Abstract
The cell cycle checkpoint mechanisms ensure the order of cell cycle events to preserve genomic integrity. Among these, the DNA-replication and DNA-damage checkpoints prevent chromosome segregation when DNA replication is inhibited or DNA is damaged. Recent studies have identified an outline of the regulatory networks for both of these controls, which apparently operate in all eukaryotes. In addition, it appears that these checkpoints have two arrest points, one is just before entry into mitosis and the other is prior to chromosome separation. The former point requires the central cell-cycle regulator Cdc2 kinase, whereas the latter involves several key regulators and substrates of the ubiquitin ligase called the anaphase promoting complex. Linkages between these cell-cycle regulators and several key checkpoint proteins are beginning to emerge. Recent findings on post-translational modifications and protein-protein interactions of the checkpoint proteins provide new insights into the checkpoint responses, although the functional significance of these biochemical properties often remains unclear. We have reviewed the molecular mechanisms acting at the DNA-replication and DNA-damage checkpoints in the fission yeast Schizosaccharomyces pombe, and the modifications of these controls during the meiotic cell cycle. We have made comparisons with the controls in fission yeast and other organisms, mainly the distantly related budding yeast.
Full Text
The Full Text of this article is available as a PDF (284.3 KB).
Selected References
These references are in PubMed. This may not be the complete list of references from this article.
- Amon A., Surana U., Muroff I., Nasmyth K. Regulation of p34CDC28 tyrosine phosphorylation is not required for entry into mitosis in S. cerevisiae. Nature. 1992 Jan 23;355(6358):368–371. doi: 10.1038/355368a0. [DOI] [PubMed] [Google Scholar]
- Aparicio O. M., Weinstein D. M., Bell S. P. Components and dynamics of DNA replication complexes in S. cerevisiae: redistribution of MCM proteins and Cdc45p during S phase. Cell. 1997 Oct 3;91(1):59–69. doi: 10.1016/s0092-8674(01)80009-x. [DOI] [PubMed] [Google Scholar]
- Aves S. J., Durkacz B. W., Carr A., Nurse P. Cloning, sequencing and transcriptional control of the Schizosaccharomyces pombe cdc10 'start' gene. EMBO J. 1985 Feb;4(2):457–463. doi: 10.1002/j.1460-2075.1985.tb03651.x. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Bailis J. M., Roeder G. S. Synaptonemal complex morphogenesis and sister-chromatid cohesion require Mek1-dependent phosphorylation of a meiotic chromosomal protein. Genes Dev. 1998 Nov 15;12(22):3551–3563. doi: 10.1101/gad.12.22.3551. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Beach D., Rodgers L., Gould J. ran1+ controls the transition from mitotic division to meiosis in fission yeast. Curr Genet. 1985;10(4):297–311. doi: 10.1007/BF00365626. [DOI] [PubMed] [Google Scholar]
- Bell S. P., Stillman B. ATP-dependent recognition of eukaryotic origins of DNA replication by a multiprotein complex. Nature. 1992 May 14;357(6374):128–134. doi: 10.1038/357128a0. [DOI] [PubMed] [Google Scholar]
- Bentley N. J., Holtzman D. A., Flaggs G., Keegan K. S., DeMaggio A., Ford J. C., Hoekstra M., Carr A. M. The Schizosaccharomyces pombe rad3 checkpoint gene. EMBO J. 1996 Dec 2;15(23):6641–6651. [PMC free article] [PubMed] [Google Scholar]
- Bickel S. E., Orr-Weaver T. L. Holding chromatids together to ensure they go their separate ways. Bioessays. 1996 Apr;18(4):293–300. doi: 10.1002/bies.950180407. [DOI] [PubMed] [Google Scholar]
- Bishop D. K., Park D., Xu L., Kleckner N. DMC1: a meiosis-specific yeast homolog of E. coli recA required for recombination, synaptonemal complex formation, and cell cycle progression. Cell. 1992 May 1;69(3):439–456. doi: 10.1016/0092-8674(92)90446-j. [DOI] [PubMed] [Google Scholar]
- Blasina A., Paegle E. S., McGowan C. H. The role of inhibitory phosphorylation of CDC2 following DNA replication block and radiation-induced damage in human cells. Mol Biol Cell. 1997 Jun;8(6):1013–1023. doi: 10.1091/mbc.8.6.1013. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Boddy M. N., Furnari B., Mondesert O., Russell P. Replication checkpoint enforced by kinases Cds1 and Chk1. Science. 1998 May 8;280(5365):909–912. doi: 10.1126/science.280.5365.909. [DOI] [PubMed] [Google Scholar]
- Booher R. N., Deshaies R. J., Kirschner M. W. Properties of Saccharomyces cerevisiae wee1 and its differential regulation of p34CDC28 in response to G1 and G2 cyclins. EMBO J. 1993 Sep;12(9):3417–3426. doi: 10.1002/j.1460-2075.1993.tb06016.x. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Borgne A., Meijer L. Sequential dephosphorylation of p34(cdc2) on Thr-14 and Tyr-15 at the prophase/metaphase transition. J Biol Chem. 1996 Nov 1;271(44):27847–27854. doi: 10.1074/jbc.271.44.27847. [DOI] [PubMed] [Google Scholar]
- Bork P., Hofmann K., Bucher P., Neuwald A. F., Altschul S. F., Koonin E. V. A superfamily of conserved domains in DNA damage-responsive cell cycle checkpoint proteins. FASEB J. 1997 Jan;11(1):68–76. [PubMed] [Google Scholar]
- Bousset K., Diffley J. F. The Cdc7 protein kinase is required for origin firing during S phase. Genes Dev. 1998 Feb 15;12(4):480–490. doi: 10.1101/gad.12.4.480. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Brown G. W., Kelly T. J. Cell cycle regulation of Dfp1, an activator of the Hsk1 protein kinase. Proc Natl Acad Sci U S A. 1999 Jul 20;96(15):8443–8448. doi: 10.1073/pnas.96.15.8443. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Caligiuri M., Beach D. Sct1 functions in partnership with Cdc10 in a transcription complex that activates cell cycle START and inhibits differentiation. Cell. 1993 Feb 26;72(4):607–619. doi: 10.1016/0092-8674(93)90079-6. [DOI] [PubMed] [Google Scholar]
- Callan H. G. DNA replication in the chromosomes of eukaryotes. Cold Spring Harb Symp Quant Biol. 1974;38:195–203. doi: 10.1101/sqb.1974.038.01.023. [DOI] [PubMed] [Google Scholar]
- Cheng L., Hunke L., Hardy C. F. Cell cycle regulation of the Saccharomyces cerevisiae polo-like kinase cdc5p. Mol Cell Biol. 1998 Dec;18(12):7360–7370. doi: 10.1128/mcb.18.12.7360. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Chikashige Y., Ding D. Q., Funabiki H., Haraguchi T., Mashiko S., Yanagida M., Hiraoka Y. Telomere-led premeiotic chromosome movement in fission yeast. Science. 1994 Apr 8;264(5156):270–273. doi: 10.1126/science.8146661. [DOI] [PubMed] [Google Scholar]
- Chikashige Y., Ding D. Q., Imai Y., Yamamoto M., Haraguchi T., Hiraoka Y. Meiotic nuclear reorganization: switching the position of centromeres and telomeres in the fission yeast Schizosaccharomyces pombe. EMBO J. 1997 Jan 2;16(1):193–202. doi: 10.1093/emboj/16.1.193. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Chu S., Herskowitz I. Gametogenesis in yeast is regulated by a transcriptional cascade dependent on Ndt80. Mol Cell. 1998 Apr;1(5):685–696. doi: 10.1016/s1097-2765(00)80068-4. [DOI] [PubMed] [Google Scholar]
- Ciosk R., Zachariae W., Michaelis C., Shevchenko A., Mann M., Nasmyth K. An ESP1/PDS1 complex regulates loss of sister chromatid cohesion at the metaphase to anaphase transition in yeast. Cell. 1998 Jun 12;93(6):1067–1076. doi: 10.1016/s0092-8674(00)81211-8. [DOI] [PubMed] [Google Scholar]
- Cohen-Fix O., Koshland D. The anaphase inhibitor of Saccharomyces cerevisiae Pds1p is a target of the DNA damage checkpoint pathway. Proc Natl Acad Sci U S A. 1997 Dec 23;94(26):14361–14366. doi: 10.1073/pnas.94.26.14361. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Cohen-Fix O., Peters J. M., Kirschner M. W., Koshland D. Anaphase initiation in Saccharomyces cerevisiae is controlled by the APC-dependent degradation of the anaphase inhibitor Pds1p. Genes Dev. 1996 Dec 15;10(24):3081–3093. doi: 10.1101/gad.10.24.3081. [DOI] [PubMed] [Google Scholar]
- Coleman T. R., Carpenter P. B., Dunphy W. G. The Xenopus Cdc6 protein is essential for the initiation of a single round of DNA replication in cell-free extracts. Cell. 1996 Oct 4;87(1):53–63. doi: 10.1016/s0092-8674(00)81322-7. [DOI] [PubMed] [Google Scholar]
- Colomina N., Garí E., Gallego C., Herrero E., Aldea M. G1 cyclins block the Ime1 pathway to make mitosis and meiosis incompatible in budding yeast. EMBO J. 1999 Jan 15;18(2):320–329. doi: 10.1093/emboj/18.2.320. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Cooper J. P., Nimmo E. R., Allshire R. C., Cech T. R. Regulation of telomere length and function by a Myb-domain protein in fission yeast. Nature. 1997 Feb 20;385(6618):744–747. doi: 10.1038/385744a0. [DOI] [PubMed] [Google Scholar]
- Cooper J. P., Watanabe Y., Nurse P. Fission yeast Taz1 protein is required for meiotic telomere clustering and recombination. Nature. 1998 Apr 23;392(6678):828–831. doi: 10.1038/33947. [DOI] [PubMed] [Google Scholar]
- D'Urso G., Grallert B., Nurse P. DNA polymerase alpha, a component of the replication initiation complex, is essential for the checkpoint coupling S phase to mitosis in fission yeast. J Cell Sci. 1995 Sep;108(Pt 9):3109–3118. doi: 10.1242/jcs.108.9.3109. [DOI] [PubMed] [Google Scholar]
- D'Urso G., Nurse P. Schizosaccharomyces pombe cdc20+ encodes DNA polymerase epsilon and is required for chromosomal replication but not for the S phase checkpoint. Proc Natl Acad Sci U S A. 1997 Nov 11;94(23):12491–12496. doi: 10.1073/pnas.94.23.12491. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Daya-Makin M., Szankasi P., Tang L., MacRae D., Pelech S. L. Regulation of p105wee1 and p34cdc2 during meiosis in Schizosaccharomyces pombe. Biochem Cell Biol. 1992 Oct-Nov;70(10-11):1088–1096. doi: 10.1139/o92-154. [DOI] [PubMed] [Google Scholar]
- Ding R., Smith G. R. Global control of meiotic recombination genes by Schizosaccharomyces pombe rec16 (rep1). Mol Gen Genet. 1998 Jun;258(6):663–670. doi: 10.1007/s004380050780. [DOI] [PubMed] [Google Scholar]
- Dirick L., Goetsch L., Ammerer G., Byers B. Regulation of meiotic S phase by Ime2 and a Clb5,6-associated kinase in Saccharomyces cerevisiae. Science. 1998 Sep 18;281(5384):1854–1857. doi: 10.1126/science.281.5384.1854. [DOI] [PubMed] [Google Scholar]
- Donaldson A. D., Blow J. J. The regulation of replication origin activation. Curr Opin Genet Dev. 1999 Feb;9(1):62–68. doi: 10.1016/s0959-437x(99)80009-4. [DOI] [PubMed] [Google Scholar]
- Donaldson A. D., Fangman W. L., Brewer B. J. Cdc7 is required throughout the yeast S phase to activate replication origins. Genes Dev. 1998 Feb 15;12(4):491–501. doi: 10.1101/gad.12.4.491. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Donovan S., Harwood J., Drury L. S., Diffley J. F. Cdc6p-dependent loading of Mcm proteins onto pre-replicative chromatin in budding yeast. Proc Natl Acad Sci U S A. 1997 May 27;94(11):5611–5616. doi: 10.1073/pnas.94.11.5611. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Dutta A., Bell S. P. Initiation of DNA replication in eukaryotic cells. Annu Rev Cell Dev Biol. 1997;13:293–332. doi: 10.1146/annurev.cellbio.13.1.293. [DOI] [PubMed] [Google Scholar]
- Edwards R. J., Bentley N. J., Carr A. M. A Rad3-Rad26 complex responds to DNA damage independently of other checkpoint proteins. Nat Cell Biol. 1999 Nov;1(7):393–398. doi: 10.1038/15623. [DOI] [PubMed] [Google Scholar]
- Enoch T., Carr A. M., Nurse P. Fission yeast genes involved in coupling mitosis to completion of DNA replication. Genes Dev. 1992 Nov;6(11):2035–2046. doi: 10.1101/gad.6.11.2035. [DOI] [PubMed] [Google Scholar]
- Enoch T., Gould K. L., Nurse P. Mitotic checkpoint control in fission yeast. Cold Spring Harb Symp Quant Biol. 1991;56:409–416. doi: 10.1101/sqb.1991.056.01.048. [DOI] [PubMed] [Google Scholar]
- Enoch T., Nurse P. Mutation of fission yeast cell cycle control genes abolishes dependence of mitosis on DNA replication. Cell. 1990 Feb 23;60(4):665–673. doi: 10.1016/0092-8674(90)90669-6. [DOI] [PubMed] [Google Scholar]
- Esashi F., Yanagida M. Cdc2 phosphorylation of Crb2 is required for reestablishing cell cycle progression after the damage checkpoint. Mol Cell. 1999 Aug;4(2):167–174. doi: 10.1016/s1097-2765(00)80364-0. [DOI] [PubMed] [Google Scholar]
- Fay D. S., Sun Z., Stern D. F. Mutations in SPK1/RAD53 that specifically abolish checkpoint but not growth-related functions. Curr Genet. 1997 Feb;31(2):97–105. doi: 10.1007/s002940050181. [DOI] [PubMed] [Google Scholar]
- Fisher D. L., Nurse P. A single fission yeast mitotic cyclin B p34cdc2 kinase promotes both S-phase and mitosis in the absence of G1 cyclins. EMBO J. 1996 Feb 15;15(4):850–860. [PMC free article] [PubMed] [Google Scholar]
- Fogarty P., Campbell S. D., Abu-Shumays R., Phalle B. S., Yu K. R., Uy G. L., Goldberg M. L., Sullivan W. The Drosophila grapes gene is related to checkpoint gene chk1/rad27 and is required for late syncytial division fidelity. Curr Biol. 1997 Jun 1;7(6):418–426. doi: 10.1016/s0960-9822(06)00189-8. [DOI] [PubMed] [Google Scholar]
- Foiani M., Nadjar-Boger E., Capone R., Sagee S., Hashimshoni T., Kassir Y. A meiosis-specific protein kinase, Ime2, is required for the correct timing of DNA replication and for spore formation in yeast meiosis. Mol Gen Genet. 1996 Dec 13;253(3):278–288. doi: 10.1007/s004380050323. [DOI] [PubMed] [Google Scholar]
- Ford J. C., al-Khodairy F., Fotou E., Sheldrick K. S., Griffiths D. J., Carr A. M. 14-3-3 protein homologs required for the DNA damage checkpoint in fission yeast. Science. 1994 Jul 22;265(5171):533–535. doi: 10.1126/science.8036497. [DOI] [PubMed] [Google Scholar]
- Fox M. E., Smith G. R. Control of meiotic recombination in Schizosaccharomyces pombe. Prog Nucleic Acid Res Mol Biol. 1998;61:345–378. doi: 10.1016/s0079-6603(08)60831-4. [DOI] [PubMed] [Google Scholar]
- Francesconi S., De Recondo A. M., Baldacci G. DNA polymerase delta is required for the replication feedback control of cell cycle progression in Schizosaccharomyces pombe. Mol Gen Genet. 1995 Mar 10;246(5):561–569. doi: 10.1007/BF00298962. [DOI] [PubMed] [Google Scholar]
- Furnari B., Blasina A., Boddy M. N., McGowan C. H., Russell P. Cdc25 inhibited in vivo and in vitro by checkpoint kinases Cds1 and Chk1. Mol Biol Cell. 1999 Apr;10(4):833–845. doi: 10.1091/mbc.10.4.833. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Furnari B., Rhind N., Russell P. Cdc25 mitotic inducer targeted by chk1 DNA damage checkpoint kinase. Science. 1997 Sep 5;277(5331):1495–1497. doi: 10.1126/science.277.5331.1495. [DOI] [PubMed] [Google Scholar]
- Furuno N., Nishizawa M., Okazaki K., Tanaka H., Iwashita J., Nakajo N., Ogawa Y., Sagata N. Suppression of DNA replication via Mos function during meiotic divisions in Xenopus oocytes. EMBO J. 1994 May 15;13(10):2399–2410. doi: 10.1002/j.1460-2075.1994.tb06524.x. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Glover D. M., Hagan I. M., Tavares A. A. Polo-like kinases: a team that plays throughout mitosis. Genes Dev. 1998 Dec 15;12(24):3777–3787. doi: 10.1101/gad.12.24.3777. [DOI] [PubMed] [Google Scholar]
- Gould K. L., Burns C. G., Feoktistova A., Hu C. P., Pasion S. G., Forsburg S. L. Fission yeast cdc24(+) encodes a novel replication factor required for chromosome integrity. Genetics. 1998 Jul;149(3):1221–1233. doi: 10.1093/genetics/149.3.1221. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Grallert B., Nurse P. The ORC1 homolog orp1 in fission yeast plays a key role in regulating onset of S phase. Genes Dev. 1996 Oct 15;10(20):2644–2654. doi: 10.1101/gad.10.20.2644. [DOI] [PubMed] [Google Scholar]
- Grallert B., Sipiczki M. Common genes and pathways in the regulation of the mitotic and meiotic cell cycles of Schizosaccharomyces pombe. Curr Genet. 1991 Aug;20(3):199–204. doi: 10.1007/BF00326233. [DOI] [PubMed] [Google Scholar]
- Grallert B., Sipiczki M. Dissociation of meiotic and mitotic roles of the fission yeast cdc2 gene. Mol Gen Genet. 1990 Jul;222(2-3):473–475. doi: 10.1007/BF00633860. [DOI] [PubMed] [Google Scholar]
- Grandin N., Reed S. I. Differential function and expression of Saccharomyces cerevisiae B-type cyclins in mitosis and meiosis. Mol Cell Biol. 1993 Apr;13(4):2113–2125. doi: 10.1128/mcb.13.4.2113. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Griffiths D. J., Barbet N. C., McCready S., Lehmann A. R., Carr A. M. Fission yeast rad17: a homologue of budding yeast RAD24 that shares regions of sequence similarity with DNA polymerase accessory proteins. EMBO J. 1995 Dec 1;14(23):5812–5823. doi: 10.1002/j.1460-2075.1995.tb00269.x. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Guacci V., Koshland D., Strunnikov A. A direct link between sister chromatid cohesion and chromosome condensation revealed through the analysis of MCD1 in S. cerevisiae. Cell. 1997 Oct 3;91(1):47–57. doi: 10.1016/s0092-8674(01)80008-8. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Hayles J., Nurse P. A pre-start checkpoint preventing mitosis in fission yeast acts independently of p34cdc2 tyrosine phosphorylation. EMBO J. 1995 Jun 15;14(12):2760–2771. doi: 10.1002/j.1460-2075.1995.tb07276.x. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Hirano T. SMC-mediated chromosome mechanics: a conserved scheme from bacteria to vertebrates? Genes Dev. 1999 Jan 1;13(1):11–19. doi: 10.1101/gad.13.1.11. [DOI] [PubMed] [Google Scholar]
- Hiraoka Y. Meiotic telomeres: a matchmaker for homologous chromosomes. Genes Cells. 1998 Jul;3(7):405–413. doi: 10.1046/j.1365-2443.1998.00205.x. [DOI] [PubMed] [Google Scholar]
- Hofmann J. F., Beach D. cdt1 is an essential target of the Cdc10/Sct1 transcription factor: requirement for DNA replication and inhibition of mitosis. EMBO J. 1994 Jan 15;13(2):425–434. doi: 10.1002/j.1460-2075.1994.tb06277.x. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Hofmann K., Bucher P. The FHA domain: a putative nuclear signalling domain found in protein kinases and transcription factors. Trends Biochem Sci. 1995 Sep;20(9):347–349. doi: 10.1016/s0968-0004(00)89072-6. [DOI] [PubMed] [Google Scholar]
- Hua X. H., 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 Jan 26;140(2):271–281. doi: 10.1083/jcb.140.2.271. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Iino Y., Hiramine Y., Yamamoto M. The role of cdc2 and other genes in meiosis in Schizosaccharomyces pombe. Genetics. 1995 Aug;140(4):1235–1245. doi: 10.1093/genetics/140.4.1235. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Jimenez G., Yucel J., Rowley R., Subramani S. The rad3+ gene of Schizosaccharomyces pombe is involved in multiple checkpoint functions and in DNA repair. Proc Natl Acad Sci U S A. 1992 Jun 1;89(11):4952–4956. doi: 10.1073/pnas.89.11.4952. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Jin P., Gu Y., Morgan D. O. Role of inhibitory CDC2 phosphorylation in radiation-induced G2 arrest in human cells. J Cell Biol. 1996 Aug;134(4):963–970. doi: 10.1083/jcb.134.4.963. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Jin P., Hardy S., Morgan D. O. Nuclear localization of cyclin B1 controls mitotic entry after DNA damage. J Cell Biol. 1998 May 18;141(4):875–885. doi: 10.1083/jcb.141.4.875. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Kelly T. J., Martin G. S., Forsburg S. L., Stephen R. J., Russo A., Nurse P. The fission yeast cdc18+ gene product couples S phase to START and mitosis. Cell. 1993 Jul 30;74(2):371–382. doi: 10.1016/0092-8674(93)90427-r. [DOI] [PubMed] [Google Scholar]
- Kishimoto T. Activation of MPF at meiosis reinitiation in starfish oocytes. Dev Biol. 1999 Oct 1;214(1):1–8. doi: 10.1006/dbio.1999.9393. [DOI] [PubMed] [Google Scholar]
- Kitamura K., Maekawa H., Shimoda C. Fission yeast Ste9, a homolog of Hct1/Cdh1 and Fizzy-related, is a novel negative regulator of cell cycle progression during G1-phase. Mol Biol Cell. 1998 May;9(5):1065–1080. doi: 10.1091/mbc.9.5.1065. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Kleckner N. Meiosis: how could it work? Proc Natl Acad Sci U S A. 1996 Aug 6;93(16):8167–8174. doi: 10.1073/pnas.93.16.8167. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Klein F., Mahr P., Galova M., Buonomo S. B., Michaelis C., Nairz K., Nasmyth K. A central role for cohesins in sister chromatid cohesion, formation of axial elements, and recombination during yeast meiosis. Cell. 1999 Jul 9;98(1):91–103. doi: 10.1016/S0092-8674(00)80609-1. [DOI] [PubMed] [Google Scholar]
- Knudsen K. E., Knudsen E. S., Wang J. Y., Subramani S. p34cdc2 kinase activity is maintained upon activation of the replication checkpoint in Schizosaccharomyces pombe. Proc Natl Acad Sci U S A. 1996 Aug 6;93(16):8278–8283. doi: 10.1073/pnas.93.16.8278. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Kominami K., Sakata Y., Sakai M., Yamashita I. Protein kinase activity associated with the IME2 gene product, a meiotic inducer in the yeast Saccharomyces cerevisiae. Biosci Biotechnol Biochem. 1993 Oct;57(10):1731–1735. doi: 10.1271/bbb.57.1731. [DOI] [PubMed] [Google Scholar]
- Kondo T., Matsumoto K., Sugimoto K. Role of a complex containing Rad17, Mec3, and Ddc1 in the yeast DNA damage checkpoint pathway. Mol Cell Biol. 1999 Feb;19(2):1136–1143. doi: 10.1128/mcb.19.2.1136. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Kostrub C. F., Knudsen K., Subramani S., Enoch T. Hus1p, a conserved fission yeast checkpoint protein, interacts with Rad1p and is phosphorylated in response to DNA damage. EMBO J. 1998 Apr 1;17(7):2055–2066. doi: 10.1093/emboj/17.7.2055. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Kumagai A., Dunphy W. G. Control of the Cdc2/cyclin B complex in Xenopus egg extracts arrested at a G2/M checkpoint with DNA synthesis inhibitors. Mol Biol Cell. 1995 Feb;6(2):199–213. doi: 10.1091/mbc.6.2.199. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Kumagai A., Guo Z., Emami K. H., Wang S. X., Dunphy W. G. The Xenopus Chk1 protein kinase mediates a caffeine-sensitive pathway of checkpoint control in cell-free extracts. J Cell Biol. 1998 Sep 21;142(6):1559–1569. doi: 10.1083/jcb.142.6.1559. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Leatherwood J., Lopez-Girona A., Russell P. Interaction of Cdc2 and Cdc18 with a fission yeast ORC2-like protein. Nature. 1996 Jan 25;379(6563):360–363. doi: 10.1038/379360a0. [DOI] [PubMed] [Google Scholar]
- Leem S. H., Chung C. N., Sunwoo Y., Araki H. Meiotic role of SWI6 in Saccharomyces cerevisiae. Nucleic Acids Res. 1998 Jul 1;26(13):3154–3158. doi: 10.1093/nar/26.13.3154. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Leu J. Y., Roeder G. S. The pachytene checkpoint in S. cerevisiae depends on Swe1-mediated phosphorylation of the cyclin-dependent kinase Cdc28. Mol Cell. 1999 Nov;4(5):805–814. doi: 10.1016/s1097-2765(00)80390-1. [DOI] [PubMed] [Google Scholar]
- Lew D. J., Reed S. I. A cell cycle checkpoint monitors cell morphogenesis in budding yeast. J Cell Biol. 1995 May;129(3):739–749. doi: 10.1083/jcb.129.3.739. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Li X., Cai M. Inactivation of the cyclin-dependent kinase Cdc28 abrogates cell cycle arrest induced by DNA damage and disassembly of mitotic spindles in Saccharomyces cerevisiae. Mol Cell Biol. 1997 May;17(5):2723–2734. doi: 10.1128/mcb.17.5.2723. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Li Y. F., Smith G. R. The Schizosaccharomyces pombe rec16 gene product regulates multiple meiotic events. Genetics. 1997 May;146(1):57–67. doi: 10.1093/genetics/146.1.57. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Lindsay H. D., Griffiths D. J., Edwards R. J., Christensen P. U., Murray J. M., Osman F., Walworth N., Carr A. M. S-phase-specific activation of Cds1 kinase defines a subpathway of the checkpoint response in Schizosaccharomyces pombe. Genes Dev. 1998 Feb 1;12(3):382–395. doi: 10.1101/gad.12.3.382. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Long K. E., Sunnerhagen P., Subramani S. The Schizosaccharomyces pombe rad1 gene consists of three exons and the cDNA sequence is partially homologous to the Ustilago maydis REC1 cDNA. Gene. 1994 Oct 11;148(1):155–159. doi: 10.1016/0378-1119(94)90250-x. [DOI] [PubMed] [Google Scholar]
- Lopez-Girona A., Furnari B., Mondesert O., Russell P. Nuclear localization of Cdc25 is regulated by DNA damage and a 14-3-3 protein. Nature. 1999 Jan 14;397(6715):172–175. doi: 10.1038/16488. [DOI] [PubMed] [Google Scholar]
- Lundgren K., Walworth N., Booher R., Dembski M., Kirschner M., Beach D. mik1 and wee1 cooperate in the inhibitory tyrosine phosphorylation of cdc2. Cell. 1991 Mar 22;64(6):1111–1122. doi: 10.1016/0092-8674(91)90266-2. [DOI] [PubMed] [Google Scholar]
- Lydall D., Nikolsky Y., Bishop D. K., Weinert T. A meiotic recombination checkpoint controlled by mitotic checkpoint genes. Nature. 1996 Oct 31;383(6603):840–843. doi: 10.1038/383840a0. [DOI] [PubMed] [Google Scholar]
- Lygerou Z., Nurse P. The fission yeast origin recognition complex is constitutively associated with chromatin and is differentially modified through the cell cycle. J Cell Sci. 1999 Nov;112(Pt 21):3703–3712. doi: 10.1242/jcs.112.21.3703. [DOI] [PubMed] [Google Scholar]
- MacNeill S. A., Moreno S., Reynolds N., Nurse P., Fantes P. A. The fission yeast Cdc1 protein, a homologue of the small subunit of DNA polymerase delta, binds to Pol3 and Cdc27. EMBO J. 1996 Sep 2;15(17):4613–4628. [PMC free article] [PubMed] [Google Scholar]
- Maiorano D., Van Assendelft G. B., Kearsey S. E. Fission yeast cdc21, a member of the MCM protein family, is required for onset of S phase and is located in the nucleus throughout the cell cycle. EMBO J. 1996 Feb 15;15(4):861–872. [PMC free article] [PubMed] [Google Scholar]
- Martin-Castellanos C., Labib K., Moreno S. B-type cyclins regulate G1 progression in fission yeast in opposition to the p25rum1 cdk inhibitor. EMBO J. 1996 Feb 15;15(4):839–849. [PMC free article] [PubMed] [Google Scholar]
- Masai H., Miyake T., Arai K. hsk1+, a Schizosaccharomyces pombe gene related to Saccharomyces cerevisiae CDC7, is required for chromosomal replication. EMBO J. 1995 Jul 3;14(13):3094–3104. doi: 10.1002/j.1460-2075.1995.tb07312.x. [DOI] [PMC free article] [PubMed] [Google Scholar]
- McMillan J. N., Sia R. A., Bardes E. S., Lew D. J. Phosphorylation-independent inhibition of Cdc28p by the tyrosine kinase Swe1p in the morphogenesis checkpoint. Mol Cell Biol. 1999 Sep;19(9):5981–5990. doi: 10.1128/mcb.19.9.5981. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Mendenhall M. D., Hodge A. E. Regulation of Cdc28 cyclin-dependent protein kinase activity during the cell cycle of the yeast Saccharomyces cerevisiae. Microbiol Mol Biol Rev. 1998 Dec;62(4):1191–1243. doi: 10.1128/mmbr.62.4.1191-1243.1998. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Michaelis C., Ciosk R., Nasmyth K. Cohesins: chromosomal proteins that prevent premature separation of sister chromatids. Cell. 1997 Oct 3;91(1):35–45. doi: 10.1016/s0092-8674(01)80007-6. [DOI] [PubMed] [Google Scholar]
- Millar J. B., Lenaers G., Russell P. Pyp3 PTPase acts as a mitotic inducer in fission yeast. EMBO J. 1992 Dec;11(13):4933–4941. doi: 10.1002/j.1460-2075.1992.tb05600.x. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Millar J. B., Russell P. The cdc25 M-phase inducer: an unconventional protein phosphatase. Cell. 1992 Feb 7;68(3):407–410. doi: 10.1016/0092-8674(92)90177-e. [DOI] [PubMed] [Google Scholar]
- Miyamoto M., Tanaka K., Okayama H. res2+, a new member of the cdc10+/SWI4 family, controls the 'start' of mitotic and meiotic cycles in fission yeast. EMBO J. 1994 Apr 15;13(8):1873–1880. doi: 10.1002/j.1460-2075.1994.tb06456.x. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Molnar M., Bähler J., Sipiczki M., Kohli J. The rec8 gene of Schizosaccharomyces pombe is involved in linear element formation, chromosome pairing and sister-chromatid cohesion during meiosis. Genetics. 1995 Sep;141(1):61–73. doi: 10.1093/genetics/141.1.61. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Moreno S., Hayles J., Nurse P. Regulation of p34cdc2 protein kinase during mitosis. Cell. 1989 Jul 28;58(2):361–372. doi: 10.1016/0092-8674(89)90850-7. [DOI] [PubMed] [Google Scholar]
- Morgan D. O. Regulation of the APC and the exit from mitosis. Nat Cell Biol. 1999 Jun;1(2):E47–E53. doi: 10.1038/10039. [DOI] [PubMed] [Google Scholar]
- Murakami H., Nurse P. Meiotic DNA replication checkpoint control in fission yeast. Genes Dev. 1999 Oct 1;13(19):2581–2593. doi: 10.1101/gad.13.19.2581. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Murakami H., Okayama H. A kinase from fission yeast responsible for blocking mitosis in S phase. Nature. 1995 Apr 27;374(6525):817–819. doi: 10.1038/374817a0. [DOI] [PubMed] [Google Scholar]
- Muzi-Falconi M., Kelly T. J. Orp1, a member of the Cdc18/Cdc6 family of S-phase regulators, is homologous to a component of the origin recognition complex. Proc Natl Acad Sci U S A. 1995 Dec 19;92(26):12475–12479. doi: 10.1073/pnas.92.26.12475. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Naito T., Matsuura A., Ishikawa F. Circular chromosome formation in a fission yeast mutant defective in two ATM homologues. Nat Genet. 1998 Oct;20(2):203–206. doi: 10.1038/2517. [DOI] [PubMed] [Google Scholar]
- Nakajo N., Yoshitome S., Iwashita J., Iida M., Uto K., Ueno S., Okamoto K., Sagata N. Absence of Wee1 ensures the meiotic cell cycle in Xenopus oocytes. Genes Dev. 2000 Feb 1;14(3):328–338. [PMC free article] [PubMed] [Google Scholar]
- Nakaseko Y., Niwa O., Yanagida M. A meiotic mutant of the fission yeast Schizosaccharomyces pombe that produces mature asci containing two diploid spores. J Bacteriol. 1984 Jan;157(1):334–336. doi: 10.1128/jb.157.1.334-336.1984. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Nakashima N., Tanaka K., Sturm S., Okayama H. Fission yeast Rep2 is a putative transcriptional activator subunit for the cell cycle 'start' function of Res2-Cdc10. EMBO J. 1995 Oct 2;14(19):4794–4802. doi: 10.1002/j.1460-2075.1995.tb00161.x. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Nasmyth K. Separating sister chromatids. Trends Biochem Sci. 1999 Mar;24(3):98–104. doi: 10.1016/s0968-0004(99)01358-4. [DOI] [PubMed] [Google Scholar]
- Newlon C. S. Putting it all together: building a prereplicative complex. Cell. 1997 Dec 12;91(6):717–720. doi: 10.1016/s0092-8674(00)80459-6. [DOI] [PubMed] [Google Scholar]
- Nigg E. A. Polo-like kinases: positive regulators of cell division from start to finish. Curr Opin Cell Biol. 1998 Dec;10(6):776–783. doi: 10.1016/s0955-0674(98)80121-x. [DOI] [PubMed] [Google Scholar]
- Nimmo E. R., Pidoux A. L., Perry P. E., Allshire R. C. Defective meiosis in telomere-silencing mutants of Schizosaccharomyces pombe. Nature. 1998 Apr 23;392(6678):825–828. doi: 10.1038/33941. [DOI] [PubMed] [Google Scholar]
- Nishitani H., Lygerou Z., Nishimoto T., Nurse P. The Cdt1 protein is required to license DNA for replication in fission yeast. Nature. 2000 Apr 6;404(6778):625–628. doi: 10.1038/35007110. [DOI] [PubMed] [Google Scholar]
- Nurse P., Bissett Y. Gene required in G1 for commitment to cell cycle and in G2 for control of mitosis in fission yeast. Nature. 1981 Aug 6;292(5823):558–560. doi: 10.1038/292558a0. [DOI] [PubMed] [Google Scholar]
- Nurse P. Checkpoint pathways come of age. Cell. 1997 Dec 26;91(7):865–867. doi: 10.1016/s0092-8674(00)80476-6. [DOI] [PubMed] [Google Scholar]
- Nurse P., Thuriaux P., Nasmyth K. Genetic control of the cell division cycle in the fission yeast Schizosaccharomyces pombe. Mol Gen Genet. 1976 Jul 23;146(2):167–178. doi: 10.1007/BF00268085. [DOI] [PubMed] [Google Scholar]
- Nurse P., Thuriaux P. Regulatory genes controlling mitosis in the fission yeast Schizosaccharomyces pombe. Genetics. 1980 Nov;96(3):627–637. doi: 10.1093/genetics/96.3.627. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Nurse P. Universal control mechanism regulating onset of M-phase. Nature. 1990 Apr 5;344(6266):503–508. doi: 10.1038/344503a0. [DOI] [PubMed] [Google Scholar]
- O'Connell M. J., Raleigh J. M., Verkade H. M., Nurse P. Chk1 is a wee1 kinase in the G2 DNA damage checkpoint inhibiting cdc2 by Y15 phosphorylation. EMBO J. 1997 Feb 3;16(3):545–554. doi: 10.1093/emboj/16.3.545. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Ohsumi K., Sawada W., Kishimoto T. Meiosis-specific cell cycle regulation in maturing Xenopus oocytes. J Cell Sci. 1994 Nov;107(Pt 11):3005–3013. doi: 10.1242/jcs.107.11.3005. [DOI] [PubMed] [Google Scholar]
- Okayama H., Nagata A., Jinno S., Murakami H., Tanaka K., Nakashima N. Cell cycle control in fission yeast and mammals: identification of new regulatory mechanisms. Adv Cancer Res. 1996;69:17–62. doi: 10.1016/s0065-230x(08)60859-3. [DOI] [PubMed] [Google Scholar]
- Osmani S. A., Ye X. S. Cell cycle regulation in Aspergillus by two protein kinases. Biochem J. 1996 Aug 1;317(Pt 3):633–641. doi: 10.1042/bj3170633. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Paciotti V., Lucchini G., Plevani P., Longhese M. P. Mec1p is essential for phosphorylation of the yeast DNA damage checkpoint protein Ddc1p, which physically interacts with Mec3p. EMBO J. 1998 Jul 15;17(14):4199–4209. doi: 10.1093/emboj/17.14.4199. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Parisi S., McKay M. J., Molnar M., Thompson M. A., van der Spek P. J., van Drunen-Schoenmaker E., Kanaar R., Lehmann E., Hoeijmakers J. H., Kohli J. Rec8p, a meiotic recombination and sister chromatid cohesion phosphoprotein of the Rad21p family conserved from fission yeast to humans. Mol Cell Biol. 1999 May;19(5):3515–3528. doi: 10.1128/mcb.19.5.3515. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Parker A. E., Van de Weyer I., Laus M. C., Oostveen I., Yon J., Verhasselt P., Luyten W. H. A human homologue of the Schizosaccharomyces pombe rad1+ checkpoint gene encodes an exonuclease. J Biol Chem. 1998 Jul 17;273(29):18332–18339. doi: 10.1074/jbc.273.29.18332. [DOI] [PubMed] [Google Scholar]
- Parker A. E., Van de Weyer I., Laus M. C., Verhasselt P., Luyten W. H. Identification of a human homologue of the Schizosaccharomyces pombe rad17+ checkpoint gene. J Biol Chem. 1998 Jul 17;273(29):18340–18346. doi: 10.1074/jbc.273.29.18340. [DOI] [PubMed] [Google Scholar]
- Picard A., Galas S., Peaucellier G., Dorée M. Newly assembled cyclin B-cdc2 kinase is required to suppress DNA replication between meiosis I and meiosis II in starfish oocytes. EMBO J. 1996 Jul 15;15(14):3590–3598. [PMC free article] [PubMed] [Google Scholar]
- Reynolds N., Fantes P. A., MacNeill S. A. A key role for replication factor C in DNA replication checkpoint function in fission yeast. Nucleic Acids Res. 1999 Jan 15;27(2):462–469. doi: 10.1093/nar/27.2.462. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Rhind N., Furnari B., Russell P. Cdc2 tyrosine phosphorylation is required for the DNA damage checkpoint in fission yeast. Genes Dev. 1997 Feb 15;11(4):504–511. doi: 10.1101/gad.11.4.504. [DOI] [PubMed] [Google Scholar]
- Rhind N., Russell P. Tyrosine phosphorylation of cdc2 is required for the replication checkpoint in Schizosaccharomyces pombe. Mol Cell Biol. 1998 Jul;18(7):3782–3787. doi: 10.1128/mcb.18.7.3782. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Roberts J. M. Evolving ideas about cyclins. Cell. 1999 Jul 23;98(2):129–132. doi: 10.1016/s0092-8674(00)81007-7. [DOI] [PubMed] [Google Scholar]
- Roeder G. S. Meiotic chromosomes: it takes two to tango. Genes Dev. 1997 Oct 15;11(20):2600–2621. doi: 10.1101/gad.11.20.2600. [DOI] [PubMed] [Google Scholar]
- Romanowski P., Madine M. A., Rowles A., Blow J. J., Laskey R. A. The Xenopus origin recognition complex is essential for DNA replication and MCM binding to chromatin. Curr Biol. 1996 Nov 1;6(11):1416–1425. doi: 10.1016/s0960-9822(96)00746-4. [DOI] [PubMed] [Google Scholar]
- Rowles A., Chong J. P., Brown L., Howell M., Evan G. I., Blow J. J. Interaction between the origin recognition complex and the replication licensing system in Xenopus. Cell. 1996 Oct 18;87(2):287–296. doi: 10.1016/s0092-8674(00)81346-x. [DOI] [PubMed] [Google Scholar]
- Rowley R., Hudson J., Young P. G. The wee1 protein kinase is required for radiation-induced mitotic delay. Nature. 1992 Mar 26;356(6367):353–355. doi: 10.1038/356353a0. [DOI] [PubMed] [Google Scholar]
- Rowley R., Subramani S., Young P. G. Checkpoint controls in Schizosaccharomyces pombe: rad1. EMBO J. 1992 Apr;11(4):1335–1342. doi: 10.1002/j.1460-2075.1992.tb05178.x. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Russell P., Moreno S., Reed S. I. Conservation of mitotic controls in fission and budding yeasts. Cell. 1989 Apr 21;57(2):295–303. doi: 10.1016/0092-8674(89)90967-7. [DOI] [PubMed] [Google Scholar]
- Sagata N. What does Mos do in oocytes and somatic cells? Bioessays. 1997 Jan;19(1):13–21. doi: 10.1002/bies.950190105. [DOI] [PubMed] [Google Scholar]
- Saka Y., Esashi F., Matsusaka T., Mochida S., Yanagida M. Damage and replication checkpoint control in fission yeast is ensured by interactions of Crb2, a protein with BRCT motif, with Cut5 and Chk1. Genes Dev. 1997 Dec 15;11(24):3387–3400. doi: 10.1101/gad.11.24.3387. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Saka Y., Yanagida M. Fission yeast cut5+, required for S phase onset and M phase restraint, is identical to the radiation-damage repair gene rad4+. Cell. 1993 Jul 30;74(2):383–393. doi: 10.1016/0092-8674(93)90428-s. [DOI] [PubMed] [Google Scholar]
- San-Segundo P. A., Roeder G. S. Pch2 links chromatin silencing to meiotic checkpoint control. Cell. 1999 Apr 30;97(3):313–324. doi: 10.1016/s0092-8674(00)80741-2. [DOI] [PubMed] [Google Scholar]
- Sanchez Y., Bachant J., Wang H., Hu F., Liu D., Tetzlaff M., Elledge S. J. Control of the DNA damage checkpoint by chk1 and rad53 protein kinases through distinct mechanisms. Science. 1999 Nov 5;286(5442):1166–1171. doi: 10.1126/science.286.5442.1166. [DOI] [PubMed] [Google Scholar]
- Schneider B. L., Yang Q. H., Futcher A. B. Linkage of replication to start by the Cdk inhibitor Sic1. Science. 1996 Apr 26;272(5261):560–562. doi: 10.1126/science.272.5261.560. [DOI] [PubMed] [Google Scholar]
- Sherr C. J., Roberts J. M. CDK inhibitors: positive and negative regulators of G1-phase progression. Genes Dev. 1999 Jun 15;13(12):1501–1512. doi: 10.1101/gad.13.12.1501. [DOI] [PubMed] [Google Scholar]
- Shimada M., Okuzaki D., Tanaka S., Tougan T., Tamai K. K., Shimoda C., Nojima H. Replication factor C3 of Schizosaccharomyces pombe, a small subunit of replication factor C complex, plays a role in both replication and damage checkpoints. Mol Biol Cell. 1999 Dec;10(12):3991–4003. doi: 10.1091/mbc.10.12.3991. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Shimomura T., Ando S., Matsumoto K., Sugimoto K. Functional and physical interaction between Rad24 and Rfc5 in the yeast checkpoint pathways. Mol Cell Biol. 1998 Sep;18(9):5485–5491. doi: 10.1128/mcb.18.9.5485. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Shuster E. O., Byers B. Pachytene arrest and other meiotic effects of the start mutations in Saccharomyces cerevisiae. Genetics. 1989 Sep;123(1):29–43. doi: 10.1093/genetics/123.1.29. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Sia R. A., Herald H. A., Lew D. J. Cdc28 tyrosine phosphorylation and the morphogenesis checkpoint in budding yeast. Mol Biol Cell. 1996 Nov;7(11):1657–1666. doi: 10.1091/mbc.7.11.1657. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Smith A. V., Roeder G. S. The yeast Red1 protein localizes to the cores of meiotic chromosomes. J Cell Biol. 1997 Mar 10;136(5):957–967. doi: 10.1083/jcb.136.5.957. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Sorger P. K., Murray A. W. S-phase feedback control in budding yeast independent of tyrosine phosphorylation of p34cdc28. Nature. 1992 Jan 23;355(6358):365–368. doi: 10.1038/355365a0. [DOI] [PubMed] [Google Scholar]
- Stern B., Nurse P. A quantitative model for the cdc2 control of S phase and mitosis in fission yeast. Trends Genet. 1996 Sep;12(9):345–350. [PubMed] [Google Scholar]
- Stern B., Nurse P. Cyclin B proteolysis and the cyclin-dependent kinase inhibitor rum1p are required for pheromone-induced G1 arrest in fission yeast. Mol Biol Cell. 1998 Jun;9(6):1309–1321. doi: 10.1091/mbc.9.6.1309. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Stern B., Nurse P. Fission yeast pheromone blocks S-phase by inhibiting the G1 cyclin B-p34cdc2 kinase. EMBO J. 1997 Feb 3;16(3):534–544. doi: 10.1093/emboj/16.3.534. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Stuart D., Wittenberg C. CLB5 and CLB6 are required for premeiotic DNA replication and activation of the meiotic S/M checkpoint. Genes Dev. 1998 Sep 1;12(17):2698–2710. doi: 10.1101/gad.12.17.2698. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Stueland C. S., Lew D. J., Cismowski M. J., Reed S. I. Full activation of p34CDC28 histone H1 kinase activity is unable to promote entry into mitosis in checkpoint-arrested cells of the yeast Saccharomyces cerevisiae. Mol Cell Biol. 1993 Jun;13(6):3744–3755. doi: 10.1128/mcb.13.6.3744. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Su T. T., Walker J., Stumpff J. Activating the DNA damage checkpoint in a developmental context. Curr Biol. 2000 Feb 10;10(3):119–126. doi: 10.1016/s0960-9822(00)00300-6. [DOI] [PubMed] [Google Scholar]
- Sugiyama A., Tanaka K., Okazaki K., Nojima H., Okayama H. A zinc finger protein controls the onset of premeiotic DNA synthesis of fission yeast in a Mei2-independent cascade. EMBO J. 1994 Apr 15;13(8):1881–1887. doi: 10.1002/j.1460-2075.1994.tb06457.x. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Sun Z., Hsiao J., Fay D. S., Stern D. F. Rad53 FHA domain associated with phosphorylated Rad9 in the DNA damage checkpoint. Science. 1998 Jul 10;281(5374):272–274. doi: 10.1126/science.281.5374.272. [DOI] [PubMed] [Google Scholar]
- Swanton C., Card G. L., Mann D., McDonald N., Jones N. Overcoming inhibitions: subversion of CKI function by viral cyclins. Trends Biochem Sci. 1999 Mar;24(3):116–120. doi: 10.1016/s0968-0004(99)01354-7. [DOI] [PubMed] [Google Scholar]
- Takeda T., Ogino K., Matsui E., Cho M. K., Kumagai H., Miyake T., Arai K., Masai H. A fission yeast gene, him1(+)/dfp1(+), encoding a regulatory subunit for Hsk1 kinase, plays essential roles in S-phase initiation as well as in S-phase checkpoint control and recovery from DNA damage. Mol Cell Biol. 1999 Aug;19(8):5535–5547. doi: 10.1128/mcb.19.8.5535. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Tanaka H., Tanaka K., Murakami H., Okayama H. Fission yeast cdc24 is a replication factor C- and proliferating cell nuclear antigen-interacting factor essential for S-phase completion. Mol Cell Biol. 1999 Feb;19(2):1038–1048. doi: 10.1128/mcb.19.2.1038. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Tanaka K., Okazaki K., Okazaki N., Ueda T., Sugiyama A., Nojima H., Okayama H. A new cdc gene required for S phase entry of Schizosaccharomyces pombe encodes a protein similar to the cdc 10+ and SWI4 gene products. EMBO J. 1992 Dec;11(13):4923–4932. doi: 10.1002/j.1460-2075.1992.tb05599.x. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Tanaka T., Knapp D., Nasmyth K. Loading of an Mcm protein onto DNA replication origins is regulated by Cdc6p and CDKs. Cell. 1997 Aug 22;90(4):649–660. doi: 10.1016/s0092-8674(00)80526-7. [DOI] [PubMed] [Google Scholar]
- Thelen M. P., Venclovas C., Fidelis K. A sliding clamp model for the Rad1 family of cell cycle checkpoint proteins. Cell. 1999 Mar 19;96(6):769–770. doi: 10.1016/s0092-8674(00)80587-5. [DOI] [PubMed] [Google Scholar]
- Toyoshima F., Moriguchi T., Wada A., Fukuda M., Nishida E. Nuclear export of cyclin B1 and its possible role in the DNA damage-induced G2 checkpoint. EMBO J. 1998 May 15;17(10):2728–2735. doi: 10.1093/emboj/17.10.2728. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Tóth A., Ciosk R., Uhlmann F., Galova M., Schleiffer A., Nasmyth K. Yeast cohesin complex requires a conserved protein, Eco1p(Ctf7), to establish cohesion between sister chromatids during DNA replication. Genes Dev. 1999 Feb 1;13(3):320–333. doi: 10.1101/gad.13.3.320. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Uhlmann F., Lottspeich F., Nasmyth K. Sister-chromatid separation at anaphase onset is promoted by cleavage of the cohesin subunit Scc1. Nature. 1999 Jul 1;400(6739):37–42. doi: 10.1038/21831. [DOI] [PubMed] [Google Scholar]
- Visintin R., Prinz S., Amon A. CDC20 and CDH1: a family of substrate-specific activators of APC-dependent proteolysis. Science. 1997 Oct 17;278(5337):460–463. doi: 10.1126/science.278.5337.460. [DOI] [PubMed] [Google Scholar]
- Walworth N., Davey S., Beach D. Fission yeast chk1 protein kinase links the rad checkpoint pathway to cdc2. Nature. 1993 May 27;363(6427):368–371. doi: 10.1038/363368a0. [DOI] [PubMed] [Google Scholar]
- Waseem N. H., Labib K., Nurse P., Lane D. P. Isolation and analysis of the fission yeast gene encoding polymerase delta accessory protein PCNA. EMBO J. 1992 Dec;11(13):5111–5120. doi: 10.1002/j.1460-2075.1992.tb05618.x. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Watanabe Y., Nurse P. Cohesin Rec8 is required for reductional chromosome segregation at meiosis. Nature. 1999 Jul 29;400(6743):461–464. doi: 10.1038/22774. [DOI] [PubMed] [Google Scholar]
- Watanabe Y., Shinozaki-Yabana S., Chikashige Y., Hiraoka Y., Yamamoto M. Phosphorylation of RNA-binding protein controls cell cycle switch from mitotic to meiotic in fission yeast. Nature. 1997 Mar 13;386(6621):187–190. doi: 10.1038/386187a0. [DOI] [PubMed] [Google Scholar]
- Watanabe Y., Yamamoto M. S. pombe mei2+ encodes an RNA-binding protein essential for premeiotic DNA synthesis and meiosis I, which cooperates with a novel RNA species meiRNA. Cell. 1994 Aug 12;78(3):487–498. doi: 10.1016/0092-8674(94)90426-x. [DOI] [PubMed] [Google Scholar]
- Weber L., Byers B. A RAD9-dependent checkpoint blocks meiosis of cdc13 yeast cells. Genetics. 1992 May;131(1):55–63. doi: 10.1093/genetics/131.1.55. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Weinert T. DNA damage checkpoints update: getting molecular. Curr Opin Genet Dev. 1998 Apr;8(2):185–193. doi: 10.1016/s0959-437x(98)80140-8. [DOI] [PubMed] [Google Scholar]
- Williamson D. H., Johnston L. H., Fennell D. J., Simchen G. The timing of the S phase and other nuclear events in yeast meiosis. Exp Cell Res. 1983 Apr 15;145(1):209–217. doi: 10.1016/s0014-4827(83)80022-6. [DOI] [PubMed] [Google Scholar]
- Willson J., Wilson S., Warr N., Watts F. Z. Isolation and characterization of the Schizosaccharomyces pombe rhp9 gene: a gene required for the DNA damage checkpoint but not the replication checkpoint. Nucleic Acids Res. 1997 Jun 1;25(11):2138–2146. doi: 10.1093/nar/25.11.2138. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Xu L., Weiner B. M., Kleckner N. Meiotic cells monitor the status of the interhomolog recombination complex. Genes Dev. 1997 Jan 1;11(1):106–118. doi: 10.1101/gad.11.1.106. [DOI] [PubMed] [Google Scholar]
- Yamaguchi S., Murakami H., Okayama H. A WD repeat protein controls the cell cycle and differentiation by negatively regulating Cdc2/B-type cyclin complexes. Mol Biol Cell. 1997 Dec;8(12):2475–2486. doi: 10.1091/mbc.8.12.2475. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Yamamoto A., Guacci V., Koshland D. Pds1p, an inhibitor of anaphase in budding yeast, plays a critical role in the APC and checkpoint pathway(s). J Cell Biol. 1996 Apr;133(1):99–110. doi: 10.1083/jcb.133.1.99. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Yamamoto M. The molecular control mechanisms of meiosis in fission yeast. Trends Biochem Sci. 1996 Jan;21(1):18–22. [PubMed] [Google Scholar]
- Yanagida M. Cell cycle mechanisms of sister chromatid separation; roles of Cut1/separin and Cut2/securin. Genes Cells. 2000 Jan;5(1):1–8. doi: 10.1046/j.1365-2443.2000.00306.x. [DOI] [PubMed] [Google Scholar]
- Ye X. S., Fincher R. R., Tang A., O'Donnell K., Osmani S. A. Two S-phase checkpoint systems, one involving the function of both BIME and Tyr15 phosphorylation of p34cdc2, inhibit NIMA and prevent premature mitosis. EMBO J. 1996 Jul 15;15(14):3599–3610. [PMC free article] [PubMed] [Google Scholar]
- Yoshida M., Kawaguchi H., Sakata Y., Kominami K., Hirano M., Shima H., Akada R., Yamashita I. Initiation of meiosis and sporulation in Saccharomyces cerevisiae requires a novel protein kinase homologue. Mol Gen Genet. 1990 Apr;221(2):176–186. doi: 10.1007/BF00261718. [DOI] [PubMed] [Google Scholar]
- Zeng Y., Forbes K. C., Wu Z., Moreno S., Piwnica-Worms H., Enoch T. Replication checkpoint requires phosphorylation of the phosphatase Cdc25 by Cds1 or Chk1. Nature. 1998 Oct 1;395(6701):507–510. doi: 10.1038/26766. [DOI] [PubMed] [Google Scholar]
- Zeng Y., Piwnica-Worms H. DNA damage and replication checkpoints in fission yeast require nuclear exclusion of the Cdc25 phosphatase via 14-3-3 binding. Mol Cell Biol. 1999 Nov;19(11):7410–7419. doi: 10.1128/mcb.19.11.7410. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Zhu Y., Takeda T., Nasmyth K., Jones N. pct1+, which encodes a new DNA-binding partner of p85cdc10, is required for meiosis in the fission yeast Schizosaccharomyces pombe. Genes Dev. 1994 Apr 15;8(8):885–898. doi: 10.1101/gad.8.8.885. [DOI] [PubMed] [Google Scholar]
- Zou L., Stillman B. Formation of a preinitiation complex by S-phase cyclin CDK-dependent loading of Cdc45p onto chromatin. Science. 1998 Apr 24;280(5363):593–596. doi: 10.1126/science.280.5363.593. [DOI] [PubMed] [Google Scholar]
- al-Khodairy F., Carr A. M. DNA repair mutants defining G2 checkpoint pathways in Schizosaccharomyces pombe. EMBO J. 1992 Apr;11(4):1343–1350. doi: 10.1002/j.1460-2075.1992.tb05179.x. [DOI] [PMC free article] [PubMed] [Google Scholar]
- al-Khodairy F., Fotou E., Sheldrick K. S., Griffiths D. J., Lehmann A. R., Carr A. M. Identification and characterization of new elements involved in checkpoint and feedback controls in fission yeast. Mol Biol Cell. 1994 Feb;5(2):147–160. doi: 10.1091/mbc.5.2.147. [DOI] [PMC free article] [PubMed] [Google Scholar]