Cell cycle machinery M G1
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1 Cell cycle machinery M G1 G2 S
2 D A cdk1 cdk1 B M G2 G1 S Cdk4/6 E E2F P P P P Rb P P A
3 cdk1 B M G1 D1 Cdk4/6 E1 D2 Cdk4/6 Cdk4/6 D3 E2 E2 A2 A1 G2 S E2 A2 A1
4 cyc A cyc E cyc D G1 S G2 M
5 E-CYCLINS cyclin E1 and E2 associate with CDK2 (CDK3 or CDK1) rapidly induced prior to the S phase entry overexpression shortens G1 phase antibodies block S phase entry cyclin E gene on 19q12 amplified in several cancers cyclin E overexpressed in ~25% of breast cancers
6 Functions for E-CYCLINS Retinoblastoma protein (prb) inactivation Cell cycle progression (p27 Kip1, CDC25) DNA replication initiation (CDC45 loading) Centrosome duplication (nucleophosmin) Histone biosynthesis (p220 NPAT) E-cyclins are absolutely essential for cell cycle progresssion
7 EXPRESSION OF CYCLINS E1 AND E2 cyclin E1 cyclin E2 BrdU E9.5 n n b m E11.5 n n
8 cdk1 B M G1 D1 Cdk4/6 E1 D2 Cdk4/6 Cdk4/6 D3 E2 E2 A2 A1 G2 S E2 A2 A1
9 Cyclin E1 knockout cyclin E1 genomic structure Xho I Asp I Xba I Xba I Xba I Spe I Sca I Not I Spe I Xho I Sca I Hpa I Not I Neo (ScaI) Spe I cyclin E1 targeting strategy Spe I Sca I Hpa I 1kb WT E1 +/- E1 -/- 7.6kb 2.8kb 2.6kb
10 Cyclin E1 -/- mice viable develop normally
11 cdk1 B M G1 D1 Cdk4/6 E1 D2 Cdk4/6 Cdk4/6 D3 E2 E2 A2 A1 G2 S E2 A2 A1
12 Cyclin E2 knockout cyclin E2 genomic structure Not I EcoR I Sph I Sph I Not I EcoR I Spe I Spe I Spe I Spe I Aat II Puro Sph I Sph I Not I Not I cyclin E2 targeting strategy Aat II 1kb WT E2 -/- E2 +/- 9.0kb with Bruno Amati
13 Cyclin E2 -/- mice viable some males sterile testicular hypoplasia reduced sperm counts
14 cdk1 B M G1 D1 Cdk4/6 E1 D2 Cdk4/6 Cdk4/6 D3 E2 E2 A2 A1 G2 S E2 A2 A1
15 Cyclin E1 -/- E2 -/- Embryonic lethal
16 Cyclin E1 -/- E2 -/- embryos Observed Expected E /111 7/111 E (13)/168 16/168 E11.5 3(6)/166 18/166 E12.5 (4)/82 10/82
17 CYCLIN E1 -/- E2 -/- EMBRYOS Control E1 -/- E2 -/- day E 10.75
18 Phenotype of trophoblast giant cells WT *
19 Phenotype of trophoblast giant cells WT * * E1 -/- E2 -/-
20 ENDOREPLICATION G S
21 % of nuclei Phenotype of trophoblast giant cells 80 WT E1 -/- E2 -/- * * E1 -/- E2 -/- WT < >900 Ploidy (n)
22 Cyclin E1 -/- E2 -/- mice E9.0 - E10.75 Normal embryo development Abnormal placental development Paucity of trophoblast giant cells Endoreplication problem
23 Mate E1 +/- E2 +/- mice E1 -/- E2 -/- blastocysts E1 -/- E2 -/- ES cells
24 2n 2n 4n 4n
25 2n 2n 4n E1 -/- E2 -/- ES Cells 4n
26 Rescued cyclin E-null neonates WT E1 -/- E2 -/-
27 Cardiac abnormalities in cyclin E-null neonates WT WT E1 -/- E2 -/- E1 -/- E2 -/-
28 Cyclin E is largely dispensable for development
29 cdk1 B M G1 D1 Cdk4/6 E1 D2 Cdk4/6 Cdk4/6 D3 E2 E2 A2 A1 G2 S E2 A2 A1
30 cdk1 B M G1 D1 Cdk4/6 E1 D2 Cdk4/6 Cdk4/6 D3 E2 E2 A1 cdk1 A2 cdk1 G2 S E2 A1 A2
31 Role of cyclin A in the cell cycle Regulate DNA synthesis induce the initiation of DNA replication prevent re-replication Plays a role in centrosome duplication Entry into mitosis
32 A1 A2
33 Expression pattern of cyclins A1 and A2 A1 A2 C. Sweeney et al., 1996
34 A1 A1 A2 A2 A1 Testis
35 cyclin A2 is an essential component of all embryonic and somatic cell cycles in mammals there are essentially two types of cyclins: those that are essential components of the cell cycle engine [cyclins A and B], and those involved in cell type specific regulation of the cycle [G1 cyclins]
36 Consequence of cyclin A2 deletion in mice Ccna2 -/- mouse fail to develop past 5.5 days pc (Murphy at al., 1997)
37 Conditional knockout of cyclin A2 A2
38 Cyclin A2 conditional construct E1 E2 E3 E4 E5 E6 E7 E8 lox P lox P pgk Neo ATG FRT FRT In vitro flp mediated recombination STOP In vitro or in vivo CRE mediated recombination E1 E8
39 Cyclin A2 expression level after Cre mediated recombination A2 A2 -actin β-actin
40 Absorbance at 595 nm In vitro proliferation of cyclin A deficient fibroblasts 0,8 0,7 A2 F/F A2 / 0,7 0,6 A1 -/- A2 F/F A1 -/- A2 / 0,6 0,5 0,5 0,4 0,4 0,3 0,3 0,2 0,2 0,1 0, Days Days
41 Re-entry of cell cycle by cyclin A deficient fibroblasts [3H]-thymidine incorporation x 10 3 [cpm] 3 2,5 2 1,5 1 0,5 A2 F/F A2 / 5 4,5 4 3,5 3 2,5 2 1,5 1 0,5 A1 -/- A2 F/F A1 -/- A2 / Hours Hours
42 Cell cycle profile of MEFs lacking cyclin A2 A2 F/F G1: 48.2% S: 19.4 % G2/M: 25 % A2 / G1: 32.2 % S: 25.9 % G2/M: 29.4 %
43 A1 -/- x A2 F/F D. Wolgemuth
44 Cyclin A2 expression level after Cre mediated recombination A2 actin
45 Absorbance at 595 nm In vitro proliferation of cyclin A deficient fibroblasts 0,8 0,7 A2 F/F A2 / 0,7 0,6 A1 -/- A2 F/F A1 -/- A2 / 0,6 0,5 0,5 0,4 0,4 0,3 0,3 0,2 0,2 0,1 0, Days Days
46 Re-entry of cell cycle by cyclin A deficient fibroblasts [3H]-thymidine incorporation x 10 3 [cpm] 3 2,5 2 1,5 1 0,5 A2 F/F A2 / 5 4,5 4 3,5 3 2,5 2 1,5 1 0,5 A1 -/- A2 F/F A1 -/- A2 / Hours Hours
47 Cell cycle profile of MEF lacking cyclins A1 & A2 A1 -/- A2 F/F G1: % S: 18.2 % G2/M: 21.6 % A1 -/- A2 G1: 39 % S: 22.6 % G2/M: 30 %
48 % of cells P< P< Control Ряд P= P= P= Ряд2 A2 / A1 -/- A2 / Ряд P= G1 S G2/M
49 G1 Control S G2/M G1 S G2/M A1 -/- A2
50 Absorbance at 595 nm In vitro proliferation of cyclin A deficient fibroblasts 0,8 0,7 A2 F/F A2 / 0,7 0,6 A1 -/- A2 F/F A1 -/- A2 / 0,6 0,5 0,5 0,4 0,4 0,3 0,3 0,2 0,2 0,1 0, Days Days
51 cdk1 B M G1 D1 Cdk4/6 E1 D2 Cdk4/6 Cdk4/6 D3 E2 E2 A1 cdk1 A2 cdk1 G2 S E2 A1 A2
52 The levels of cell cycle regulators in cyclin A-deficient fibroblasts D1 D2 Cdk1 Cdk2 Cdk4 Cdk6 p27 E1 A2 β-actin actin E1 A2 β-actin
53 cyc A cyc E cyc D G1 S G2 M
54 Relative level The levels of cell cycle regulators in cyclin A-deficient fibroblasts D1 D2 D3 E1 A2 Control A1 -/- A hours 1,2 1 0,8 0,6 0,4 0,2 Cyclin E level in control Cyclin E level in A1 -/- A2 Cyclin A2 level in control Ряд1 Ряд2 Ряд Hours
55 Cyclin A-Cdk and E-cdk complexes E E cdk1 A A cdk1
56 Cyclin A-Cdk and E-cdk complexes E E cdk1 Cdk1-kinase Cdk2-kinase A A cdk1
57 prb T821 NPM T199 B-Myb S581 B-Myb T490 Cdc6 S54 Smad3 T8 Smad3 S212 RRM2 p107 PCNA Mcm2 E2F1 B-Myb β-actin
58 cdk1 B M G1 D1 Cdk4/6 E1 D2 Cdk4/6 Cdk4/6 D3 E2 E2 A1 cdk1 A2 cdk1 G2 S E2 A1 A2
59 -/F/F A1 A2 -/-/E1 E2
60 E1 E2
61 E1 E2
62 Cyclin E1 -/- E2 -/- day E Yan Geng
63 CYCLIN E1 -/- E2 -/- MICE Control E1 -/- E2 -/-
64 Mate E1 +/- E2 +/- mice E1 -/- E2 -/- blastocysts E1 -/- E2 -/- ES cells
65 2n 2n 4n 4n
66 2n 2n 4n E1 -/- E2 -/- ES Cells 4n
67 A1 -/- A2 F/F E1 -/- E2 +/- x A1 +/- A2 F/F E1 +/- E2 +/- A1 -/- A2 F/F E1 -/- E2 -/- A1 -/- A2 F/F E1 -/- E2 -/- ES cells 4n blastocyst A1 -/- A2 F/F E1 -/- E2 -/- MEFs + Cre A1 -/- A2 E1 -/- E2 -/- MEFs A2
68 BrdU + cells [%] Analysis of fibroblasts lacking all A- and E-type cyclins Cell number x 10 5 [3H]-thymidine incorporation x 10 3 [cpm] Control A1 + Cre + Control + Cre -/- A2 / E1 -/- E2 -/- A1 -/- A2 f/f E1 -/- E2 -/- A1 -/- A2 f/f E1 -/- E2 -/- A1 -/- A2 f/f E1 -/- E2 -/- 44% % 14% % 40% 16%
69 Analysis of fibroblasts lacking all A- and E-type cyclins 7 A1 -/- A2 f/f E1 -/- E2 -/- A1 -/- A2 / E1 -/- E2 -/- 6 Cell number x Days
70 Cdk1-kinase Cdk1-kinase Cdk2-kinase Cdk2-kinase
71 cdk1 B M G1 D1 Cdk4/6 E1 D2 Cdk4/6 Cdk4/6 D3 E2 E2 A1 cdk1 A2 cdk1 G2 S E2 A1 A2
72 Debra Wolgemuth Columbia Hirokazu Shigematsu Tadafumi Iino Shin-ichi Mizuno Koichi Akashi
73 Consequence of cyclin A deletion in vivo Bone marrow specific deletion in Mx A1 -/- A2 F/F mice
74 Cyclin A2 expression level in A2 F/F Mx1 mice Before and after poly(i) poly (C) treatment A2 A2 β-actin A2 F/F WT
75 % Survival Kaplan-Meier survival graph 100 Mx control n=18 25 Mx P<0,0001 A2 Mx A1 -/- A Days Time after 6x pipc injection
76 Cell number x 10 6 /ul Cell number x 10 3 /ul Cell number x 10 5 /ul Сотни Peripheral blood analysis 12 RBC WBC PLT Mx control Mx A2 Mx A1 -/- A n=
77 Analysis of bone marrow cells Cell number x 10 6 / 2 femurs Mx control Mx A2 Mx A1 -/- A2 n= H&E
78 Cell number x 10 6 / femur Analysis of BM subpopulations Mx control 25 Mx A Mx A1 -/- A2 n= Gr-1 Mac-1 Ter119 B220 CD3 Granulocytes Monocytes Erythroid lineage B cells T cells
79 At which stage of hematopoiesis cyclin A is rate-limiting?
80 Cell number x 10 3 / 2 femurs Cell number x 10 3 / 2 femurs Analysis of hematopoetic stem cells and progenitors Mx control Mx A2 80 Mx A1 -/- A HSC CMP GMP MEP Tadafumi Iino Shin-ichi Mizuno Koichi Akashi
81 Gr-1 Gr-1 Gr-1 Analysis of progenitors Mx control Mx A2 Mx A1 -/- A Mac Mac Mac-1 Tadafumi Iino Shin-ichi Mizuno Koichi Akashi
82 Number of colonies Analysis of Hematopoetic Stem Cells Before Cre A1 +/- A2 f/+ A1 -/- A2 f/f A A1 +/- A1 +/- A2 /+ A1 -/- A1 -/- A2 F/+ A2 F/ A2 A2 + After Cre A1 +/- A2 /+ A1 -/- A2 f/ A2 f A1 +/- A2 F/+ +Cre A1 -/- A2 F/F Tadafumi Iino Shin-ichi Mizuno Koichi Akashi
83 Embryonic stem cells A1 -/- A2 F/F
84 Embryonal stem cells A1 -/- A2 F/F +Cre
85 Number of colonies Analysis of embryonal stem cells A1 +/- A2 f/+ A1 +/- A2 /+ A1 -/- A1 -/- A2 f/ 0 0 A1 -/- A2 / A1 -/- A1 -/- A2 f/f A2 f/ A2 / +Cre +Cre A1 +/- A2 f/+ A1 -/- A2 f/f A1 -/- A2 f/
86 Proliferation of MEFs does NOT depend on cyclin A but Proliferation (survival) of HSC and ES cells critically depends on cyclin A What is the molecular basis of this difference?
87 Mammalian cell cycle in stem cells Rb E2F A cdk1 cdk1 B M G2 G1 S D Cdk4/6 E E2F P P P Rb E2F P P P P Rb P P A
88 Debra Wolgemuth Columbia Hirokazu Shigematsu Tadafumi Iino Shin-ichi Mizuno Koichi Akashi
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