Preventing re-replication of chromosomal DNA (original) (raw)

1. Rao PN, Johnson RT. Mammalian cell fusion: studies on the regulation of DNA synthesis and mitosis. Nature. 1970;225:159–164. [PubMed] [Google Scholar]

2. Blow JJ, Laskey RA. A role for the nuclear envelope in controlling DNA replication within the cell cycle. Nature. 1988;332:546–548. [PubMed] [Google Scholar]

3. Blow JJ, Hodgson B. Replication licensing - defining the proliferative state? Trends Cell Biol. 2002;12:72–78. [PMC free article] [PubMed] [Google Scholar]

4. Nishitani H, Lygerou Z. DNA replication licensing. Frontiers in Bioscience. 2004;9:2115–2132. [PubMed] [Google Scholar]

5. Ishimi Y. A DNA helicase activity is associated with an MCM4, -6, and -7 protein complex. J. Biol. Chem. 1997;272:24508–24513. [PubMed] [Google Scholar]

6. Prokhorova TA, Blow JJ. Sequential MCM/P1 subcomplex assembly is required to form a heterohexamer with replication licensing activity. J. Biol. Chem. 2000;275:2491–2498. [PMC free article] [PubMed] [Google Scholar]

7. Schwacha A, Bell SP. Interactions between two catalytically distinct MCM subgroups are essential for coordinated ATP hydrolysis and DNA replication. Mol. Cell. 2001;8:1093–1104. [PubMed] [Google Scholar]

8. Aparicio OM, Weinstein DM, Bell SP. Components and dynamics of DNA replication complexes in S- cerevisiae: Redistribution of MCM proteins and Cdc45p during S phase. Cell. 1997;91:59–69. [PubMed] [Google Scholar]

9. Labib K, Tercero JA, Diffley JFX. Uninterrupted MCM2-7 function required for DNA replication fork progression. Science. 2000;288:1643–1647. [PubMed] [Google Scholar]

10. Pacek M, Walter JC. A requirement for MCM7 and Cdc45 in chromosome unwinding during eukaryotic DNA replication. EMBO J. 2004;23:3667–76. Epub 2004 Aug 26. [PMC free article] [PubMed] [Google Scholar]

11. Shechter D, Ying CY, Gautier J. DNA unwinding is an Mcm complex-dependent and ATP hydrolysis-dependent process. J Biol Chem. 2004;279:45586–93. Epub 2004 Aug 23. [PubMed] [Google Scholar]

12. Kelman Z, Lee JK, Hurwitz J. The single minichromosome maintenance protein of Methanobacterium thermoautotrophicum DeltaH contains DNA helicase activity. Proc. Natl. Acad. Sci. USA. 1999;96:14783–14788. [PMC free article] [PubMed] [Google Scholar]

13. Chong JP, Hayashi MK, Simon MN, Xu RM, Stillman B. A double-hexamer archaeal minichromosome maintenance protein is an ATP-dependent DNA helicase. Proc. Natl. Acad. Sci. USA. 2000;97:1530–1535. [PMC free article] [PubMed] [Google Scholar]

14. Shechter DF, Ying CY, Gautier J. The intrinsic DNA helicase activity of Methanobacterium thermoautotrophicum delta H minichromosome maintenance protein. J. Biol. Chem. 2000;275:15049–15059. [PubMed] [Google Scholar]

15. Fletcher RJ, et al. The structure and function of MCM from archaeal M. Thermoautotrophicum. Nature Structural Biology. 2003;10:160–167. [PubMed] [Google Scholar]The crystal structure of an archaeal MCM reveals a dodecameric complex providing a positively-charged central channel capable of accommodating double-stranded DNA.

16. Pape T, et al. Hexameric ring structure of the full-length archaeal MCM protein complex. EMBO Rep. 2003;4:1079–1083. [PMC free article] [PubMed] [Google Scholar]A 3-dimensional reconstruction of an archaeal MCM shows a structure with a central channel, consistent both with the MCM crystal structure of Fletcher et al and the structure of SV40 T antigen (Li et al).

17. Li D, et al. Structure of the replicative helicase of the oncoprotein SV40 large tumour antigen. Nature. 2003;423:512–518. [PubMed] [Google Scholar]The crystal structure of SV40 T antigen shows hexamers organised into two tiers that enclose a positively-charged channel capable of accommodating double-stranded DNA.

18. Wessel R, Schweizer J, Stahl H. Simian virus 40 T-antigen DNA helicase is a hexamer which forms a binary complex during bidirectional unwinding from the viral origin of DNA replication. J. Virol. 1992;66:804–815. [PMC free article] [PubMed] [Google Scholar]

19. Burkhart R, et al. Interactions of human nuclear proteins P1Mcm3 and P1Cdc46. Eur. J. Biochem. 1995;228:431–438. [PubMed] [Google Scholar]

20. Lei M, Kawasaki Y, Tye BK. Physical interactions among Mcm proteins and effects of Mcm dosage on DNA replication in Saccharomyces cerevisiae. Mol. Cell. Biol. 1996;16:5081–5090. [PMC free article] [PubMed] [Google Scholar]

21. Mahbubani HM, Chong JP, Chevalier S, Thömmes P, Blow JJ. Cell cycle regulation of the replication licensing system: involvement of a Cdk-dependent inhibitor. J. Cell Biol. 1997;136:125–135. [PMC free article] [PubMed] [Google Scholar]

22. Edwards MC, et al. MCM2-7 complexes bind chromatin in a distributed pattern surrounding the origin recognition complex in Xenopus egg extracts. J. Biol. Chem. 2002;277:33049–33057. [PubMed] [Google Scholar]

23. Ritzi M, et al. Human minichromosome maintenance proteins and human origin recognition complex 2 protein on chromatin. J. Biol. Chem. 1998;273:24543–24549. [PubMed] [Google Scholar]

24. Romanowski P, Madine MA, Rowles A, Blow JJ, Laskey RA. The Xenopus origin recognition complex is essential for DNA replication and MCM binding to chromatin. Curr. Biol. 1996;6:1416–1425. [PubMed] [Google Scholar]

25. Harvey KJ, Newport J. CpG methylation of DNA restricts prereplication complex assembly in Xenopus egg extracts. Mol. Cell. Biol. 2003;23:6769–6779. [PMC free article] [PubMed] [Google Scholar]

26. Danis E, et al. Specification of a DNA replication origin by a transcription complex. Nat. Cell Biol. 2004;6:721–730. [PubMed] [Google Scholar]Shows that initiation sites in Xenopus egg extracts can be induced by creating a transcription domain, possibly by inducing histone acetylation at that site.

27. Bowers JL, Randell JC, Chen S, Bell SP. ATP hydrolysis by ORC catalyzes reiterative Mcm2-7 assembly at a defined origin of replication. Mol. Cell. 2004;16:967–978. [PubMed] [Google Scholar]

28. Madine MA, Khoo CY, Mills AD, Musahl C, Laskey RA. The nuclear envelope prevents reinitiation of replication by regulating the binding of MCM3 to chromatin in Xenopus egg extracts. Curr. Biol. 1995;5:1270–1279. [PubMed] [Google Scholar]

29. Krude T, Musahl C, Laskey RA, Knippers R. Human replication proteins hCdc21, hCdc46 and P1Mcm3 bind chromatin uniformly before S-phase and are displaced locally during DNA replication. J. Cell Sci. 1996;109:309–318. [PubMed] [Google Scholar]

30. Dimitrova DS, Todorov IT, Melendy T, Gilbert DM. Mcm2, but not RPA, is a component of the mammalian early G1-phase prereplication complex. J. Cell Biol. 1999;146:709–722. [PMC free article] [PubMed] [Google Scholar]

31. Laskey RA, Madine MA. A rotary pumping model for helicase function of MCM proteins at a distance from replication forks. EMBO Rep. 2003;4:26–30. [PMC free article] [PubMed] [Google Scholar] Proposes a solution to the ‘MCM paradox’ whereby Mcm2-7 act at a distance from the replication fork unwinding DNA by a rotary pump mechanism.

32. Yankulov K, et al. MCM proteins are associated with RNA polymerase II holoenzyme. Mol. Cell. Biol. 1999;19:6154–6163. [PMC free article] [PubMed] [Google Scholar]

33. Dziak R, Leishman D, Radovic M, Tye BK, Yankulov K. Evidence for a role of MCM (mini-chromosome maintenance)5 in transcriptional repression of sub-telomeric and Ty-proximal genes in Saccharomyces cerevisiae. J. Biol. Chem. 2003;278:27372–27381. [PubMed] [Google Scholar]

34. Oehlmann M, Score AJ, Blow JJ. The role of Cdc6 in ensuring complete genome licensing and S phase checkpoint activation. J. Cell Biol. 2004;165:181–190. [PMC free article] [PubMed] [Google Scholar] Shows that the affinity of Xenopus Cdc6 for chromatin drops once the origin has loaded the first Mcm2-7 hexamers, potentially providing a mechanism to ensure that all origins are licensed.

35. Cortez D, Glick G, Elledge SJ. Minichromosome maintenance proteins are direct targets of the ATM and ATR checkpoint kinases. Proc. Natl. Acad. Sci. USA. 2004;101:10078–10083. [PMC free article] [PubMed] [Google Scholar]

37. Mendez J, Stillman B. Perpetuating the double helix: molecular machines at eukaryotic DNA replication origins. Bioessays. 2003;25:1158–1167. [PubMed] [Google Scholar]

38. Gillespie PJ, Li A, Blow JJ. Reconstitution of licensed replication origins on Xenopus sperm nuclei using purified proteins. BMC Biochem. 2001;2:15. [PMC free article] [PubMed] [Google Scholar]

39. Donovan S, Harwood J, Drury LS, Diffley JF. Cdc6p-dependent loading of Mcm proteins onto pre-replicative chromatin in budding yeast. Proc. Natl. Acad. Sci. USA. 1997;94:5611–5616. [PMC free article] [PubMed] [Google Scholar]

40. Hua XH, Newport J. Identification of a preinitiation step in DNA replication that is independent of origin recognition complex and cdc6, but dependent on cdk2. J. Cell Biol. 1998;140:271–281. [PMC free article] [PubMed] [Google Scholar]

41. Rowles A, Tada S, Blow JJ. Changes in association of the Xenopus origin recognition complex with chromatin on licensing of replication origins. J. Cell Sci. 1999;112:2011–2018. [PMC free article] [PubMed] [Google Scholar]

42. Maiorano D, Moreau J, Mechali M. XCDT1 is required for the assembly of pre-replicative complexes in Xenopus laevis. Nature. 2000;404:622–625. [PubMed] [Google Scholar]

43. Tanaka S, Diffley JF. Interdependent nuclear accumulation of budding yeast Cdt1 and Mcm2-7 during G1 phase. Nat. Cell Biol. 2002;4:198–207. [PubMed] [Google Scholar]

44. Yanagi K, Mizuno T, You Z, Hanaoka F. Mouse geminin inhibits not only Cdt1-MCM6 interactions but also a novel intrinsic Cdt1 DNA binding activity. J. Biol. Chem. 2002;277:40871–40880. [PubMed] [Google Scholar]

45. Cook JG, Chasse DA, Nevins JR. The regulated association of Cdt1 with minichromosome maintenance proteins and Cdc6 in mammalian cells. J. Biol. Chem. 2004;279:9625–9633. [PubMed] [Google Scholar]

46. Tsuyama T, Tada S, Watanabe S, Seki M, Enomoto T. Licensing for DNA replication requires a strict sequential assembly of Cdc6 and Cdt1 onto chromatin in Xenopus egg extracts. Nucl. Acids Res. 2005;33:765–775. [PMC free article] [PubMed] [Google Scholar]

47. Tada S, Li A, Maiorano D, Mechali M, Blow JJ. Repression of origin assembly in metaphase depends on inhibition of RLF-B/Cdt1 by geminin. Nat. Cell Biol. 2001;3:107–113. [PMC free article] [PubMed] [Google Scholar]

48. Wohlschlegel JA, et al. Inhibition of eukaryotic replication by geminin binding to Cdt1. Science. 2000;290:2309–2312. [PubMed] [Google Scholar]

49. Lee C, et al. Structural basis for inhibition of the replication licensing factor Cdt1 by geminin. Nature. 2004;430:913–917. [PubMed] [Google Scholar]Shows the crystal structure of a complex between portions of Cdt1 and geminin, and suggests how the binding of geminin hinders the association of Mcm2-7 with the C-terminus of Cdt1.

50. Leatherwood J, Vas A. Connecting ORC and heterochromatin: why? Cell Cycle. 2003;2:573–575. [PubMed] [Google Scholar]

51. Clay-Farrace L, Pelizon C, Santamaria D, Pines J, Laskey RA. Human replication protein Cdc6 prevents mitosis through a checkpoint mechanism that implicates Chk1. EMBO J. 2003;22:704–712. [PMC free article] [PubMed] [Google Scholar]

52. Murakami H, Yanow SK, Griffiths D, Nakanishi M, Nurse P. Maintenance of replication forks and the S-phase checkpoint by Cdc18p and Orp1p. Nat. Cell Biol. 2002;4:384–388. [PubMed] [Google Scholar]

53. Broek D, Bartlett R, Crawford K, Nurse P. Involvement of p34cdc2 in establishing the dependency of S phase on mitosis. Nature. 1991;349:388–393. [PubMed] [Google Scholar]

54. Hayles J, Fisher D, Woollard A, Nurse P. Temporal order of S phase and mitosis in fission yeast is determined by the state of the p34cdc2-mitotic B cyclin complex. Cell. 1994;78:813–822. [PubMed] [Google Scholar]

55. Dahmann C, Diffley J, Nasmyth K. S-phase-promoting cyclin-dependent kinases prevent re-replication by inhibiting the transition of replication origins to a pre-replicative state. Curr. Biol. 1995;5:1257–1269. [PubMed] [Google Scholar]

56. Diffley JF. Once and only once upon a time: specifying and regulating origins of DNA replication in eukaryotic cells. Genes and Development. 1996;10:2819–2830. [PubMed] [Google Scholar]

57. Jallepalli PV, Brown GW, MuziFalconi M, Tien D, Kelly TJ. Regulation of the replication initiator protein p65(cdc18) by CDK phosphorylation. Genes Dev. 1997;11:2767–2779. [PMC free article] [PubMed] [Google Scholar]

58. Elsasser S, Chi Y, Yang P, Campbell JL. Phosphorylation controls timing of Cdc6p destruction: A biochemical analysis. Mol. Biol. Cell. 1999;10:3263–3277. [PMC free article] [PubMed] [Google Scholar]

59. Drury LS, Perkins G, Diffley JFX. The cyclin-dependent kinase Cdc28p regulates distinct modes of Cdc6p proteolysis during the budding yeast cell cycle. Curr. Biol. 2000;10:231–240. [PubMed] [Google Scholar]

60. Nguyen VQ, Co C, Li JJ. Cyclin-dependent kinases prevent DNA re-replication through multiple mechanisms. Nature. 2001;411:1068–1073. [PubMed] [Google Scholar]

61. Vas A, Mok W, Leatherwood J. Control of DNA rereplication via Cdc2 phosphorylation sites in the origin recognition complex. Mol Cell Biol. 2001;21:5767–5777. [PMC free article] [PubMed] [Google Scholar]

62. Wuarin J, Buck V, Nurse P, Millar JB. Stable association of mitotic cyclin B/Cdc2 to replication origins prevents endoreduplication. Cell. 2002;111:419–431. [PubMed] [Google Scholar]

63. Wilmes GM, et al. Interaction of the S-phase cyclin Clb5 with an “RXL” docking sequence in the initiator protein Orc6 provides an origin-localized replication control switch. Genes Dev. 2004;18:981–991. [PMC free article] [PubMed] [Google Scholar] Shows that in S. cerevisiae, Orc6 can bind the Clb5 cyclin, and that blocking the interaction permits re-replication of DNA.

64. Mimura S, Seki T, Tanaka S, Diffley JF. Phosphorylation-dependent binding of mitotic cyclins to Cdc6 contributes to DNA replication control. Nature. 2004;431:1118–23. Epub 2004 Oct 20. [PubMed] [Google Scholar]

65. Nishitani H, Lygerou Z, Nishimoto T, Nurse P. The Cdt1 protein is required to license DNA for replication in fission yeast. Nature. 2000;404:625–628. [PubMed] [Google Scholar]

66. Labib K, Diffley JFX, Kearsey SE. G1-phase and B-type cyclins exclude the DNA-replication factor Mcm4 from the nucleus. Nat. Cell Biol. 1999;1:415–422. [PubMed] [Google Scholar]

67. Nguyen VQ, Co C, Irie K, Li JJ. Clb/Cdc28 kinases promote nuclear export of the replication initiator proteins Mcm2-7. Curr. Biol. 2000;10:195–205. [PubMed] [Google Scholar]

68. Follette PJ, Duronio RJ, O’Farrell PH. Fluctuations in cyclin E levels are required for multiple rounds of endocycle S phase in Drosophila. Curr. Biol. 1998;8:235–238. [PMC free article] [PubMed] [Google Scholar]

69. Su TT, O’Farrell PH. Chromosome association of minichromosome maintenance proteins in Drosophila endoreplication cycles. J. Cell Biol. 1998;140:451–460. [PMC free article] [PubMed] [Google Scholar]

70. Dhar SK, Delmolino L, Dutta A. Architecture of the human origin recognition complex. J. Biol. Chem. 2001;276:29067–29071. [PubMed] [Google Scholar]

71. Vashee S, Simancek P, Challberg MD, Kelly TJ. Assembly of the human origin recognition complex. J. Biol. Chem. 2001;276:26666–26673. [PubMed] [Google Scholar]

72. Mendez J, et al. Human origin recognition complex large subunit is degraded by ubiquitin-mediated proteolysis after initiation of DNA replication. Mol. Cell. 2002;9:481–491. [PubMed] [Google Scholar]

73. Li CJ, DePamphilis ML. Mammalian Orc1 protein is selectively released from chromatin and ubiquitinated during the S-to-M transition in the cell division cycle. Mol. Cell. Biol. 2002;22:105–116. [PMC free article] [PubMed] [Google Scholar]

74. Li CJ, Vassilev A, DePamphilis ML. Role for Cdk1 (Cdc2)/cyclin A in preventing the mammalian origin recognition complex’s largest subunit (Orc1) from binding to chromatin during mitosis. Molecular and Cellular Biology. 2004;24:5875–5886. [PMC free article] [PubMed] [Google Scholar]

75. Coverley D, Pelizon C, Trewick S, Laskey RA. Chromatin-bound Cdc6 persists in S and G(2) phases in human cells, while soluble Cdc6 is destroyed in a cyclin A-cdk2 dependent process. J. Cell Sci. 2000;113:1929–1938. [PubMed] [Google Scholar]

76. Mendez J, Stillman B. Chromatin association of human origin recognition complex, Cdc6, and minichromosome maintenance proteins during the cell cycle: Assembly of prereplication complexes in late mitosis. Mol. Cell. Biol. 2000;20:8602–8612. [PMC free article] [PubMed] [Google Scholar]

77. Saha P, et al. Human CDC6/Cdc18 associates with Orc1 and cyclin-cdk and is selectively eliminated from the nucleus at the onset of S phase. Mol. Cell. Biol. 1998;18:2758–2767. [PMC free article] [PubMed] [Google Scholar]

78. Petersen BO, Lukas J, Sorensen CS, Bartek J, Helin K. Phosphorylation of mammalian CDC6 by Cyclin A/CDK2 regulates its subcellular localization. EMBO J. 1999;18:396–410. [PMC free article] [PubMed] [Google Scholar]

79. Vaziri C, et al. A p53-dependent checkpoint pathway prevents rereplication. Mol. Cell. 2003;11:997–1008. [PubMed] [Google Scholar]Shows that overexpression of Cdt1 and Cdc6 can cause extensive re-replication in human cells, and that as a consequence, a ATM/ATR and p53-dependent checkpoint pathways are activated.

80. Alexandrow MG, Hamlin JL. Cdc6 chromatin affinity is unaffected by serine-54 phosphorylation, S-phase progression, and overexpression of cyclin A. Mol. Cell. Biol. 2004;24:1614–1627. [PMC free article] [PubMed] [Google Scholar]

81. Ballabeni A, et al. Human Geminin promotes pre-RC formation and DNA replication by stabilizing CDT1 in mitosis. EMBO J. 2004;23:3122–3132. [PMC free article] [PubMed] [Google Scholar] Describes the effect of geminin in protecting Cdt1 from proteasome-mediated degradation on exit from mitosis, and shows that CDK inhibition during mitosis is sufficient to promote premature licensing.

82. Nishitani H, Taraviras S, Lygerou Z, Nishimoto T. The human licensing factor for DNA replication Cdt1 accumulates in G1 and is destabilized after initiation of S-phase. J. Biol. Chem. 2001;276:44905–44911. [PubMed] [Google Scholar]

83. Quinn LM, Herr A, McGarry TJ, Richardson H. The Drosophila Geminin homolog: roles for Geminin in limiting DNA replication, in anaphase and in neurogenesis. Genes Dev. 2001;15:2741–2754. [PMC free article] [PubMed] [Google Scholar]

84. Hodgson B, Li A, Tada S, Blow JJ. Geminin becomes activated as an inhibitor of Cdt1/RLF-B following nuclear import. Curr. Biol. 2002;12:678–683. [PMC free article] [PubMed] [Google Scholar]

85. Li X, Zhao Q, Liao R, Sun P, Wu X. The SCF(Skp2) ubiquitin ligase complex interacts with the human replication licensing factor Cdt1 and regulates Cdt1 degradation. J. Biol. Chem. 2003;278:30854–30858. [PubMed] [Google Scholar] Provides evidence that down-regulation of Cdt1 late in the cell cycle involves SCF-dependent ubiquitination of phosphorylated Cdt1.

86. Zhong W, Feng H, Santiago FE, Kipreos ET. CUL-4 ubiquitin ligase maintains genome stability by restraining DNA-replication licensing. Nature. 2003;423:885–889. [PubMed] [Google Scholar] Shows that the C. elegans CUL-4 ubiquitin ligase is responsible for down-regulating Cdt1 late in the cell cycle, and that failure of this process can cause re-replication of DNA.

87. Liu E, Li X, Yan F, Zhao Q, Wu X. Cyclin-dependent kinases phosphorylate human Cdt1 and induce its degradation. J. Biol. Chem. 2004;279:17283–17288. [PubMed] [Google Scholar]

88. Nishitani H, Lygerou Z, Nishimoto T. Proteolysis of DNA replication licensing factor Cdt1 in S-phase is performed independently of geminin through its N-terminal region. J. Biol. Chem. 2004;279:30807–30816. [PubMed] [Google Scholar]

89. Sugimoto N, et al. Cdt1 phosphorylation by cyclin A-dependent kinases negatively regulates its function without affecting geminin binding. J. Biol. Chem. 2004;279:19691–19697. [PubMed] [Google Scholar]

90. Arias EE, Walter JC. Replication-dependent destruction of Cdt1 limits DNA replication to a single round per cell cycle in Xenopus egg extracts. Genes Dev. 2005;19:114–26. Epub 2004 Dec 14. [PMC free article] [PubMed] [Google Scholar]

91. Thomer M, May NR, Aggarwal BD, Kwok G, Calvi BR. Drosophila double-parked is sufficient to induce re-replication during development and is regulated by cyclin E/CDK2. Development. 2004;131:4807–18. Epub 2004 Sep 01. [PubMed] [Google Scholar]

92. Li A, Blow JJ. Cdt1 downregulation by proteolysis and geminin inhibition prevents DNA re-replication in Xenopus. EMBO J. 2005;24:395–404. [PMC free article] [PubMed] [Google Scholar]Shows that in Xenopus, activation of the licensing system on exit from metaphase depends on the ubiquitination of geminin which renders it incapable of inhibiting Cdt1.

93. Maiorano D, Krasinska L, Lutzmann M, Mechali M. Recombinant cdt1 induces rereplication of G2 nuclei in Xenopus egg extracts. Curr Biol. 2005;15:146–53. [PubMed] [Google Scholar]

94. Yoshida K, Takisawa H, Kubota Y. Intrinsic nuclear import activity of geminin is essential to prevent re-initiation of DNA replication in Xenopus eggs. Genes Cells. 2005;10:63–73. [PubMed] [Google Scholar]

95. McGarry TJ, Kirschner MW. Geminin, an inhibitor of DNA replication, is degraded during mitosis. Cell. 1998;93:1043–1053. [PubMed] [Google Scholar]

96. Li A, Blow JJ. Non-proteolytic inactivation of geminin requires CDK-dependent ubiquitination. Nature Cell Biology. 2004;6:260–267. [PMC free article] [PubMed] [Google Scholar]

97. Mihaylov IS, et al. Control of DNA replication and chromosome ploidy by geminin and cyclin A. Mol. Cell. Biol. 2002;22:1868–1880. [PMC free article] [PubMed] [Google Scholar]

98. Melixetian M, et al. Loss of Geminin induces rereplication in the presence of functional p53. J. Cell Biol. 2004;165:473–482. [PMC free article] [PubMed] [Google Scholar] This paper, along with that of Zhu et al, shows that in human cells, ablation of geminin is sufficient to induce re-replication of DNA, and that as a consequence p53-independent checkpoint pathways are activated that block subsequent entry into mitosis.

99. Zhu W, Chen Y, Dutta A. Rereplication by depletion of geminin is seen regardless of p53 status and activates a G2/M checkpoint. Mol. Cell. Biol. 2004;24:7140–7150. [PMC free article] [PubMed] [Google Scholar]See Melixetian et al.

100. Del Bene F, Tessmar-Raible K, Wittbrodt J. Direct interaction of geminin and Six3 in eye development. Nature. 2004;427:745–749. [PubMed] [Google Scholar]

101. Luo L, Yang X, Takihara Y, Knoetgen H, Kessel M. The cell-cycle regulator geminin inhibits Hox function through direct and polycomb-mediated interactions. Nature. 2004;427:749–753. [PubMed] [Google Scholar]

102. Benjamin JM, Torke SJ, Demeler B, McGarry TJ. Geminin has dimerization, Cdt1-binding, and destruction domains that are required for biological activity. J. Biol. Chem. 2004;279:45957–45968. [PubMed] [Google Scholar]

103. Saxena S, et al. A dimerized coiled-coil domain and an adjoining part of geminin interact with two sites on Cdt1 for replication inhibition. Mol. Cell. 2004;15:245–258. [PubMed] [Google Scholar]

104. Thepaut M, et al. Crystal structure of the coiled-coil dimerization motif of geminin: structural and functional insights on DNA replication regulation. J. Mol. Biol. 2004;342:275–287. [PubMed] [Google Scholar]

105. Okorokov AL, et al. Molecular structure of human geminin. Nature Structural and Molecular Biology. 2004;11:1021–1022. [PubMed] [Google Scholar]

106. Green BM, Li JJ. Loss of rereplication control in Saccharomyces cerevisiae results in extensive DNA damage. Mol Biol Cell. 2005;16:421–32. Epub 2004 Nov 10. [PMC free article] [PubMed] [Google Scholar]

107. Higa LA, Mihaylov IS, Banks DP, Zheng J, Zhang H. Radiation-mediated proteolysis of CDT1 by CUL4-ROC1 and CSN complexes constitutes a new checkpoint. Nat. Cell Biol. 2003;5:1008–1015. [PubMed] [Google Scholar]Shows that ionising radiation induces the degradation of Cdt1 in a process depending on the CUL-4-ubiquitin ligase and the COP9-signalosome.

108. Kondo T, et al. Rapid degradation of Cdt1 upon UV-induced DNA damage is mediated by SCFSkp2 complex. J. Biol. Chem. 2004;279:27315–27319. [PubMed] [Google Scholar]

109. Williams GH, et al. Improved cervical smear assessment using antibodies against proteins that regulate DNA replication. Proc. Natl. Acad. Sci. USA. 1998;95:14932–14937. [PMC free article] [PubMed] [Google Scholar]

110. Stoeber K, et al. DNA replication licensing and human cell proliferation. J. Cell Sci. 2001;114:2027–2041. [PubMed] [Google Scholar]

111. Leone G, et al. E2F3 activity is regulated during the cell cycle and is required for the induction of S phase. Genes Dev. 1998;12:2120–2130. [PMC free article] [PubMed] [Google Scholar]

112. Ohtani K, Tsujimoto A, Ikeda M, Nakamura M. Regulation of cell growth-dependent expression of mammalian CDC6 gene by the cell cycle transcription factor E2F. Oncogene. 1998;17:1777–1785. [PubMed] [Google Scholar]

113. Ohtani K, et al. Cell growth-regulated expression of mammalian MCM5 and MCM6 genes mediated by the transcription factor E2F. Oncogene. 1999;18:2299–2309. [PubMed] [Google Scholar]

114. Geng Y, et al. Cyclin E ablation in the mouse. Cell. 2003;114:431–443. [PubMed] [Google Scholar] Shows two specific defects in mice lacking both cyclins E1 and E2: a failure to form various endoreduplicated cell types and an inability of cells to relicense DNA during progression from G0 into S phase.

115. Parisi T, et al. Cyclins E1 and E2 are required for endoreplication in placental trophoblast giant cells. EMBO J. 2003;22:4794–4803. [PMC free article] [PubMed] [Google Scholar]

116. Su TT, O’Farrell PH. Chromosome association of minichromosome maintenance proteins in Drosophila mitotic cycles. J. Cell Biol. 1997;139:13–21. [PMC free article] [PubMed] [Google Scholar]

117. Ortega S, et al. Cyclin-dependent kinase 2 is essential for meiosis but not for mitotic cell division in mice. Nat. Genet. 2003;35:25–31. [PubMed] [Google Scholar]

118. Berthet C, Aleem E, Coppola V, Tessarollo L, Kaldis P. Cdk2 knockout mice are viable. Curr. Biol. 2003;13:1775–1785. [PubMed] [Google Scholar]

119. Coverley D, Laman H, Laskey RA. Distinct roles for cyclins E and A during DNA replication complex assembly and activation. Nat. Cell Biol. 2002;4:523–528. [PubMed] [Google Scholar]

120. Wohlschlegel JA, Kutok JL, Weng AP, Dutta A. Expression of geminin as a marker of cell proliferation in normal tissues and malignancies. Am. J. Pathol. 2002;161:267–273. [PMC free article] [PubMed] [Google Scholar]

121. Xouri G, et al. Cdt1 and geminin are down-regulated upon cell cycle exit and are over-expressed in cancer-derived cell lines. Eur. J. Biochem. 2004;271:3368–3378. [PubMed] [Google Scholar]

122. Shreeram S, Sparks A, Lane DP, Blow JJ. Cell type-specific responses of human cells to inhibition of replication licensing. Oncogene. 2002;21:6624–6632. [PMC free article] [PubMed] [Google Scholar]

123. Feng D, Tu Z, Wu W, Liang C. Inhibiting the expression of DNA replication-initiation proteins induces apoptosis in human cancer cells. Cancer Research. 2003;63:7356–7364. [PubMed] [Google Scholar]