Contents 1 Types 2 CDKs and cyclins in the cell cycle 3 Regulation of activity 3.1 Cyclin binding 3.2 Phosphorylation 3.3 CDK inhibitors 3.4 Suk1 or Cks 3.5 Non-cyclin activators 3.5.1 Viral cyclins 3.5.2 CDK5 activators 3.5.3 RINGO/Speedy 4 History 5 Medical significance 6 References 7 External links

Types[edit] Table 1: Known CDKs, their cyclin partners, and their functions in the human[3] and consequences of deletion in mice[4] CDK Cyclin partner Function Deletion Phenotype in Mice Cdk1 Cyclin B M phase None Cdk2 Cyclin E G1/S transition Reduced size, imparted neural progenitor cell proliferation. Viable, but both males & females sterile. Cdk2 Cyclin A S phase, G2 phase Cdk3 Cyclin C G1 phase ? No defects. Viable, fertile. Cdk4 Cyclin D G1 phase Reduced size, insulin deficient diabetes. Viable, but both male & female infertile.

CDKs and cyclins in the cell cycle[edit] Most of the known cyclin-CDK complexes regulate the progression through the cell cycle. Animal cells contain at least nine CDKs, four of which, CDK1, 2, 3, and 4, are directly involved in cell cycle regulation.[1] In mammalian cells, CDK1, with its partners cyclin A2 and B1, alone can drive the cell cycle.[4] Another one, CDK7, is involved indirectly as the CDK-activating kinase.[1] Cyclin-CDK complexes phosphorylate substrates appropriate for the particular cell cycle phase.[3] Cyclin-CDK complexes of earlier cell-cycle phase help activate cyclin-CDK complexes in later phase.[1] Table 2: Cyclins and CDKs by Cell-Cycle Phase Phase Cyclin CDK G0 C Cdk3 G1 D, E Cdk4, Cdk2, Cdk6 S A, E Cdk2 G2 A Cdk2, Cdk1 M B Cdk1 Table 3: Cyclin-dependent kinases that control the cell cycle in model organisms[1] Species Name Original name Size (amino acids) Function Saccharomyces cerevisiae Cdk1 Cdc28 298 All cell-cycle stages Schizosaccharomyces pombe Cdk1 Cdc2 297 All cell-cycle stages Drosophila melanogaster Cdk1 Cdc2 297 M Cdk2 Cdc2c 314 G1/S, S, possibly M Cdk4 Cdk4/6 317 G1, promotes growth Xenopus laevis Cdk1 Cdc2 301 M Cdk2 297 S, possibly M Homo sapiens Cdk1 Cdc2 297 M Cdk2 298 G1, S, possibly M Cdk4 301 G1 Cdk6 326 G1 A list of CDKs with their regulator protein, cyclin or other: CDK1; cyclin A, cyclin B CDK2; cyclin A, cyclin E CDK3; cyclin C CDK4; cyclin D1, cyclin D2, cyclin D3 CDK5; CDK5R1, CDK5R2. See also CDKL5. CDK6; cyclin D1, cyclin D2, cyclin D3 CDK7; cyclin H CDK8; cyclin C CDK9; cyclin T1, cyclin T2a, cyclin T2b, cyclin K CDK10 CDK11 (CDC2L2) ; cyclin L CDK12; cyclin L CDK13 (CDC2L5) ; cyclin L

Regulation of activity[edit] CDK levels remain relatively constant throughout the cell cycle and most regulation is post-translational. Most knowledge of CDK structure and function is based on CDKs of S. pombe (Cdc2), S. cerevisiae (CDC28), and vertebrates (CDC2 and CDK2). The four major mechanisms of CDK regulation are cyclin binding, CAK phosphorylation, regulatory inhibitory phosphorylation, and binding of CDK inhibitory subunits (CKIs).[5] Cyclin binding[edit] The active site, or ATP-binding site, of all kinases is a cleft between a small amino-terminal lobe and a larger carboxy-terminal lobe.[1] The structure of human Cdk2 revealed that CDKs have a modified ATP-binding site that can be regulated by cyclin binding.[1] Phosphorylation by CDK-activating kinase (CAK) at Thr 161 on the T-loop increases the complex activity. Without cyclin, a flexible loop called the activation loop or T-loop blocks the cleft, and the position of several key amino acid residues is not optimal for ATP-binding.[1] With cyclin, two alpha helices change position to permit ATP binding. One of them, the L12 helix that comes just before the T-loop in the primary sequence, becomes a beta strand and helps rearrange the T-loop, so it no longer blocks the active site.[1] The other alpha helix called the PSTAIRE helix rearranges and helps change the position of the key amino acid residues in the active site.[1] There is considerable specificity in which cyclin binds with CDK.[3] Furthermore, cyclin binding determines the specificity of the cyclin-CDK complex for particular substrates.[3] Cyclins can directly bind the substrate or localize the CDK to a subcellular area where the substrate is found. Substrate specificity of S cyclins is imparted by the hydrophobic batch (centered on the MRAIL sequence), which has affinity for substrate proteins that contain a hydrophobic RXL (or Cy) motif. Cyclin B1 and B2 can localize Cdk1 to the nucleus and the Golgi, respectively, through a localization sequence outside the CDK-binding region.[1] Phosphorylation[edit] Full kinase activity requires an activating phosphorylation on a threonine adjacent to the active site.[1] The identity of the CDK-activating kinase (CAK) that performs this phosphorylation varies across the model organisms.[1] The timing of this phosphorylation varies as well. In mammalian cells, the activating phosphorylation occurs after cyclin binding.[1] In yeast cells, it occurs before cyclin binding.[1] CAK activity is not regulated by known cell-cycle pathways and cyclin binding is the limiting step for CDK activation.[1] Unlike activating phosphorylation, CDK inhibitory phosphorylation is vital for regulation of the cell cycle. Various kinases and phosphatases regulate their phosphorylation state. One of the kinases that place the tyrosine phosphate is Wee1, a kinase conserved in all eukaryotes.[1] Fission yeast also contains a second kinase Mik1 that can phosphorylate the tyrosine.[1] Vertebrates contain a different second kinase called Myt1 that is related to Wee1 but can phosphorylate both the threonine and the tyrosine.[1] Phosphatases from the Cdc25 family dephosphorylate both the threonine and the tyrosine.[1] CDK inhibitors[edit] A cyclin-dependent kinase inhibitor (CKI) is a protein that interacts with a cyclin-CDK complex to block kinase activity, usually during G1 or in response to signals from the environment or from damaged DNA.[1] In animal cells, there are two major CKI families: the INK4 family and the CIP/KIP family.[1] The INK4 family proteins are strictly inhibitory and bind CDK monomers. Crystal structures of CDK6-INK4 complexes show that INK4 binding twists the CDK to distort cyclin binding and kinase activity. The CIP/KIP family proteins bind both the cyclin and the CDK of a complex and can be inhibitory or activating. CIP/KIP family proteins activate cyclin D and CDK4 or CDK6 complexes by enhancing complex formation.[1] In yeast and Drosophila, CKIs are strong inhibitors of S- and M-CDK, but do not inhibit G1/S-CDKs. During G1, high levels of CKIs prevent cell cycle events from occurring out of order, but do not prevent transition through the Start checkpoint, which is initiated through G1/S-CDKs. Once the cell cycle is initiated, phosphorylation by early G1/S-CDKs leads to destruction of CKIs, relieving inhibition on later cell cycle transitions. In mammalian cells, the CKI regulation works differently. Mammalian protein p27 (Dacapo in Drosophila) inhibits G1/S- and S-CDKs, but does not inhibit S- and M-CDKs.[1] Suk1 or Cks[edit] The CDKs directly involved in the regulation of the cell cycle associate with small, 9- to 13-kiloDalton proteins called Suk1 or Cks.[3] These proteins are required for CDK function, but their precise role is unknown.[3] Cks1 binds the carboxy lobe of the CDK, and recognizes phosphorylated residues. It may help the cyclin-CDK complex with substrates that have multiple phosphorylation sites by increasing affinity for the substrate.[3] Non-cyclin activators[edit] Viral cyclins[edit] Viruses can encode proteins with sequence homology to cyclins. One much-studied example is K-cyclin (or v-cyclin) from Kaposi sarcoma herpes virus (see Kaposi’s sarcoma), which activates CDK6. Viral cyclin-CDK complexes have different substrate specificities and regulation sensitivities.[6] CDK5 activators[edit] The proteins p35 and p39 activate CDK5. Although they lack cyclin sequence homology, crystal structures show that p35 folds in a similar way as the cyclins. However, activation of CDK5 does not require activation loop phosphorylation.[6] RINGO/Speedy[edit] Proteins with no homology to the cyclin family can be direct activators of CDKs.[7] One family of such activators is the RINGO/Speedy family,[7] which was originally discovered in Xenopus. All five members discovered so far directly activate Cdk1 and Cdk2, but the RINGO/Speedy-CDK2 complex recognizes different substrates than cyclin A-CDK2 complex.[6]

History[edit] Leland H. Hartwell, J. Hoonhorst, R. Timothy Hunt, and Paul M. Nurse received the 2001 Nobel Prize in Physiology or Medicine for their complete description of cyclin and cyclin-dependent kinase mechanisms, which are central to the regulation of the cell cycle.

Medical significance[edit] Main article: CDK inhibitor CDKs are considered a potential target for anti-cancer medication. If it is possible to selectively interrupt the cell cycle regulation in cancer cells by interfering with CDK action, the cell will die. At present, some CDK inhibitors such as seliciclib are undergoing clinical trials. Although it was originally developed as a potential anti-cancer drug, seliciclib has also proven to induce apoptosis in neutrophil granulocytes, which mediate inflammation.[8] This means that novel drugs for treatment of chronic inflammation diseases such as arthritis and cystic fibrosis could be developed. Flavopiridol (alvocidib) is the first CDK inhibitor to be tested in clinical trials after being identified in an anti-cancer agent screen in 1992. It competes for the ATP site of the CDKs.[9] Palbociclib and abemaciclib have been approved for the management of hormone receptor (estrogen receptor/progestogen receptor) expressing metastatic breast cancer in combination with endocrine therapy. [10][11] More research is required, however, because disruption of the CDK-mediated pathway has potentially serious consequences; while CDK inhibitors seem promising, it has to be determined how side-effects can be limited so that only target cells are affected. As such diseases are currently treated with glucocorticoids, which have often serious side-effects, even a minor success would be an improvement.[12] Complications of developing a CDK drug include the fact that many CDKs are not involved in the cell cycle, but other processes such as transcription, neural physiology, and glucose homeostasis.[13] Table 4: Cyclin-dependent kinase inhibitor drugs[13] Drug CDKs Inhibited Flavopiridol (alvocidib) 1, 2, 4, 6, 7, 9 Olomoucine 1, 2, 5 Roscovitine 1, 2, 5 Purvalanol 1, 2, 5 Paullones 1, 2, 5 Butryolactone 1, 2, 5 Palbociclib 4, 6 Thio/oxoflavopiridols 1 Oxindoles 2 Aminothiazoles 4 Benzocarbazoles 4 Pyrimidines 4 Seliciclib ?

References[edit] ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab Morgan, David O. (2007). The Cell Cycle: Principles of Control. London: New Science Press, 1st ed. ^ Lee, M. G.; Nurse, P (1987). "Complementation used to clone a human homologue of the fission yeast cell cycle control gene cdc2". Nature. 327 (6117): 31–5. doi:10.1038/327031a0. PMID 3553962.  ^ a b c d e f g Morgan, D. O. (1997). "Cyclin-dependent kinases: Engines, clocks, and microprocessors". Annual Review of Cell and Developmental Biology. 13: 261–91. doi:10.1146/annurev.cellbio.13.1.261. PMID 9442875.  ^ a b Satyanarayana, A; Kaldis, P (2009). "Mammalian cell-cycle regulation: Several Cdks, numerous cyclins and diverse compensatory mechanisms". Oncogene. 28 (33): 2925–39. doi:10.1038/onc.2009.170. PMID 19561645.  ^ Morgan, D. O. (1995). "Principles of CDK regulation". Nature. 374 (6518): 131–4. doi:10.1038/374131a0. PMID 7877684.  ^ a b c Nebreda, A. R. (2006). "CDK activation by non-cyclin proteins". Current Opinion in Cell Biology. 18 (2): 192–8. doi:10.1016/ PMID 16488127.  ^ a b Mourón, S; De Cárcer, G; Seco, E; Fernández-Miranda, G; Malumbres, M; Nebreda, A. R. (2010). "RINGO C is required to sustain the spindle-assembly checkpoint". Journal of Cell Science. 123 (Pt 15): 2586–95. doi:10.1242/jcs.059964. PMID 20605920.  ^ Rossi, A. G.; Sawatzky, D. A.; Walker, A; Ward, C; Sheldrake, T. A.; Riley, N. A.; Caldicott, A; Martinez-Losa, M; Walker, T. R.; Duffin, R; Gray, M; Crescenzi, E; Martin, M. C.; Brady, H. J.; Savill, J. S.; Dransfield, I; Haslett, C (2006). "Cyclin-dependent kinase inhibitors enhance the resolution of inflammation by promoting inflammatory cell apoptosis". Nature Medicine. 12 (9): 1056–64. doi:10.1038/nm1468. PMID 16951685.  ^ Senderowicz, A. M. (1999). "Flavopiridol: The first cyclin-dependent kinase inhibitor in human clinical trials". Investigational new drugs. 17 (3): 313–20. PMID 10665481.  ^ "FDA Grants Palbociclib Accelerated Approval for Advanced Breast Cancer". National Cancer Institute. Retrieved 2017-11-30.  ^ Research, Center for Drug Evaluation and. "Approved Drugs - FDA approves abemaciclib for HR-positive, HER2-negative breast cancer". Retrieved 2017-11-30.  ^ "FDA Grants Palbociclib Accelerated Approval for Advanced Breast Cancer". National Cancer Institute. Retrieved 2017-11-30.  ^ a b Sausville, E. A. (2002). "Complexities in the development of cyclin-dependent kinase inhibitor drugs". Trends in Molecular Medicine. 8 (4 Suppl): S32–7. PMID 11927285. 

External links[edit] Cyclin-Dependent Kinases at the US National Library of Medicine Medical Subject Headings (MeSH) EC KEGG – Human Cell Cycle Loyer P, Trembley J, Katona R, Kidd V, Lahti J (2005). "Role of CDK/cyclin complexes in transcription and RNA splicing". Cell Signal. 17 (9): 1033–51. doi:10.1016/j.cellsig.2005.02.005. PMID 15935619.  v t e Intracellular signaling peptides and proteins MAP see MAP kinase pathway Calcium Intracellular calcium-sensing proteins Calcineurin Ca2+/calmodulin-dependent protein kinase G protein Heterotrimeric cAMP: Heterotrimeric G protein Gs/Gi Adenylate cyclase cAMP 3',5'-cyclic-AMP phosphodiesterase Protein kinase A cGMP: Transducin Gustducin Guanylate cyclase cGMP 3',5'-cyclic-GMP phosphodiesterase Protein kinase G G alpha subunit Gα GNAO1 GNAI1 GNAI2 GNAI3 GNAT1 GNAT2 GNAT3 GNAZ GNAS GNAL GNAQ GNA11 GNA12 GNA13 GNA14 GNA15/GNA16 G beta-gamma complex Gβ GNB1 GNB2 GNB3 GNB4 GNB5 Gγ GNGT1 GNGT2 GNG2 GNG3 GNG4 GNG5 GNG7 GNG8 GNG10 GNG11 GNG12 GNG13 BSCL2 G protein-coupled receptor kinase AMP-activated protein kinase Monomeric ARFs Rabs Ras HRAS KRAS NRAS Rhos Arfs Ran Rhebs Raps RGKs Cyclin Cyclin-dependent kinase inhibitor protein Cyclin-dependent kinase Cyclin Lipid Phosphoinositide phospholipase C Phospholipase C Other protein kinase Serine/threonine: Casein kinase 1 2 eIF-2 kinase EIF2AK3 Glycogen synthase kinase GSK1 GSK2 GSK-3 GSK3A GSK3B IκB kinase CHUK IKK2 IKBKG Interleukin-1 receptor associated kinase IRAK1 IRAK2 IRAK3 IRAK4 Lim kinase LIMK1 LIMK2 p21-activated kinases PAK1 PAK2 PAK3 PAK4 Rho-associated protein kinase ROCK1 ROCK2 Ribosomal s6 kinase RPS6KA1 Tyrosine: ZAP70 Focal adhesion protein-tyrosine kinase PTK2 PTK2B BTK both Dual-specificity kinase Other protein phosphatase Serine/threonine: Protein phosphatase 2 Tyrosine: protein tyrosine phosphatase: Receptor-like protein tyrosine phosphatase Sh2 domain-containing protein tyrosine phosphatase both: Dual-specificity phosphatase Apoptosis see apoptosis signaling pathway GTP-binding protein regulators see GTP-binding protein regulators Other Activating transcription factor 6 Signal transducing adaptor protein I-kappa B protein Mucin-4 Olfactory marker protein Phosphatidylethanolamine binding protein EDARADD PRKCSH see also deficiencies of intracellular signaling peptides and proteins 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 Non-specific serine/threonine protein kinases (EC 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 PDK1 PDK2 PDK3 Dephospho-(reductase kinase) kinase (EC AMP-activated protein kinase α PRKAA1 PRKAA2 β PRKAB1 PRKAB2 γ PRKAG1 PRKAG2 PRKAG3 3-methyl-2-oxobutanoate dehydrogenase (acetyl-transferring) kinase (EC BCKDK BCKDHA BCKDHB (isocitrate dehydrogenase (NADP+)) kinase (EC IDH2 IDH3A IDH3B IDH3G (tyrosine 3-monooxygenase) kinase (EC STK4 Myosin-heavy-chain kinase (EC Aurora kinase Aurora A kinase Aurora B kinase Fas-activated serine/threonine kinase (EC FASTK STK10 Goodpasture-antigen-binding protein kinase (EC - IκB kinase (EC CHUK IKK2 TBK1 IKBKE IKBKG IKBKAP cAMP-dependent protein kinase (EC Protein kinase A PRKACG PRKACB PRKACA PRKY cGMP-dependent protein kinase (EC Protein kinase G PRKG1 Protein kinase C (EC Protein kinase C Protein kinase Cζ PKC alpha PRKCB1 PRKCD PRKCE PRKCH PRKCG PRKCI PRKCQ Protein kinase N1 PKN2 PKN3 Rhodopsin kinase (EC Rhodopsin kinase Beta adrenergic receptor kinase (EC Beta adrenergic receptor kinase Beta adrenergic receptor kinase-2 G-protein coupled receptor kinases (EC GRK4 GRK5 GRK6 Ca2+/calmodulin-dependent (EC 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 MYLK MYLK2 MYLK3 MYLK4 Phosphorylase kinase (EC PHKA1 PHKA2 PHKB PHKG1 PHKG2 Elongation factor 2 kinase (EC EEF2K STK19 Polo kinase (EC PLK1 PLK2 PLK3 PLK4 Serine/threonine-specific protein kinases (EC Polo kinase (EC PLK1 PLK2 PLK3 PLK4 Cyclin-dependent kinase (EC CDK1 CDK2 CDKL2 CDK3 CDK4 CDK5 CDKL5 CDK6 CDK7 CDK8 CDK9 CDK10 CDK12 CDC2L5 PCTK1 PCTK2 PCTK3 PFTK1 CDC2L1 (RNA-polymerase)-subunit kinase (EC RPS6KA5 RPS6KA4 P70S6 kinase P70-S6 Kinase 1 RPS6KB2 RPS6KA2 RPS6KA3 RPS6KA1 RPS6KC1 Mitogen-activated protein kinase (EC 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 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 TPK1 TTK GSK-3 (acetyl-CoA carboxylase) kinase (EC - Tropomyosin kinase (EC - Low-density-lipoprotein receptor kinase (EC - Receptor protein serine/threonine kinase (EC 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 "" Categories: Cell cycleProtein familiesEC 2.7.11

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Enzyme Commission NumberIntEnzBRENDAExPASyKEGGMetaCycPRIAM Enzyme Specific ProfilesProtein Data BankPubMed CentralPubMedNational Center For Biotechnology InformationEnlargeInterphaseMitosisG1 PhaseS PhaseG2 PhaseSugarKinasesCell CycleTranscription (genetics)EukaryotesCell CycleCyclinCyclinConsensus SequencePhosphorylationAmino AcidSerineThreonineProlineLysineArginineCyclinCyclin BCyclin ECyclin ACyclin CCyclin DCell CycleSaccharomyces CerevisiaeSchizosaccharomyces PombeDrosophila MelanogasterXenopus LaevisHomo SapiensCDK1Cyclin ACyclin BCyclin-dependent Kinase 2Cyclin ACyclin ECyclin CCDK4Cyclin D1Cyclin D2Cyclin D3CDK5CDK5R1CDK5R2CDKL5CDK6Cyclin D1Cyclin D2Cyclin D3CDK7Cyclin HCyclin CCDK9Cyclin T1Cyclin KCDK12CDK-activating KinaseActive SiteActivation LoopKinasePhosphorylationThreonineActive SiteWee1Fission YeastWee1Cdc25Cyclin-dependent Kinase Inhibitor ProteinInk4CIP/KIPCKS1BKaposi’s SarcomaLeland H. HartwellR. Timothy HuntPaul M. NurseNobel Prize In Physiology Or MedicineCyclinCDK InhibitorCDK InhibitorSeliciclibApoptosisNeutrophil GranulocytesInflammationChronic (medicine)ArthritisCystic FibrosisAlvocidibGlucocorticoidAlvocidibOlomoucineRoscovitinePalbociclibPyrimidineSeliciclibDigital Object IdentifierPubMed IdentifierDigital Object IdentifierPubMed IdentifierDigital Object IdentifierPubMed IdentifierDigital Object IdentifierPubMed IdentifierDigital Object IdentifierPubMed IdentifierDigital Object IdentifierPubMed IdentifierDigital Object IdentifierPubMed IdentifierPubMed IdentifierPubMed IdentifierMedical Subject HeadingsEnzyme Commission NumberDigital Object IdentifierPubMed IdentifierTemplate:Intracellular Signaling Peptides And ProteinsTemplate Talk:Intracellular Signaling Peptides And ProteinsIntracellular Signaling Peptides And ProteinsTemplate:MAP Kinase PathwayIntracellular Calcium-sensing ProteinsCalcineurinCa2+/calmodulin-dependent Protein KinaseG ProteinHeterotrimeric G ProteinCAMP-dependent PathwayHeterotrimeric G ProteinGs Alpha SubunitGi Alpha SubunitAdenylate CyclaseCyclic Adenosine Monophosphate3',5'-cyclic-AMP PhosphodiesteraseProtein Kinase ACyclic Guanosine MonophosphateTransducinGustducinGuanylate CyclaseCyclic Guanosine Monophosphate3',5'-cyclic-GMP PhosphodiesteraseCGMP-dependent Protein KinaseG Alpha SubunitGNAO1GNAI1GNAI2GNAI3GNAT1GNAT2GNAT3GNAZGNAS Complex LocusGNALGNAQGNA11GNA12GNA13G Beta-gamma ComplexGNB1GNB2GNB3GNB4GNB5GNGT1GNGT2GNG2GNG3GNG4GNG5GNG7GNG11GNG12GNG13BSCL2G Protein-coupled Receptor KinaseAMP-activated Protein KinaseSmall GTPaseADP Ribosylation FactorRab (G-protein)Ras SubfamilyHRASKRASNeuroblastoma RAS Viral Oncogene HomologRho Family Of GTPasesADP Ribosylation FactorRan (gene)RHEBRap GTP-binding ProteinRRADCyclin-dependent Kinase Inhibitor ProteinCyclinPhosphoinositide Phospholipase CPhospholipase CProtein KinaseSerine/threonine-specific Protein KinaseCasein KinaseCasein Kinase 1Casein Kinase 2EIF-2 KinaseEIF2AK3Glycogen Synthase KinaseGSK-3GSK3AGSK3BIκB KinaseCHUKIKK2IKBKGInterleukin-1 Receptor Associated KinaseIRAK1IRAK2IRAK3IRAK4Lim KinaseLIMK1LIMK2P21-activated KinasesPAK1PAK2PAK3PAK4Rho-associated Protein KinaseROCK1ROCK2Ribosomal S6 KinaseRPS6KA1Non-receptor Tyrosine KinaseZAP70PTK2PTK2BBruton's Tyrosine KinaseDual-specificity KinaseProtein PhosphataseProtein Serine/threonine PhosphataseProtein Phosphatase 2Protein Tyrosine PhosphataseProtein Tyrosine PhosphataseDual-specificity PhosphataseApoptosisTemplate:Apoptosis Signaling PathwayGTP-binding Protein RegulatorsTemplate:GTP-binding Protein RegulatorsATF6Signal Transducing Adaptor ProteinIκBαMUC4Olfactory Marker ProteinPhosphatidylethanolamine Binding ProteinEDARADDPRKCSHTemplate:Deficiencies Of Intracellular Signaling Peptides And ProteinsTemplate:Cell Cycle ProteinsTemplate Talk:Cell Cycle ProteinsCell CycleProteinCyclinCyclin ACyclin A1Cyclin A2Cyclin BCyclin B1Cyclin B2Cyclin DCyclin D1Cyclin D2Cyclin D3Cyclin ECyclin E1Cyclin E2Cyclin-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 FactorWee1CullinCUL7InterphaseG1 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 REIF2AK1EIF2AK3EIF2AK4Wee1Wee1-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 KinasePLK1PLK2PLK3PLK4Cdk1Cyclin-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:Cell CycleCategory:Protein FamiliesCategory:EC 2.7.11Discussion About Edits From This IP Address [n]A List Of Edits Made From This IP Address [y]View The Content Page [c]Discussion About The Content Page [t]Edit This Page 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