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Protein

Nucleolin

Gene

NCL

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

Functioni

Nucleolin is the major nucleolar protein of growing eukaryotic cells. It is found associated with intranucleolar chromatin and pre-ribosomal particles. It induces chromatin decondensation by binding to histone H1. It is thought to play a role in pre-rRNA transcription and ribosome assembly. May play a role in the process of transcriptional elongation. Binds RNA oligonucleotides with 5'-UUAGGG-3' repeats more tightly than the telomeric single-stranded DNA 5'-TTAGGG-3' repeats.1 Publication

GO - Molecular functioni

  • DNA topoisomerase binding Source: CAFA
  • identical protein binding Source: IntAct
  • mRNA 5'-UTR binding Source: CAFA
  • protein C-terminus binding Source: UniProtKB
  • RNA binding Source: UniProtKB
  • telomeric DNA binding Source: UniProtKB

GO - Biological processi

Keywordsi

Molecular functionDNA-binding, RNA-binding

Enzyme and pathway databases

ReactomeiR-HSA-6791226 Major pathway of rRNA processing in the nucleolus and cytosol
SIGNORiP19338

Protein family/group databases

MoonProtiP19338

Names & Taxonomyi

Protein namesi
Recommended name:
Nucleolin
Alternative name(s):
Protein C23
Gene namesi
Name:NCL
OrganismiHomo sapiens (Human)
Taxonomic identifieri9606 [NCBI]
Taxonomic lineageiEukaryotaMetazoaChordataCraniataVertebrataEuteleostomiMammaliaEutheriaEuarchontogliresPrimatesHaplorrhiniCatarrhiniHominidaeHomo
Proteomesi
  • UP000005640 Componenti: Chromosome 2

Organism-specific databases

EuPathDBiHostDB:ENSG00000115053.15
HGNCiHGNC:7667 NCL
MIMi164035 gene
neXtProtiNX_P19338

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

Organism-specific databases

DisGeNETi4691
OpenTargetsiENSG00000115053
PharmGKBiPA31469

Polymorphism and mutation databases

BioMutaiNCL
DMDMi90110781

PTM / Processingi

Molecule processing

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Initiator methionineiRemoved1 Publication
ChainiPRO_00000816912 – 710NucleolinAdd BLAST709

Amino acid modifications

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Modified residuei9N6-acetyllysineCombined sources1
Modified residuei15N6-acetyllysineBy similarity1
Modified residuei16N6-acetyllysineBy similarity1
Modified residuei28PhosphoserineCombined sources1
Modified residuei34PhosphoserineCombined sources1
Modified residuei41PhosphoserineCombined sources1
Modified residuei42PhosphoserineCombined sources1
Modified residuei67PhosphoserineCombined sources1
Modified residuei69PhosphothreonineCombined sources1
Modified residuei76PhosphothreonineCombined sources1
Modified residuei84PhosphothreonineCombined sources1
Modified residuei92PhosphothreonineCombined sources1
Modified residuei96N6-acetyllysineBy similarity1
Modified residuei99PhosphothreonineCombined sources1
Modified residuei102N6-acetyllysineCombined sources1
Modified residuei106PhosphothreonineCombined sources1
Modified residuei109N6-acetyllysineBy similarity1
Modified residuei113PhosphothreonineCombined sources1
Modified residuei116N6-acetyllysineCombined sources1
Modified residuei121PhosphothreonineCombined sources1
Modified residuei124N6-acetyllysineCombined sources1
Modified residuei145PhosphoserineCombined sources1
Modified residuei153PhosphoserineCombined sources1
Modified residuei184PhosphoserineCombined sources1
Modified residuei206PhosphoserineCombined sources1
Modified residuei214PhosphothreonineCombined sources1
Cross-linki297Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in SUMO1); alternateCombined sources
Cross-linki297Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in SUMO2); alternateCombined sources
Modified residuei301PhosphothreonineCombined sources1
Modified residuei318N6-acetyllysineCombined sources1
Cross-linki324Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in SUMO1); alternateCombined sources
Cross-linki324Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in SUMO2); alternateCombined sources
Modified residuei348N6-acetyllysineBy similarity1
Modified residuei356PhosphoserineCombined sources1
Modified residuei367PhosphothreonineCombined sources1
Cross-linki370Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in SUMO2)Combined sources
Modified residuei377N6-acetyllysine; alternateCombined sources1
Cross-linki377Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in SUMO2); alternateCombined sources
Modified residuei398N6-acetyllysineCombined sources1
Modified residuei403N6-acetyllysineCombined sources1
Modified residuei405PhosphothreonineCombined sources1
Modified residuei427N6-acetyllysineBy similarity1
Modified residuei444N6-acetyllysineBy similarity1
Modified residuei458PhosphoserineCombined sources1
Modified residuei460PhosphoserineCombined sources1
Modified residuei467N6-acetyllysineBy similarity1
Modified residuei477N6-acetyllysineBy similarity1
Modified residuei513N6-acetyllysine; alternateCombined sources1
Cross-linki513Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in SUMO2); alternateCombined sources
Modified residuei521N6-acetyllysineBy similarity1
Modified residuei563PhosphoserineCombined sources1
Modified residuei572N6-acetyllysineCombined sources1
Modified residuei577N6-acetyllysine; alternateCombined sources1
Cross-linki577Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in SUMO2); alternateCombined sources
Modified residuei580PhosphoserineCombined sources1
Cross-linki589Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in SUMO1); alternateCombined sources
Cross-linki589Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in SUMO2); alternateCombined sources
Modified residuei591PhosphoserineCombined sources1
Modified residuei619PhosphoserineCombined sources1
Cross-linki624Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in SUMO2)Combined sources
Modified residuei646N6-acetyllysineCombined sources1
Modified residuei656Asymmetric dimethylarginineBy similarity1
Modified residuei660Asymmetric dimethylarginineBy similarity1
Modified residuei666Asymmetric dimethylarginineBy similarity1
Modified residuei670Asymmetric dimethylarginineBy similarity1
Modified residuei673Asymmetric dimethylarginineBy similarity1
Modified residuei679Asymmetric dimethylarginineBy similarity1
Modified residuei681Asymmetric dimethylarginineBy similarity1
Modified residuei687Asymmetric dimethylarginineBy similarity1
Modified residuei691Asymmetric dimethylarginineBy similarity1
Modified residuei694Asymmetric dimethylarginine; alternateBy similarity1
Modified residuei694Omega-N-methylarginine; alternateBy similarity1

Post-translational modificationi

Some glutamate residues are glycylated by TTLL8. This modification occurs exclusively on glutamate residues and results in a glycine chain on the gamma-carboxyl group (By similarity).By similarity
Symmetrically methylated by PRMT5.1 Publication

Keywords - PTMi

Acetylation, Isopeptide bond, Methylation, Phosphoprotein, Ubl conjugation

Proteomic databases

EPDiP19338
MaxQBiP19338
PaxDbiP19338
PeptideAtlasiP19338
PRIDEiP19338
ProteomicsDBi53647
TopDownProteomicsiP19338

2D gel databases

DOSAC-COBS-2DPAGEiP19338
SWISS-2DPAGEiP19338

PTM databases

iPTMnetiP19338
PhosphoSitePlusiP19338
SwissPalmiP19338

Miscellaneous databases

PMAP-CutDBiP19338

Expressioni

Gene expression databases

BgeeiENSG00000115053 Expressed in 103 organ(s), highest expression level in testis
CleanExiHS_NCL
ExpressionAtlasiP19338 baseline and differential
GenevisibleiP19338 HS

Organism-specific databases

HPAiCAB004210
HPA023981

Interactioni

Subunit structurei

Identified in a IGF2BP1-dependent mRNP granule complex containing untranslated mRNAs (PubMed:17289661). Component of the SWAP complex that consists of NPM1, NCL/nucleolin, PARP1 and SWAP70 (By similarity). Component of a complex which is at least composed of HTATSF1/Tat-SF1, the P-TEFb complex components CDK9 and CCNT1, RNA polymerase II, SUPT5H, and NCL/nucleolin (PubMed:10393184). Interacts with AICDA (By similarity). Interacts with APTX (PubMed:15044383). Interacts with C1QBP (PubMed:21536856). Interacts with ERBB4 (PubMed:20858735). Interacts (via C-terminus) with FMR1 isoform 6 (via N-terminus) (PubMed:24658146). Interacts with GZF1; this interaction is important for nucleolar localization of GZF1 (PubMed:17674968). Interacts with NSUN2 (PubMed:17215513). Interacts with NVL (PubMed:21474449). Interacts (via N-terminus domain) with SETX (PubMed:21700224). Interacts (via RRM1 and C-terminal RRM4/Arg/Gly-rich domains) with TERT; the interaction is important for nucleolar localization of TERT (PubMed:15371412). Interacts with WDR46 (PubMed:23848194). Interacts with ZFP36 (PubMed:20221403). Interacts with LRRC34 (By similarity). Interacts with RRP1B (PubMed:19710015). Interacts with HNRNPU; this interaction occurs during mitosis (PubMed:21242313). Interacts with RIOK1; RIOK1 recruits NCL to PRMT5 for symmetrically methylation (PubMed:21081503). Interacts with ZBTB7B (By similarity).By similarity16 Publications

Binary interactionsi

GO - Molecular functioni

Protein-protein interaction databases

BioGridi110771, 370 interactors
CORUMiP19338
DIPiDIP-89N
ELMiP19338
IntActiP19338, 103 interactors
MINTiP19338
STRINGi9606.ENSP00000318195

Structurei

Secondary structure

1710
Legend: HelixTurnBeta strandPDB Structure known for this area
Show more details

3D structure databases

ProteinModelPortaliP19338
SMRiP19338
ModBaseiSearch...
MobiDBiSearch...

Miscellaneous databases

EvolutionaryTraceiP19338

Family & Domainsi

Domains and Repeats

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Repeati58 – 6518
Repeati75 – 8228
Repeati83 – 9038
Repeati91 – 9848
Repeati99 – 1045; truncated6
Repeati105 – 11268
Repeati120 – 12778
Repeati128 – 13588
Domaini307 – 383RRM 1PROSITE-ProRule annotationAdd BLAST77
Domaini393 – 466RRM 2PROSITE-ProRule annotationAdd BLAST74
Domaini486 – 560RRM 3PROSITE-ProRule annotationAdd BLAST75
Domaini572 – 647RRM 4PROSITE-ProRule annotationAdd BLAST76

Region

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Regioni58 – 1358 X 8 AA tandem repeats of X-T-P-X-K-K-X-XAdd BLAST78

Compositional bias

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Compositional biasi143 – 171Asp/Glu-rich (acidic)Add BLAST29
Compositional biasi185 – 209Asp/Glu-rich (acidic)Add BLAST25
Compositional biasi234 – 271Asp/Glu-rich (acidic)Add BLAST38
Compositional biasi649 – 698Arg/Gly/Phe-richAdd BLAST50

Keywords - Domaini

Repeat

Phylogenomic databases

eggNOGiENOG410IRS7 Eukaryota
ENOG410XSFV LUCA
GeneTreeiENSGT00900000141052
HOGENOMiHOG000113885
HOVERGENiHBG002295
InParanoidiP19338
KOiK11294
OMAiDMESDEP
OrthoDBiEOG091G15MK
PhylomeDBiP19338
TreeFamiTF328499

Family and domain databases

CDDicd12403 RRM1_NCL, 1 hit
cd12404 RRM2_NCL, 1 hit
cd12405 RRM3_NCL, 1 hit
cd12406 RRM4_NCL, 1 hit
Gene3Di3.30.70.330, 4 hits
InterProiView protein in InterPro
IPR034230 Nucleolin_RRM1
IPR034233 Nucleolin_RRM2
IPR034234 Nucleolin_RRM3
IPR034235 Nucleolin_RRM4
IPR012677 Nucleotide-bd_a/b_plait_sf
IPR035979 RBD_domain_sf
IPR000504 RRM_dom
IPR003954 RRM_dom_euk
PfamiView protein in Pfam
PF00076 RRM_1, 4 hits
SMARTiView protein in SMART
SM00360 RRM, 4 hits
SM00361 RRM_1, 2 hits
SUPFAMiSSF54928 SSF54928, 3 hits
PROSITEiView protein in PROSITE
PS50102 RRM, 4 hits

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 5 potential isoforms that are computationally mapped.Show allAlign All

P19338-1 [UniParc]FASTAAdd to basket
« Hide
        10         20         30         40         50
MVKLAKAGKN QGDPKKMAPP PKEVEEDSED EEMSEDEEDD SSGEEVVIPQ
60 70 80 90 100
KKGKKAAATS AKKVVVSPTK KVAVATPAKK AAVTPGKKAA ATPAKKTVTP
110 120 130 140 150
AKAVTTPGKK GATPGKALVA TPGKKGAAIP AKGAKNGKNA KKEDSDEEED
160 170 180 190 200
DDSEEDEEDD EDEDEDEDEI EPAAMKAAAA APASEDEDDE DDEDDEDDDD
210 220 230 240 250
DEEDDSEEEA METTPAKGKK AAKVVPVKAK NVAEDEDEEE DDEDEDDDDD
260 270 280 290 300
EDDEDDDDED DEEEEEEEEE EPVKEAPGKR KKEMAKQKAA PEAKKQKVEG
310 320 330 340 350
TEPTTAFNLF VGNLNFNKSA PELKTGISDV FAKNDLAVVD VRIGMTRKFG
360 370 380 390 400
YVDFESAEDL EKALELTGLK VFGNEIKLEK PKGKDSKKER DARTLLAKNL
410 420 430 440 450
PYKVTQDELK EVFEDAAEIR LVSKDGKSKG IAYIEFKTEA DAEKTFEEKQ
460 470 480 490 500
GTEIDGRSIS LYYTGEKGQN QDYRGGKNST WSGESKTLVL SNLSYSATEE
510 520 530 540 550
TLQEVFEKAT FIKVPQNQNG KSKGYAFIEF ASFEDAKEAL NSCNKREIEG
560 570 580 590 600
RAIRLELQGP RGSPNARSQP SKTLFVKGLS EDTTEETLKE SFDGSVRARI
610 620 630 640 650
VTDRETGSSK GFGFVDFNSE EDAKAAKEAM EDGEIDGNKV TLDWAKPKGE
660 670 680 690 700
GGFGGRGGGR GGFGGRGGGR GGRGGFGGRG RGGFGGRGGF RGGRGGGGDH
710
KPQGKKTKFE
Length:710
Mass (Da):76,614
Last modified:January 23, 2007 - v3
Checksum:iC97F6E34E5CA6727
GO

Computationally mapped potential isoform sequencesi

There are 5 potential isoforms mapped to this entry.BLASTAlignShow allAdd to basket
EntryEntry nameProtein names
Gene namesLengthAnnotation
H7BY16H7BY16_HUMAN
Nucleolin
NCL
296Annotation score:
C9JYW2C9JYW2_HUMAN
Nucleolin
NCL
141Annotation score:
C9JLB1C9JLB1_HUMAN
Nucleolin
NCL
151Annotation score:
C9J1H7C9J1H7_HUMAN
Nucleolin
NCL
166Annotation score:
C9JWL1C9JWL1_HUMAN
Nucleolin
NCL
170Annotation score:

Sequence cautioni

The sequence BAC03738 differs from that shown. Unlikely isoform. Aberrant splice sites.Curated

Experimental Info

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Sequence conflicti549E → G in BAC03738 (PubMed:14702039).Curated1
Sequence conflicti623 – 626Missing in AAA59954 (PubMed:2394707).Curated4
Sequence conflicti625 – 627Missing (PubMed:2737305).Curated3

Natural variant

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Natural variantiVAR_04635368P → L. Corresponds to variant dbSNP:rs11542691Ensembl.1
Natural variantiVAR_046354122P → L. Corresponds to variant dbSNP:rs11542687Ensembl.1
Natural variantiVAR_046355174A → V. Corresponds to variant dbSNP:rs11542689Ensembl.1

Sequence databases

Select the link destinations:
EMBLi
GenBanki
DDBJi
Links Updated
M60858 Genomic DNA Translation: AAA59954.1
AK091742 mRNA Translation: BAC03738.1 Sequence problems.
AC017104 Genomic DNA Translation: AAY24247.1
CCDSiCCDS33397.1
PIRiA35804
S53728
RefSeqiNP_005372.2, NM_005381.2
UniGeneiHs.79110

Genome annotation databases

EnsembliENST00000322723; ENSP00000318195; ENSG00000115053
GeneIDi4691
KEGGihsa:4691
UCSCiuc002vru.4 human

Keywords - Coding sequence diversityi

Polymorphism

Similar proteinsi

Cross-referencesi

Sequence databases

Select the link destinations:
EMBLi
GenBanki
DDBJi
Links Updated
M60858 Genomic DNA Translation: AAA59954.1
AK091742 mRNA Translation: BAC03738.1 Sequence problems.
AC017104 Genomic DNA Translation: AAY24247.1
CCDSiCCDS33397.1
PIRiA35804
S53728
RefSeqiNP_005372.2, NM_005381.2
UniGeneiHs.79110

3D structure databases

Select the link destinations:
PDBei
RCSB PDBi
PDBji
Links Updated
PDB entryMethodResolution (Å)ChainPositionsPDBsum
2FC8NMR-A564-652[»]
2FC9NMR-A478-565[»]
2KRRNMR-A300-466[»]
ProteinModelPortaliP19338
SMRiP19338
ModBaseiSearch...
MobiDBiSearch...

Protein-protein interaction databases

BioGridi110771, 370 interactors
CORUMiP19338
DIPiDIP-89N
ELMiP19338
IntActiP19338, 103 interactors
MINTiP19338
STRINGi9606.ENSP00000318195

Protein family/group databases

MoonProtiP19338

PTM databases

iPTMnetiP19338
PhosphoSitePlusiP19338
SwissPalmiP19338

Polymorphism and mutation databases

BioMutaiNCL
DMDMi90110781

2D gel databases

DOSAC-COBS-2DPAGEiP19338
SWISS-2DPAGEiP19338

Proteomic databases

EPDiP19338
MaxQBiP19338
PaxDbiP19338
PeptideAtlasiP19338
PRIDEiP19338
ProteomicsDBi53647
TopDownProteomicsiP19338

Protocols and materials databases

DNASUi4691
Structural Biology KnowledgebaseSearch...

Genome annotation databases

EnsembliENST00000322723; ENSP00000318195; ENSG00000115053
GeneIDi4691
KEGGihsa:4691
UCSCiuc002vru.4 human

Organism-specific databases

CTDi4691
DisGeNETi4691
EuPathDBiHostDB:ENSG00000115053.15
GeneCardsiNCL
HGNCiHGNC:7667 NCL
HPAiCAB004210
HPA023981
MIMi164035 gene
neXtProtiNX_P19338
OpenTargetsiENSG00000115053
PharmGKBiPA31469
GenAtlasiSearch...

Phylogenomic databases

eggNOGiENOG410IRS7 Eukaryota
ENOG410XSFV LUCA
GeneTreeiENSGT00900000141052
HOGENOMiHOG000113885
HOVERGENiHBG002295
InParanoidiP19338
KOiK11294
OMAiDMESDEP
OrthoDBiEOG091G15MK
PhylomeDBiP19338
TreeFamiTF328499

Enzyme and pathway databases

ReactomeiR-HSA-6791226 Major pathway of rRNA processing in the nucleolus and cytosol
SIGNORiP19338

Miscellaneous databases

ChiTaRSiNCL human
EvolutionaryTraceiP19338
GenomeRNAii4691
PMAP-CutDBiP19338
PROiPR:P19338
SOURCEiSearch...

Gene expression databases

BgeeiENSG00000115053 Expressed in 103 organ(s), highest expression level in testis
CleanExiHS_NCL
ExpressionAtlasiP19338 baseline and differential
GenevisibleiP19338 HS

Family and domain databases

CDDicd12403 RRM1_NCL, 1 hit
cd12404 RRM2_NCL, 1 hit
cd12405 RRM3_NCL, 1 hit
cd12406 RRM4_NCL, 1 hit
Gene3Di3.30.70.330, 4 hits
InterProiView protein in InterPro
IPR034230 Nucleolin_RRM1
IPR034233 Nucleolin_RRM2
IPR034234 Nucleolin_RRM3
IPR034235 Nucleolin_RRM4
IPR012677 Nucleotide-bd_a/b_plait_sf
IPR035979 RBD_domain_sf
IPR000504 RRM_dom
IPR003954 RRM_dom_euk
PfamiView protein in Pfam
PF00076 RRM_1, 4 hits
SMARTiView protein in SMART
SM00360 RRM, 4 hits
SM00361 RRM_1, 2 hits
SUPFAMiSSF54928 SSF54928, 3 hits
PROSITEiView protein in PROSITE
PS50102 RRM, 4 hits
ProtoNetiSearch...

Entry informationi

Entry nameiNUCL_HUMAN
AccessioniPrimary (citable) accession number: P19338
Secondary accession number(s): Q53SK1
, Q8NB06, Q9UCF0, Q9UDG1
Entry historyiIntegrated into UniProtKB/Swiss-Prot: November 1, 1990
Last sequence update: January 23, 2007
Last modified: November 7, 2018
This is version 210 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. Human chromosome 2
    Human chromosome 2: 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. PDB cross-references
    Index of Protein Data Bank (PDB) cross-references
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