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Protein

Methylcrotonoyl-CoA carboxylase subunit alpha, mitochondrial

Gene

MCCC1

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

Functioni

Biotin-attachment subunit of the 3-methylcrotonyl-CoA carboxylase, an enzyme that catalyzes the conversion of 3-methylcrotonyl-CoA to 3-methylglutaconyl-CoA, a critical step for leucine and isovaleric acid catabolism.1 Publication

Catalytic activityi

ATP + 3-methylcrotonoyl-CoA + HCO3- = ADP + phosphate + 3-methylglutaconyl-CoA.1 Publication

Cofactori

Pathwayi: L-leucine degradation

This protein is involved in step 2 of the subpathway that synthesizes (S)-3-hydroxy-3-methylglutaryl-CoA from 3-isovaleryl-CoA.
Proteins known to be involved in the 3 steps of the subpathway in this organism are:
  1. Isovaleryl-CoA dehydrogenase, mitochondrial (IVD)
  2. Methylcrotonoyl-CoA carboxylase beta chain, mitochondrial (MCCC2), Methylcrotonoyl-CoA carboxylase subunit alpha, mitochondrial (MCCC1)
  3. Methylglutaconyl-CoA hydratase, mitochondrial (AUH)
This subpathway is part of the pathway L-leucine degradation, which is itself part of Amino-acid degradation.
View all proteins of this organism that are known to be involved in the subpathway that synthesizes (S)-3-hydroxy-3-methylglutaryl-CoA from 3-isovaleryl-CoA, the pathway L-leucine degradation and in Amino-acid degradation.

Sites

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Binding sitei163ATPBy similarity1
Binding sitei247ATPBy similarity1
Binding sitei282ATPBy similarity1
Active sitei339By similarity1

GO - Molecular functioni

GO - Biological processi

  • biotin metabolic process Source: Reactome
  • leucine catabolic process Source: ParkinsonsUK-UCL
  • protein heterooligomerization Source: ParkinsonsUK-UCL

Keywordsi

Molecular functionLigase
LigandATP-binding, Biotin, Nucleotide-binding

Enzyme and pathway databases

BioCyciMetaCyc:ENSG00000078070-MONOMER
ReactomeiR-HSA-196780 Biotin transport and metabolism
R-HSA-3371599 Defective HLCS causes multiple carboxylase deficiency
R-HSA-70895 Branched-chain amino acid catabolism
UniPathwayi
UPA00363;UER00861

Names & Taxonomyi

Protein namesi
Recommended name:
Methylcrotonoyl-CoA carboxylase subunit alpha, mitochondrial (EC:6.4.1.4)
Short name:
MCCase subunit alpha
Alternative name(s):
3-methylcrotonyl-CoA carboxylase 1
3-methylcrotonyl-CoA carboxylase biotin-containing subunit
3-methylcrotonyl-CoA:carbon dioxide ligase subunit alpha
Gene namesi
Name:MCCC1
Synonyms:MCCA
OrganismiHomo sapiens (Human)
Taxonomic identifieri9606 [NCBI]
Taxonomic lineageiEukaryotaMetazoaChordataCraniataVertebrataEuteleostomiMammaliaEutheriaEuarchontogliresPrimatesHaplorrhiniCatarrhiniHominidaeHomo
Proteomesi
  • UP000005640 Componenti: Chromosome 3

Organism-specific databases

EuPathDBiHostDB:ENSG00000078070.11
HGNCiHGNC:6936 MCCC1
MIMi609010 gene
neXtProtiNX_Q96RQ3

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

Mitochondrion

Pathology & Biotechi

Involvement in diseasei

3-methylcrotonoyl-CoA carboxylase 1 deficiency (MCC1D)12 Publications
The disease is caused by mutations affecting the gene represented in this entry.
Disease descriptionAn autosomal recessive disorder of leucine catabolism. The phenotype is variable, ranging from neonatal onset with severe neurological involvement to asymptomatic adults. There is a characteristic organic aciduria with massive excretion of 3-hydroxyisovaleric acid and 3-methylcrotonylglycine, usually in combination with a severe secondary carnitine deficiency.
See also OMIM:210200
Feature keyPosition(s)DescriptionActionsGraphical viewLength
Natural variantiVAR_07248646G → E in MCC1D. 2 PublicationsCorresponds to variant dbSNP:rs199517715EnsemblClinVar.1
Natural variantiVAR_07248756N → K in MCC1D. 2 PublicationsCorresponds to variant dbSNP:rs1057520695Ensembl.1
Natural variantiVAR_07248865M → L in MCC1D. 1 Publication1
Natural variantiVAR_07728479Y → C in MCC1D. 1 Publication1
Natural variantiVAR_077285120S → F in MCC1D. 1 Publication1
Natural variantiVAR_072489123Q → H in MCC1D. 1 Publication1
Natural variantiVAR_072490125I → M in MCC1D. 1 Publication1
Natural variantiVAR_077286130G → S in MCC1D; clinically asymptomatic form. 1 PublicationCorresponds to variant dbSNP:rs202197951EnsemblClinVar.1
Natural variantiVAR_072491134E → K in MCC1D. 2 Publications1
Natural variantiVAR_072492160M → R in MCC1D. 1 Publication1
Natural variantiVAR_072493180G → V in MCC1D. 1 Publication1
Natural variantiVAR_072494187S → P in MCC1D. 2 PublicationsCorresponds to variant dbSNP:rs757362635Ensembl.1
Natural variantiVAR_077287209G → V in MCC1D. 1 PublicationCorresponds to variant dbSNP:rs186209189Ensembl.1
Natural variantiVAR_072495232R → W in MCC1D. 2 PublicationsCorresponds to variant dbSNP:rs727504004EnsemblClinVar.1
Natural variantiVAR_072496268A → D in MCC1D. 1 Publication1
Natural variantiVAR_067197276C → R in MCC1D. 1 PublicationCorresponds to variant dbSNP:rs773433541Ensembl.1
Natural variantiVAR_067198281R → Q in MCC1D. 3 PublicationsCorresponds to variant dbSNP:rs754437245Ensembl.1
Natural variantiVAR_072497288E → G in MCC1D; shows no residual activity. 2 PublicationsCorresponds to variant dbSNP:rs746500530EnsemblClinVar.1
Natural variantiVAR_012785289A → V in MCC1D; mild form. 2 Publications1
Natural variantiVAR_072498291A → V in MCC1D; associated with a reduction of wild-type residual activity. 2 PublicationsCorresponds to variant dbSNP:rs201041864EnsemblClinVar.1
Natural variantiVAR_012786325M → R in MCC1D. 2 PublicationsCorresponds to variant dbSNP:rs119103212EnsemblClinVar.1
Natural variantiVAR_077288366E → K in MCC1D; unknown pathological significance. 1 PublicationCorresponds to variant dbSNP:rs201386261Ensembl.1
Natural variantiVAR_072499372Q → P in MCC1D. 1 PublicationCorresponds to variant dbSNP:rs755328329Ensembl.1
Natural variantiVAR_072500379G → D in MCC1D. 1 Publication1
Natural variantiVAR_072501379G → S in MCC1D. 1 PublicationCorresponds to variant dbSNP:rs887877405Ensembl.1
Natural variantiVAR_072502380H → P in MCC1D. 2 PublicationsCorresponds to variant dbSNP:rs794727036EnsemblClinVar.1
Natural variantiVAR_077289383E → K in MCC1D; unknown pathological significance. 1 Publication1
Natural variantiVAR_012787385R → S in MCC1D; severe form. 6 PublicationsCorresponds to variant dbSNP:rs119103213EnsemblClinVar.1
Natural variantiVAR_072503434I → M in MCC1D; shows some wild-type residual activity. 1 PublicationCorresponds to variant dbSNP:rs376289130Ensembl.1
Natural variantiVAR_012788437L → P in MCC1D; severe form. 1 PublicationCorresponds to variant dbSNP:rs119103215EnsemblClinVar.1
Natural variantiVAR_072504439V → M in MCC1D. 1 PublicationCorresponds to variant dbSNP:rs398124352EnsemblClinVar.1
Natural variantiVAR_077290444R → H in MCC1D. 1 PublicationCorresponds to variant dbSNP:rs768785753Ensembl.1
Natural variantiVAR_072505460I → M in MCC1D. 2 PublicationsCorresponds to variant dbSNP:rs119103218EnsemblClinVar.1
Natural variantiVAR_012790532D → H in MCC1D; severe form. 2 PublicationsCorresponds to variant dbSNP:rs119103214EnsemblClinVar.1
Natural variantiVAR_012791535S → F in MCC1D; asymptomatic form. 2 PublicationsCorresponds to variant dbSNP:rs119103216EnsemblClinVar.1
Natural variantiVAR_072506566 – 567Missing in MCC1D. 1 Publication2

Keywords - Diseasei

Disease mutation

Organism-specific databases

DisGeNETi56922
MalaCardsiMCCC1
MIMi210200 phenotype
OpenTargetsiENSG00000078070
Orphaneti6 Isolated 3-methylcrotonyl-CoA carboxylase deficiency
PharmGKBiPA30680

Chemistry databases

DrugBankiDB00121 Biotin

Polymorphism and mutation databases

BioMutaiMCCC1
DMDMi108861983

PTM / Processingi

Molecule processing

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Transit peptidei1 – 41Mitochondrion1 PublicationAdd BLAST41
ChainiPRO_000000283342 – 725Methylcrotonoyl-CoA carboxylase subunit alpha, mitochondrialAdd BLAST684

Amino acid modifications

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Modified residuei237N6-acetyllysineBy similarity1
Modified residuei494N6-acetyllysineBy similarity1
Modified residuei581N6-acetyllysine; alternateBy similarity1
Modified residuei581N6-succinyllysine; alternateBy similarity1
Modified residuei681N6-biotinyllysinePROSITE-ProRule annotationBy similarity1

Post-translational modificationi

Acetylated.By similarity

Keywords - PTMi

Acetylation

Proteomic databases

EPDiQ96RQ3
MaxQBiQ96RQ3
PaxDbiQ96RQ3
PeptideAtlasiQ96RQ3
PRIDEiQ96RQ3
ProteomicsDBi78003

PTM databases

iPTMnetiQ96RQ3
PhosphoSitePlusiQ96RQ3
SwissPalmiQ96RQ3

Expressioni

Gene expression databases

BgeeiENSG00000078070 Expressed in 218 organ(s), highest expression level in body of pancreas
CleanExiHS_MCCC1
ExpressionAtlasiQ96RQ3 baseline and differential
GenevisibleiQ96RQ3 HS

Organism-specific databases

HPAiHPA008310

Interactioni

Subunit structurei

Probably a dodecamer composed of six biotin-containing alpha subunits (MCCC1) and six beta (MCCC2) subunits (PubMed:17360195). Interacts (via the biotin carboxylation domain) with SIRT4 (PubMed:23438705).2 Publications

Protein-protein interaction databases

BioGridi121249, 36 interactors
IntActiQ96RQ3, 8 interactors
MINTiQ96RQ3
STRINGi9606.ENSP00000265594

Structurei

Secondary structure

1725
Legend: HelixTurnBeta strandPDB Structure known for this area
Show more details

3D structure databases

ProteinModelPortaliQ96RQ3
SMRiQ96RQ3
ModBaseiSearch...
MobiDBiSearch...

Miscellaneous databases

EvolutionaryTraceiQ96RQ3

Family & Domainsi

Domains and Repeats

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Domaini48 – 494Biotin carboxylationAdd BLAST447
Domaini167 – 364ATP-graspPROSITE-ProRule annotationAdd BLAST198
Domaini643 – 715Biotinyl-bindingPROSITE-ProRule annotationAdd BLAST73

Keywords - Domaini

Transit peptide

Phylogenomic databases

eggNOGiKOG0238 Eukaryota
COG4770 LUCA
GeneTreeiENSGT00550000074675
HOGENOMiHOG000008989
HOVERGENiHBG000555
InParanoidiQ96RQ3
KOiK01968
OMAiFVEICSH
OrthoDBiEOG091G06RG
PhylomeDBiQ96RQ3
TreeFamiTF105650

Family and domain databases

Gene3Di3.30.1490.20, 1 hit
InterProiView protein in InterPro
IPR011761 ATP-grasp
IPR013815 ATP_grasp_subdomain_1
IPR005481 BC-like_N
IPR001882 Biotin_BS
IPR011764 Biotin_carboxylation_dom
IPR005482 Biotin_COase_C
IPR000089 Biotin_lipoyl
IPR005479 CbamoylP_synth_lsu-like_ATP-bd
IPR016185 PreATP-grasp_dom_sf
IPR011054 Rudment_hybrid_motif
IPR011053 Single_hybrid_motif
PfamiView protein in Pfam
PF02785 Biotin_carb_C, 1 hit
PF00289 Biotin_carb_N, 1 hit
PF00364 Biotin_lipoyl, 1 hit
PF02786 CPSase_L_D2, 1 hit
SMARTiView protein in SMART
SM00878 Biotin_carb_C, 1 hit
SUPFAMiSSF51230 SSF51230, 1 hit
SSF51246 SSF51246, 1 hit
SSF52440 SSF52440, 1 hit
PROSITEiView protein in PROSITE
PS50975 ATP_GRASP, 1 hit
PS50979 BC, 1 hit
PS00188 BIOTIN, 1 hit
PS50968 BIOTINYL_LIPOYL, 1 hit
PS00867 CPSASE_2, 1 hit

Sequence (1+)i

Sequence statusi: Complete.

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

This entry has 1 described isoform and 8 potential isoforms that are computationally mapped.iShow all

Q96RQ3-1 [UniParc]FASTAAdd to basket
« Hide
        10         20         30         40         50
MAAASAVSVL LVAAERNRWH RLPSLLLPPR TWVWRQRTMK YTTATGRNIT
60 70 80 90 100
KVLIANRGEI ACRVMRTAKK LGVQTVAVYS EADRNSMHVD MADEAYSIGP
110 120 130 140 150
APSQQSYLSM EKIIQVAKTS AAQAIHPGCG FLSENMEFAE LCKQEGIIFI
160 170 180 190 200
GPPPSAIRDM GIKSTSKSIM AAAGVPVVEG YHGEDQSDQC LKEHARRIGY
210 220 230 240 250
PVMIKAVRGG GGKGMRIVRS EQEFQEQLES ARREAKKSFN DDAMLIEKFV
260 270 280 290 300
DTPRHVEVQV FGDHHGNAVY LFERDCSVQR RHQKIIEEAP APGIKSEVRK
310 320 330 340 350
KLGEAAVRAA KAVNYVGAGT VEFIMDSKHN FCFMEMNTRL QVEHPVTEMI
360 370 380 390 400
TGTDLVEWQL RIAAGEKIPL SQEEITLQGH AFEARIYAED PSNNFMPVAG
410 420 430 440 450
PLVHLSTPRA DPSTRIETGV RQGDEVSVHY DPMIAKLVVW AADRQAALTK
460 470 480 490 500
LRYSLRQYNI VGLHTNIDFL LNLSGHPEFE AGNVHTDFIP QHHKQLLLSR
510 520 530 540 550
KAAAKESLCQ AALGLILKEK AMTDTFTLQA HDQFSPFSSS SGRRLNISYT
560 570 580 590 600
RNMTLKDGKN NVAIAVTYNH DGSYSMQIED KTFQVLGNLY SEGDCTYLKC
610 620 630 640 650
SVNGVASKAK LIILENTIYL FSKEGSIEID IPVPKYLSSV SSQETQGGPL
660 670 680 690 700
APMTGTIEKV FVKAGDKVKA GDSLMVMIAM KMEHTIKSPK DGTVKKVFYR
710 720
EGAQANRHTP LVEFEEEESD KRESE
Length:725
Mass (Da):80,473
Last modified:May 30, 2006 - v3
Checksum:iB84AD23806035A40
GO

Computationally mapped potential isoform sequencesi

There are 8 potential isoforms mapped to this entry.BLASTAlignShow allAdd to basket
EntryEntry nameProtein names
Gene namesLengthAnnotation
E9PHF7E9PHF7_HUMAN
Methylcrotonoyl-CoA carboxylase sub...
MCCC1
616Annotation score:
F5GYT8F5GYT8_HUMAN
Methylcrotonoyl-CoA carboxylase sub...
MCCC1
575Annotation score:
E9PG35E9PG35_HUMAN
Methylcrotonoyl-CoA carboxylase sub...
MCCC1
598Annotation score:
G5E9X5G5E9X5_HUMAN
Methylcrotonoyl-CoA carboxylase sub...
MCCC1 hCG_1811721
434Annotation score:
F8WDI3F8WDI3_HUMAN
Methylcrotonoyl-CoA carboxylase sub...
MCCC1
112Annotation score:
F8WF46F8WF46_HUMAN
Methylcrotonoyl-CoA carboxylase sub...
MCCC1
56Annotation score:
F2Z2Z7F2Z2Z7_HUMAN
Methylcrotonoyl-CoA carboxylase sub...
MCCC1
61Annotation score:
F2Z3E2F2Z3E2_HUMAN
Methylcrotonoyl-CoA carboxylase sub...
MCCC1
47Annotation score:

Sequence cautioni

The sequence BAD92974 differs from that shown. Reason: Erroneous initiation. Translation N-terminally shortened.Curated

Experimental Info

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Sequence conflicti469F → L in AAK67986 (PubMed:11406611).Curated1

Natural variant

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Natural variantiVAR_07248646G → E in MCC1D. 2 PublicationsCorresponds to variant dbSNP:rs199517715EnsemblClinVar.1
Natural variantiVAR_07248756N → K in MCC1D. 2 PublicationsCorresponds to variant dbSNP:rs1057520695Ensembl.1
Natural variantiVAR_07248865M → L in MCC1D. 1 Publication1
Natural variantiVAR_07728479Y → C in MCC1D. 1 Publication1
Natural variantiVAR_077285120S → F in MCC1D. 1 Publication1
Natural variantiVAR_072489123Q → H in MCC1D. 1 Publication1
Natural variantiVAR_072490125I → M in MCC1D. 1 Publication1
Natural variantiVAR_077286130G → S in MCC1D; clinically asymptomatic form. 1 PublicationCorresponds to variant dbSNP:rs202197951EnsemblClinVar.1
Natural variantiVAR_072491134E → K in MCC1D. 2 Publications1
Natural variantiVAR_072492160M → R in MCC1D. 1 Publication1
Natural variantiVAR_072493180G → V in MCC1D. 1 Publication1
Natural variantiVAR_072494187S → P in MCC1D. 2 PublicationsCorresponds to variant dbSNP:rs757362635Ensembl.1
Natural variantiVAR_077287209G → V in MCC1D. 1 PublicationCorresponds to variant dbSNP:rs186209189Ensembl.1
Natural variantiVAR_072495232R → W in MCC1D. 2 PublicationsCorresponds to variant dbSNP:rs727504004EnsemblClinVar.1
Natural variantiVAR_072496268A → D in MCC1D. 1 Publication1
Natural variantiVAR_067197276C → R in MCC1D. 1 PublicationCorresponds to variant dbSNP:rs773433541Ensembl.1
Natural variantiVAR_067198281R → Q in MCC1D. 3 PublicationsCorresponds to variant dbSNP:rs754437245Ensembl.1
Natural variantiVAR_072497288E → G in MCC1D; shows no residual activity. 2 PublicationsCorresponds to variant dbSNP:rs746500530EnsemblClinVar.1
Natural variantiVAR_012785289A → V in MCC1D; mild form. 2 Publications1
Natural variantiVAR_072498291A → V in MCC1D; associated with a reduction of wild-type residual activity. 2 PublicationsCorresponds to variant dbSNP:rs201041864EnsemblClinVar.1
Natural variantiVAR_012786325M → R in MCC1D. 2 PublicationsCorresponds to variant dbSNP:rs119103212EnsemblClinVar.1
Natural variantiVAR_077288366E → K in MCC1D; unknown pathological significance. 1 PublicationCorresponds to variant dbSNP:rs201386261Ensembl.1
Natural variantiVAR_072499372Q → P in MCC1D. 1 PublicationCorresponds to variant dbSNP:rs755328329Ensembl.1
Natural variantiVAR_072500379G → D in MCC1D. 1 Publication1
Natural variantiVAR_072501379G → S in MCC1D. 1 PublicationCorresponds to variant dbSNP:rs887877405Ensembl.1
Natural variantiVAR_072502380H → P in MCC1D. 2 PublicationsCorresponds to variant dbSNP:rs794727036EnsemblClinVar.1
Natural variantiVAR_077289383E → K in MCC1D; unknown pathological significance. 1 Publication1
Natural variantiVAR_012787385R → S in MCC1D; severe form. 6 PublicationsCorresponds to variant dbSNP:rs119103213EnsemblClinVar.1
Natural variantiVAR_072503434I → M in MCC1D; shows some wild-type residual activity. 1 PublicationCorresponds to variant dbSNP:rs376289130Ensembl.1
Natural variantiVAR_012788437L → P in MCC1D; severe form. 1 PublicationCorresponds to variant dbSNP:rs119103215EnsemblClinVar.1
Natural variantiVAR_072504439V → M in MCC1D. 1 PublicationCorresponds to variant dbSNP:rs398124352EnsemblClinVar.1
Natural variantiVAR_077290444R → H in MCC1D. 1 PublicationCorresponds to variant dbSNP:rs768785753Ensembl.1
Natural variantiVAR_072505460I → M in MCC1D. 2 PublicationsCorresponds to variant dbSNP:rs119103218EnsemblClinVar.1
Natural variantiVAR_012789464H → P5 PublicationsCorresponds to variant dbSNP:rs2270968EnsemblClinVar.1
Natural variantiVAR_012790532D → H in MCC1D; severe form. 2 PublicationsCorresponds to variant dbSNP:rs119103214EnsemblClinVar.1
Natural variantiVAR_012791535S → F in MCC1D; asymptomatic form. 2 PublicationsCorresponds to variant dbSNP:rs119103216EnsemblClinVar.1
Natural variantiVAR_038631560N → T. Corresponds to variant dbSNP:rs35219417Ensembl.1
Natural variantiVAR_072506566 – 567Missing in MCC1D. 1 Publication2
Natural variantiVAR_079752632P → S1 PublicationCorresponds to variant dbSNP:rs142867987EnsemblClinVar.1

Sequence databases

Select the link destinations:
EMBLi
GenBanki
DDBJi
Links Updated
AF310972 mRNA Translation: AAG53095.1
AB029826 mRNA Translation: BAA99407.1
AF297332 mRNA Translation: AAK67986.1
AF310339 mRNA Translation: AAG50245.1
AK023051 mRNA Translation: BAB14377.1
AB209737 mRNA Translation: BAD92974.1 Different initiation.
BC004214 mRNA Translation: AAH04214.1
BC004187 mRNA Translation: AAH04187.1
CCDSiCCDS3241.1
RefSeqiNP_001280202.1, NM_001293273.1
NP_064551.3, NM_020166.4
UniGeneiHs.47649

Genome annotation databases

EnsembliENST00000265594; ENSP00000265594; ENSG00000078070
GeneIDi56922
KEGGihsa:56922
UCSCiuc003fle.4 human

Keywords - Coding sequence diversityi

Polymorphism

Similar proteinsi

Entry informationi

Entry nameiMCCA_HUMAN
AccessioniPrimary (citable) accession number: Q96RQ3
Secondary accession number(s): Q59ES4, Q9H959, Q9NS97
Entry historyiIntegrated into UniProtKB/Swiss-Prot: March 5, 2002
Last sequence update: May 30, 2006
Last modified: September 12, 2018
This is version 191 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
UniProt is an ELIXIR core data resource
Main funding by: National Institutes of Health

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