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Protein NCL PDB 2fc8.png
PDB rendering based on 2fc8.
Available structures
PDB Ortholog search: PDBe, RCSB
Symbols NCL ; C23
External IDs OMIM164035 HomoloGene136488 GeneCards: NCL Gene
RNA expression pattern
PBB GE NCL 200610 s at tn.png
More reference expression data
Species Human Mouse
Entrez 4691 17975
Ensembl ENSG00000115053 ENSMUSG00000026234
UniProt P19338 P09405
RefSeq (mRNA) NM_005381 NM_010880
RefSeq (protein) NP_005372 NP_035010
Location (UCSC) Chr 2:
232.32 – 232.35 Mb
Chr 1:
86.34 – 86.36 Mb
PubMed search [1] [2]

Nucleolin is a protein that in humans is encoded by the NCL gene.[1][2]


The human NCL gene is located on chromosome 2 and consists of 14 exons with 13 introns and spans approximately 11kb. The intron 11 of the NCL gene encodes a small nucleolar RNA, termed U20.[3]


Nucleolin (NCL), a eukaryotic nucleolar phosphoprotein, is involved in the synthesis and maturation of ribosomes.[3] The protein is located mainly in dense fibrillar regions of the nucleolus.

Nucleolin is also able to act as a transcriptional coactivator with Chicken Ovalbumin Upstream Promoter Transcription Factor II (COUP-TFII).[4]

Clinical significance[edit]

Midkine and pleiotrophin bind to cell-surface nucleolin as a low affinity receptor. This binding can inhibit HIV infection.[5][6]


Nucleolin has been shown to interact with:


  1. ^ Srivastava M, McBride O, Fleming P, Pollard H, Burns A (October 1990). "Genomic organization and chromosomal localization of the human nucleolin gene". J Biol Chem 265 (25): 14922–31. PMID 2394707. 
  2. ^ Erard M, Belenguer P, Caizergues-Ferrer M, Pantaloni A, Amalric F (October 1988). "A major nucleolar protein, nucleolin, induces chromatin decondensation by binding to histone H1". Eur J Biochem 175 (3): 525–30. doi:10.1111/j.1432-1033.1988.tb14224.x. PMID 3409881. 
  3. ^ a b "Entrez Gene: NCL nucleolin". 
  4. ^ Litchfield L, Riggs K, Hockenberry A, Oliver L, Barnhart K, Cai J et al. (May 2012). "Identification and characterization of nucleolin as a COUP-TFII coactivator of retinoic acid receptor β transcription in breast cancer cells.". PLoS ONE 7 (5). doi:10.1371/journal.pone.0038278. PMC 3365040. PMID 22693611. 
  5. ^ Said E, Krust B, Nisole S, Svab J, Briand J, Hovanessian A (October 2002). "The anti-HIV cytokine midkine binds the cell surface-expressed nucleolin as a low affinity receptor". J. Biol. Chem. 277 (40): 37492–502. doi:10.1074/jbc.M201194200. PMID 12147681. 
  6. ^ Said E, Courty J, Svab J, Delbé J, Krust B, Hovanessian A (September 2005). "Pleiotrophin inhibits HIV infection by binding the cell surface-expressed nucleolin". FEBS J. 272 (18): 4646–59. doi:10.1111/j.1742-4658.2005.04870.x. PMID 16156786. 
  7. ^ Li D, Dobrowolska G, Krebs E (June 1996). "The physical association of casein kinase 2 with nucleolin". J. Biol. Chem. 271 (26): 15662–8. PMID 8663258. 
  8. ^ Dubois T, Zemlickova E, Howell S, Aitken A (February 2003). "Centaurin-alpha 1 associates in vitro and in vivo with nucleolin". Biochem. Biophys. Res. Commun. 301 (2): 502–8. PMID 12565890. 
  9. ^ Tominaga K, Srikantan S, Lee E, Subaran S, Martindale J, Abdelmohsen K et al. (August 2011). "Competitive regulation of nucleolin expression by HuR and miR-494". Mol. Cell. Biol. 31 (20): 4219–31. doi:10.1128/MCB.05955-11. PMC 3187287. PMID 21859890. 
  10. ^ Li Y, Busch R, Valdez B, Busch H (April 1996). "C23 interacts with B23, a putative nucleolar-localization-signal-binding protein". Eur. J. Biochem. 237 (1): 153–8. PMID 8620867. 
  11. ^ Daniely Y, Dimitrova D, Borowiec J (August 2002). "Stress-dependent nucleolin mobilization mediated by p53-nucleolin complex formation". Mol. Cell. Biol. 22 (16): 6014–22. PMC 133981. PMID 12138209. 
  12. ^ Morimoto H, Okamura H, Haneji T (September 2002). "Interaction of protein phosphatase 1 delta with nucleolin in human osteoblastic cells". J. Histochem. Cytochem. 50 (9): 1187–93. PMID 12185196. 
  13. ^ Sakaguchi M, Miyazaki M, Takaishi M, Sakaguchi Y, Makino E, Kataoka N et al. (November 2003). "S100C/A11 is a key mediator of Ca(2+)-induced growth inhibition of human epidermal keratinocytes". J. Cell Biol. 163 (4): 825–35. doi:10.1083/jcb.200304017. PMC 2173690. PMID 14623863. 
  14. ^ Fouraux M, Bouvet P, Verkaart S, van Venrooij W, Pruijn G (July 2002). "Nucleolin associates with a subset of the human Ro ribonucleoprotein complexes". J. Mol. Biol. 320 (3): 475–88. PMID 12096904. 
  15. ^ Haluska P, Saleem A, Edwards T, Rubin E (April 1998). "Interaction between the N-terminus of human topoisomerase I and SV40 large T antigen". Nucleic Acids Res. 26 (7): 1841–7. PMC 147454. PMID 9512561. 
  16. ^ Bharti A, Olson M, Kufe D, Rubin E (January 1996). "Identification of a nucleolin binding site in human topoisomerase I". J. Biol. Chem. 271 (4): 1993–7. PMID 8567649. 
  17. ^ Khurts S, Masutomi K, Delgermaa L, Arai K, Oishi N, Mizuno H et al. (December 2004). "Nucleolin interacts with telomerase". J. Biol. Chem. 279 (49): 51508–15. doi:10.1074/jbc.M407643200. PMID 15371412. 

Further reading[edit]