Atlas of Genetics and Cytogenetics in Oncology and Haematology

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CKS2 (CDC28 protein kinase regulatory subunit 2)

Written2010-02Yongyou Zhang
Case Western Reserve University, WRB-3101, 2103 Cornell Rd, Cleveland, OH 44106, USA

(Note : for Links provided by Atlas : click)


HGNC (Hugo) CKS2
HGNC Previous nameCDC28 protein kinase 2
LocusID (NCBI) 1164
Atlas_Id 40093
Location 9q22.2  [Link to chromosome band 9q22]
Location_base_pair Starts at 89311195 and ends at 89316703 bp from pter ( according to GRCh38/hg38-Dec_2013)  [Mapping CKS2.png]
Fusion genes
(updated 2017)
Data from Atlas, Mitelman, Cosmic Fusion, Fusion Cancer, TCGA fusion databases with official HUGO symbols (see references in chromosomal bands)
CKS2 (9q22.2)::PCSK5 (9q21.13)
Note There is no evidence that CKS2 gene has different transcript variant.


  Genomic organization of the CKS2 gene.
Description Three exons, spans approximately 5.5 kb of genomic DNA in the centromere-to-telomere orientation. The translation initiation codon ATG is located in exon 1, and the stop codon in exon 3.
Transcription mRNA is 627 bp.
Pseudogene 1 processed, non-expressed, pseudogene in human genome.


Note The Cks2 protein can form a special homohexamer structure. Six kinase subunits can bind the assembled hexamer, and therefore this Cks2 hexamer may participate in cell cycle control by acting as the hub for Cdk multimerization in vivo.
Description The open reading frame encodes a 79 amino acid protein, with an estimated molecular weight of approximately 9860 Da.
Expression Basic level expression in all mammalian cells and aberrant expression in cancer cells.
Localisation Cytoplasm and nucleus.
Function CKS2 protein binds to the catalytic subunit of the cyclin-dependent kinases and is essential for their biological function of cell cycle control. Especially, CKS2 is required for the first metaphase/anaphase transition of mammalian meiosis. The mice ablated of Cks2 are viable but sterile in both sexes. Sterility is due to failure of both male and female germ cells to progress from the first meiotic metaphase to anaphase. In cancer cells, CKS2 may protect the cells from apoptosis.
Homology The CKS2 protein is evolutionary conserved. Mammalian cells express two well-conserved CKS members, like the human CKS2 and CKS1B proteins. CKS2 and CKS1B may have redundant function in some context and have different functions in other context. The CKS2 protein is highly conserved cross species.


Note Mutation of glutamine for glutamate 63 (E63Q), disrupted the essential biological function of the protein and significantly reduced its ability to bind to cyclin-dependent kinases, but preserves protein structure and assembly.

Implicated in

Entity Various cancers
Note Emerging evidence showed that the expression of CKS2 is elevated in multiple cancers, including prostate cancer, breast cancer, gastric cancer, colorectal cancer, uterine cervical cancer, bladder cancer, nasopharyngeal carcinoma, melanoma, lymphoma, lung cancer, esophageal squamous cell carcinoma et al. The expression of CKS2 is correlated with poor survival rate of the patients of some cancers.
Prognosis Overexpression of CKS2 has been reported to be associated with high aggressiveness and a poor prognosis in multiple cancers, including breast cancer, prostate cancer, colon cancer, hepatocellular carcinoma and meningiomas et al.
Entity Hepatocellular carcinoma (HCC)
Note Expressions of CKS2 were significantly higher in HCC compared with the adjacent noncancerous tissues (including chronic hepatitis and cirrhosis) and normal liver tissues. Overexpression of CKS2 in HCC were closely associated with poor differentiation features (Shen et al., 2010).
Entity Gastric cancer
Note CKS2 was showed to be significantly unregulated in gastric cancers. The high level of CKS2 was highly correlated with tumor differentiation and pathological grade of the tumor size, lymph node, and metastasis stage (Kang et al., 2009).
Entity Prostate cancer
Note CKS2 were significantly unregulated in prostate tumors of human and animal models, as well as prostatic cancer cell lines. Forced expression of CKS2 in benign prostate tumor epithelial cells promoted cell population growth. Inhibition of CKS2 expression can induce programmed cell death and inhibit the tumorigenesis. (Lan et al., 2008). Over expression of CKS2 may linked with androgen-independent prostate cancer progression (Stanbrough et al., 2006).
Entity Colon cancer
Note CKS2 was reported significantly overexpressed in microdissected invasive colon tumor cells compared with adjacent normal epithelial cells (Wiese et al., 2007). CKS2 was also showed to be higher in liver metastasis compared with primary colon cancer (Lin et al., 2007).
Oncogenesis Amplification and overexpression of CKS2 were associated with liver metastasis and poor prognosis in colon cancer. CKS2 is required for germ cell to go past the first meiotic metaphase and enter anaphase. In cancer cell, overexpression of CKS2 can accelerate the cell cycles and promote the cell proliferation. Recently research showed that CKS2 may also be involved in apoptosis and metabolism since it can protect mitochondrial genome integrity via interaction with mitochondrial single-stranded DNA-binding protein. Study also showed CKS2 as a transcriptional target downregulated by the tumor suppressor p53. CKS2 expression was found to be repressed by p53 both at the mRNA and the protein levels, which may provide a mechanism that explain why CKS2 is upregulated in many types of cancer. All of these suggest that CKS2 alterations may have a significant biological role in the tumorigenesis in different tissue. The novel therapeutic strategy for cancer though may be developed via inhibiting the CKS2 activity. Therefore, disruption of CKS2-Cyclin Complex assembly or down-regulation of CKS2 expression may be used for cancer therapy.
Entity Esophageal squamous cell carcinoma
Note Gene expression profiling of lymph node metastasis by oligomicroarray analysis and Real-time RT-PCR confirmed that CKS2 is unregulated in laser microdissection of esophageal squamous cell carcinoma compared with adjacent normal tissue (Uchikado et al., 2006).
Entity Uterine cervical cancer
Note CKS2 was showed significantly higher in node positive tumor compared with negative one. The CKS2 expression is correlated with metastatic phenotypes and progression free survival. (Lyng et al., 2006).
Entity Bladder cancer
Note Large-scale gene expression profiling and Real-Time RT-PCR confirmed that a the CKS2 expression is elevated in invasive bladder cancer compared with superficial cancer (Kawakami et al., 2006).
Entity Glioblastoma
Note CKS2 was significantly up-regulated in primary glioblastomas compared with the non-neoplastic brain tissues (Scrideli et al., 2008).
Entity Meningioma
Note This microarray-based expression profiling study showed CKS2 is unregulated in atypical and anaplastic meningiomas compared with benign meningiomas (Fevre-Montange et al., 2009).


Identification of RASSF1A modulated genes in nasopharyngeal carcinoma.
Chow LS, Lam CW, Chan SY, Tsao SW, To KF, Tong SF, Hung WK, Dammann R, Huang DP, Lo KW.
Oncogene. 2006 Jan 12;25(2):310-6.
PMID 16116475
Chromosomal mapping of the human genes CKS1 to 8q21 and CKS2 to 9q22.
Demetrick DJ, Zhang H, Beach DH.
Cytogenet Cell Genet. 1996;73(3):250-4.
PMID 8697818
Germline exclusion of Cks1 in the mouse reveals a metaphase I role for Cks proteins in male and female meiosis.
Donovan PJ, Reed SI.
Cell Cycle. 2003 Jul-Aug;2(4):275-6. (REVIEW)
PMID 12851469
Cyclin-stimulated binding of Cks proteins to cyclin-dependent kinases.
Egan EA, Solomon MJ.
Mol Cell Biol. 1998 Jul;18(7):3659-67.
PMID 9632748
Microarray gene expression profiling in meningiomas: differential expression according to grade or histopathological subtype.
Fevre-Montange M, Champier J, Durand A, Wierinckx A, Honnorat J, Guyotat J, Jouvet A.
Int J Oncol. 2009 Dec;35(6):1395-407.
PMID 19885562
Myeloma cell expression of 10 candidate genes for osteolytic bone disease. Only overexpression of DKK1 correlates with clinical bone involvement at diagnosis.
Haaber J, Abildgaard N, Knudsen LM, Dahl IM, Lodahl M, Thomassen M, Kerndrup GB, Rasmussen T.
Br J Haematol. 2008 Jan;140(1):25-35. Epub 2007 Nov 12.
PMID 18005268
Upregulation of the cycline kinase subunit CKS2 increases cell proliferation rate in gastric cancer.
Kang MA, Kim JT, Kim JH, Kim SY, Kim YH, Yeom YI, Lee Y, Lee HG.
J Cancer Res Clin Oncol. 2009 Jun;135(6):761-9. Epub 2008 Nov 26.
PMID 19034516
Identification of differentially expressed genes in human bladder cancer through genome-wide gene expression profiling.
Kawakami K, Enokida H, Tachiwada T, Gotanda T, Tsuneyoshi K, Kubo H, Nishiyama K, Takiguchi M, Nakagawa M, Seki N.
Oncol Rep. 2006 Sep;16(3):521-31.
PMID 16865252
Aberrant expression of Cks1 and Cks2 contributes to prostate tumorigenesis by promoting proliferation and inhibiting programmed cell death.
Lan Y, Zhang Y, Wang J, Lin C, Ittmann MM, Wang F.
Int J Cancer. 2008 Aug 1;123(3):543-51.
PMID 18498131
Genes associated with liver metastasis of colon cancer, identified by genome-wide cDNA microarray.
Li M, Lin YM, Hasegawa S, Shimokawa T, Murata K, Kameyama M, Ishikawa O, Katagiri T, Tsunoda T, Nakamura Y, Furukawa Y.
Int J Oncol. 2004 Feb;24(2):305-12.
PMID 14719106
Inhibitory effects of estrogen receptor beta on specific hormone-responsive gene expression and association with disease outcome in primary breast cancer.
Lin CY, Strom A, Li Kong S, Kietz S, Thomsen JS, Tee JB, Vega VB, Miller LD, Smeds J, Bergh J, Gustafsson JA, Liu ET.
Breast Cancer Res. 2007;9(2):R25.
PMID 17428314
Genome wide expression profiling identifies genes associated with colorectal liver metastasis.
Lin HM, Chatterjee A, Lin YH, Anjomshoaa A, Fukuzawa R, McCall JL, Reeve AE.
Oncol Rep. 2007 Jun;17(6):1541-9.
PMID 17487416
PC-SPES inhibits cell proliferation by modulating p21, cyclins D, E and B and multiple cell cycle-related genes in prostate cancer cells.
Lu X, Guo J, Hsieh TC.
Cell Cycle. 2003 Jan-Feb;2(1):59-63.
PMID 12695690
Gene expressions and copy numbers associated with metastatic phenotypes of uterine cervical cancer.
Lyng H, Brovig RS, Svendsrud DH, Holm R, Kaalhus O, Knutstad K, Oksefjell H, Sundfor K, Kristensen GB, Stokke T.
BMC Genomics. 2006 Oct 20;7:268.
PMID 17054779
Cyclin-dependent kinase-associated proteins Cks1 and Cks2 are essential during early embryogenesis and for cell cycle progression in somatic cells.
Martinsson-Ahlzen HS, Liberal V, Grunenfelder B, Chaves SR, Spruck CH, Reed SI.
Mol Cell Biol. 2008 Sep;28(18):5698-709. Epub 2008 Jul 14.
PMID 18625720
Clinical, pathological, proliferative and molecular responses associated with neoadjuvant aromatase inhibitor treatment in breast cancer.
Miller WR.
J Steroid Biochem Mol Biol. 2010 Feb 28;118(4-5):273-6. Epub 2009 Oct 27.
PMID 19837160
Human CksHs2 atomic structure: a role for its hexameric assembly in cell cycle control.
Parge HE, Arvai AS, Murtari DJ, Reed SI, Tainer JA.
Science. 1993 Oct 15;262(5132):387-95.
PMID 8211159
Cell cycle: reaching for a role for the Cks proteins.
Pines J.
Curr Biol. 1996 Nov 1;6(11):1399-402. (REVIEW)
PMID 8939596
CKS proteins protect mitochondrial genome integrity by interacting with mitochondrial single-stranded DNA-binding protein.
Radulovic M, Crane E, Crawford M, Godovac-Zimmermann J, Yu VP.
Mol Cell Proteomics. 2010 Jan;9(1):145-52. Epub 2009 Sep 28.
PMID 19786724
Gene expression of cyclin-dependent kinase subunit Cks2 is repressed by the tumor suppressor p53 but not by the related proteins p63 or p73.
Rother K, Dengl M, Lorenz J, Tschop K, Kirschner R, Mossner J, Engeland K.
FEBS Lett. 2007 Mar 20;581(6):1166-72. Epub 2007 Feb 26.
PMID 17336302
Gene expression profile analysis of primary glioblastomas and non-neoplastic brain tissue: identification of potential target genes by oligonucleotide microarray and real-time quantitative PCR.
Scrideli CA, Carlotti CG Jr, Okamoto OK, Andrade VS, Cortez MA, Motta FJ, Lucio-Eterovic AK, Neder L, Rosemberg S, Oba-Shinjo SM, Marie SK, Tone LG.
J Neurooncol. 2008 Jul;88(3):281-91. Epub 2008 Apr 9.
PMID 18398573
Weak cooperativity in the core causes a switch in folding mechanism between two proteins of the cks family.
Seeliger MA, Breward SE, Itzhaki LS.
J Mol Biol. 2003 Jan 3;325(1):189-99.
PMID 12473461
Clinical significance and expression of cyclin kinase subunits 1 and 2 in hepatocellular carcinoma.
Shen DY, Fang ZX, You P, Liu PG, Wang F, Huang CL, Yao XB, Chen ZX, Zhang ZY.
Liver Int. 2010 Jan;30(1):119-25. Epub 2009 Oct 21.
PMID 19845855
Requirement of Cks2 for the first metaphase/anaphase transition of mammalian meiosis.
Spruck CH, de Miguel MP, Smith AP, Ryan A, Stein P, Schultz RM, Lincoln AJ, Donovan PJ, Reed SI.
Science. 2003 Apr 25;300(5619):647-50.
PMID 12714746
Increased expression of genes converting adrenal androgens to testosterone in androgen-independent prostate cancer.
Stanbrough M, Bubley GJ, Ross K, Golub TR, Rubin MA, Penning TM, Febbo PG, Balk SP.
Cancer Res. 2006 Mar 1;66(5):2815-25.
PMID 16510604
Gene expression profiling of lymph node metastasis by oligomicroarray analysis using laser microdissection in esophageal squamous cell carcinoma.
Uchikado Y, Inoue H, Haraguchi N, Mimori K, Natsugoe S, Okumura H, Aikou T, Mori M.
Int J Oncol. 2006 Dec;29(6):1337-47.
PMID 17088971
ckshs expression is linked to cell proliferation in normal and malignant human lymphoid cells.
Urbanowicz-Kachnowicz I, Baghdassarian N, Nakache C, Gracia D, Mekki Y, Bryon PA, Ffrench M.
Int J Cancer. 1999 Jul 2;82(1):98-104.
PMID 10360827
Cross-species hybridization of woodchuck hepatitis viral infection-induced woodchuck hepatocellular carcinoma using human, rat and mouse oligonucleotide microarrays.
Wang F, Kuang Y, Salem N, Anderson PW, Lee Z.
J Gastroenterol Hepatol. 2009 Apr;24(4):605-17. Epub 2008 Oct 21.
PMID 19175833
A mutation in the human cyclin-dependent kinase interacting protein, CksHs2, interferes with cyclin-dependent kinase binding and biological function, but preserves protein structure and assembly.
Watson MH, Bourne Y, Arvai AS, Hickey MJ, Santiago A, Bernstein SL, Tainer JA, Reed SI.
J Mol Biol. 1996 Sep 6;261(5):646-57.
PMID 8800213
Identification of gene signatures for invasive colorectal tumor cells.
Wiese AH, Auer J, Lassmann S, Nahrig J, Rosenberg R, Hofler H, Ruger R, Werner M.
Cancer Detect Prev. 2007;31(4):282-95.
PMID 17936523
Genome-wide gene expression profiling of cervical cancer in Hong Kong women by oligonucleotide microarray.
Wong YF, Cheung TH, Tsao GS, Lo KW, Yim SF, Wang VW, Heung MM, Chan SC, Chan LK, Ho TW, Wong KW, Li C, Guo Y, Chung TK, Smith DI.
Int J Cancer. 2006 May 15;118(10):2461-9.
PMID 16353136
Analysis of differential gene expression in human melanocytic tumour lesions by custom made oligonucleotide arrays.
de Wit NJ, Rijntjes J, Diepstra JH, van Kuppevelt TH, Weidle UH, Ruiter DJ, van Muijen GN.
Br J Cancer. 2005 Jun 20;92(12):2249-61.
PMID 15900300


This paper should be referenced as such :
Zhang, Y
CKS2 (CDC28 protein kinase regulatory subunit 2)
Atlas Genet Cytogenet Oncol Haematol. 2010;14(12):1100-1103.
Free journal version : [ pdf ]   [ DOI ]

External links


HGNC (Hugo)CKS2   2000
Entrez_Gene (NCBI)CKS2    CDC28 protein kinase regulatory subunit 2
GeneCards (Weizmann)CKS2
Ensembl hg19 (Hinxton)ENSG00000123975 [Gene_View]
Ensembl hg38 (Hinxton)ENSG00000123975 [Gene_View]  ENSG00000123975 [Sequence]  chr9:89311195-89316703 [Contig_View]  CKS2 [Vega]
ICGC DataPortalENSG00000123975
TCGA cBioPortalCKS2
AceView (NCBI)CKS2
Genatlas (Paris)CKS2
SOURCE (Princeton)CKS2
Genetics Home Reference (NIH)CKS2
Genomic and cartography
GoldenPath hg38 (UCSC)CKS2  -     chr9:89311195-89316703 +  9q22.2   [Description]    (hg38-Dec_2013)
GoldenPath hg19 (UCSC)CKS2  -     9q22.2   [Description]    (hg19-Feb_2009)
GoldenPathCKS2 - 9q22.2 [CytoView hg19]  CKS2 - 9q22.2 [CytoView hg38]
Genome Data Viewer NCBICKS2 [Mapview hg19]  
Gene and transcription
Genbank (Entrez)BC006458 BM838320 BM841681 BT006630 CR407630
RefSeq transcript (Entrez)NM_001827
Consensus coding sequences : CCDS (NCBI)CKS2
Gene ExpressionCKS2 [ NCBI-GEO ]   CKS2 [ EBI - ARRAY_EXPRESS ]   CKS2 [ SEEK ]   CKS2 [ MEM ]
Gene Expression Viewer (FireBrowse)CKS2 [ Firebrowse - Broad ]
GenevisibleExpression of CKS2 in : [tissues]  [cell-lines]  [cancer]  [perturbations]  
BioGPS (Tissue expression)1164
GTEX Portal (Tissue expression)CKS2
Human Protein AtlasENSG00000123975-CKS2 [pathology]   [cell]   [tissue]
Protein : pattern, domain, 3D structure
UniProt/SwissProtP33552   [function]  [subcellular_location]  [family_and_domains]  [pathology_and_biotech]  [ptm_processing]  [expression]  [interaction]
NextProtP33552  [Sequence]  [Exons]  [Medical]  [Publications]
With graphics : InterProP33552
Domaine pattern : Prosite (Expaxy)CKS_1 (PS00944)    CKS_2 (PS00945)   
Domains : Interpro (EBI)Cyclin-dep_kinase_reg-sub    Cyclin-dep_kinase_reg-sub_sf   
Domain families : Pfam (Sanger)CKS (PF01111)   
Domain families : Pfam (NCBI)pfam01111   
Domain families : Smart (EMBL)CKS (SM01084)  
Conserved Domain (NCBI)CKS2
PDB (RSDB)1CKS    4Y72    4YC3    5HQ0    5LQF    6GU2    6GU3    6GU4    6GU6    6GU7   
PDB Europe1CKS    4Y72    4YC3    5HQ0    5LQF    6GU2    6GU3    6GU4    6GU6    6GU7   
PDB (PDBSum)1CKS    4Y72    4YC3    5HQ0    5LQF    6GU2    6GU3    6GU4    6GU6    6GU7   
PDB (IMB)1CKS    4Y72    4YC3    5HQ0    5LQF    6GU2    6GU3    6GU4    6GU6    6GU7   
Structural Biology KnowledgeBase1CKS    4Y72    4YC3    5HQ0    5LQF    6GU2    6GU3    6GU4    6GU6    6GU7   
SCOP (Structural Classification of Proteins)1CKS    4Y72    4YC3    5HQ0    5LQF    6GU2    6GU3    6GU4    6GU6    6GU7   
CATH (Classification of proteins structures)1CKS    4Y72    4YC3    5HQ0    5LQF    6GU2    6GU3    6GU4    6GU6    6GU7   
AlphaFold pdb e-kbP33552   
Human Protein Atlas [tissue]ENSG00000123975-CKS2 [tissue]
Protein Interaction databases
IntAct (EBI)P33552
Ontologies - Pathways
Ontology : AmiGOcyclin-dependent protein kinase holoenzyme complex  chromatin binding  protein binding  regulation of transcription by RNA polymerase II  meiosis I  regulation of mitotic cell cycle  cell population proliferation  SCF ubiquitin ligase complex  protein kinase binding  histone binding  ubiquitin binding  mitotic cell cycle phase transition  positive regulation of cyclin-dependent protein serine/threonine kinase activity  cell division  cyclin-dependent protein serine/threonine kinase activator activity  
Ontology : EGO-EBIcyclin-dependent protein kinase holoenzyme complex  chromatin binding  protein binding  regulation of transcription by RNA polymerase II  meiosis I  regulation of mitotic cell cycle  cell population proliferation  SCF ubiquitin ligase complex  protein kinase binding  histone binding  ubiquitin binding  mitotic cell cycle phase transition  positive regulation of cyclin-dependent protein serine/threonine kinase activity  cell division  cyclin-dependent protein serine/threonine kinase activator activity  
Pathways : KEGGPathways in cancer    Small cell lung cancer   
NDEx NetworkCKS2
Atlas of Cancer Signalling NetworkCKS2
Wikipedia pathwaysCKS2
Orthology - Evolution
GeneTree (enSembl)ENSG00000123975
Phylogenetic Trees/Animal Genes : TreeFamCKS2
Homologs : HomoloGeneCKS2
Homology/Alignments : Family Browser (UCSC)CKS2
Gene fusions - Rearrangements
Fusion : MitelmanCKS2::PCSK5 [9q22.2/9q21.13]  
Fusion : QuiverCKS2
Polymorphisms : SNP and Copy number variants
NCBI Variation ViewerCKS2 [hg38]
dbSNP Single Nucleotide Polymorphism (NCBI)CKS2
Exome Variant ServerCKS2
GNOMAD BrowserENSG00000123975
Varsome BrowserCKS2
ACMGCKS2 variants
Genomic Variants (DGV)CKS2 [DGVbeta]
DECIPHERCKS2 [patients]   [syndromes]   [variants]   [genes]  
CONAN: Copy Number AnalysisCKS2 
ICGC Data PortalCKS2 
TCGA Data PortalCKS2 
Broad Tumor PortalCKS2
OASIS PortalCKS2 [ Somatic mutations - Copy number]
Somatic Mutations in Cancer : COSMICCKS2  [overview]  [genome browser]  [tissue]  [distribution]  
Somatic Mutations in Cancer : COSMIC3DCKS2
Mutations and Diseases : HGMDCKS2
LOVD (Leiden Open Variation Database)[gene] [transcripts] [variants]
DgiDB (Drug Gene Interaction Database)CKS2
DoCM (Curated mutations)CKS2
CIViC (Clinical Interpretations of Variants in Cancer)CKS2
NCG (London)CKS2
Impact of mutations[PolyPhen2] [Provean] [Buck Institute : MutDB] [Mutation Assessor] [Mutanalyser]
Genetic Testing Registry CKS2
NextProtP33552 [Medical]
Target ValidationCKS2
Huge Navigator CKS2 [HugePedia]
Clinical trials, drugs, therapy
Protein Interactions : CTDCKS2
Pharm GKB GenePA26536
Clinical trialCKS2
DataMed IndexCKS2
PubMed52 Pubmed reference(s) in Entrez
GeneRIFsGene References Into Functions (Entrez)
REVIEW articlesautomatic search in PubMed
Last year publicationsautomatic search in PubMed

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