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Protein

Histone deacetylase 5

Gene

HDAC5

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

Functioni

Responsible for the deacetylation of lysine residues on the N-terminal part of the core histones (H2A, H2B, H3 and H4). Histone deacetylation gives a tag for epigenetic repression and plays an important role in transcriptional regulation, cell cycle progression and developmental events. Histone deacetylases act via the formation of large multiprotein complexes. Involved in muscle maturation by repressing transcription of myocyte enhancer MEF2C. During muscle differentiation, it shuttles into the cytoplasm, allowing the expression of myocyte enhancer factors. Involved in the MTA1-mediated epigenetic regulation of ESR1 expression in breast cancer.1 Publication

Catalytic activityi

Hydrolysis of an N6-acetyl-lysine residue of a histone to yield a deacetylated histone.

Sites

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Metal bindingi696ZincBy similarity1
Metal bindingi698ZincBy similarity1
Metal bindingi704ZincBy similarity1
Metal bindingi781ZincBy similarity1
Active sitei833By similarity1

GO - Molecular functioni

GO - Biological processi

Keywordsi

Molecular functionChromatin regulator, Hydrolase, Repressor
Biological processTranscription, Transcription regulation
LigandMetal-binding, Zinc

Enzyme and pathway databases

ReactomeiR-HSA-2122947 NOTCH1 Intracellular Domain Regulates Transcription
R-HSA-2644606 Constitutive Signaling by NOTCH1 PEST Domain Mutants
R-HSA-2894862 Constitutive Signaling by NOTCH1 HD+PEST Domain Mutants
R-HSA-8943724 Regulation of PTEN gene transcription
SIGNORiQ9UQL6

Names & Taxonomyi

Protein namesi
Recommended name:
Histone deacetylase 5 (EC:3.5.1.98)
Short name:
HD5
Alternative name(s):
Antigen NY-CO-9
Gene namesi
Name:HDAC5
Synonyms:KIAA0600
OrganismiHomo sapiens (Human)
Taxonomic identifieri9606 [NCBI]
Taxonomic lineageiEukaryotaMetazoaChordataCraniataVertebrataEuteleostomiMammaliaEutheriaEuarchontogliresPrimatesHaplorrhiniCatarrhiniHominidaeHomo
Proteomesi
  • UP000005640 Componenti: Chromosome 17

Organism-specific databases

EuPathDBiHostDB:ENSG00000108840.15
HGNCiHGNC:14068 HDAC5
MIMi605315 gene
neXtProtiNX_Q9UQL6

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

Cytoplasm, Nucleus

Pathology & Biotechi

Mutagenesis

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Mutagenesisi259S → A: Reduces AMPK- and caMK-dependent phosphorylation and the subsequent nuclear export. Abolishes nuclear export; when associated with A-498. Does not affect phosphorylation by PKN1 and PKN2. 3 Publications1
Mutagenesisi279S → A: No effect. 1 Publication1
Mutagenesisi291S → A: Does not affect phosphorylation by PKC. 1 Publication1
Mutagenesisi292T → A: Abolishes phosphorylation by PKC. 1 Publication1
Mutagenesisi498S → A: Reduces AMPK- and CaMK-dependent phosphorylation and the subsequent nuclear export. Abolishes nuclear export; when associated with A-259. 2 Publications1
Mutagenesisi661S → A: No effect. 1 Publication1
Mutagenesisi713S → A: No effect. 1 Publication1
Mutagenesisi1086V → A: Reduces CaMK-dependent nuclear export. 1 Publication1
Mutagenesisi1092L → A: Reduces CaMK-dependent nuclear export. 1 Publication1

Organism-specific databases

DisGeNETi10014
OpenTargetsiENSG00000108840
PharmGKBiPA29230

Chemistry databases

ChEMBLiCHEMBL2563
DrugBankiDB05015 Belinostat
DB06603 Panobinostat
GuidetoPHARMACOLOGYi2660

Polymorphism and mutation databases

BioMutaiHDAC5
DMDMi296434519

PTM / Processingi

Molecule processing

Feature keyPosition(s)DescriptionActionsGraphical viewLength
ChainiPRO_00001147011 – 1122Histone deacetylase 5Add BLAST1122

Amino acid modifications

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Cross-linki35Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in SUMO2)Combined sources
Modified residuei259Phosphoserine; by AMPK, CaMK1, SIK1 and PKD/PRKD13 Publications1
Modified residuei292Phosphothreonine; by PKC1 Publication1
Modified residuei498Phosphoserine; by AMPK, CaMK1, SIK1 and PKD/PRKD13 Publications1
Modified residuei533N6-acetyllysineCombined sources1
Modified residuei611PhosphoserineCombined sources1
Modified residuei661PhosphoserineCombined sources1
Modified residuei1108PhosphoserineCombined sources1

Post-translational modificationi

Phosphorylated by AMPK, CaMK1, SIK1 and PRKD1 at Ser-259 and Ser-498. The phosphorylation is required for the export to the cytoplasm and inhibition. Phosphorylated by the PKC kinases PKN1 and PKN2, impairing nuclear import. Phosphorylated by GRK5, leading to nuclear export of HDAC5 and allowing MEF2-mediated transcription (By similarity).By similarity
Ubiquitinated. Polyubiquitination however does not lead to its degradation.1 Publication

Keywords - PTMi

Acetylation, Isopeptide bond, Phosphoprotein, Ubl conjugation

Proteomic databases

EPDiQ9UQL6
PaxDbiQ9UQL6
PeptideAtlasiQ9UQL6
PRIDEiQ9UQL6
ProteomicsDBi85556
85557 [Q9UQL6-2]
85558 [Q9UQL6-3]

PTM databases

iPTMnetiQ9UQL6
PhosphoSitePlusiQ9UQL6

Expressioni

Tissue specificityi

Ubiquitous.

Gene expression databases

BgeeiENSG00000108840 Expressed in 202 organ(s), highest expression level in thoracic aorta
CleanExiHS_HDAC5
ExpressionAtlasiQ9UQL6 baseline and differential
GenevisibleiQ9UQL6 HS

Organism-specific databases

HPAiCAB019400
HPA030991

Interactioni

Subunit structurei

Interacts with AHRR, BAHD1, BCOR, HDAC7, HDAC9, CTBP1, MEF2C, NCOR2, NRIP1, PHB2 and a 14-3-3 chaperone protein. Interacts with BCL6, DDIT3/CHOP, GRK5, KDM5B and MYOCD. Interacts with EP300 in the presence of TFAP2C. Interacts with ANKRA2. Interacts with CUL7 (as part of the 3M complex); negatively regulated by ANKRA2. Interacts with ZBTB7B; the interaction allows the recruitment of HDAC4 on CD8 loci for deacetylation and possible inhibition of CD8 genes expression (By similarity).By similarity9 Publications

Binary interactionsi

GO - Molecular functioni

Protein-protein interaction databases

BioGridi115331, 352 interactors
CORUMiQ9UQL6
DIPiDIP-38260N
ELMiQ9UQL6
IntActiQ9UQL6, 34 interactors
MINTiQ9UQL6
STRINGi9606.ENSP00000225983

Chemistry databases

BindingDBiQ9UQL6

Structurei

Secondary structure

11122
Legend: HelixTurnBeta strandPDB Structure known for this area
Show more details

3D structure databases

ProteinModelPortaliQ9UQL6
SMRiQ9UQL6
ModBaseiSearch...
MobiDBiSearch...

Family & Domainsi

Region

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Regioni684 – 1028Histone deacetylaseAdd BLAST345

Motif

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Motifi1081 – 1122Nuclear export signalAdd BLAST42

Compositional bias

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Compositional biasi47 – 52Poly-Gly6
Compositional biasi85 – 92Poly-Gln8
Compositional biasi596 – 599Poly-Glu4
Compositional biasi1099 – 1104Poly-Ala6

Domaini

The nuclear export sequence mediates the shuttling between the nucleus and the cytoplasm.By similarity

Sequence similaritiesi

Phylogenomic databases

eggNOGiKOG1343 Eukaryota
COG0123 LUCA
GeneTreeiENSGT00530000062809
HOGENOMiHOG000232065
HOVERGENiHBG057100
InParanoidiQ9UQL6
KOiK11406
OMAiEHHTLLY
OrthoDBiEOG091G0EQO
PhylomeDBiQ9UQL6
TreeFamiTF106174

Family and domain databases

Gene3Di3.40.800.20, 1 hit
InterProiView protein in InterPro
IPR000286 His_deacetylse
IPR023801 His_deacetylse_dom
IPR037138 His_deacetylse_dom_sf
IPR024643 Hist_deacetylase_Gln_rich_N
IPR017320 Histone_deAcase_II_euk
IPR030703 Histone_deacetylase_5
IPR023696 Ureohydrolase_dom_sf
PANTHERiPTHR10625 PTHR10625, 1 hit
PTHR10625:SF57 PTHR10625:SF57, 1 hit
PfamiView protein in Pfam
PF12203 HDAC4_Gln, 1 hit
PF00850 Hist_deacetyl, 1 hit
PIRSFiPIRSF037911 HDAC_II_euk, 1 hit
PRINTSiPR01270 HDASUPER
SUPFAMiSSF52768 SSF52768, 1 hit

Sequences (3+)i

Sequence statusi: Complete.

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: Q9UQL6-1) [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
MNSPNESDGM SGREPSLEIL PRTSLHSIPV TVEVKPVLPR AMPSSMGGGG
60 70 80 90 100
GGSPSPVELR GALVGSVDPT LREQQLQQEL LALKQQQQLQ KQLLFAEFQK
110 120 130 140 150
QHDHLTRQHE VQLQKHLKQQ QEMLAAKQQQ EMLAAKRQQE LEQQRQREQQ
160 170 180 190 200
RQEELEKQRL EQQLLILRNK EKSKESAIAS TEVKLRLQEF LLSKSKEPTP
210 220 230 240 250
GGLNHSLPQH PKCWGAHHAS LDQSSPPQSG PPGTPPSYKL PLPGPYDSRD
260 270 280 290 300
DFPLRKTASE PNLKVRSRLK QKVAERRSSP LLRRKDGTVI STFKKRAVEI
310 320 330 340 350
TGAGPGASSV CNSAPGSGPS SPNSSHSTIA ENGFTGSVPN IPTEMLPQHR
360 370 380 390 400
ALPLDSSPNQ FSLYTSPSLP NISLGLQATV TVTNSHLTAS PKLSTQQEAE
410 420 430 440 450
RQALQSLRQG GTLTGKFMST SSIPGCLLGV ALEGDGSPHG HASLLQHVLL
460 470 480 490 500
LEQARQQSTL IAVPLHGQSP LVTGERVATS MRTVGKLPRH RPLSRTQSSP
510 520 530 540 550
LPQSPQALQQ LVMQQQHQQF LEKQKQQQLQ LGKILTKTGE LPRQPTTHPE
560 570 580 590 600
ETEEELTEQQ EVLLGEGALT MPREGSTESE STQEDLEEED EEDDGEEEED
610 620 630 640 650
CIQVKDEEGE SGAEEGPDLE EPGAGYKKLF SDAQPLQPLQ VYQAPLSLAT
660 670 680 690 700
VPHQALGRTQ SSPAAPGGMK SPPDQPVKHL FTTGVVYDTF MLKHQCMCGN
710 720 730 740 750
THVHPEHAGR IQSIWSRLQE TGLLSKCERI RGRKATLDEI QTVHSEYHTL
760 770 780 790 800
LYGTSPLNRQ KLDSKKLLGP ISQKMYAVLP CGGIGVDSDT VWNEMHSSSA
810 820 830 840 850
VRMAVGCLLE LAFKVAAGEL KNGFAIIRPP GHHAEESTAM GFCFFNSVAI
860 870 880 890 900
TAKLLQQKLN VGKVLIVDWD IHHGNGTQQA FYNDPSVLYI SLHRYDNGNF
910 920 930 940 950
FPGSGAPEEV GGGPGVGYNV NVAWTGGVDP PIGDVEYLTA FRTVVMPIAH
960 970 980 990 1000
EFSPDVVLVS AGFDAVEGHL SPLGGYSVTA RCFGHLTRQL MTLAGGRVVL
1010 1020 1030 1040 1050
ALEGGHDLTA ICDASEACVS ALLSVELQPL DEAVLQQKPN INAVATLEKV
1060 1070 1080 1090 1100
IEIQSKHWSC VQKFAAGLGR SLREAQAGET EEAETVSAMA LLSVGAEQAQ
1110 1120
AAAAREHSPR PAEEPMEQEP AL
Length:1,122
Mass (Da):121,978
Last modified:May 18, 2010 - v2
Checksum:iCF4BE774E3588FEC
GO
Isoform 2 (identifier: Q9UQL6-2) [UniParc]FASTAAdd to basket

The sequence of this isoform differs from the canonical sequence as follows:
     684-768: Missing.

Note: No experimental confirmation available.
Show »
Length:1,037
Mass (Da):112,191
Checksum:iD4F09B907D2C1457
GO
Isoform 3 (identifier: Q9UQL6-3) [UniParc]FASTAAdd to basket

The sequence of this isoform differs from the canonical sequence as follows:
     7-7: S → SA

Show »
Length:1,123
Mass (Da):122,049
Checksum:i909ED5CCFEE3FB4F
GO

Computationally mapped potential isoform sequencesi

There are 4 potential isoforms mapped to this entry.BLASTAlignShow allAdd to basket
EntryEntry nameProtein names
Gene namesLengthAnnotation
K7EJL4K7EJL4_HUMAN
Histone deacetylase 5
HDAC5
282Annotation score:
K7EJL6K7EJL6_HUMAN
Histone deacetylase 5
HDAC5
79Annotation score:
K7ES01K7ES01_HUMAN
Histone deacetylase 5
HDAC5
68Annotation score:
K7EJZ7K7EJZ7_HUMAN
Histone deacetylase 5
HDAC5
40Annotation score:

Sequence cautioni

The sequence AAC18040 differs from that shown. Reason: Frameshift at position 1085.Curated
The sequence BAA25526 differs from that shown. Reason: Erroneous initiation. Translation N-terminally shortened.Curated

Experimental Info

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Sequence conflicti37V → L in BX458255 (Ref. 6) Curated1
Sequence conflicti139Q → R in BX458255 (Ref. 6) Curated1
Sequence conflicti147R → G in BX458255 (Ref. 6) Curated1
Sequence conflicti593D → E in AAD29047 (PubMed:10220385).Curated1
Sequence conflicti593D → E in AAH51824 (PubMed:15489334).Curated1
Sequence conflicti593D → E in AAC18040 (PubMed:9610721).Curated1
Sequence conflicti671S → N in AAC18040 (PubMed:9610721).Curated1
Sequence conflicti684G → S in AAC18040 (PubMed:9610721).Curated1
Sequence conflicti1026E → K in AAC18040 (PubMed:9610721).Curated1
Sequence conflicti1074E → G in AAC18040 (PubMed:9610721).Curated1
Sequence conflicti1093S → L in AAC18040 (PubMed:9610721).Curated1

Natural variant

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Natural variantiVAR_055903137R → Q. Corresponds to variant dbSNP:rs438096Ensembl.1
Natural variantiVAR_055904565G → A. Corresponds to variant dbSNP:rs33916560Ensembl.1

Alternative sequence

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Alternative sequenceiVSP_0391807S → SA in isoform 3. 1 Publication1
Alternative sequenceiVSP_002081684 – 768Missing in isoform 2. 1 PublicationAdd BLAST85

Sequence databases

Select the link destinations:
EMBLi
GenBanki
DDBJi
Links Updated
AF132608 mRNA Translation: AAD29047.1
AB011172 mRNA Translation: BAA25526.2 Different initiation.
AC023855 Genomic DNA No translation available.
BC013140 mRNA Translation: AAH13140.1 Different termination.
BC051824 mRNA Translation: AAH51824.1
BX458255 mRNA No translation available.
AF039691 mRNA Translation: AAC18040.1 Frameshift.
BK000028 Genomic DNA Translation: DAA00017.1
CCDSiCCDS32663.1 [Q9UQL6-3]
CCDS45696.1 [Q9UQL6-1]
RefSeqiNP_001015053.1, NM_001015053.1 [Q9UQL6-3]
NP_005465.2, NM_005474.4 [Q9UQL6-1]
XP_005256963.1, XM_005256906.4 [Q9UQL6-1]
UniGeneiHs.438782

Genome annotation databases

EnsembliENST00000225983; ENSP00000225983; ENSG00000108840 [Q9UQL6-3]
ENST00000336057; ENSP00000337290; ENSG00000108840 [Q9UQL6-2]
ENST00000586802; ENSP00000468004; ENSG00000108840 [Q9UQL6-1]
GeneIDi10014
KEGGihsa:10014
UCSCiuc002ifd.2 human [Q9UQL6-1]

Keywords - Coding sequence diversityi

Alternative splicing, Polymorphism

Similar proteinsi

Cross-referencesi

Sequence databases

Select the link destinations:
EMBLi
GenBanki
DDBJi
Links Updated
AF132608 mRNA Translation: AAD29047.1
AB011172 mRNA Translation: BAA25526.2 Different initiation.
AC023855 Genomic DNA No translation available.
BC013140 mRNA Translation: AAH13140.1 Different termination.
BC051824 mRNA Translation: AAH51824.1
BX458255 mRNA No translation available.
AF039691 mRNA Translation: AAC18040.1 Frameshift.
BK000028 Genomic DNA Translation: DAA00017.1
CCDSiCCDS32663.1 [Q9UQL6-3]
CCDS45696.1 [Q9UQL6-1]
RefSeqiNP_001015053.1, NM_001015053.1 [Q9UQL6-3]
NP_005465.2, NM_005474.4 [Q9UQL6-1]
XP_005256963.1, XM_005256906.4 [Q9UQL6-1]
UniGeneiHs.438782

3D structure databases

Select the link destinations:
PDBei
RCSB PDBi
PDBji
Links Updated
PDB entryMethodResolution (Å)ChainPositionsPDBsum
5UWIX-ray2.14D1078-1095[»]
ProteinModelPortaliQ9UQL6
SMRiQ9UQL6
ModBaseiSearch...
MobiDBiSearch...

Protein-protein interaction databases

BioGridi115331, 352 interactors
CORUMiQ9UQL6
DIPiDIP-38260N
ELMiQ9UQL6
IntActiQ9UQL6, 34 interactors
MINTiQ9UQL6
STRINGi9606.ENSP00000225983

Chemistry databases

BindingDBiQ9UQL6
ChEMBLiCHEMBL2563
DrugBankiDB05015 Belinostat
DB06603 Panobinostat
GuidetoPHARMACOLOGYi2660

PTM databases

iPTMnetiQ9UQL6
PhosphoSitePlusiQ9UQL6

Polymorphism and mutation databases

BioMutaiHDAC5
DMDMi296434519

Proteomic databases

EPDiQ9UQL6
PaxDbiQ9UQL6
PeptideAtlasiQ9UQL6
PRIDEiQ9UQL6
ProteomicsDBi85556
85557 [Q9UQL6-2]
85558 [Q9UQL6-3]

Protocols and materials databases

Structural Biology KnowledgebaseSearch...

Genome annotation databases

EnsembliENST00000225983; ENSP00000225983; ENSG00000108840 [Q9UQL6-3]
ENST00000336057; ENSP00000337290; ENSG00000108840 [Q9UQL6-2]
ENST00000586802; ENSP00000468004; ENSG00000108840 [Q9UQL6-1]
GeneIDi10014
KEGGihsa:10014
UCSCiuc002ifd.2 human [Q9UQL6-1]

Organism-specific databases

CTDi10014
DisGeNETi10014
EuPathDBiHostDB:ENSG00000108840.15
GeneCardsiHDAC5
H-InvDBiHIX0013862
HGNCiHGNC:14068 HDAC5
HPAiCAB019400
HPA030991
MIMi605315 gene
neXtProtiNX_Q9UQL6
OpenTargetsiENSG00000108840
PharmGKBiPA29230
HUGEiSearch...
GenAtlasiSearch...

Phylogenomic databases

eggNOGiKOG1343 Eukaryota
COG0123 LUCA
GeneTreeiENSGT00530000062809
HOGENOMiHOG000232065
HOVERGENiHBG057100
InParanoidiQ9UQL6
KOiK11406
OMAiEHHTLLY
OrthoDBiEOG091G0EQO
PhylomeDBiQ9UQL6
TreeFamiTF106174

Enzyme and pathway databases

ReactomeiR-HSA-2122947 NOTCH1 Intracellular Domain Regulates Transcription
R-HSA-2644606 Constitutive Signaling by NOTCH1 PEST Domain Mutants
R-HSA-2894862 Constitutive Signaling by NOTCH1 HD+PEST Domain Mutants
R-HSA-8943724 Regulation of PTEN gene transcription
SIGNORiQ9UQL6

Miscellaneous databases

ChiTaRSiHDAC5 human
GeneWikiiHistone_deacetylase_5
GenomeRNAii10014
PROiPR:Q9UQL6
SOURCEiSearch...

Gene expression databases

BgeeiENSG00000108840 Expressed in 202 organ(s), highest expression level in thoracic aorta
CleanExiHS_HDAC5
ExpressionAtlasiQ9UQL6 baseline and differential
GenevisibleiQ9UQL6 HS

Family and domain databases

Gene3Di3.40.800.20, 1 hit
InterProiView protein in InterPro
IPR000286 His_deacetylse
IPR023801 His_deacetylse_dom
IPR037138 His_deacetylse_dom_sf
IPR024643 Hist_deacetylase_Gln_rich_N
IPR017320 Histone_deAcase_II_euk
IPR030703 Histone_deacetylase_5
IPR023696 Ureohydrolase_dom_sf
PANTHERiPTHR10625 PTHR10625, 1 hit
PTHR10625:SF57 PTHR10625:SF57, 1 hit
PfamiView protein in Pfam
PF12203 HDAC4_Gln, 1 hit
PF00850 Hist_deacetyl, 1 hit
PIRSFiPIRSF037911 HDAC_II_euk, 1 hit
PRINTSiPR01270 HDASUPER
SUPFAMiSSF52768 SSF52768, 1 hit
ProtoNetiSearch...

Entry informationi

Entry nameiHDAC5_HUMAN
AccessioniPrimary (citable) accession number: Q9UQL6
Secondary accession number(s): C9JFV9
, O60340, O60528, Q96DY4
Entry historyiIntegrated into UniProtKB/Swiss-Prot: December 1, 2000
Last sequence update: May 18, 2010
Last modified: November 7, 2018
This is version 195 of the entry and version 2 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, Reference proteome

Documents

  1. SIMILARITY comments
    Index of protein domains and families
  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 chromosome 17
    Human chromosome 17: entries, gene names and cross-references to MIM
  6. PDB cross-references
    Index of Protein Data Bank (PDB) cross-references
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