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KSR1 (kinase suppressor of ras 1)

Written2011-02Mario Fernandez, Robert Lewis
Eppley Institute for Cancer Research, University of Nebraska Medical Center, Omaha, NE 68198, USA

(Note : for Links provided by Atlas : click)


HGNC (Hugo) KSR1
HGNC Alias symbRSU2
HGNC Previous nameKSR
HGNC Previous namekinase suppressor of ras
LocusID (NCBI) 8844
Atlas_Id 41107
Location 17q11.1  [Link to chromosome band 17q11]
Location_base_pair Starts at 27471964 and ends at 27626435 bp from pter ( according to GRCh38/hg38-Dec_2013)  [Mapping KSR1.png]
Local_order From centromere to telomere on 17q11.2, the KSR1 gene is flanked by: EOE1, EVI2A, JJAZ1, KNO3, KRT24, KSR1, LGALS9, LYZL6, MIR144, MIR451 (NCBI).
Fusion genes
(updated 2017)
Data from Atlas, Mitelman, Cosmic Fusion, Fusion Cancer, TCGA fusion databases with official HUGO symbols (see references in chromosomal bands)
AGPAT3 (21q22.3) / KSR1 (17q11.1)AGPAT3 (21q22.3) / KSR1 (17q11.2)CDR2L (17q25.1) / KSR1 (17q11.1)
CDR2L (17q25.1) / KSR1 (17q11.2)CFAP46 (10q26.3) / KSR1 (17q11.1)CHD9 (16q12.2) / KSR1 (17q11.1)
CHD9 (16q12.2) / KSR1 (17q11.2)CYTH1 (17q25.3) / KSR1 (17q11.1)CYTH1 (17q25.3) / KSR1 (17q11.2)
KSR1 (17q11.1) / EFCAB5 (17q11.2)KSR1 (17q11.1) / KSR1 (17q11.1)KSR1 (17q11.1) / LGALS9 (17q11.2)
KSR1 (17q11.1) / MSI2 (17q22)KSR1 (17q11.1) / PIPOX (17q11.2)KSR1 (17q11.1) / TENM1 (Xq25)
KSR1 (17q11.2) / EFCAB5 (17q11.2)KSR1 (17q11.2) / MSI2 (17q22)KSR1 (17q11.2) / PIPOX (17q11.2)
KSR1 (17q11.2) / TENM1 (Xq25)MKS1 (17q22) / KSR1 (17q11.1)MKS1 (17q22) / KSR1 (17q11.2)


  KSR1 contains 17 exons, 154426 base pairs.
Description The KSR1 gene was discovered in D. melanogaster and is highly conserved along the evolutionary tree through H. sapiens.
Transcription Transcription can produce two different mRNA transcripts. The KSR1 is expressed in the thymus, bone marrow, brain, heart, kidney, lung, liver, pancreas, mammary gland, ovary, testis and muscle.
Pseudogene No known pseudogenes.


Note Belongs to the KSR family of proteins.
  The KSR1 protein contains five conserved areas (CA). The CA1 region is unique to KSR1 protein; the CA2 region is a proline-rich region; the CA3 region is implicated in the Ras-induced plasma membrane localization; the CA4 region is a serine/threonine rich region and contains the MAPK docking site (FXFP); and the putative kinase domain (CA5), which contains an amino acid variation in subdomain II that suggests that this protein is catalytically inert. The KSR1 protein is composed of 921 amino acids.
Description The KSR1 protein is 921 kDa. There have been 15 phosphorylation sites identified.
Expression Brain, kidney, lung, pancreas, ovaries, testis, breast.
Localisation Sequestered in the cytoplasm in quiescents cells. Upon stimulation, KSR1 protein localizes to the plasma membrane.
Function Acts as a scaffold for the Raf/MEK/ERK kinase cascade.
Homology R. norvegicus KSR1 (91.85%), M. musculus KSR1 (91.3%), C. familians KSR1 (93.62%), G. gallus KSR1 (76.39%), D. reno KSR1 (67.85%), D. Melanogaster KSR (41.06%), C. elegans KSR1 (35.37%), X. laevis KSR1 (76.06%).


Note There have been no prominent germinal or somatic mutations identified.

Implicated in

Entity Cancer
Note KSR1 has not been found to be upregulated or downregulated in tumors. However, studies in KSR1 knockout mice showed that the mice were less susceptible to papilloma virus driven tumors, indicating that KSR1 is required for Ras-mediated oncogenesis.

To be noted

KSR1 is primarily recognized as a scaffold for the Raf/MEK/ERK kinase cascade. However, there are several published studies that sustain that KSR1 has catalytic activity. KSR1 is recognized as a pseudokinase, since mammalian KSR1 does not possess the lysine responsible for ATP orientation and hydrolysis in the putative kinase domain. This lysine is present in C. elegans and D. melanogaster, but mutation of this site did not affect activation of the Raf/MEK/ERK pathway. A recent publication (Rajakulendran et al., 2009) showed that KSR1 dimerization via its kinase domain to Raf facilitated the activation of Raf without KSR1 catalytic activity. It remains possible that KSR1 proteins have catalytic activity based on a yet to be identified novel mechanism.


Ras subcellular localization defines extracellular signal-regulated kinase 1 and 2 substrate specificity through distinct utilization of scaffold proteins.
Casar B, Arozarena I, Sanz-Moreno V, Pinto A, Agudo-Ibanez L, Marais R, Lewis RE, Berciano MT, Crespo P.
Mol Cell Biol. 2009 Mar;29(5):1338-53. Epub 2008 Dec 29.
PMID 19114553
IMP modulates KSR1-dependent multivalent complex formation to specify ERK1/2 pathway activation and response thresholds.
Chen C, Lewis RE, White MA.
J Biol Chem. 2008 May 9;283(19):12789-96. Epub 2008 Mar 10.
PMID 18332145
The MAPK scaffold kinase suppressor of Ras is involved in ERK activation by stress and proinflammatory cytokines and induction of arthritis.
Fusello AM, Mandik-Nayak L, Shih F, Lewis RE, Allen PM, Shaw AS.
J Immunol. 2006 Nov 1;177(9):6152-8.
PMID 17056543
Expression of kinase suppressor of Ras in the normal adult and embryonic mouse.
Giblett SM, Lloyd DJ, Light Y, Marais R, Pritchard CA.
Cell Growth Differ. 2002 Jul;13(7):307-13.
PMID 12133899
The mitogen-activated protein kinase scaffold KSR1 is required for recruitment of extracellular signal-regulated kinase to the immunological synapse.
Giurisato E, Lin J, Harding A, Cerutti E, Cella M, Lewis RE, Colonna M, Shaw AS.
Mol Cell Biol. 2009 Mar;29(6):1554-64. Epub 2009 Jan 12.
PMID 19139278
Expression of kinase suppressor of Ras1 enhances cisplatin-induced extracellular signal-regulated kinase activation and cisplatin sensitivity.
Kim M, Yan Y, Kortum RL, Stoeger SM, Sgagias MK, Lee K, Lewis RE, Cowan KH.
Cancer Res. 2005 May 15;65(10):3986-92.
PMID 15899786
The ksr-1 gene encodes a novel protein kinase involved in Ras-mediated signaling in C. elegans.
Kornfeld K, Hom DB, Horvitz HR.
Cell. 1995 Dec 15;83(6):903-13.
PMID 8521514
The molecular scaffold kinase suppressor of Ras 1 (KSR1) regulates adipogenesis.
Kortum RL, Costanzo DL, Haferbier J, Schreiner SJ, Razidlo GL, Wu MH, Volle DJ, Mori T, Sakaue H, Chaika NV, Chaika OV, Lewis RE.
Mol Cell Biol. 2005 Sep;25(17):7592-604.
PMID 16107706
The molecular scaffold kinase suppressor of Ras 1 is a modifier of RasV12-induced and replicative senescence.
Kortum RL, Johnson HJ, Costanzo DL, Volle DJ, Razidlo GL, Fusello AM, Shaw AS, Lewis RE.
Mol Cell Biol. 2006 Mar;26(6):2202-14.
PMID 16507997
The molecular scaffold KSR1 regulates the proliferative and oncogenic potential of cells.
Kortum RL, Lewis RE.
Mol Cell Biol. 2004 May;24(10):4407-16.
PMID 15121859
Kinase suppressor of Ras couples Ras to the ERK cascade during T cell development.
Laurent MN, Ramirez DM, Alberola-Ila J.
J Immunol. 2004 Jul 15;173(2):986-92.
PMID 15240686
Deficiency of kinase suppressor of Ras1 prevents oncogenic ras signaling in mice.
Lozano J, Xing R, Cai Z, Jensen HL, Trempus C, Mark W, Cannon R, Kolesnick R.
Cancer Res. 2003 Jul 15;63(14):4232-8.
PMID 12874031
Caspase-dependent cleavage disrupts the ERK cascade scaffolding function of KSR1.
McKay MM, Morrison DK.
J Biol Chem. 2007 Sep 7;282(36):26225-34. Epub 2007 Jul 5.
PMID 17613518
Signaling dynamics of the KSR1 scaffold complex.
McKay MM, Ritt DA, Morrison DK.
Proc Natl Acad Sci U S A. 2009 Jul 7;106(27):11022-7. Epub 2009 Jun 18.
PMID 19541618
KSR stimulates Raf-1 activity in a kinase-independent manner.
Michaud NR, Therrien M, Cacace A, Edsall LC, Spiegel S, Rubin GM, Morrison DK.
Proc Natl Acad Sci U S A. 1997 Nov 25;94(24):12792-6.
PMID 9371754
Identification of B-KSR1, a novel brain-specific isoform of KSR1 that functions in neuronal signaling.
Muller J, Cacace AM, Lyons WE, McGill CB, Morrison DK.
Mol Cell Biol. 2000 Aug;20(15):5529-39.
PMID 10891492
C-TAK1 regulates Ras signaling by phosphorylating the MAPK scaffold, KSR1.
Muller J, Ory S, Copeland T, Piwnica-Worms H, Morrison DK.
Mol Cell. 2001 Nov;8(5):983-93.
PMID 11741534
Kinase suppressor of Ras (KSR) is a scaffold which facilitates mitogen-activated protein kinase activation in vivo.
Nguyen A, Burack WR, Stock JL, Kortum R, Chaika OV, Afkarian M, Muller WJ, Murphy KM, Morrison DK, Lewis RE, McNeish J, Shaw AS.
Mol Cell Biol. 2002 May;22(9):3035-45.
PMID 11940661
Protein phosphatase 2A positively regulates Ras signaling by dephosphorylating KSR1 and Raf-1 on critical 14-3-3 binding sites.
Ory S, Zhou M, Conrads TP, Veenstra TD, Morrison DK.
Curr Biol. 2003 Aug 19;13(16):1356-64.
PMID 12932319
A dimerization-dependent mechanism drives RAF catalytic activation.
Rajakulendran T, Sahmi M, Lefrancois M, Sicheri F, Therrien M.
Nature. 2009 Sep 24;461(7263):542-5. Epub 2009 Sep 2.
PMID 19727074
KSR1 is required for cell cycle reinitiation following DNA damage.
Razidlo GL, Johnson HJ, Stoeger SM, Cowan KH, Bessho T, Lewis RE.
J Biol Chem. 2009 Mar 13;284(11):6705-15. Epub 2009 Jan 15.
PMID 19147494
CK2 Is a component of the KSR1 scaffold complex that contributes to Raf kinase activation.
Ritt DA, Zhou M, Conrads TP, Veenstra TD, Copeland TD, Morrison DK.
Curr Biol. 2007 Jan 23;17(2):179-84. Epub 2006 Dec 14.
PMID 17174095
KSR is a scaffold required for activation of the ERK/MAPK module.
Roy F, Laberge G, Douziech M, Ferland-McCollough D, Therrien M.
Genes Dev. 2002 Feb 15;16(4):427-38.
PMID 11850406
Nm23-H1 metastasis suppressor expression level influences the binding properties, stability, and function of the kinase suppressor of Ras1 (KSR1) Erk scaffold in breast carcinoma cells.
Salerno M, Palmieri D, Bouadis A, Halverson D, Steeg PS.
Mol Cell Biol. 2005 Feb;25(4):1379-88.
PMID 15684389
The C. elegans ksr-1 gene encodes a novel Raf-related kinase involved in Ras-mediated signal transduction.
Sundaram M, Han M.
Cell. 1995 Dec 15;83(6):889-901.
PMID 8521513
KSR modulates signal propagation within the MAPK cascade.
Therrien M, Michaud NR, Rubin GM, Morrison DK.
Genes Dev. 1996 Nov 1;10(21):2684-95.
PMID 8946910
Pharmacologic inactivation of kinase suppressor of ras-1 abrogates Ras-mediated pancreatic cancer.
Xing HR, Cordon-Cardo C, Deng X, Tong W, Campodonico L, Fuks Z, Kolesnick R.
Nat Med. 2003 Oct;9(10):1266-8. Epub 2003 Sep 7.
PMID 12960962
Kinase suppressor of Ras-1 protects intestinal epithelium from cytokine-mediated apoptosis during inflammation.
Yan F, John SK, Wilson G, Jones DS, Washington MK, Polk DB.
J Clin Invest. 2004 Nov;114(9):1272-80.
PMID 15520859


This paper should be referenced as such :
Fernandez, M ; Lewis, R
KSR1 (kinase suppressor of ras 1)
Atlas Genet Cytogenet Oncol Haematol. 2011;15(9):721-723.
Free journal version : [ pdf ]   [ DOI ]

Other Leukemias implicated (Data extracted from papers in the Atlas) [ 1 ]
  t(X;17)(q25;q11) KSR1/TENM1

Other Solid tumors implicated (Data extracted from papers in the Atlas) [ 8 ]
  t(16;17)(q12;q11) CHD9/KSR1
KSR1/EFCAB5 (17q11)
KSR1/PIPOX (17q11)
t(17;17)(q11;q22) KSR1/MSI2
t(17;17)(q11;q22) MKS1/KSR1
t(17;17)(q11;q25) CDR2L/KSR1
t(17;17)(q11;q25) CYTH1/KSR1
t(17;21)(q11;q22) AGPAT3/KSR1

External links

HGNC (Hugo)KSR1   6465
Entrez_Gene (NCBI)KSR1    kinase suppressor of ras 1
AliasesKSR; RSU2
GeneCards (Weizmann)KSR1
Ensembl hg19 (Hinxton)ENSG00000141068 [Gene_View]
Ensembl hg38 (Hinxton)ENSG00000141068 [Gene_View]  ENSG00000141068 [Sequence]  chr17:27471964-27626435 [Contig_View]  KSR1 [Vega]
ICGC DataPortalENSG00000141068
TCGA cBioPortalKSR1
AceView (NCBI)KSR1
Genatlas (Paris)KSR1
SOURCE (Princeton)KSR1
Genetics Home Reference (NIH)KSR1
Genomic and cartography
GoldenPath hg38 (UCSC)KSR1  -     chr17:27471964-27626435 +  17q11.2   [Description]    (hg38-Dec_2013)
GoldenPath hg19 (UCSC)KSR1  -     17q11.2   [Description]    (hg19-Feb_2009)
GoldenPathKSR1 - 17q11.2 [CytoView hg19]  KSR1 - 17q11.2 [CytoView hg38]
genome Data Viewer NCBIKSR1 [Mapview hg19]  
Gene and transcription
Genbank (Entrez)AK095721 AK124344 AK130719 BC035488 BC037586
RefSeq transcript (Entrez)NM_001367810 NM_014238
RefSeq genomic (Entrez)
Consensus coding sequences : CCDS (NCBI)KSR1
Alternative Splicing GalleryENSG00000141068
Gene ExpressionKSR1 [ NCBI-GEO ]   KSR1 [ EBI - ARRAY_EXPRESS ]   KSR1 [ SEEK ]   KSR1 [ MEM ]
Gene Expression Viewer (FireBrowse)KSR1 [ Firebrowse - Broad ]
GenevisibleExpression of KSR1 in : [tissues]  [cell-lines]  [cancer]  [perturbations]  
BioGPS (Tissue expression)8844
GTEX Portal (Tissue expression)KSR1
Human Protein AtlasENSG00000141068-KSR1 [pathology]   [cell]   [tissue]
Protein : pattern, domain, 3D structure
UniProt/SwissProtQ8IVT5   [function]  [subcellular_location]  [family_and_domains]  [pathology_and_biotech]  [ptm_processing]  [expression]  [interaction]
NextProtQ8IVT5  [Sequence]  [Exons]  [Medical]  [Publications]
With graphics : InterProQ8IVT5
Splice isoforms : SwissVarQ8IVT5
Domaine pattern : Prosite (Expaxy)PROTEIN_KINASE_DOM (PS50011)    PROTEIN_KINASE_ST (PS00108)    ZF_DAG_PE_1 (PS00479)    ZF_DAG_PE_2 (PS50081)   
Domains : Interpro (EBI)Kinase-like_dom_sf    KSR_SAM-like_dom    PE/DAG-bd    Prot_kinase_dom    SAM/pointed_sf    Ser-Thr/Tyr_kinase_cat_dom    Ser/Thr_kinase_AS   
Domain families : Pfam (Sanger)Pkinase_Tyr (PF07714)    SAM_KSR1 (PF13543)   
Domain families : Pfam (NCBI)pfam07714    pfam13543   
Domain families : Smart (EMBL)C1 (SM00109)  S_TKc (SM00220)  
Conserved Domain (NCBI)KSR1
Blocks (Seattle)KSR1
PDB Europe5VYK   
Structural Biology KnowledgeBase5VYK   
SCOP (Structural Classification of Proteins)5VYK   
CATH (Classification of proteins structures)5VYK   
Human Protein Atlas [tissue]ENSG00000141068-KSR1 [tissue]
Peptide AtlasQ8IVT5
IPIIPI01015914   IPI00217124   IPI01018676   IPI00013384   IPI00747688   IPI01026015   
Protein Interaction databases
IntAct (EBI)Q8IVT5
Ontologies - Pathways
Ontology : AmiGOMAPK cascade  activation of MAPKKK activity  protein kinase activity  protein serine/threonine kinase activity  MAP-kinase scaffold activity  protein binding  ATP binding  cytoplasm  endoplasmic reticulum  endoplasmic reticulum  endoplasmic reticulum membrane  cytosol  signal transduction  Ras protein signal transduction  Ras protein signal transduction  Ras protein signal transduction  protein C-terminus binding  membrane  membrane  membrane  cAMP-mediated signaling  mitogen-activated protein kinase kinase binding  ruffle membrane  protein-containing complex  regulation of cell proliferation  regulation of MAP kinase activity  positive regulation of MAPK cascade  metal ion binding  chaperone binding  Hsp90 protein binding  14-3-3 protein binding  
Ontology : EGO-EBIMAPK cascade  activation of MAPKKK activity  protein kinase activity  protein serine/threonine kinase activity  MAP-kinase scaffold activity  protein binding  ATP binding  cytoplasm  endoplasmic reticulum  endoplasmic reticulum  endoplasmic reticulum membrane  cytosol  signal transduction  Ras protein signal transduction  Ras protein signal transduction  Ras protein signal transduction  protein C-terminus binding  membrane  membrane  membrane  cAMP-mediated signaling  mitogen-activated protein kinase kinase binding  ruffle membrane  protein-containing complex  regulation of cell proliferation  regulation of MAP kinase activity  positive regulation of MAPK cascade  metal ion binding  chaperone binding  Hsp90 protein binding  14-3-3 protein binding  
Pathways : KEGGRas signaling pathway    Tuberculosis   
REACTOMEQ8IVT5 [protein]
REACTOME PathwaysR-HSA-6802955 [pathway]   
NDEx NetworkKSR1
Atlas of Cancer Signalling NetworkKSR1
Wikipedia pathwaysKSR1
Orthology - Evolution
GeneTree (enSembl)ENSG00000141068
Phylogenetic Trees/Animal Genes : TreeFamKSR1
Homologs : HomoloGeneKSR1
Homology/Alignments : Family Browser (UCSC)KSR1
Gene fusions - Rearrangements
Fusion : MitelmanAGPAT3/KSR1 [21q22.3/17q11.1]  
Fusion : MitelmanCDR2L/KSR1 [17q25.1/17q11.1]  
Fusion : MitelmanCHD9/KSR1 [16q12.2/17q11.1]  
Fusion : MitelmanCYTH1/KSR1 [17q25.3/17q11.1]  
Fusion : MitelmanKSR1/EFCAB5 [17q11.1/17q11.2]  
Fusion : MitelmanKSR1/MSI2 [17q11.1/17q22]  
Fusion : MitelmanKSR1/PIPOX [17q11.1/17q11.2]  
Fusion : MitelmanKSR1/TENM1 [17q11.1/Xq25]  
Fusion : MitelmanMKS1/KSR1 [17q22/17q11.1]  
Fusion PortalAGPAT3 21q22.3 KSR1 17q11.1 BRCA
Fusion PortalCDR2L 17q25.1 KSR1 17q11.1 LUSC
Fusion PortalCHD9 16q12.2 KSR1 17q11.1 BRCA
Fusion PortalCYTH1 17q25.3 KSR1 17q11.1 LUAD
Fusion PortalKSR1 17q11.1 EFCAB5 17q11.2 LUAD
Fusion PortalKSR1 17q11.1 MSI2 17q22 BRCA
Fusion PortalKSR1 17q11.1 ODZ1 LAML
Fusion PortalKSR1 17q11.1 PIPOX 17q11.2 BRCA
Fusion PortalMKS1 17q22 KSR1 17q11.1 BRCA
Fusion : Fusion_HubAGPAT3--KSR1    CDR2L--KSR1    CHD9--KSR1    CYTH1--KSR1    DDX60L--KSR1    DSP--KSR1    EFCAB5--KSR1    INTS4L1--KSR1    INTS4L2--KSR1    KSR1--BSCL2    KSR1--C17ORF108    KSR1--ECI1    KSR1--EFCAB5    KSR1--GP2    KSR1--IL1R2   
KSR1--KSR1    KSR1--LGALS9    KSR1--LGALS9C    KSR1--MSI2    KSR1--ODZ1    KSR1--PIPOX    KSR1--PKN3    KSR1--PLA2G16    KSR1--PMP22    KSR1--PRKAR1A    KSR1--TRB@    KSR1--TUBGCP6    KSR1--UTP18    MKS1--KSR1    NLK--KSR1   
PLEC--KSR1    R3HCC1L--KSR1    SPECC1--KSR1    SRCIN1--KSR1    TTC40--KSR1   
Fusion : QuiverKSR1
Polymorphisms : SNP and Copy number variants
NCBI Variation ViewerKSR1 [hg38]
Exome Variant ServerKSR1
GNOMAD BrowserENSG00000141068
Varsome BrowserKSR1
Genomic Variants (DGV)KSR1 [DGVbeta]
DECIPHERKSR1 [patients]   [syndromes]   [variants]   [genes]  
CONAN: Copy Number AnalysisKSR1 
ICGC Data PortalKSR1 
TCGA Data PortalKSR1 
Broad Tumor PortalKSR1
OASIS PortalKSR1 [ Somatic mutations - Copy number]
Somatic Mutations in Cancer : COSMICKSR1  [overview]  [genome browser]  [tissue]  [distribution]  
Somatic Mutations in Cancer : COSMIC3DKSR1
Mutations and Diseases : HGMDKSR1
LOVD (Leiden Open Variation Database)Whole genome datasets
LOVD (Leiden Open Variation Database)LOVD - Leiden Open Variation Database
LOVD (Leiden Open Variation Database)LOVD 3.0 shared installation
BioMutasearch KSR1
DgiDB (Drug Gene Interaction Database)KSR1
DoCM (Curated mutations)KSR1 (select the gene name)
CIViC (Clinical Interpretations of Variants in Cancer)KSR1 (select a term)
NCG6 (London) select KSR1
Cancer3DKSR1(select the gene name)
Impact of mutations[PolyPhen2] [Provean] [Buck Institute : MutDB] [Mutation Assessor] [Mutanalyser]
Genetic Testing Registry KSR1
NextProtQ8IVT5 [Medical]
Target ValidationKSR1
Huge Navigator KSR1 [HugePedia]
Clinical trials, drugs, therapy
Protein Interactions : CTD
Pharm GKB GenePA30254
Clinical trialKSR1
canSAR (ICR)KSR1 (select the gene name)
DataMed IndexKSR1
PubMed70 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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indexed on : Fri Feb 19 17:53:39 CET 2021

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