Regulation of the cell cycle
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1 Regulation of the cell cycle
2 Dr. F. Neuschäfer-Rube Cyclin-dependent Kinases (CDKs)
3 Cyclin-dependent kinases: Motors and switches of the cell cycle Dr. F. Neuschäfer-Rube
4 The cell cyle M S
5 The cell cyle M S = DNA-Synthesis (Replication)
6 The cell cyle M = Mitosis Dissection of the chromosoms S = DNA-Synthesis
7 The cell cycle: Steps of the Mitosis Prophase Metaphase Anaphase Telophase
8 The cell cycle M = Mitosis G2 G1 S = DNA-Synthesis
9 The cell cycle G2 = Control of DNA-Synthesis M = Mitosis G1 S = DNA-Synthesis
10 The cell cycle G2 = Control of DNA-Synthesis M = Mitosis G1 = Cell growth S = DNA-Synthesis
11 The cell cycle G2 = Control of DNA-Synthesis M = Mitosis G1 = Cell growth Go = Quiescent state of differentiated cells S = DNA-Synthesis
12 The cell cycle G2 = Control of DNA-Synthesis M = Mitosis G1 = Cell growth How is the correct sequence of the cell cycle controlled? Go = Quiescent state S = DNA-Synthesis
13 The cell cycle G2 = Control of DNA-Synthesis M = Mitosis G1 = Cell growth How is the correct sequence of the cell cycle controlled? Checkpoints Go = Quiescent state S = DNA-Synthesis
14 The cell cycle G2 = Control of DNA-Synthesis M = Mitosis G1 = Cell growth How is the correct sequence of the cell cycle controlled? Go = Quiescent state S = DNA-Synthesis Restriction point intrinsic: final cell size reached?
15 The cell cycle G2 = Control of DNA-Synthesis M = Mitosis G1 = Cell growth How is the correct sequence of the cell cycle controlled? Go = Quiescent state S = DNA-Synthesis Restriction point intrinsic: final cell size reached? external: growth factors?
16 The cell cycle G2 = Control of DNA-Synthesis M = Mitosis G1 = Cell growth G2-Control Replication complete? DNA intact? Go = Quiescent state S = DNA-Synthesis Restriction point intrinsic: final cell size reached? external: growth factors?
17 G2 = Control of DNA-Synthesis The cell cycle M = Mitosis Metaphase-Control correct annealing of chromosoms? G1 = Cell growth S = DNA-Synthesis G2-Control Replication complete? DNA intact? Go = Quiescent state Restriction point intrinsic: final cell size reached? external: growth factors?
18 Checkpoints of the cell cycle M = Mitosis G2 G1 Which biochemical factors regulate the transition of cell cycle steps? Go S
19 Cellfusion experiments with synchronised HeLa cells S-Phase G1 + S-Phase G2 + G2 G1 +
20 Cellfusion experiments with synchronised HeLa cells S-Phase G1 + S-Phase G2 + G2 G1 +
21 Cellfusion experiments with synchronised HeLa cells S-Phase G1 + S-Phase G2 + G2 G1 +
22 Cell fusion experiments show three results: 1. Only cells in the G1-phase are competent to enter the S-phase (DNA-replication) 2. Only cells in the S-Phase express a factor which stimulates DNA-replication in competent cells in the G1-phase. 3. Cells in the G2-phase can`t execute DNA-replication until mitosis is not completed. Initiation of DNA-synthesis by a soluble factor in the early S-phase G1/S-Phase: S-Phase-promoting- factor (SPF) G2/M-Phase: Mitosis-promoting-factor (MPF)
23 Cell fusion experiments show three results: 1. Only cells in the G1-phase are competent to enter the S-phase (DNA-replication) 2. Only cells in the S-Phase express a factor which stimulates DNA-replication in competent cells in the G1-phase. 3. Cells in the G2-phase can`t execute DNA-replication until mitosis is not completed. Initiation of DNA-synthesis by a soluble factor in the early S-phase G1/S-Phase: S-Phase-promoting- factor (SPF) G2/M-Phase: Mitosis-promoting-factor (MPF)
24 Cell fusion experiments show three results: 1. Only cells in the G1-phase are competent to enter the S-phase (DNA-replication) 2. Only cells in the S-Phase express a factor which stimulates DNA-replication in competent cells in the G1-phase. 3. Cells in the G2-phase can`t execute DNA-replication until mitosis is not completed. Initiation of DNA-synthesis by a soluble factor in the early S-phase G1/S-Phase: S-Phase-promoting- factor (SPF) G2/M-Phase: Mitosis-promoting-factor (MPF)
25 Cell fusion experiments show three results: 1. Only cells in the G1-phase are competent to enter the S-phase (DNA-replication) 2. Only cells in the S-Phase express a factor which stimulates DNA-replication in competent cells in the G1-phase. 3. Cells in the G2-phase can`t execute DNA-replication until mitosis is not completed. Initiation of DNA-synthesis by a soluble factor in the early S-phase G1/S-Phase: S-Phase-promoting- factor (SPF) G2/M-Phase: Mitosis-promoting-factor (MPF)
26 Cell fusion experiments show three results: 1. Only cells in the G1-phase are competent to enter the S-phase (DNA-replication) 2. Only cells in the S-Phase express a factor which stimulates DNA-replication in competent cells in the G1-phase. 3. Cells in the G2-phase can`t execute DNA-replication until mitosis is not completed. Initiation of DNA-synthesis by a soluble factor in the early S-phase G1/S-Phase: S-Phase-promoting- factor (SPF) G2/M-Phase: Mitosis-promoting-factor (MPF)
27 Identification of cell cycle regulators: CDKs Leland H Hartwell
28 Identification of cell cycle regulators: CDKs Identification of temperature-sensitive CDC (Cell-Division-Cycle) Mutants Model: Saccharomyces sereviciae Mutagenized cells 25 C 35 C
29 Identification of cell cycle regulators: CDKs Identification of temperature-sensitive CDC (Cell-Division-Cycle) Mutants Model: Saccharomyces sereviciae Mutagenized cells 25 C non CDC-Mutants 35 C 35 C budding Entry into the Cell cycle
30 Identification of cell cycle regulators: CDKs Identification of temperature-sensitive CDC (Cell-Division-Cycle) Mutants Model: Saccharomyces sereviciae Mutagenized cells 25 C non CDC-Mutants 35 C 35 C CDC-START-Mutants no entry into the cell cycle
31 Identification of cell cycle regulators: CDKs Identification of the CDC-START gene CDC-START-Mutant 35 C Wt Gen X 35 C Wt CDC-START-Gen
32 Identification of cell cycle regulators: CDKs Identification of the CDC-START gene CDC-START-Mutant 35 C Wt Gen X 35 C Wt CDC START Gen Analysis Cyclin-dependent Kinase (CDK)
33 Discovery of Cyclin-dependent Kinases (CDKs): Nobel price in medicine 2001 Leland H Hartwell Paul Nurse Tim Hunt CDKs in yeast Cyclins in sea urch
34 Cyclin-dependent kinases (CDKs): Heterodimeric proteins C.L. Card et al., EMBO Journal 2000 katalytische Untereinheit: CDK regulatorische Untereinheit: Cyclin
35 Cyclin-dependent kinases (CDKs): Heterodimeric proteins C.L. Card et.al., EMBO Journal 2000 catalytic subunit: CDK - Serine/Threonine-Kinases - Yeast: one CDK - Mammals: CDK1 - CDK7 - high identity - conserved cyclin-binding site regulatorische Untereinheit: Cyclin
36 Cyclin-dependent kinases (CDKs): Heterodimeric proteins C.L. Card et.al., EMBO Journal 2000 catalytic subunit: CDK - Serine/Threonine-Kinases - Yeast: one CDK - Mammals: CDK1 - CDK7 - high identity - conserved cyclin-binding site regulatory subunit: Cyclin - Cyclin A-H - heterogeneous protein family - cyclic concentration changes in the cell cycle - Nuclear localisation
37 CDK/Cyclin-complexes in the cell cycle M Go G2 G1 CDK2/CyclinD CDK4/CyclinD CDK6/CyclinD Restriction point G1/S-Phase transition S CDK2/CyclinE
38 CDK/Cyclin-complexes in the cell cycle M Go G2 CDK2/CyclinD CDK4/CyclinD CDK6/CyclinD Restriction point G1/S-Phase transition S CDK2/CyclinA CDK2/CyclinE
39 CDK/Cyclin-complexes in the cell cycle CDK1/CyclinB G2/M-Phase transition G2 M Go CDK2/CyclinD CDK4/CyclinD CDK6/CyclinD G1 Restriction point G1/S-Phase transition S CDK2/CyclinA CDK2/CyclinE
40 CDKs: Motors of the cellcycle How act CDKs as motors of te cell cycle? Which substrates are phophorylated by CDKs during the cell cycle?
41 CDK Substrates: Initiation of the S-Phase Fuction of CDKs in initiation of the S-Phase
42 CDK Substrates: Initiation of the S-Phase Retinoblastom-Protein (Rb) - Key - substrate of the S-Phase - nuclear protein, 110 kda NH2- A B -COOH
43 CDK Substrates: Initiation of the S-Phase Retinoblastom-Protein (Rb) - Key - substrate of the S-Phase - nuclear protein, 110 kda NH2- A B -COOH Binding of transcription factor E2F
44 CDK Substrates: Initiation of the S-Phase Retinoblastom-Protein (Rb) - Key - substrate of the S-Phase - nuclear protein, 110 kda NH2- A B -COOH Binding of transcription factor E2F E2F: central transcription factor for the induction of S-Phase genes
45 CDK Substrates: Initiation of the S-Phase Retinoblastom-Protein (Rb) - Key - substrate of the S-Phase - nuclear protein, 110 kda NH2- A B -COOH Binding of transcription factor E2F Rb act as a tumor supressor gene
46 CDK Substrates: Initiation of the S-Phase Retinoblastom-Protein (Rb) - Key - substrate of the S-Phase - nuclear protein, 110 kda P P P P P P P P P P NH2- A B -COOH Binding of transcription factor E2F - Rb act as a tumor supressor gene - Rb can be hyperphosphorylated
47 CDK Substrates: Initiation of the S-Phase Rb Rb E2F Repression of E2F-controlled genes
48 CDK Substrates: Initiation of the S-Phase Cyclin E CDK 2 Rb P P P Rb Rb E2F Repression of E2F-controlled genes E2F Induktion of E2F-controlled genes
49 E2F: Initiator of the S-Phase Cyclin E E2F-controlled genes CDK 2 P P P Rb E2F DNA-Pol I dntp-synth.
50 E2F: Initiator of the S-Phase Cyclin E E2F-controlled genes CDK 2 Cyclin E P P P Rb E2F DNA-Pol I dntp-synth. E2F positive autoregulatory feedback loop!
51 E2F: Initiator of the S-Phase Cyclin E E2F-controlled genes CDK 2 Cyclin E P P P Rb E2F positive autoregulatory feedback loop! DNA-Pol I dntp-synth. E2F forced" transition of the restriction point S-Phase
52 CDK substrate: Mitosis Function of CDKs in the mitosis
53 CDK substrate: Mitosis An important step in mitosis is the desintegration of the nuclear membrane Interphase Mitose
54 CDK substrate: Mitosis chromatin nuclear lamina inner nuclear membrane
55 CDK substrate: Mitosis chromatin nuclear lamina mitosis desintegration of the nuclear lamina desintegration of the der nuclear membran
56 CDK substrate: Mitosis chromatin nuclear lamina Mitose ATP Cyclin B CDK 1 desintegration of the nuclear lamina desintegration of the der nuclear membran
57 CDK substrate: Mitosis chromatin nuclear lamina lamin network Mitose ATP Cyclin B CDK 1 lamin tetramer desintegration of the nuclear lamina desintegration of the der nuclear membran
58 CDK substrate: Mitosis chromatin nuclear lamina lamin netork Mitose ATP Cyclin B CDK 1 lamin tetramer ATP desintegration of the nuclear lamina desintegration of the der nuclear membran -P P- -P P- phosphorylated lamin dimers
59 CDK/Cyclin complexes in the cell cycle CDK1/CyclinB Lamin-P G2/M-phase transition CDK2/CyclinA G2 S M restriction point G1 Go CDK2/CyclinD CDK4/CyclinD CDK6/CyclinD G1/S-phase transition Rb-P E2F CDK2/CyclinE
60 CDKs: Switches of the cell cycle How is the activity of CDKs switched on- and off in the cell cycle?
61 Regulation of CDK activity inactive CDK
62 Regulation of CDK activity: Cyclin concentration Cyclin Cyclin inactive CDK CDK inactive
63 Regulation of CDK activity: Cyclin concentration Cyclin Cyclin inactive CDK CDK inactive ATP P Threonin-Kinase (CAK) Cyclin CDK active T160 P only CDK-Cyclin complexes are substrates P
64 Conformation change in CDK structure by cyclin binding
65 Conformation change in CDK structure by cyclin binding + CAK
66 Regulation of CDK activity: Cyclin concentration Cyclin Cyclin inactive CDK CDK inactive Cyclin concentration ATP Threonin-Kinase (CAK) Cyclin CDK T160 P active P
67 Regulation of CDK activity: Cyclin concentration How is the concentration of cyclins regulated in the cell cycle? Cyclin
68 Regulation of CDK activity: Cyclin concentration How is the concentration of cyclins regulated in the cell cycle? De novo synthesis by transcriptional induction Cyclin
69 Regulation of cyclin concentration: transcriptional Induktion Example: Induction by growth factor signal chains P
70 GF-R Regulation of cyclin concentration: transcriptional Induktion GF Example: Induction by growth factor signal chains P
71 GF-R Regulation of cyclin concentration: transcriptional induction GF Example: Induction by growth factor signal chains Protein kinases TF-OH TF-O-P P
72 GF-R Regulation of cyclin concentration: transcriptional induction GF Example: Induction by growth factor signal chains Protein kinases TF-OH TF-O-P immediate early genes P
73 GF-R Regulation of cyclin concentration: transcriptional induction GF Example: Induction by growth factor signal chains Protein kinases c-jun/c-fos (TF) TF-OH TF-O-P immediate early genes P
74 GF-R Regulation of cyclin concentration: transcriptional induction GF Example: Induction by growth factor signal chains Protein kinases c-jun/c-fos (TF) TF-OH TF-O-P immediate early genes c-jun c-fos delayed genes P
75 GF-R Regulation of cyclin concentration: transcriptional induction GF Example: Induction by growth factor signal chains Protein kinases c-jun/c-fos (TF) Cyclin D Cyclin E CDK2 CDK4 TF-OH TF-O-P immediate early genes c-jun c-fos delayed genes P
76 GF-R Regulation of cyclin concentration: transcriptional induction GF Example: Induction by growth factor signal chains Protein kinases c-jun/c-fos (TF) Cyclin D Cyclin E CDK2 CDK4 TF-OH TF-O-P immediate early genes c-jun c-fos delayed genes P Restriction point (G1 S-Phase)
77 Regulation of CDK activity: Cyclin concentration How is the concentration of cyclins regulated in the cell cycle? De novo synthesis by transcriptional induction Degradation by proteolysis Restriktion point (G1 S-Phase) Cyclin
78 Regulation of cyclin concentration: proteolytic degradation Example: Regulation of CDK1 activity by degradation of cyclin B CDK1-activity Metaphase G1 S G2 M G1 S G2 M Cell cycle-phase
79 Regulation of cyclin concentration: proteolytic degradation Example: Regulation of CDK1 activity by degradation of cyclin B CDK1-activity CDK1 concentration Metaphase G1 S G2 M G1 S G2 M Cell cycle-phase
80 Cyclin B concentration Regulation of cyclin concentration: proteolytic degradation Example: Regulation of CDK1 activity by degradation of cyclin B CDK1-activity CDK1 concentration Metaphase Degradation of cyclinb G1 S G2 M G1 S G2 M Cell cycle-phase
81 Regulation of cyclin concentration: proteolytic degradation NH 2 Cyclin A,B COOH Destruction-Box
82 Regulation of cyclin concentration: proteolytic degradation NH 2 Cyclin A,B COOH Destruction-Box Ubiquitin NH 2 COOH
83 Regulation of cyclin concentration: proteolytic degradation NH 2 Cyclin A,B COOH Destruction-Box Ubiquitin Anaphase-promoting-complex (APC, Ubiquitin-Ligase-complex) NH 2 COOH
84 Regulation of cyclin concentration: proteolytic degradation NH 2 Cyclin A,B COOH Destruction-Box Ubiquitin Anaphase-promoting-complex (APC, Ubiquitin-Ligase-Komplex) NH 2 COOH Labeling for proteolytic degradation
85 Regulation of cyclin concentration: proteolytic degradation NH 2 Cyclin A,B COOH Destruction-Box Ubiquitin Anaphase-promoting-complex (APC, Ubiquitin-Ligase-Komplex) NH 2 COOH Labeling for proteolytic degradation degradation NH 2 Proteasom COOH
86 Regulation of cyclin concentration: proteolytic degradation How is the proteolytic degradation of cyclin B regulated?
87 Regulation of cyclin concentration: proteolytic degradation APC inactive Cyclin B CDK1 G1-Phase
88 Regulation of cyclin concentration: proteolytic degradation Cyclin B synthesis APC inactive S, G2-Phase Cyclin B CDK1 G1-Phase
89 Regulation of cyclin concentration: proteolytic degradation Metaphase Cyclin B CDK1 Cyclin B synthesis APC inactiv P APC active S, G2-Phase Cyclin B CDK1 G1-Phase
90 Regulation of cyclin concentration: proteolytic degradation Metaphase Cyclin B CDK1 ATP Cyclin B synthesis APC inactive P APC active S, G2-Phase Cyclin B CDK1 G1-Phase
91 Regulation of cyclin concentration: proteolytic degradation Metaphase Cyclin B CDK1 polyubiquitinylation ATP Cyclin B synthesis APC inactive P APC active Cyclin B CDK1 proteasomal degradation S, G2-Phase Cyclin B CDK1 G1-Phase
92 Regulation of cyclin concentration: proteolytic degradation Metaphase Cyclin B CDK1 polyubiquitinylation ATP Cyclin B synthesis APC inactive P APC active Cyclin B CDK1 proteasomal degradation Cyclin B S, G2-Phase CDK1 G1-Phase Anaphase
93 Regulation of cyclin concentration: proteolytic degradation Metaphase Cyclin B CDK1 negative autoregulatory feedbackloop polyubiquitinylation ATP Cyclin B synthesis APC inactive P APC active Cyclin B CDK1 proteasomal degradation Cyclin B S, G2-Phase CDK1 G1-Phase Anaphase
94 Regulation of CDK activity: Cyclin concentration Growth factors Autoregulation De novo synthesis by transcriptional induktion Degradation by proteolysis Restriktion point (G1 S-Phase) Cyclin Metaphase Anaphase completion of mitosis
95 Regulation of CDK activity: Phosphorylation Cyclin Cyclin inactive CDK CDK inactive Cyclin concentration ATP Threonin-Kinase (CAK) Cyclin CDK T160 P aktiv ATP Threonin/Tyrosin- Kinase P P P T14 Y15 Cyclin CDK inactive T160 P
96 Regulation of CDK activity: Dephosphorylation Cyclin Cyclin inactive CDK CDK inactive Cyclin concentration ATP Threonin-Kinase (CAK) Cyclin CDK T160 P aktiv ATP Threonin/Tyrosin- Kinase Threonin/Tyrosin Phosphatase P Pi P P T14 Y15 Cyclin CDK inactive T160 P
97 Regulation of CDK activity: Dephosphorylation How is the activation of CDKs by Dephosphorylation regulated?
98 Regulation of CDK activity: Dephosphorylation of CDK1 G2-Phase P P T14 Y15 CyclinB CDK1 inactive T160 P P Threonin/Tyrosin Phosphatase G2-Phase inactive
99 Regulation of CDK activity: Dephosphorylation of CDK1 G2-Phase P P T14 Y15 CyclinB CDK1 inactive T160 P P Mitosis P Pi Threonin/Tyrosin Phosphatase active Mitosis CyclinB CDK1 T160 P P active Threonin/Tyrosin Phosphatase inactive G2-Phase
100 Regulation of CDK activity: Dephosphorylation of CDK1 G2-Phase P P T14 Y15 CyclinB CDK1 inactive T160 P P Mitosis P Pi P Threonin/Tyrosin Phosphatase active Mitosis CyclinB CDK1 T160 P activation active ATP Threonin/Tyrosin Phosphatase inactive G2-Phase
101 Regulation of CDK activity: Dephosphorylation of CDK1 G2-Phase P P T14 Y15 CyclinB CDK1 inactive T160 P P Mitosis P Pi P Threonin/Tyrosin Phosphatase active Mitosis CyclinB CDK1 T160 P activation positive autoregulatory feedbackloop active ATP Threonin/Tyrosin Phosphatase G2-Phase inactive
102 Regulation of CDK activity: Inhibitors Cyclin Cyclin inactive CDK CDK inactive Cyclinconcentration ATP CDK-Inhibitors CKI Cyclin CDK T160 P Phosphorylation Dephosphorylation active ATP CDK CKI inactive T160 P P Pi P P T14 Y15 Cyclin CDK inactive T160 P
103 Regulation of CDK activity: Inhibitors Example: CKI p21 Isosteric inhibition of CDK activity by binding in the active centre Cyclin E CDK 2 CKI p21 G1 S-Phase
104 Regulation of CDK activity: Inhibitors How is the concentration of CKI p21 regulated?
105 Regulation of CDK activity: Induction of the inhibitor p21 p53 Transcription factor, tumor supressor gene
106 Regulation of CDK activity: Induction of the inhibitor p21 p53 Transcription factor, tumor supressor gene CKI p21 p53
107 Regulation of CDK activity: Induction of the inhibitor p21 p53 Transcription factor, tumor supressor gene CKI p21 p53 Cyclin E CDK 2 CKI p21
108 T1/2 = 30 min Regulation of CDK activity: Induction of the inhibitor p21 p53 Transcription factor, tumor supressor gene CKI p21 p53 Cyclin E CDK 2 p53 CKI p21 degradation
109 T1/2 = 30 min T1/2 = 150 min Regulation of CDK activity: Induction of the inhibitor p21 p53 Transcription factor, tumor supressor gene CKI p21 DNA damage p53 Cyclin E CDK 2 p53 CKI p21 degradation
110 T1/2 = 30 min T1/2 = 150 min Regulation of CDK activity: Induction of the inhibitor p21 p53 Transcription factor, tumor supressor gene CKI p21 DNA damage p53 Cyclin E CDK 2 p53 degradation CKI p21 G1-Phase arrest
111 T1/2 = 30 min T1/2 = 150 min Regulation of CDK activity: Induction of the inhibitor p21 p53 Transkriptiorfactor, tumor supressorgene CKI p21 DNA damage p53 p53 Degradation Time for DNA repair before replication Cyclin E CDK 2 CKI p21 G1-Phase Arrest
112 Regulation of CDK activity: regulation of p53 p53 Mdm2 p53 Mdm2 Ubiquitin ligase Degradation
113 Regulation of CDK activity: regulation of p53 P p53 p53 Mdm2 p53 Mdm2 Degradation
114 Regulation der CDK-Aktivität: Regulation von p53 Wachstumsfaktorhyperstimulation P MAPK p53 p53 Mdm2 p53 Mdm2 Abbau
115 Regulation of CDK activity: regulation of p53 Growth factorhyperstimulation DNA-damage P p53 MAPK DNA-PK ATM p53 Mdm2 protein kinase p53 Mdm2 Degradation
116 Regulation of CDK activity: regulation of p53 Growth factorhyperstimulation DNA-damage P P MAPK DNA-PK ATM ATR Mdm2 protein kinase p53 p53 Mdm2 p53 Mdm2 Degradation
117 Regulation of CDK activity: regulation of p53 Growth factorhyperstimulation DNA-damage P P MAPK DNA-PK ATM ATR Mdm2 p53 p53 Mdm2 p19/arf p53 Mdm2 Degradation
118 Regulation of CDK activity: regulation of p53 Growth factor hyperstimulation DNA-damage P P MAPK DNA-PK ATM ATR Mdm2 p53 p53 Mdm2 p19/arf p53 Mdm2 Mdm2 p19/arf Dgradation
119 Regulation of CDK activity: regulation of p53 Growth factor hyperstimulation DNA-damage P P MAPK DNA-PK ATM ATR Mdm2 p53 p53 Mdm2 p19/arf induktion of p21 cell cyle arrest p53 Mdm2 Dgradation Mdm2 p19/arf
120 Regulation of CDK activity: summary Cyclin Cyclin inactive CDK CDK inactive P G1 S Cyclin concentration M G1 ATP Cyclin CDK T160 P active P
121 Regulation of CDK activity: summary Cyclin Cyclin inactive CDK CDK inactive ATP G1 S Cyclin concentration M G1 G2 M Cyclin CDK active T160 P ATP Phosphorylation Dephosphorylation G2 M P Pi P P T14 Y15 Cyclin CDK inactive T160 P
122 Regulation of CDK activity: summary Cyclin Cyclin inactive CDK CDK inactive ATP G1 S Cyclin concentration M G1 G2 M CDK-Inhibitors G1 S CKI Cyclin CDK active T160 P ATP Phosphorylation Dephosphorylation G2 M CDK CKI inactive T160 P P Pi P P T14 Y15 Cyclin CDK inactive T160 P
123 Cyclin-dependent kinases: Motors and switches of the cell cycle M Dephosphorylation Cyclin degradation G2 Lamin-P G1 prb/e2f R Cyclin synthesis S Inhibitors
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