Contents 1 Introduction 2 Function 3 Homologues 4 Regulation 5 Role in cancer 6 Mutant phenotype 7 See also 8 Notes 9 References 10 External links


Introduction[edit] The regulation of cell size is critical to ensure functionality of a cell. Besides environmental factors such as nutrients, growth factors and functional load, cell size is also controlled by a cellular cell size checkpoint. Wee1 is a component of this checkpoint. It is a kinase determining the timepoint of entry into mitosis, thus influencing the size of the daughter cells. Loss of Wee1 function will produce smaller than normal daughter cell, because cell division occurs prematurely. Its name is derived from the Scottish dialect word wee, meaning small - its discoverer Paul Nurse was working at the University of Edinburgh in Scotland at the time of discovery.[1][2]


Function[edit] Fig. 1 Role and regulation of Wee1 Wee1 inhibits Cdk1 by phosphorylating it on two different sites, Tyr15 and Thr14.[3] Cdk1 is crucial for the cyclin-dependent passage of the various cell cycle checkpoints. At least three checkpoints exist for which the inhibition of Cdk1 by Wee1 is important: G2/M checkpoint: Wee1 phosphorylates the amino acids Tyr15 and Thr14 of Cdk1, which keeps the kinase activity of Cdk1 low and prevents entry into mitosis; in S. pombe further cell growth can occur. Wee1 mediated inactivation of Cdk1 has been shown to be ultrasensitive as a result of substrate competition.[4] During mitotic entry the activity of Wee1 is decreased by several regulators and thus Cdk1 activity is increased. In S. pombe, Pom1, a protein kinase, localizes to the cell poles. This activates a pathway in which Cdr2 inhibits Wee1 through Cdr1. Cdk1 itself negatively regulates Wee1 by phosphorylation, which leads to a positive feedback loop. The decreased Wee1 activity alone is not sufficient for mitotic entry: Synthesis of cyclins and an activating phosphorylation by a Cdk activating kinase (CAK) are also required.[5] Cell size checkpoint: There is evidence for the existence of a cell size checkpoint, which prevents small cells from entering mitosis. Wee1 plays a role in this checkpoint by coordinating cell size and cell cycle progression.[6] DNA damage checkpoint: This checkpoint also controls the G2/M transition. In S. pombe this checkpoint delays the mitosis entry of cells with DNA damage (for example induced by gamma radiation). The lengthening of the G2 phase depends on Wee1; wee1 mutants have no prolonged G2 phase after gamma irradiation.[7] Epigenetic function of Wee1 kinase has also been reported. Wee1 was shown to phosphorylate histone H2B at tyrosine 37 residue which regulated global expression of histones.[8] [9]


Homologues[edit] human WEE1 homolog (S. pombe) Identifiers Symbol WEE1 Entrez 7465 HUGO 12761 OMIM 193525 RefSeq NM_003390 UniProt P30291 Other data Locus Chr. 11 p15.3-15.1 human WEE1 homolog 2 (S. pombe) Identifiers Symbol WEE2 Entrez 494551 HUGO 19684 RefSeq NM_001105558 UniProt P0C1S8 Other data Locus Chr. 7 q32-q32 The WEE1 gene has two known homologues in humans, WEE1 (also known as WEE1A) and WEE2 (WEE1B). The corresponding proteins are Wee1-like protein kinase and Wee1-like protein kinase 2 which act on the human Cdk1 homologue Cdk1. The homologue to Wee1 in budding yeast Saccharomyces cerevisiae is called Swe1.


Regulation[edit] In S. pombe, Wee1 is phosphorylated Cdk1 and cyclin B make up the maturation promoting factor (MPF) which promotes the entry into mitosis. It is inactivated by phosphorylation through Wee1 and activated by the phosphatase Cdc25C. Cdc25C in turn is activated by Polo kinase and inactivated by Chk1.[6] Thus in S. pombe Wee1 regulation is mainly under the control of phosphorylation through the polarity kinase, Pom1's, pathway including Cdr2 and Cdr1.[10][11][12][13] At the G2/M transition, Cdk1 is activated by Cdc25 through dephosphorylation of Tyr15. At the same time, Wee1 is inactivated through phosphorylation at its C-terminal catalytic domain by Nim1/Cdr1.[12] Also, the active MPF will promote its own activity by activating Cdc25 and inactivating Wee1, creating a positive feedback loop, though this is not yet understood in detail.[6] Higher eukaryotes regulate Wee1 via phosphorylation and degradation In higher eukaryotes, Wee1 inactivation occurs both by phosphorylation and degradation.[14] The protein complex[nb 1] SCFβ-TrCP1/2 is an E3 ubiquitin ligase that functions in Wee1A ubiquitination. The M-phase kinases Polo-like kinase (Plk1) and Cdc2 phosphorylate two serine residues in Wee1A which are recognized by SCFβ-TrCP1/2.[15] S. cerevisiae homologue Swe1 In S. cerevisiae, cyclin-dependent kinase Cdc28 (Cdk1 homologue) is phosphorylated by Swe1 (Wee1 homologue) and dephosphorylated by Mih1 (Cdc25 homologue). Nim1/Cdr1 homologue in S. cerevisiae, Hsl1, together with its related kinases Gin4 and Kcc4 localize Swe1 to the bud-neck. Bud-neck associating kinases Cla4 and Cdc5 (polo kinase homologue) phosphorylate Swe1 at different stages of the cell cycle. Swe1 is also phosphorylated by Clb2-Cdc28 which serves as a recognition for further phosphorylation by Cdc5. The S. cerevisiae protein Swe1 is also regulated by degradation. Swe1 is hyperphosphorylated by Clb2-Cdc28 and Cdc5 which may be a signal for ubiquitination and degradation by SCF E3 ubiquitin ligase complex as in higher eukaryotes.[16]


Role in cancer[edit] The mitosis promoting factor MPF also regulates DNA-damage induced apoptosis. Negative regulation of MPF by WEE1 causes aberrant mitosis and thus resistance to DNA-damage induced apoptosis. Kruppel-like factor 2 (KLF2) negatively regulates human WEE1, thus increasing sensitivity to DNA-damage induced apoptosis in cancer cells.[17]


Mutant phenotype[edit] Wee1 acts as a dosage-dependent inhibitor of mitosis.[18] Thus, the amount of Wee1 protein correlates with the size of the cells: The fission yeast mutant wee1, also called wee1−, divides at a significantly smaller cell size than wildtype cells. Since Wee1 inhibits entry into mitosis, its absence will lead to division at a premature stage and sub-normal cell size. Conversely, when Wee1 expression is increased, mitosis is delayed and cells grow to a large size before dividing.


See also[edit] Wee1-like protein kinase Cell cycle


Notes[edit] ^ β-transducin repeat-containing protein 1/2 (β-TrCP1/2) F-box protein-containing SKP1/Cul1/F-box protein complex


References[edit] ^ Nurse P (December 2004). "Wee beasties". Nature. 432 (7017): 557. doi:10.1038/432557a. PMID 15577889.  ^ Nurse P, Thuriaux P (November 1980). "Regulatory genes controlling mitosis in the fission yeast Schizosaccharomyces pombe". Genetics. 96 (3): 627–37. PMC 1214365 . PMID 7262540.  ^ Den Haese GJ, Walworth N, Carr AM, Gould KL (1995). "The Wee1 protein kinase regulates T14 phosphorylation of fission yeast Cdc2". Mol Biol Cell. 6 (4): 371–85. doi:10.1091/mbc.6.4.371. PMC 301198 . PMID 7626804.  ^ Kim, SY; Ferrell JE, Jr (23 March 2007). "Substrate competition as a source of ultrasensitivity in the inactivation of Wee1". Cell. 128 (6): 1133–45. doi:10.1016/j.cell.2007.01.039. PMID 17382882.  ^ Coleman TR, Dunphy WG (1994). "Cdc2 regulatory factors". Current Opinion in Cell Biology. 6 (6): 877–82. doi:10.1016/0955-0674(94)90060-4. PMID 7880537.  ^ a b c Kellogg DR (2003). "Wee1-dependent mechanisms required for coordination of cell growth and cell division". J Cell Sci. 116 (24): 4883–90. doi:10.1242/jcs.00908. PMID 14625382.  ^ Rowley R, Hudson J, Young PG (1992). "The wee1 protein kinase is required for radiation-induced mitotic delay". Nature. 356 (6367): 353–5. doi:10.1038/356353a0. PMID 1549179.  ^ Mahajan K, Fang B, Koomen JM, Mahajan NP (2012). "H2B Tyr37 phosphorylation suppresses expression of replication-dependent core histone genes". Nature Structural & Molecular Biology. 19 (9): 930–7. doi:10.1038/nsmb.2356. PMC 4533924 . PMID 22885324.  ^ Mahajan K, Mahajan NP (2013). "WEE1 tyrosine kinase, a novel epigenetic modifier". Trends Genet. 29 (7): 394–402. doi:10.1016/j.tig.2013.02.003. PMC 3700603 . PMID 23537585.  ^ Boddy MN, Furnari B, Mondesert O, Russell P (May 1998). "Replication checkpoint enforced by kinases Cds1 and Chk1". Science. 280 (5365): 909–12. doi:10.1126/science.280.5365.909. PMID 9572736.  ^ Wu L, Russell P (June 1993). "Nim1 kinase promotes mitosis by inactivating Wee1 tyrosine kinase". Nature. 363 (6431): 738–41. doi:10.1038/363738a0. PMID 8515818.  ^ a b Coleman TR, Tang Z, Dunphy WG (March 1993). "Negative regulation of the wee1 protein kinase by direct action of the nim1/cdr1 mitotic inducer". Cell. 72 (6): 919–29. doi:10.1016/0092-8674(93)90580-J. PMID 7681363.  ^ Tang Z, Coleman TR, Dunphy WG (September 1993). "Two distinct mechanisms for negative regulation of the Wee1 protein kinase". EMBO J. 12 (9): 3427–36. PMC 413619 . PMID 7504624.  ^ Watanabe N, Broome M, Hunter T (May 1995). "Regulation of the human WEE1Hu CDK tyrosine 15-kinase during the cell cycle". EMBO J. 14 (9): 1878–91. PMC 398287 . PMID 7743995.  ^ Watanabe N, Arai H, Nishihara Y, et al. (March 2004). "M-phase kinases induce phospho-dependent ubiquitination of somatic Wee1 by SCFbeta-TrCP". Proc. Natl. Acad. Sci. U.S.A. 101 (13): 4419–24. doi:10.1073/pnas.0307700101. PMC 384762 . PMID 15070733.  ^ Lee KS, Asano S, Park JE, Sakchaisri K, Erikson RL (October 2005). "Monitoring the cell cycle by multi-kinase-dependent regulation of Swe1/Wee1 in budding yeast". Cell Cycle. 4 (10): 1346–9. doi:10.4161/cc.4.10.2049. PMID 16123596.  ^ Wang F, Zhu Y, Huang Y, et al. (June 2005). "Transcriptional repression of WEE1 by Kruppel-like factor 2 is involved in DNA damage-induced apoptosis". Oncogene. 24 (24): 3875–85. doi:10.1038/sj.onc.1208546. PMID 15735666.  ^ Russell P, Nurse P (May 1987). "Negative regulation of mitosis by wee1+, a gene encoding a protein kinase homolog". Cell. 49 (4): 559–67. doi:10.1016/0092-8674(87)90458-2. PMID 3032459. 


External links[edit] Drosophila wee - The Interactive Fly v t e Cell cycle proteins Cyclin A (A1, A2) B (B1, B2, B3) D (D1, D2, D3) E (E1, E2) CDK 1 2 3 4 5 6 7 8 9 10 CDK-activating kinase CDK inhibitor INK4a/ARF (p14arf/p16, p15, p18, p19) cip/kip (p21, p27, p57) P53 p63 p73 family p53 p63 p73 Other Cdc2 Cdc25 Cdc42 Cellular apoptosis susceptibility protein E2F Maturation promoting factor Wee Cullin (CUL7) Phases and checkpoints Interphase G1 phase S phase G2 phase M phase Mitosis (Preprophase Prophase Prometaphase Metaphase Anaphase Telophase) Cytokinesis Cell cycle checkpoints Restriction point Spindle checkpoint Postreplication checkpoint Other cellular phases Apoptosis G0 phase Meiosis v t e Kinases: Serine/threonine-specific protein kinases (EC 2.7.11-12) Serine/threonine-specific protein kinases (EC 2.7.11.1-EC 2.7.11.20) Non-specific serine/threonine protein kinases (EC 2.7.11.1) LATS1 LATS2 MAST1 MAST2 STK38 STK38L CIT ROCK1 SGK SGK2 SGK3 Protein kinase B AKT1 AKT2 AKT3 Ataxia telangiectasia mutated Mammalian target of rapamycin EIF-2 kinases PKR HRI EIF2AK3 EIF2AK4 Wee1 WEE1 Pyruvate dehydrogenase kinase (EC 2.7.11.2) PDK1 PDK2 PDK3 Dephospho-(reductase kinase) kinase (EC 2.7.11.3) AMP-activated protein kinase α PRKAA1 PRKAA2 β PRKAB1 PRKAB2 γ PRKAG1 PRKAG2 PRKAG3 3-methyl-2-oxobutanoate dehydrogenase (acetyl-transferring) kinase (EC 2.7.11.4) BCKDK BCKDHA BCKDHB (isocitrate dehydrogenase (NADP+)) kinase (EC 2.7.11.5) IDH2 IDH3A IDH3B IDH3G (tyrosine 3-monooxygenase) kinase (EC 2.7.11.6) STK4 Myosin-heavy-chain kinase (EC 2.7.11.7) Aurora kinase Aurora A kinase Aurora B kinase Fas-activated serine/threonine kinase (EC 2.7.11.8) FASTK STK10 Goodpasture-antigen-binding protein kinase (EC 2.7.11.9) - IκB kinase (EC 2.7.11.10) CHUK IKK2 TBK1 IKBKE IKBKG IKBKAP cAMP-dependent protein kinase (EC 2.7.11.11) Protein kinase A PRKACG PRKACB PRKACA PRKY cGMP-dependent protein kinase (EC 2.7.11.12) Protein kinase G PRKG1 Protein kinase C (EC 2.7.11.13) Protein kinase C Protein kinase Cζ PKC alpha PRKCB1 PRKCD PRKCE PRKCH PRKCG PRKCI PRKCQ Protein kinase N1 PKN2 PKN3 Rhodopsin kinase (EC 2.7.11.14) Rhodopsin kinase Beta adrenergic receptor kinase (EC 2.7.11.15) Beta adrenergic receptor kinase Beta adrenergic receptor kinase-2 G-protein coupled receptor kinases (EC 2.7.11.16) GRK4 GRK5 GRK6 Ca2+/calmodulin-dependent (EC 2.7.11.17) BRSK2 CAMK1 CAMK2A CAMK2B CAMK2D CAMK2G CAMK4 MLCK CASK CHEK1 CHEK2 DAPK1 DAPK2 DAPK3 STK11 MAPKAPK2 MAPKAPK3 MAPKAPK5 MARK1 MARK2 MARK3 MARK4 MELK MKNK1 MKNK2 NUAK1 NUAK2 OBSCN PASK PHKG1 PHKG2 PIM1 PIM2 PKD1 PRKD2 PRKD3 PSKH1 SNF1LK2 KIAA0999 STK40 SNF1LK SNRK SPEG TSSK2 Kalirin TRIB1 TRIB2 TRIB3 TRIO Titin DCLK1 Myosin light-chain kinase (EC 2.7.11.18) MYLK MYLK2 MYLK3 MYLK4 Phosphorylase kinase (EC 2.7.11.19) PHKA1 PHKA2 PHKB PHKG1 PHKG2 Elongation factor 2 kinase (EC 2.7.11.20) EEF2K STK19 Polo kinase (EC 2.7.11.21) PLK1 PLK2 PLK3 PLK4 Serine/threonine-specific protein kinases (EC 2.7.11.21-EC 2.7.11.30) Polo kinase (EC 2.7.11.21) PLK1 PLK2 PLK3 PLK4 Cyclin-dependent kinase (EC 2.7.11.22) CDK1 CDK2 CDKL2 CDK3 CDK4 CDK5 CDKL5 CDK6 CDK7 CDK8 CDK9 CDK10 CDK12 CDC2L5 PCTK1 PCTK2 PCTK3 PFTK1 CDC2L1 (RNA-polymerase)-subunit kinase (EC 2.7.11.23) RPS6KA5 RPS6KA4 P70S6 kinase P70-S6 Kinase 1 RPS6KB2 RPS6KA2 RPS6KA3 RPS6KA1 RPS6KC1 Mitogen-activated protein kinase (EC 2.7.11.24) Extracellular signal-regulated MAPK1 MAPK3 MAPK4 MAPK6 MAPK7 MAPK12 MAPK15 C-Jun N-terminal MAPK8 MAPK9 MAPK10 P38 mitogen-activated protein MAPK11 MAPK13 MAPK14 MAP3K (EC 2.7.11.25) MAP kinase kinase kinases MAP3K1 MAP3K2 MAP3K3 MAP3K4 MAP3K5 MAP3K6 MAP3K7 MAP3K8 RAFs ARAF BRAF KSR1 KSR2 MLKs MAP3K12 MAP3K13 MAP3K9 MAP3K10 MAP3K11 MAP3K7 ZAK CDC7 MAP3K14 Tau-protein kinase (EC 2.7.11.26) TPK1 TTK GSK-3 (acetyl-CoA carboxylase) kinase (EC 2.7.11.27) - Tropomyosin kinase (EC 2.7.11.28) - Low-density-lipoprotein receptor kinase (EC 2.7.11.29) - Receptor protein serine/threonine kinase (EC 2.7.11.30) Bone morphogenetic protein receptors BMPR1 BMPR1A BMPR1B BMPR2 ACVR1 ACVR1B ACVR1C ACVR2A ACVR2B ACVRL1 Anti-Müllerian hormone receptor Dual-specificity kinases (EC 2.7.12) MAP2K MAP2K1 MAP2K2 MAP2K3 MAP2K4 MAP2K5 MAP2K6 MAP2K7 v t e Enzymes Activity Active site Binding site Catalytic triad Oxyanion hole Enzyme promiscuity Catalytically perfect enzyme Coenzyme Cofactor Enzyme catalysis Regulation Allosteric regulation Cooperativity Enzyme inhibitor Classification EC number Enzyme superfamily Enzyme family List of enzymes Kinetics Enzyme kinetics Eadie–Hofstee diagram Hanes–Woolf plot Lineweaver–Burk plot Michaelis–Menten kinetics Types EC1 Oxidoreductases (list) EC2 Transferases (list) EC3 Hydrolases (list) EC4 Lyases (list) EC5 Isomerases (list) EC6 Ligases (list) Molecular and Cellular Biology portal Retrieved from "https://en.wikipedia.org/w/index.php?title=Wee1&oldid=778232248" Categories: Genes on human chromosome 11Genes on human chromosome 7Cell cycleFungal proteinsEC 2.7.11Hidden categories: Protein pages needing a picture


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EntrezSwiss-ProtEnzyme Commission NumberProtein KinaseSerine/threonine-specific Protein KinaseSchizosaccharomyces PombeMolecular MassAtomic Mass UnitCell CycleMitosisCdk1Homology (biology)Cell SizeKinaseScottish EnglishPaul NurseEnlargeCdk1Cell Cycle CheckpointMitosisSchizosaccharomyces PombeUltrasensitivityPom1CyclinGamma RayWee1-like Protein KinaseWee1-like Protein KinaseEntrezHuman Genome OrganisationOMIMNational Center For Biotechnology InformationSwiss-ProtLocus (genetics)Chromosome 11 (human)EntrezHuman Genome OrganisationNational Center For Biotechnology InformationSwiss-ProtLocus (genetics)Chromosome 7 (human)GeneWee1-like Protein KinaseCdk1Saccharomyces CerevisiaeCdk1Cyclin BMaturation Promoting FactorCdc25CPolo KinaseCHEK1PhosphorylationPom1C-terminusPositive FeedbackEukaryoteProteolysisSCF ComplexUbiquitin LigaseWee1-like Protein KinaseMitosisPolo-like KinaseS. CerevisiaeCDC28 Protein KinaseBuddingSCF ComplexDNA RepairApoptosisKruppel-like FactorsGene DosageMutantWee1-like Protein KinaseCell CycleBTRC (gene)Digital Object IdentifierPubMed IdentifierPubMed CentralPubMed IdentifierDigital Object IdentifierPubMed CentralPubMed IdentifierDigital Object IdentifierPubMed IdentifierDigital Object IdentifierPubMed IdentifierDigital Object IdentifierPubMed IdentifierDigital Object IdentifierPubMed IdentifierDigital Object IdentifierPubMed CentralPubMed IdentifierDigital Object IdentifierPubMed CentralPubMed IdentifierDigital Object IdentifierPubMed IdentifierDigital Object IdentifierPubMed IdentifierDigital Object IdentifierPubMed IdentifierPubMed CentralPubMed IdentifierPubMed CentralPubMed IdentifierDigital Object IdentifierPubMed CentralPubMed IdentifierDigital Object IdentifierPubMed IdentifierDigital Object IdentifierPubMed IdentifierDigital Object IdentifierPubMed IdentifierTemplate:Cell Cycle ProteinsTemplate Talk:Cell Cycle ProteinsCell CycleProteinCyclinCyclin ACyclin A1Cyclin A2Cyclin BCyclin B1Cyclin B2Cyclin DCyclin D1Cyclin D2Cyclin D3Cyclin ECyclin E1Cyclin E2Cyclin-dependent KinaseCyclin-dependent Kinase 1Cyclin-dependent Kinase 2Cyclin-dependent Kinase 3Cyclin-dependent Kinase 4Cyclin-dependent Kinase 5Cyclin-dependent Kinase 6Cyclin-dependent Kinase 7Cyclin-dependent Kinase 8Cyclin-dependent Kinase 9Cyclin-dependent Kinase 10CDK-activating KinaseCyclin-dependent Kinase Inhibitor ProteinCell CycleP14arfP16CDKN2BCDKN2CCDKN2DCell CycleP21CDKN1BCyclin-dependent Kinase Inhibitor 1CP53 P63 P73 FamilyP53TP63P73Cdk1Cdc25CDC42Cellular Apoptosis Susceptibility ProteinE2FMaturation Promoting FactorCullinCUL7InterphaseG1 PhaseS PhaseG2 PhaseCell DivisionMitosisPreprophaseProphasePrometaphaseMetaphaseAnaphaseTelophaseCytokinesisCell Cycle CheckpointRestriction PointSpindle CheckpointPostreplication CheckpointApoptosisG0 PhaseMeiosisTemplate:Serine/threonine-specific Protein KinasesTemplate Talk:Serine/threonine-specific Protein KinasesKinaseSerine/threonine-specific Protein KinaseEnzyme Commission NumberSerine/threonine-specific Protein KinaseNon-specific Serine/threonine Protein KinaseLATS1LATS2MAST1MAST2STK38STK38LCIT (gene)ROCK1SGKSGK2SGK3AKTAKT1AKT2AKT3Ataxia Telangiectasia MutatedMammalian Target Of RapamycinEIF-2 KinaseProtein Kinase REIF2AK1EIF2AK3EIF2AK4Wee1-like Protein KinasePyruvate Dehydrogenase KinasePyruvate Dehydrogenase Lipoamide Kinase Isozyme 1PDK2PDK3Dephospho-(reductase Kinase) KinaseAMP-activated Protein KinaseProtein Kinase, AMP-activated, Alpha 1PRKAA2PRKAB1PRKAB2PRKAG1PRKAG2PRKAG33-methyl-2-oxobutanoate Dehydrogenase (acetyl-transferring) KinaseBCKDKBCKDHABCKDHB(isocitrate Dehydrogenase (NADP+)) KinaseIDH2IDH3AIDH3BIDH3G(tyrosine 3-monooxygenase) KinaseSTK4Myosin-heavy-chain KinaseAurora KinaseAurora A KinaseAurora B KinaseFas-activated Serine/threonine KinaseFASTKSTK10Goodpasture-antigen-binding Protein KinaseIκB KinaseCHUKIKK2TANK-binding Kinase 1IKBKEIKBKGIKBKAPProtein Kinase AProtein Kinase APRKACGPRKACBPRKACAPRKYCGMP-dependent Protein KinaseCGMP-dependent Protein KinasePRKG1Protein Kinase CProtein Kinase CProtein Kinase C Zeta TypePKC AlphaPRKCB1PRKCDPRKCEPRKCHPRKCGPRKCIPRKCQProtein Kinase N1PKN2PKN3 (gene)Rhodopsin KinaseRhodopsin KinaseBeta Adrenergic Receptor KinaseBeta Adrenergic Receptor KinaseBeta Adrenergic Receptor Kinase-2G Protein-coupled Receptor KinaseGRK4GRK5GRK6Ca2+/calmodulin-dependent Protein KinaseBRSK2CAMK1CAMK2ACAMK2BCAMK2DCAMK2GCAMK4Myosin Light-chain KinaseCASKCHEK1CHEK2DAPK1DAPK2DAPK3STK11MAPKAPK2MAPKAPK3MAPKAPK5MARK1MARK2MARK3MARK4MELKMKNK1MKNK2NUAK1NUAK2OBSCNPASKPHKG1PHKG2PIM1PIM2 (gene)Protein Kinase D1PRKD2PRKD3PSKH1SNF1LK2KIAA0999STK40SNF1LKSNRKSPEGTSSK2KalirinTRIB1TRIB2TRIB3TRIO (gene)TitinDCLK1Myosin Light-chain KinaseMYLKMYLK2MYLK3MYLK4Phosphorylase KinasePhosphorylase Kinase, Alpha 1PHKA2PHKBPHKG1PHKG2Elongation Factor 2 KinaseEEF2KSTK19Polo KinasePLK1PLK2PLK3PLK4Serine/threonine-specific Protein KinasePolo KinasePLK1PLK2PLK3PLK4Cyclin-dependent KinaseCdk1Cyclin-dependent Kinase 2CDKL2Cyclin-dependent Kinase 3Cyclin-dependent Kinase 4Cyclin-dependent Kinase 5CDKL5Cyclin-dependent Kinase 6Cyclin-dependent Kinase 7Cyclin-dependent Kinase 8CDK9Cyclin-dependent Kinase 10CDK12CDC2L5PCTK1PCTK2PCTK3PFTK1CDC2L1(RNA-polymerase)-subunit KinaseRPS6KA5RPS6KA4P70S6 KinaseP70-S6 Kinase 1RPS6KB2RPS6KA2RPS6KA3RPS6KA1RPS6KC1Mitogen-activated Protein KinaseExtracellular Signal-regulated KinasesMAPK1MAPK3MAPK4MAPK6MAPK7MAPK12MAPK15C-Jun N-terminal KinasesMAPK8Mitogen-activated Protein Kinase 9MAPK10P38 Mitogen-activated Protein KinasesMAPK11MAPK13MAPK14MAP Kinase Kinase KinaseMAP Kinase Kinase KinaseMAP3K1MAP3K2MAP3K3MAP3K4ASK1MAP3K7MAP3K8C-RafARAFBRAF (gene)KSR1KSR2MAP3K12MAP3K13MAP3K9MAP3K10MAP3K11MAP3K7ZAKCell Division Cycle 7-related Protein KinaseMAP3K14Tau-protein KinaseTPK1TTK (gene)GSK-3(acetyl-CoA Carboxylase) KinaseTropomyosin KinaseLow-density-lipoprotein Receptor KinaseReceptor Protein Serine/threonine KinaseBone Morphogenetic Protein ReceptorsBone Morphogenetic Protein Receptor, Type 1BMPR1ABMPR1BBMPR2ACVR1ACVR1BACVR1CACVR2AACVR2BACVRL1Anti-Müllerian Hormone ReceptorDual-specificity KinaseMitogen-activated Protein Kinase KinaseMAP2K1MAP2K2MAP2K3MAP2K4MAP2K5MAP2K6MAP2K7Template:EnzymesTemplate Talk:EnzymesEnzymeActive SiteBinding SiteCatalytic TriadOxyanion HoleEnzyme PromiscuityCatalytically Perfect EnzymeCoenzymeCofactor (biochemistry)Enzyme CatalysisAllosteric RegulationCooperativityEnzyme InhibitorEnzyme Commission NumberEnzyme SuperfamilyEnzyme FamilyList Of EnzymesEnzyme KineticsEadie–Hofstee DiagramHanes–Woolf PlotLineweaver–Burk PlotMichaelis–Menten KineticsOxidoreductaseList Of EC Numbers (EC 1)TransferaseList Of EC Numbers (EC 2)HydrolaseList Of EC Numbers (EC 3)LyaseList Of EC Numbers (EC 4)IsomeraseList Of EC Numbers (EC 5)LigaseList Of EC Numbers (EC 6)Portal:Molecular And Cellular BiologyHelp:CategoryCategory:Genes On Human Chromosome 11Category:Genes 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