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Protein

Golgi reassembly-stacking protein 2

Gene

GORASP2

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

Functioni

Plays a role in the assembly and membrane stacking of the Golgi cisternae, and in the process by which Golgi stacks reform after breakdown during mitosis and meiosis (PubMed:10487747, PubMed:21515684, PubMed:22523075). May regulate the intracellular transport and presentation of a defined set of transmembrane proteins, such as transmembrane TGFA (PubMed:11101516). Required for normal acrosome formation during spermiogenesis and normal male fertility, probably by promoting colocalization of JAM2 and JAM3 at contact sites between germ cells and Sertoli cells (By similarity). Mediates ER stress-induced unconventional (ER/Golgi-independent) trafficking of core-glycosylated CFTR to cell membrane (PubMed:21884936, PubMed:27062250, PubMed:28067262).By similarity7 Publications

GO - Biological processi

  • cell differentiation Source: UniProtKB-KW
  • Golgi organization Source: GO_Central
  • organelle assembly Source: UniProtKB
  • organelle organization Source: UniProtKB
  • spermatogenesis Source: UniProtKB-KW

Keywordsi

Biological processDifferentiation, Spermatogenesis

Enzyme and pathway databases

ReactomeiR-HSA-162658 Golgi Cisternae Pericentriolar Stack Reorganization
SIGNORiQ9H8Y8

Names & Taxonomyi

Protein namesi
Recommended name:
Golgi reassembly-stacking protein 2
Short name:
GRS2
Alternative name(s):
Golgi phosphoprotein 6
Short name:
GOLPH6
Golgi reassembly-stacking protein of 55 kDa
Short name:
GRASP55
p59
Gene namesi
Name:GORASP2
Synonyms:GOLPH6
OrganismiHomo sapiens (Human)
Taxonomic identifieri9606 [NCBI]
Taxonomic lineageiEukaryotaMetazoaChordataCraniataVertebrataEuteleostomiMammaliaEutheriaEuarchontogliresPrimatesHaplorrhiniCatarrhiniHominidaeHomo
Proteomesi
  • UP000005640 Componenti: Chromosome 2

Organism-specific databases

EuPathDBiHostDB:ENSG00000115806.12
HGNCiHGNC:17500 GORASP2
MIMi608693 gene
neXtProtiNX_Q9H8Y8

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

Keywords - Cellular componenti

Endoplasmic reticulum, Golgi apparatus, Membrane

Pathology & Biotechi

Mutagenesis

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Mutagenesisi2G → A: Abolishes myristoylation. Loss of its ability to mediate unconventional (ER/Golgi-independent) trafficking of CFTR to the cell membrane. No effect on ER stress-induced relocalization from Golgi to ER. 2 Publications1
Mutagenesisi59L → A: Abolishes organelle clustering; when associated with S-100. 1 Publication1
Mutagenesisi100I → S: Abolishes organelle clustering; when associated with A-59. 1 Publication1
Mutagenesisi148D → N: Enhances homodimerization. Loss of ER stress-induced relocalization from Golgi to ER. 1 Publication1
Mutagenesisi189S → D: Phosphomimetic mutation that decreases the ability to promote organelle clustering. 1 Publication1
Mutagenesisi222T → A: Abolishes mitotic phosphorylation; when associated with A-225. 1 Publication1
Mutagenesisi225T → A: Abolishes mitotic phosphorylation; when associated with A-222. 1 Publication1
Mutagenesisi441S → A: Loss of phosphorylation and its ability to mediate unconventional (ER/Golgi-independent) trafficking of CFTR to the cell membrane. Inhibits ER stress-mediated disruption of homodimerization. Loss of ER stress-induced relocalization from Golgi to ER. 2 Publications1
Mutagenesisi441S → D: Phosphomimetic mutant. No loss of its ability to mediate unconventional (ER/Golgi-independent) trafficking of CFTR to the cell membrane. Disrupts homodimerization. No loss of ER stress-induced relocalization from Golgi to ER. 2 Publications1
Mutagenesisi449S → A: Loss of phosphorylation. Does not inhibit ER stress-mediated disruption of homodimerization. 1 Publication1
Mutagenesisi449S → D: Phosphomimetic mutant. No disruption of homodimerization. 1 Publication1
Mutagenesisi451S → A: Loss of phosphorylation. Does not inhibit ER stress-mediated disruption of homodimerization. 1 Publication1
Mutagenesisi451S → D: Phosphomimetic mutant. Partial disruption of homodimerization. 1 Publication1

Organism-specific databases

DisGeNETi26003
OpenTargetsiENSG00000115806
PharmGKBiPA38457

Polymorphism and mutation databases

BioMutaiGORASP2
DMDMi51316097

PTM / Processingi

Molecule processing

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Initiator methionineiRemoved2 Publications
ChainiPRO_00000875452 – 452Golgi reassembly-stacking protein 2Add BLAST451

Amino acid modifications

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Lipidationi2N-myristoyl glycine1 Publication3 Publications1
Modified residuei30Dimethylated arginine; alternateBy similarity1
Modified residuei30Omega-N-methylated arginine; alternateBy similarity1
Modified residuei47Dimethylated arginine; alternateBy similarity1
Modified residuei47Omega-N-methylated arginine; alternateBy similarity1
Modified residuei214PhosphoserineCombined sources1
Modified residuei222PhosphothreonineCombined sources1
Modified residuei225PhosphothreonineCombined sources1 Publication1
Modified residuei409PhosphoserineBy similarity1
Modified residuei415PhosphothreonineCombined sources1
Modified residuei433PhosphothreonineCombined sources1
Modified residuei436PhosphoserineCombined sources1
Modified residuei441Phosphoserine2 Publications1
Modified residuei449PhosphoserineCombined sources1 Publication1
Modified residuei451PhosphoserineCombined sources1 Publication1

Post-translational modificationi

Myristoylated (PubMed:11101516). Myristoylation is essential for the Golgi targeting (By similarity).By similarity1 Publication
Palmitoylated.1 Publication
Phosphorylated in mitotic cells (PubMed:11408587). ER stress-induced phosphorylation at Ser-441 induces monomerization and subsequent relocalization from Golgi to ER which is essential for mediating unconventional (ER/Golgi-independent) trafficking of CFTR to the cell membrane (PubMed:21884936, PubMed:27062250).3 Publications

Keywords - PTMi

Lipoprotein, Methylation, Myristate, Palmitate, Phosphoprotein

Proteomic databases

EPDiQ9H8Y8
MaxQBiQ9H8Y8
PaxDbiQ9H8Y8
PeptideAtlasiQ9H8Y8
PRIDEiQ9H8Y8
ProteomicsDBi81257
81258 [Q9H8Y8-2]

PTM databases

iPTMnetiQ9H8Y8
PhosphoSitePlusiQ9H8Y8
SwissPalmiQ9H8Y8

Expressioni

Gene expression databases

BgeeiENSG00000115806 Expressed in 237 organ(s), highest expression level in sperm
CleanExiHS_GORASP2
ExpressionAtlasiQ9H8Y8 baseline and differential
GenevisibleiQ9H8Y8 HS

Organism-specific databases

HPAiCAB075758
HPA035274
HPA035275

Interactioni

Subunit structurei

Homodimer. Homooligomer. ER stress induces phosphorylation-dependent monomerization (PubMed:27062250). Interacts with BLZF1/Golgin 45 (PubMed:11739401). Identified in a complex with RAB2 and GORASP2 (PubMed:11739401). Interacts with JAM2 and JAM3 (By similarity). Interacts with members of the p24 cargo receptors (PubMed:11739402). Intebracts with CNIH1 and the cytoplasmic domain of transmembrane TGFA, prior its transit in the trans-Golgi (PubMed:11101516, PubMed:17607000). Interacts with KCTD5 (PubMed:19361449). Interacts with TMED2 and TMED3 (By similarity). Interacts with SEC16A in response to ER stress (PubMed:28067262). Interacts (via PDZ GRASP-type 1 domain) with core-glycosylated CFTR in response to ER stress (PubMed:21884936).By similarity8 Publications

Binary interactionsi

WithEntry#Exp.IntActNotes
Q99IB83EBI-739467,EBI-6927928From Hepatitis C virus genotype 2a (isolate JFH-1).
AAMDCQ9H7C93EBI-739467,EBI-10308705
ACY3Q96HD910EBI-739467,EBI-3916242
AESQ081173EBI-739467,EBI-717810
ARL6IP1Q150413EBI-739467,EBI-714543
BEND5Q7L4P63EBI-739467,EBI-724373
C1orf94Q6P1W55EBI-739467,EBI-946029
CBLBQ131916EBI-739467,EBI-744027
CCDC153Q494R43EBI-739467,EBI-10241443
CDC23Q9UJX27EBI-739467,EBI-396137
CFTRP135693EBI-739467,EBI-349854
CRYAAP024898EBI-739467,EBI-6875961
DCTDP323215EBI-739467,EBI-739870
DMC1Q145655EBI-739467,EBI-930865
DPYSL2Q165557EBI-739467,EBI-1104711
DUSP21Q9H5965EBI-739467,EBI-7357329
EIF2B1Q142325EBI-739467,EBI-491065
ENOX1Q8TC923EBI-739467,EBI-713221
GABPB2Q8TAK55EBI-739467,EBI-8468945
GOLGA2Q083796EBI-739467,EBI-618309
JADE2Q9NQC1-23EBI-739467,EBI-10311936
KCTD9Q7L2735EBI-739467,EBI-4397613
KIFC3Q9BVG83EBI-739467,EBI-2125614
LONRF1Q17RB87EBI-739467,EBI-2341787
MAP1LC3BQ9GZQ85EBI-739467,EBI-373144
MID2Q9UJV3-25EBI-739467,EBI-10172526
MORN2Q502X03EBI-739467,EBI-725982
NQO2P160833EBI-739467,EBI-358466
ODC1P119269EBI-739467,EBI-1044287
PCBD1P614573EBI-739467,EBI-740475
PDE9AO760833EBI-739467,EBI-742764
PNMA5Q96PV45EBI-739467,EBI-10171633
PRDX3P300483EBI-739467,EBI-748336
PRPS1P608913EBI-739467,EBI-749195
PSMA3P257883EBI-739467,EBI-348380
RGS3P49796-83EBI-739467,EBI-10211517
RPIAP492476EBI-739467,EBI-744831
SCAND1P570865EBI-739467,EBI-745846
STOX1Q6ZVD73EBI-739467,EBI-3923644
TCF4P158843EBI-739467,EBI-533224
TEKT3Q9BXF94EBI-739467,EBI-8644516
TRAF1Q130775EBI-739467,EBI-359224
TRAF2Q129336EBI-739467,EBI-355744
TRAF4Q9BUZ43EBI-739467,EBI-3650647
TRAF5O004633EBI-739467,EBI-523498
TRIM38O006355EBI-739467,EBI-2130415
TSC22D1Q15714-24EBI-739467,EBI-12034704
TSNQ156314EBI-739467,EBI-1044160
TXLNAP402225EBI-739467,EBI-359793
ZBTB14O438293EBI-739467,EBI-10176632
ZNF148Q9UQR13EBI-739467,EBI-2688184

Protein-protein interaction databases

BioGridi117479, 117 interactors
IntActiQ9H8Y8, 143 interactors
MINTiQ9H8Y8
STRINGi9606.ENSP00000234160

Structurei

Secondary structure

1452
Legend: HelixTurnBeta strandPDB Structure known for this area
Show more details

3D structure databases

ProteinModelPortaliQ9H8Y8
SMRiQ9H8Y8
ModBaseiSearch...
MobiDBiSearch...

Miscellaneous databases

EvolutionaryTraceiQ9H8Y8

Family & Domainsi

Domains and Repeats

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Domaini15 – 105PDZ GRASP-type 1PROSITE-ProRule annotationAdd BLAST91
Domaini111 – 199PDZ GRASP-type 2PROSITE-ProRule annotationAdd BLAST89

Region

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Regioni15 – 215GRASPPROSITE-ProRule annotationAdd BLAST201
Regioni194 – 199Important for membrane binding6

Compositional bias

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Compositional biasi277 – 372Pro-richAdd BLAST96

Sequence similaritiesi

Belongs to the GORASP family.Curated

Keywords - Domaini

Repeat

Phylogenomic databases

eggNOGiKOG3834 Eukaryota
COG5233 LUCA
GeneTreeiENSGT00390000008686
HOGENOMiHOG000231920
HOVERGENiHBG051826
InParanoidiQ9H8Y8
OMAiMPSEFLP
OrthoDBiEOG091G0E90
PhylomeDBiQ9H8Y8
TreeFamiTF314053

Family and domain databases

InterProiView protein in InterPro
IPR024958 GRASP55/65_PDZ
IPR007583 GRASP55_65
IPR036034 PDZ_sf
PANTHERiPTHR12893 PTHR12893, 1 hit
PfamiView protein in Pfam
PF04495 GRASP55_65, 1 hit
SUPFAMiSSF50156 SSF50156, 2 hits
PROSITEiView protein in PROSITE
PS51865 PDZ_GRASP, 2 hits

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

This entry has 3 described isoforms and 4 potential isoforms that are computationally mapped.Show allAlign All

Isoform 1 (identifier: Q9H8Y8-1) [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
MGSSQSVEIP GGGTEGYHVL RVQENSPGHR AGLEPFFDFI VSINGSRLNK
60 70 80 90 100
DNDTLKDLLK ANVEKPVKML IYSSKTLELR ETSVTPSNLW GGQGLLGVSI
110 120 130 140 150
RFCSFDGANE NVWHVLEVES NSPAALAGLR PHSDYIIGAD TVMNESEDLF
160 170 180 190 200
SLIETHEAKP LKLYVYNTDT DNCREVIITP NSAWGGEGSL GCGIGYGYLH
210 220 230 240 250
RIPTRPFEEG KKISLPGQMA GTPITPLKDG FTEVQLSSVN PPSLSPPGTT
260 270 280 290 300
GIEQSLTGLS ISSTPPAVSS VLSTGVPTVP LLPPQVNQSL TSVPPMNPAT
310 320 330 340 350
TLPGLMPLPA GLPNLPNLNL NLPAPHIMPG VGLPELVNPG LPPLPSMPPR
360 370 380 390 400
NLPGIAPLPL PSEFLPSFPL VPESSSAASS GELLSSLPPT SNAPSDPATT
410 420 430 440 450
TAKADAASSL TVDVTPPTAK APTTVEDRVG DSTPVSEKPV SAAVDANASE

SP
Length:452
Mass (Da):47,145
Last modified:January 23, 2007 - v3
Checksum:i326EA6C107D2EA8B
GO
Isoform 2 (identifier: Q9H8Y8-2) [UniParc]FASTAAdd to basket

The sequence of this isoform differs from the canonical sequence as follows:
     1-68: Missing.

Show »
Length:384
Mass (Da):39,768
Checksum:i707E11F2C55B7BDA
GO
Isoform 3 (identifier: Q9H8Y8-3) [UniParc]FASTAAdd to basket

The sequence of this isoform differs from the canonical sequence as follows:
     1-21: MGSSQSVEIPGGGTEGYHVLR → MREGSSTLSEIRKLKPGIMVCTCNPSYSNQETE

Note: No experimental confirmation available.
Show »
Length:464
Mass (Da):48,674
Checksum:iF6A0C68893A94434
GO

Computationally mapped potential isoform sequencesi

There are 4 potential isoforms mapped to this entry.BLASTAlignShow allAdd to basket
EntryEntry nameProtein names
Gene namesLengthAnnotation
F8WEG2F8WEG2_HUMAN
Golgi reassembly-stacking protein 2
GORASP2
32Annotation score:
F8WBR9F8WBR9_HUMAN
Golgi reassembly-stacking protein 2
GORASP2
73Annotation score:
F8WF79F8WF79_HUMAN
Golgi reassembly-stacking protein 2
GORASP2
23Annotation score:
F8WEH9F8WEH9_HUMAN
Golgi reassembly-stacking protein 2
GORASP2
38Annotation score:

Experimental Info

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Sequence conflicti20L → Q in BAB14459 (PubMed:14702039).Curated1
Sequence conflicti34E → G in BAB14395 (PubMed:14702039).Curated1
Sequence conflicti83S → P AA sequence (PubMed:11101516).Curated1
Sequence conflicti299A → T in BAB14395 (PubMed:14702039).Curated1

Natural variant

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Natural variantiVAR_051013432S → F1 PublicationCorresponds to variant dbSNP:rs3770436Ensembl.1

Alternative sequence

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Alternative sequenceiVSP_0113001 – 68Missing in isoform 2. 1 PublicationAdd BLAST68
Alternative sequenceiVSP_0543641 – 21MGSSQ…YHVLR → MREGSSTLSEIRKLKPGIMV CTCNPSYSNQETE in isoform 3. 1 PublicationAdd BLAST21

Sequence databases

Select the link destinations:
EMBLi
GenBanki
DDBJi
Links Updated
AK027349 mRNA Translation: BAB55054.1
AK023082 mRNA Translation: BAB14395.1
AK023201 mRNA Translation: BAB14459.1
AK296698 mRNA Translation: BAG59292.1
AL117430 mRNA Translation: CAB55919.1
AC010092 Genomic DNA Translation: AAY15076.1
CH471058 Genomic DNA Translation: EAX11227.1
BC007770 mRNA Translation: AAH07770.1
CCDSiCCDS33325.1 [Q9H8Y8-1]
PIRiT17229
RefSeqiNP_001188357.1, NM_001201428.1 [Q9H8Y8-2]
NP_056345.3, NM_015530.4 [Q9H8Y8-1]
XP_006712471.1, XM_006712408.2
UniGeneiHs.431317

Genome annotation databases

EnsembliENST00000234160; ENSP00000234160; ENSG00000115806 [Q9H8Y8-1]
GeneIDi26003
KEGGihsa:26003
UCSCiuc002ugk.4 human [Q9H8Y8-1]

Keywords - Coding sequence diversityi

Alternative splicing, Polymorphism

Similar proteinsi

Cross-referencesi

Sequence databases

Select the link destinations:
EMBLi
GenBanki
DDBJi
Links Updated
AK027349 mRNA Translation: BAB55054.1
AK023082 mRNA Translation: BAB14395.1
AK023201 mRNA Translation: BAB14459.1
AK296698 mRNA Translation: BAG59292.1
AL117430 mRNA Translation: CAB55919.1
AC010092 Genomic DNA Translation: AAY15076.1
CH471058 Genomic DNA Translation: EAX11227.1
BC007770 mRNA Translation: AAH07770.1
CCDSiCCDS33325.1 [Q9H8Y8-1]
PIRiT17229
RefSeqiNP_001188357.1, NM_001201428.1 [Q9H8Y8-2]
NP_056345.3, NM_015530.4 [Q9H8Y8-1]
XP_006712471.1, XM_006712408.2
UniGeneiHs.431317

3D structure databases

Select the link destinations:
PDBei
RCSB PDBi
PDBji
Links Updated
PDB entryMethodResolution (Å)ChainPositionsPDBsum
3RLEX-ray1.65A2-208[»]
4EDJX-ray1.90A/B1-208[»]
ProteinModelPortaliQ9H8Y8
SMRiQ9H8Y8
ModBaseiSearch...
MobiDBiSearch...

Protein-protein interaction databases

BioGridi117479, 117 interactors
IntActiQ9H8Y8, 143 interactors
MINTiQ9H8Y8
STRINGi9606.ENSP00000234160

PTM databases

iPTMnetiQ9H8Y8
PhosphoSitePlusiQ9H8Y8
SwissPalmiQ9H8Y8

Polymorphism and mutation databases

BioMutaiGORASP2
DMDMi51316097

Proteomic databases

EPDiQ9H8Y8
MaxQBiQ9H8Y8
PaxDbiQ9H8Y8
PeptideAtlasiQ9H8Y8
PRIDEiQ9H8Y8
ProteomicsDBi81257
81258 [Q9H8Y8-2]

Protocols and materials databases

DNASUi26003
Structural Biology KnowledgebaseSearch...

Genome annotation databases

EnsembliENST00000234160; ENSP00000234160; ENSG00000115806 [Q9H8Y8-1]
GeneIDi26003
KEGGihsa:26003
UCSCiuc002ugk.4 human [Q9H8Y8-1]

Organism-specific databases

CTDi26003
DisGeNETi26003
EuPathDBiHostDB:ENSG00000115806.12
GeneCardsiGORASP2
HGNCiHGNC:17500 GORASP2
HPAiCAB075758
HPA035274
HPA035275
MIMi608693 gene
neXtProtiNX_Q9H8Y8
OpenTargetsiENSG00000115806
PharmGKBiPA38457
GenAtlasiSearch...

Phylogenomic databases

eggNOGiKOG3834 Eukaryota
COG5233 LUCA
GeneTreeiENSGT00390000008686
HOGENOMiHOG000231920
HOVERGENiHBG051826
InParanoidiQ9H8Y8
OMAiMPSEFLP
OrthoDBiEOG091G0E90
PhylomeDBiQ9H8Y8
TreeFamiTF314053

Enzyme and pathway databases

ReactomeiR-HSA-162658 Golgi Cisternae Pericentriolar Stack Reorganization
SIGNORiQ9H8Y8

Miscellaneous databases

ChiTaRSiGORASP2 human
EvolutionaryTraceiQ9H8Y8
GeneWikiiGORASP2
GenomeRNAii26003
PROiPR:Q9H8Y8
SOURCEiSearch...

Gene expression databases

BgeeiENSG00000115806 Expressed in 237 organ(s), highest expression level in sperm
CleanExiHS_GORASP2
ExpressionAtlasiQ9H8Y8 baseline and differential
GenevisibleiQ9H8Y8 HS

Family and domain databases

InterProiView protein in InterPro
IPR024958 GRASP55/65_PDZ
IPR007583 GRASP55_65
IPR036034 PDZ_sf
PANTHERiPTHR12893 PTHR12893, 1 hit
PfamiView protein in Pfam
PF04495 GRASP55_65, 1 hit
SUPFAMiSSF50156 SSF50156, 2 hits
PROSITEiView protein in PROSITE
PS51865 PDZ_GRASP, 2 hits
ProtoNetiSearch...

Entry informationi

Entry nameiGORS2_HUMAN
AccessioniPrimary (citable) accession number: Q9H8Y8
Secondary accession number(s): B4DKT0
, Q53TE3, Q96I74, Q96K84, Q9H946, Q9UFW4
Entry historyiIntegrated into UniProtKB/Swiss-Prot: August 16, 2004
Last sequence update: January 23, 2007
Last modified: November 7, 2018
This is version 158 of the entry and version 3 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

3D-structure, Complete proteome, Direct protein sequencing, Reference proteome

Documents

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