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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 PPP1CB,[7] Centaurin, alpha 1,[8] CSNK2A2,[9] Sjogren syndrome antigen B,[10] TOP1,[11][12] P53,[13] S100A11,[14] NPM1,[15] Telomerase reverse transcriptase,[16] and HuR.[17]


  1. ^ Srivastava M, McBride OW, Fleming PJ, Pollard HB, Burns AL (October 1990). "Genomic organization and chromosomal localization of the human nucleolin gene". J Biol Chem 265 (25): 14922–31. PMID 2394707. 
  2. ^ Erard MS, 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 LM, Riggs KA, Hockenberry AM, Oliver LD, Barnhart KG, Cai J, Pierce WM Jr, Ivanova MM, Bates PJ, Appana SN, Datta S, Kulesza P, McBryan J, Young LS, Klinge CM. (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 EA, Krust B, Nisole S, Svab J, Briand JP, Hovanessian AG (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 EA, Courty J, Svab J, Delbé J, Krust B, Hovanessian AG (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. ^ Morimoto, Hiroyuki; Okamura Hirohiko; Haneji Tatsuji (September 2002). "Interaction of protein phosphatase 1 delta with nucleolin in human osteoblastic cells". J. Histochem. Cytochem. (United States) 50 (9): 1187–93. doi:10.1177/002215540205000905. ISSN 0022-1554. PMID 12185196. 
  8. ^ Dubois, Thierry; Zemlickova Eva; Howell Steven; Aitken Alastair (February 2003). "Centaurin-alpha 1 associates in vitro and in vivo with nucleolin". Biochem. Biophys. Res. Commun. (United States) 301 (2): 502–8. doi:10.1016/S0006-291X(02)03010-3. ISSN 0006-291X. PMID 12565890. 
  9. ^ Li, D; Dobrowolska G; Krebs E G (June 1996). "The physical association of casein kinase 2 with nucleolin". J. Biol. Chem. (UNITED STATES) 271 (26): 15662–8. doi:10.1074/jbc.271.26.15662. ISSN 0021-9258. PMID 8663258. 
  10. ^ Fouraux, Michael A; Bouvet Philippe; Verkaart Sjoerd; van Venrooij Walther J; Pruijn Ger J M (July 2002). "Nucleolin associates with a subset of the human Ro ribonucleoprotein complexes". J. Mol. Biol. (England) 320 (3): 475–88. doi:10.1016/S0022-2836(02)00518-1. ISSN 0022-2836. PMID 12096904. 
  11. ^ Haluska, P; Saleem A; Edwards T K; Rubin E H (April 1998). "Interaction between the N-terminus of human topoisomerase I and SV40 large T antigen". Nucleic Acids Res. (ENGLAND) 26 (7): 1841–7. doi:10.1093/nar/26.7.1841. ISSN 0305-1048. PMC 147454. PMID 9512561. 
  12. ^ Bharti, A K; Olson M O; Kufe D W; Rubin E H (January 1996). "Identification of a nucleolin binding site in human topoisomerase I". J. Biol. Chem. (UNITED STATES) 271 (4): 1993–7. doi:10.1074/jbc.271.4.1993. ISSN 0021-9258. PMID 8567649. 
  13. ^ Daniely, Yaron; Dimitrova Diana D; Borowiec James A (August 2002). "Stress-Dependent Nucleolin Mobilization Mediated by p53-Nucleolin Complex Formation". Mol. Cell. Biol. (United States) 22 (16): 6014–22. doi:10.1128/MCB.22.16.6014-6022.2002. ISSN 0270-7306. PMC 133981. PMID 12138209. 
  14. ^ Sakaguchi, Masakiyo; Miyazaki Masahiro; Takaishi Mikiro; Sakaguchi Yoshihiko; Makino Eiichi; Kataoka Noriyuki; Yamada Hidenori; Namba Masayoshi; Huh Nam-ho (November 2003). "S100C/A11 is a key mediator of Ca2+-induced growth inhibition of human epidermal keratinocytes". J. Cell Biol. (United States) 163 (4): 825–35. doi:10.1083/jcb.200304017. ISSN 0021-9525. PMC 2173690. PMID 14623863. 
  15. ^ Li, Y P; Busch R K; Valdez B C; Busch H (April 1996). "C23 interacts with B23, a putative nucleolar-localization-signal-binding protein". Eur. J. Biochem. (GERMANY) 237 (1): 153–8. doi:10.1111/j.1432-1033.1996.0153n.x. ISSN 0014-2956. PMID 8620867. 
  16. ^ Khurts, Shilagardi; Masutomi Kenkichi; Delgermaa Luvsanjav; Arai Kuniaki; Oishi Naoki; Mizuno Hideki; Hayashi Naoyuki; Hahn William C; Murakami Seishi (December 2004). "Nucleolin interacts with telomerase". J. Biol. Chem. (United States) 279 (49): 51508–15. doi:10.1074/jbc.M407643200. ISSN 0021-9258. PMID 15371412. 
  17. ^ Tominaga, K; Srikantan S; Lee EK; Subaran SS; Martindale JL; Abdelmohsen K; Gorospe M (August 2011). "Competitive Regulation of Nucleolin Expression by HuR and miR-494". Mol. Cell. Biol. (United States) 31 (20): 4219–31. doi:10.1128/MCB.05955-11. PMC 3187287. PMID 21859890. 

Further reading[edit]