Cell cycle control (mammalian)

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1 Apr. 21, 2005 Cell cycle control (mammalian) Basic mechanisms & protein components Checkpoints Chap. 21, by Lodish et al., 5 th ed Chap. 17, by Alberts et al., 4 th ed 鍾明怡 mychung@vghtpe.gov.tw

2 Advanced readings (not all required) By mechanism SCF APC Nat Rev MCB 2004;5: Nat Rev MCB 2005;6:9-20. Oncogene 2005;24: Oncogene 2005; Ralph Wasch & Dirk Engelbert (published online ahead of print). Polo-like kinases Nat Rev MCB 2004; 5: Oncogene 2005; 24: Role of translation Nat Rev MCB 2005; 6: By checkpoint DNA damage checkpoint Nat Rev MCB 2004; 5: Nat Rev MCB 2005;6: Spindle checkpoint Chromosome Research 2004;12: Others Connecting proliferation and apoptosis in development and disease. Nat Rev MCB 2004; 5: Cancer genes and the pathways they control. Nat Med 2004; 10: Haploinsufficiency for tumor suppressor genes: when you don t need to go all the way. Biochimica et Biophysica Acta 2004;1654:

3 Mammalian cell cycle control

4 The basic organization of cell cycle and its control system are essentially the same in all eukaryotic cells. Why discuss mammals separately? Differences between yeast and mammals.

5 Cell division in multicellular organisms In addition to availability of nutrients, it must also receive stimulatory extracellular signals for a multicellular organism to divide. Mitogens vs. growth factors Proliferation vs. growth e.g. PDGF Cell cycle control and human diseases.

6 Cell growth and cell cycle control N/C ratio Only cycle control N/C ratio both

7 A simplified model of one way that mitogens stimulate cell division.

8 Study of cell cycle in mammalian systems Cell-culture based. Normal tissues, e.g. fibroblast. Complication: replicative cell senescence after divisions. Tumor tissues Reveal characteristics of normal cells. Diploid. Anchorage dependence. Contact inhibition. Replicative cell senescence.

9 Cell cycle control why & how? Basic programs: timed events in the correct order; triggered only once in each cycle; irreversible; must complete prior to initiation of the next event. Sensor. Adaptability: adjust to cell type, environment, etc. Robustness: backup mechanisms.

10 Cell cycle checkpoints The two most important things in a cell cycle: 1. 2.

11 Mechanisms of cell cycle control (1) Generally operate through negative intracellular signals. Default setting: STOP. Gradual decrease in the negative signal to zero results in GO. Inactivation of checkpoints results in progression of cell cycle without surveillance. Based on cyclically activated protein kinases, cdks. Cyclin-dependent kinase (cdk). Regulated by a bunch of enzymes and proteins, e.g. cyclin.

12 Cyclins and cdks in vertebrates Cyclins activate cdks; also direct their targets. Another level of regulation: accessibility of the substrate, or target site on the substrate. B-type cyclins Target of APC

13 Mechanisms of cell cycle control (2) Transcriptional regulation. Translational regulation. Post-translational regulation. Activating and inhibitory phosphorylation. Inhibitory proteins. Protein degradation. SCF mediated. APC mediated. Subcellular localization.

14 Transcriptional regulation Level of cyclin is regulated at the level of transcription, translation and degradation. In embryonic cell cycles, M-cyclin is synthesized at a constant rate; M-cyclin accumulates due to decreased degradation. In most cells, however, M-cyclin increases during G2 and M by increasing transcription. Will be discussed later. G 0 G 1

15 Translational regulation Basal cell hyperplasia in prostate eif4 JBC 1998;273: Normal basal cell Webpathology.com

16 cyclins and cdks amount vs. activity (Cdk7-cyclin H-Mat1)

17 Mat1 Cdk7/cyclin H Cdk7/cyclin H/Mat1 Cdk7( Ser 170 Thr 176 )/cyclin H Cdk2/cyclin A Active CAK Cdk2 ( Thr 160 ) /cyclin A Wee1 Cdc25A Cdk2 ( Tyr 15 Thr 160 ) /cyclin A Cannot activate oneself via autophosphorylation

18 Activating and inhibitory phosphorylations of M-Cdk Cyclin A/Cdk2 Cdc25A Cyclin A/Cdk2 Cyclin B/Cdk1 Cdc25B Cyclin B/Cdk1

19 Cdk inhibitory proteins (CKIs) Two classes of Cdk inhibitors Cdk Inhibitory proteins (CIP/KIP) Inhibit all Cdk1,2,4,6 complexes. p21, p27, p57. Inhibitors of kinase 4 (INK4) Inhibit Cdk4&6. p16, p15.

20 Proteolysis in cell cycle control (1) SCF mediated Named after its three subunits. Regulation of G1/S-cyclins and CKI levels. SCF activity is constant throughout cell cycle (?). Controlled by phosphorylation of the target protein. p27 Cyclin E Cdk2 p27

21 SCF ubiquitin ligases Skp1-Cul1-F box (Skp2) = SCF Skp2 E2 Substrate interaction E2/E3 interaction

22 Proteolysis and CDK activity 3 Nat Rev Mol Cell Biol 2004;5:

23 Nat Rev Mol Cell Biol 2004;5:

24 Proteolysis in cell cycle control (2) APC mediated Anaphase-promoting complex. Degrades M-cyclins and other proteins at mitosis. Oncogene (2005) 24,

25 APC/C (cyclosome) APC/C is a multisubunit E3 ubiquitin ligase with at least two alternative forms, one activated by Cdc20 (APC/C CDC20 ) and the other by Cdh1 (APC/C CDH1 ). Cdh1=Hct1. The homolog of the largest subunit of APC, Apc1, in fission yeast has also been described as a constituent of the 20S cyclosome. Required for Entry into anaphase (APC/C CDC20 ). Exit from mitosis (APC/C CDH1 ). Maintenance of low Cdk activity in early G1 (APC/C CDH1 ).

26 Subunits of APC Oncogene (2005) 24,

27 active inactive inactive inactive active active Role of APC in cell cycle 4 Oncogene (2005) 24,

28 Checkpoints (by cell-cycle phases; by specific aims) Molecular signaling cascades that promote cellcycle delay or arrest in response to unfavorable conditions, e.g. DNA damage, thereby providing more time to overcome the unfavorable condition or to pursue an alternative. -- Adapted from Nat Rev Mol Cell Biol 2004; 5:

29 2 Lodish, et al. 5th ed.

30 Passage through restriction point : G1 progression & initiation of S phase Growth factor cyclin D ( txc) => active G1-Cdk (cyclin D:cdk4/6) (cyclin E:cdk2) Restriction point

31 Intracellular control of cell cycle Re-replication block Cells at S phase contain signals for DNA replication and initiate DNA synthesis in G1 cell after fusion. S + G2 => no DNA replication in G2 => DNA can be replicated only once in a cycle. G2 + G1 => cannot induce G1 cells into S phase.

32 Control of entry into S phase (1) Origin of replication. ORC. Associated with Ori throughout the cell cycle. Regulatory proteins Cdc6. Transient in early G1. Mcm (Minichromosome maintenance ) proteins, etc. Form pre-replicative complex. Ready for DNA replication.

33 Control of entry into S phase (2) Activation of cyclin A/Cdk2 in late G1 triggers S phase. Phosphorylation of ORC => initiate DNA replication. phosphorylation of cdc6 => degradation by SCF. Phosphorylation of Mcm => export from the nucleus. geminin occupies Ori, cannot re-initiate until cycle is completed. cannot reassemble pre-rc. How to initiate DNA synthesis in the next cycle?

34 Activated ATM Chk2 (active) Cdc25A Ser 126 (inactive) (degradation) G1/S block DNA repair Apoptosis DNA damage checkpoint G1, G2 arrest

35 DNA damage checkpoint DSB, or Replication checkpoint ATM= ataxia telangiectasia mutated. ATR= ataxia telangiectasia and RAD related CHK1/2= checkpoint kinase 1/2 Cdc25C (inactive) G2 arrest Nat Rev Mol Cell Biol 2005;6:44-55.

36 Role of p53 Overexpression of a mutated form of the p53 tumor suppressor gene in a hepatic metastasis in colorectal cancer. The brown stain (immunohistochemistry, PAP) shows that the mutated p53 localizes to the nuclei of the cancer cells where it fails to transduce the apoptotic signal. This renders cancer cells refractory to classical radio- or chemotherapy. tion.html

37 Both Rb and p53 are tumor suppressor genes and show dominant inheritance in affected families. What are the mechanisms for these phenomena? How about p27 Kip1?

38 DNA replication checkpoint Late S/G2. Sensor mechanisms are not fully understood. Unreplicated DNA (?). Unfinished replication fork (?). Final target: the pathway for M-Cdk activation. Inhibits DNA synthesis high doses

39 DNA replication checkpoint RPA = replication protein A Nat Rev Mol Cell Biol 2005;6: Cdc25C (inactive) G2 arrest p21 CIP p53 Repair Apoptosis G2 arrest

40 Summary of S phase checkpoints Nat Rev MCB 2004;5:792.

41 Nat Rev MCB 2004;5:792.

42 Control of entry into M-phase Mitosis Prophase Pro-metaphase Metaphase Anaphase Telophase Cytokinesis

43 Control of M-Cyclin quantity M-cyclin transcript S M ~30min/cycle fairly synchronous (transcript and protein) G1 allows Cell growth Regulation by extracellular signals.

44 Control of M-Cdk activity Cyclin B concentration Polo kinase

45 The roles of M-Cdk Chromosome condensation (condensin complex; phosphorylated by M-cyclin => change DNA coiling => initiating condensation.; MCB). Induces the assembly of mitotic spindle. Ensures attachment of chromosomes to mitotic spindle. Nuclear envelop breakdown. Actin rearrangement. Re-organization of Golgi and ER. Details of these events are not all fully understood.

46 Cohesion of sister chromatids Sister chromatids are joined together by cohesins, deposited along the each sister chromatid as DNA is replicated. Cohesion of sister chromatids are broken at anaphase.

47 Chrosome condensation M-Cdk phosphorylates condensin subunits, triggering the assembly of condensin complex and the progressive condensation of the chromosomes. Prometaphase Metaphase Blue: DNA Red: anti-condensin

48 Nuclear envelope breakdown in mitosis

49 Assembly of mitotic spindle M-Cdk phosphorylates MAP and promotes stabilization of microtubule.

50 Sister chromatid separation APC-mediated degradation of M-cyclin => inactivate M-Cdk => initiation of anaphase.

51 Unattached kinetochore txc - Spindle checkpoint

52 Spindle checkpoint Sensor monitors the status of kinetochore. Mechanism not yet clear. Mad2 at unattached kinetochore sends out (-) signal; (-) APC activation. Defective spindle checkpoint => earlier anaphase initiation. Spindle checkpoint is not required in frog embryo and yeast; some other mechanism(s) is controlling the timing of anaphase.

53 Mad2 and spindle checkpoint Mad2 - tubulin DNA (A) Just after nuclear envelope breakdown. (B) Early prometaphase. (C) Mid-prometaphase. (D) Late prometaphase. (E) Metaphase. (F) Anaphase.

54 Chromosome Research 2004;12:

55 Chromosome Research 2004;12:

56 Non-disjunction

57 Cytokinesis Contractile ring: composed of actin and myosin filaments, formed around the equator of the cell. Control mechanisms to assure that mitosis occurs prior to cytokinesis.

58 Exit from mitosis Degradation of M-cyclin => inactivation of M-Cdk => decrease of kinases activation of phosphatases. Events Disassembly of mitotic spindle. Decondensation of chromosomes. Formation of nuclear envelope.

59 Control of M-cyclin quantity M-cyclin transcript S M ~30min/cycle fairly synchronous (transcript and protein) G1 allows Cell growth Regulation by extracellular signals.

60 Suppressed Cdk activities in early G1 CKI: Sic1 in yeast; p21/p27/p57 in mammals; transcription. G1 cyclins are not inactivated by APC/C CDH1 and CKI. Inactive Txc factor active Txc factor CKI CKI CKI CKI Cdh1 Cdh1 Cdh1 w/ cdc20 CKI Cdh1 S-cyclin CKI Cdh1 Txc

61 (1)

62 (2)

63 1

64

65 Try to give an organized general overview of mammalian cell cycle control with updated information. Some of the details are not yet fully explored. Please do not be misled by the cartoons.

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