageiEukaryotaMetazoaChordataCraniataVertebrataEuteleostomiMammaliaEutheriaEuarchontogliresPrimatesHaplorrhiniCatarrhiniHominidaeHomo
Proteomesi
  • UP000005640 Componenti: Chromosome 9

Organism-specific databases

EuPathDBiHostDB:ENSG00000030304.13
HGNCiHGNC:7525 MUSK
MIMi601296 gene
neXtProtiNX_O15146

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; Source: COMPARTMENTS

Topology

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Topological domaini24 – 495ExtracellularSequence analysisAdd BLAST472
Transmembranei496 – 516HelicalSequence analysisAdd BLAST21
Topological domaini517 – 869CytoplasmicSequence analysisAdd BLAST353

Keywords - Cellular componenti

Cell junction, Cell membrane, Membrane, Postsynaptic cell membrane, Synapse

Pathology & Biotechi

Involvement in diseasei

Myasthenic syndrome, congenital, 9, associated with acetylcholine receptor deficiency (CMS9)5 Publications
The disease is caused by mutations affecting the gene represented in this entry. MUSK mutations lead to decreased agrin-dependent AChR aggregation, a critical step in the formation of the neuromuscular junction.
Disease descriptionA form of congenital myasthenic syndrome, a group of disorders characterized by failure of neuromuscular transmission, including pre-synaptic, synaptic, and post-synaptic disorders that are not of autoimmune origin. Clinical features are easy fatigability and muscle weakness affecting the axial and limb muscles (with hypotonia in early-onset forms), the ocular muscles (leading to ptosis and ophthalmoplegia), and the facial and bulbar musculature (affecting sucking and swallowing, and leading to dysphonia). The symptoms fluctuate and worsen with physical effort. CMS9 is a disorder of postsynaptic neuromuscular transmission, due to deficiency of AChR at the endplate that results in low amplitude of the miniature endplate potential and current.
See also OMIM:616325
Feature keyPosition(s)DescriptionActionsGraphical viewLength
Natural variantiVAR_07278538D → E in CMS9. 1 PublicationCorresponds to variant dbSNP:rs775587809Ensembl.1
Natural variantiVAR_072786344P → R in CMS9. 1 PublicationCorresponds to variant dbSNP:rs387906803EnsemblClinVar.1
Natural variantiVAR_066604605M → I in CMS9; affects interaction with DOK7 and impairs MUSK phosphorylation; altered AChR clustering. 1 PublicationCorresponds to variant dbSNP:rs766640370Ensembl.1
Natural variantiVAR_066605727A → V in CMS9; affects interaction with DOK7 and impairs MUSK phosphorylation; altered AChR clustering. 1 PublicationCorresponds to variant dbSNP:rs397515450EnsemblClinVar.1
Natural variantiVAR_023046790V → M in CMS9; does not affect catalytic kinase activity; reduces protein expression and stability. 1 PublicationCorresponds to variant dbSNP:rs199476083EnsemblClinVar.1
Natural variantiVAR_072788835M → V in CMS9; reduces AChR aggregation in developing neuromuscular junction. 1 Publication1
Fetal akinesia deformation sequence (FADS)2 Publications
The disease is caused by mutations affecting the gene represented in this entry.
Disease descriptionA clinically and genetically heterogeneous group of disorders with congenital malformations related to impaired fetal movement. Clinical features include fetal akinesia, intrauterine growth retardation, polyhydramnios, arthrogryposis, pulmonary hypoplasia, craniofacial abnormalities, and cryptorchidism.
See also OMIM:208150
Feature keyPosition(s)DescriptionActionsGraphical viewLength
Natural variantiVAR_072787575I → T in FADS; reduces agrin-dependent AChR aggregation and tyrosin kinase activity in developing neuromuscular junction. 1 PublicationCorresponds to variant dbSNP:rs751889864EnsemblClinVar.1

Mutagenesis

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Mutagenesisi584G → C or D: Mild decrease in kinase activity. 1 Publication1
Mutagenesisi609K → R: Severe loss of kinase activity. 1 Publication1
Mutagenesisi743D → N: Severe loss of kinase activity. 1 Publication1

Keywords - Diseasei

Congenital myasthenic syndrome, Disease mutation

Organism-specific databases

DisGeNETi4593
GeneReviewsiMUSK
MalaCardsiMUSK
MIMi208150 phenotype
616325 phenotype
OpenTargetsiENSG00000030304
Orphaneti98913 Postsynaptic congenital myasthenic syndromes
PharmGKBiPA31326

Chemistry databases

ChEMBLiCHEMBL5684
GuidetoPHARMACOLOGYi1847

Polymorphism and mutation databases

BioMutaiMUSK

PTM / Processingi

Molecule processing

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Signal peptidei1 – 23Sequence analysisAdd BLAST23
ChainiPRO_000002444624 – 869Muscle, skeletal receptor tyrosine-protein kinaseAdd BLAST846

Amino acid modifications

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Disulfide bondi49 ↔ 99By similarity
Disulfide bondi98 ↔ 112By similarity
Disulfide bondi142 ↔ 190By similarity
Glycosylationi222N-linked (GlcNAc...) asparagineSequence analysis1
Disulfide bondi233 ↔ 282By similarity
Disulfide bondi317 ↔ 382By similarity
Disulfide bondi325 ↔ 375By similarity
Glycosylationi338N-linked (GlcNAc...) asparagineBy similarity1
Disulfide bondi366 ↔ 406By similarity
Disulfide bondi394 ↔ 447By similarity
Disulfide bondi398 ↔ 434By similarity
Modified residuei554Phosphotyrosine; by autocatalysisBy similarity1
Modified residuei681Phosphoserine; by CK2By similarity1
Modified residuei698Phosphoserine; by CK2By similarity1
Modified residuei755Phosphotyrosine; by autocatalysisBy similarity1

Post-translational modificationi

Ubiquitinated by PDZRN3. Ubiquitination promotes endocytosis and lysosomal degradation (By similarity).By similarity
Phosphorylated. Phosphorylation is induced by AGRIN in a LRP4-dependent manner (By similarity). Autophosphorylated (PubMed:25029443). Autophosphorylation at Tyr-554 is required for interaction with DOK7 which in turn stimulates the phosphorylation and the activation of MUSK (By similarity).By similarity1 Publication
Neddylated.1 Publication

Keywords - PTMi

Disulfide bond, Glycoprotein, Phosphoprotein, Ubl conjugation

Proteomic databases

PaxDbiO15146
PeptideAtlasiO15146
PRIDEiO15146
ProteomicsDBi48471
48472 [O15146-2]
48473 [O15146-3]

PTM databases

iPTMnetiO15146
PhosphoSitePlusiO15146

Expressioni

Gene expression databases

BgeeiENSG00000030304
CleanExiHS_MUSK
ExpressionAtlasiO15146 baseline and differential
GenevisibleiO15146 HS

Interactioni

Subunit structurei

Monomer (By similarity). Homodimer (Probable). Interacts with LRP4; the heterodimer forms an AGRIN receptor complex that binds AGRIN resulting in activation of MUSK (By similarity). Forms a heterotetramer composed of 2 DOK7 and 2 MUSK molecules which facilitates MUSK trans-autophosphorylation on tyrosine residue and activation. Interacts (via cytoplasmic part) with DOK7 (via IRS-type PTB domain); requires MUSK phosphorylation. Interacts with DVL1 (via DEP domain); the interaction is direct and mediates the formation of a DVL1, MUSK and PAK1 ternary complex involved in AChR clustering (By similarity). Interacts with PDZRN3; this interaction is enhanced by agrin (By similarity). Interacts with FNTA; the interaction is direct and mediates AGRIN-induced phosphorylation and activation of FNTA (By similarity). Interacts with CSNK2B; mediates regulation by CK2 (By similarity). Interacts (via the cytoplasmic domain) with DNAJA3 (By similarity). Interacts with NSF; may regulate MUSK endocytosis and activity (By similarity). Interacts with CAV3; may regulate MUSK signaling (By similarity). Interacts with RNF31 (By similarity).By similarityCurated

Binary interactionsi

WithEntry#Exp.IntActNotes
HSP90AB1P082382EBI-6423196,EBI-352572

Protein-protein interaction databases

BioGridi110679, 8 interactors
IntActiO15146, 6 interactors
MINTiO15146
STRINGi9606.ENSP00000363571

Chemistry databases

BindingDBiO15146

Structurei

3D structure databases

ProteinModelPortaliO15146
SMRiO15146
ModBaseiSearch...
MobiDBiSearch...

Family & Domainsi

Domains and Repeats

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Domaini28 – 116Ig-like 1Add BLAST89
Domaini121 – 205Ig-like 2Add BLAST85
Domaini212 – 302Ig-like 3Add BLAST91
Domaini312 – 450FZPROSITE-ProRule annotationAdd BLAST139
Domaini575 – 856Protein kinasePROSITE-ProRule annotationAdd BLAST282

Sequence similaritiesi

Belongs to the protein kinase superfamily. Tyr protein kinase family.PROSITE-ProRule annotation

Keywords - Domaini

Immunoglobulin domain, Repeat, Signal, Transmembrane, Transmembrane helix

Phylogenomic databases

eggNOGiENOG410IMMJ Eukaryota
COG0515 LUCA
GeneTreeiENSGT00760000118818
HOGENOMiHOG000044461
HOVERGENiHBG052539
InParanoidiO15146
KOiK05129
OMAiKGYCAQY
OrthoDBiEOG091G016V
PhylomeDBiO15146
TreeFamiTF106465

Family and domain databases

Gene3Di1.10.2000.10, 1 hit
2.60.40.10, 3 hits
InterProiView protein in InterPro
IPR020067 Frizzled_dom
IPR036790 Frizzled_dom_sf
IPR007110 Ig-like_dom
IPR036179 Ig-like_dom_sf
IPR013783 Ig-like_fold
IPR013098 Ig_I-set
IPR003599 Ig_sub
IPR003598 Ig_sub2
IPR011009 Kinase-like_dom_sf
IPR000719 Prot_kinase_dom
IPR017441 Protein_kinase_ATP_BS
IPR001245 Ser-Thr/Tyr_kinase_cat_dom
IPR008266 Tyr_kinase_AS
IPR020635 Tyr_kinase_cat_dom
PfamiView protein in Pfam
PF01392 Fz, 1 hit
PF07679 I-set, 2 hits
PF07714 Pkinase_Tyr, 1 hit
PRINTSiPR00109 TYRKINASE
SMARTiView protein in SMART
SM00409 IG, 3 hits
SM00408 IGc2, 3 hits
SM00219 TyrKc, 1 hit
SUPFAMiSSF48726 SSF48726, 3 hits
SSF56112 SSF56112, 1 hit
PROSITEiView protein in PROSITE
PS50038 FZ, 1 hit
PS50835 IG_LIKE, 3 hits
PS00107 PROTEIN_KINASE_ATP, 1 hit
PS50011 PROTEIN_KINASE_DOM, 1 hit
PS00109 PROTEIN_KINASE_TYR, 1 hit

Sequences (3)i

Sequence statusi: Complete.

Sequence processingi: The displayed sequence is further processed into a mature form.

This entry describes 3 isoformsi produced by alternative splicing. AlignAdd to basket

Isoform 1 (identifier: O15146-1) [UniParc]FASTAAdd to basket

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
MRELVNIPLV HILTLVAFSG TEKLPKAPVI TTPLETVDAL VEEVATFMCA
60 70 80 90 100
VESYPQPEIS WTRNKILIKL FDTRYSIREN GQLLTILSVE DSDDGIYCCT
110 120 130 140 150
ANNGVGGAVE SCGALQVKMK PKITRPPINV KIIEGLKAVL PCTTMGNPKP
160 170 180 190 200
SVSWIKGDSP LRENSRIAVL ESGSLRIHNV QKEDAGQYRC VAKNSLGTAY
210 220 230 240 250
SKVVKLEVEV FARILRAPES HNVTFGSFVT LHCTATGIPV PTITWIENGN
260 270 280 290 300
AVSSGSIQES VKDRVIDSRL QLFITKPGLY TCIATNKHGE KFSTAKAAAT
310 320 330 340 350
ISIAEWSKPQ KDNKGYCAQY RGEVCNAVLA KDALVFLNTS YADPEEAQEL
360 370 380 390 400
LVHTAWNELK VVSPVCRPAA EALLCNHIFQ ECSPGVVPTP IPICREYCLA
410 420 430 440 450
VKELFCAKEW LVMEEKTHRG LYRSEMHLLS VPECSKLPSM HWDPTACARL
460 470 480 490 500
PHLDYNKENL KTFPPMTSSK PSVDIPNLPS SSSSSFSVSP TYSMTVIISI
510 520 530 540 550
MSSFAIFVLL TITTLYCCRR RKQWKNKKRE SAAVTLTTLP SELLLDRLHP
560 570 580 590 600
NPMYQRMPLL LNPKLLSLEY PRNNIEYVRD IGEGAFGRVF QARAPGLLPY
610 620 630 640 650
EPFTMVAVKM LKEEASADMQ ADFQREAALM AEFDNPNIVK LLGVCAVGKP
660 670 680 690 700
MCLLFEYMAY GDLNEFLRSM SPHTVCSLSH SDLSMRAQVS SPGPPPLSCA
710 720 730 740 750
EQLCIARQVA AGMAYLSERK FVHRDLATRN CLVGENMVVK IADFGLSRNI
760 770 780 790 800
YSADYYKANE NDAIPIRWMP PESIFYNRYT TESDVWAYGV VLWEIFSYGL
810 820 830 840 850
QPYYGMAHEE VIYYVRDGNI LSCPENCPVE LYNLMRLCWS KLPADRPSFT
860
SIHRILERMC ERAEGTVSV
Length:869
Mass (Da):97,056
Last modified:January 1, 1998 - v1
Checksum:i3DDC20E179FA010C
GO
Isoform 2 (identifier: O15146-2) [UniParc]FASTAAdd to basket

The sequence of this isoform differs from the canonical sequence as follows:
     209-209: E → EEESEPEQDTK
     307-394: Missing.
     454-462: DYNKENLKT → A

Show »
Length:783
Mass (Da):87,598
Checksum:i695BD37016C0D980
GO
Isoform 3 (identifier: O15146-3) [UniParc]FASTAAdd to basket

The sequence of this isoform differs from the canonical sequence as follows:
     307-394: Missing.
     454-462: DYNKENLKT → A

Show »
Length:773
Mass (Da):86,425
Checksum:i3BEA481E3C84D000
GO

Sequence cautioni

The sequence CAH69977 differs from that shown. Reason: Erroneous gene model prediction.Curated
The sequence CAH69978 differs from that shown. Reason: Erroneous gene model prediction.Curated
The sequence CAI17349 differs from that shown. Reason: Erroneous gene model prediction.Curated
The sequence CAI17350 differs from that shown. Reason: Erroneous gene model prediction.Curated

Natural variant

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Natural variantiVAR_04174827A → G1 PublicationCorresponds to variant dbSNP:rs56054734Ensembl.1
Natural variantiVAR_07278538D → E in CMS9. 1 PublicationCorresponds to variant dbSNP:rs775587809Ensembl.1
Natural variantiVAR_041749100T → M1 PublicationCorresponds to variant dbSNP:rs35142681EnsemblClinVar.1
Natural variantiVAR_041750107G → E1 PublicationCorresponds to variant dbSNP:rs55786136EnsemblClinVar.1
Natural variantiVAR_041751159S → G1 PublicationCorresponds to variant dbSNP:rs35176182EnsemblClinVar.1
Natural variantiVAR_041752222N → S1 PublicationCorresponds to variant dbSNP:rs55826142EnsemblClinVar.1
Natural variantiVAR_072786344P → R in CMS9. 1 PublicationCorresponds to variant dbSNP:rs387906803EnsemblClinVar.1
Natural variantiVAR_021930413M → I1 PublicationCorresponds to variant dbSNP:rs2274419EnsemblClinVar.1
Natural variantiVAR_072787575I → T in FADS; reduces agrin-dependent AChR aggregation and tyrosin kinase activity in developing neuromuscular junction. 1 PublicationCorresponds to variant dbSNP:rs751889864EnsemblClinVar.1
Natural variantiVAR_066604605M → I in CMS9; affects interaction with DOK7 and impairs MUSK phosphorylation; altered AChR clustering. 1 PublicationCorresponds to variant dbSNP:rs766640370Ensembl.1
Natural variantiVAR_041753629L → F1 PublicationCorresponds to variant dbSNP:rs34267283Ensembl.1
Natural variantiVAR_041754644V → A1 PublicationCorresponds to variant dbSNP:rs41279055EnsemblClinVar.1
Natural variantiVAR_041755664N → S1 PublicationCorresponds to variant dbSNP:rs55963442EnsemblClinVar.1
Natural variantiVAR_041756696P → L1 PublicationCorresponds to variant dbSNP:rs56126328Ensembl.1
Natural variantiVAR_066605727A → V in CMS9; affects interaction with DOK7 and impairs MUSK phosphorylation; altered AChR clustering. 1 PublicationCorresponds to variant dbSNP:rs397515450EnsemblClinVar.1
Natural variantiVAR_041757782E → D1 PublicationCorresponds to variant dbSNP:rs34614566Ensembl.1
Natural variantiVAR_023046790V → M in CMS9; does not affect catalytic kinase activity; reduces protein expression and stability. 1 PublicationCorresponds to variant dbSNP:rs199476083EnsemblClinVar.1
Natural variantiVAR_041758819N → S in a lung neuroendocrine carcinoma sample; somatic mutation. 1 PublicationCorresponds to variant dbSNP:rs757577755Ensembl.1
Natural variantiVAR_033837829V → L1 PublicationCorresponds to variant dbSNP:rs578430EnsemblClinVar.1
Natural variantiVAR_072788835M → V in CMS9; reduces AChR aggregation in developing neuromuscular junction. 1 Publication1
Natural variantiVAR_041759858R → H1 PublicationCorresponds to variant dbSNP:rs34115159EnsemblClinVar.1

Alternative sequence

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Alternative sequenceiVSP_035958209E → EEESEPEQDTK in isoform 2. 1 Publication1
Alternative sequenceiVSP_035959307 – 394Missing in isoform 2 and isoform 3. 1 PublicationAdd BLAST88
Alternative sequenceiVSP_035960454 – 462DYNKENLKT → A in isoform 2 and isoform 3. 1 Publication9

Sequence databases

Select the link destinations:
EMBLi
GenBanki
DDBJi
Links Updated
AF006464 mRNA Translation: AAB63044.1
AL157881, AL513328 Genomic DNA Translation: CAH69977.1 Sequence problems.
AL513328, AL157881 Genomic DNA Translation: CAI17349.1 Sequence problems.
AL157881, AL513328 Genomic DNA Translation: CAH69978.1 Sequence problems.
AL513328, AL157881 Genomic DNA Translation: CAI17350.1 Sequence problems.
BC109098 mRNA Translation: AAI09099.1
BC109099 mRNA Translation: AAI09100.1
CCDSiCCDS48005.1 [O15146-1]
CCDS75874.1 [O15146-2]
RefSeqiNP_001159752.1, NM_001166280.1 [O15146-2]
NP_001159753.1, NM_001166281.1 [O15146-3]
NP_005583.1, NM_005592.3 [O15146-1]
UniGeneiHs.521653

Genome annotation databases

EnsembliENST00000189978; ENSP00000189978; ENSG00000030304 [O15146-2]
ENST00000374448; ENSP00000363571; ENSG00000030304 [O15146-1]
GeneIDi4593
KEGGihsa:4593
UCSCiuc064vai.1 human [O15146-1]

Keywords - Coding sequence diversityi

Alternative splicing, Polymorphism

Similar proteinsi

Entry informationi

Entry nameiMUSK_HUMAN
AccessioniPrimary (citable) accession number: O15146
Secondary accession number(s): Q32MJ8
, Q32MJ9, Q5VZW7, Q5VZW8
Entry historyiIntegrated into UniProtKB/Swiss-Prot: July 19, 2004
Last sequence update: January 1, 1998
Last modified: June 20, 2018
This is version 162 of the entry and version 1 of the sequence. See complete history.
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.

Miscellaneousi

Keywords - Technical termi

Complete proteome, Reference proteome

Documents

  1. Human chromosome 9
    Human chromosome 9: entries, gene names and cross-references to MIM
  2. Human entries with polymorphisms or disease mutations
    List of human entries with polymorphisms or disease mutations
  3. Human polymorphisms and disease mutations
    Index of human polymorphisms and disease mutations
  4. MIM cross-references
    Online Mendelian Inheritance in Man (MIM) cross-references in UniProtKB/Swiss-Prot
  5. Human and mouse protein kinases
    Human and mouse protein kinases: classification and index
  6. SIMILARITY comments
    Index of protein domains and families

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