Biochemical and cytogenetic changes in postovulatory and in vitro aged mammalian oocytes: a predisposition to aneuploidy

John B. Mailhes


Aneuploidy represents the most prevalent genetic disorder of man. Its association with spontaneous abortions, mental and physical retardation, and numerous malignant cells is well-known. Unfortunately, little is known about the causes and even less about the underlying molecular mechanisms of aneuploidy, especially in mammalian germ cells. Although several etiologies have been proposed for describing human aneuploidy, the only consistent finding remains its positive correlation with maternal age. At the outset, it is essential to point out that there exist numerous potential causes and mechanisms for the etiology of aneuploidy. Nevertheless, information about the molecular mechanisms of chromosome segregation in various species is providing a foundation for research designed to investigate the causes and mechanisms of aneuploidy. The intent of this review is to propose that the biochemical reactions and cellular organelles responsible for accurate chromosome segregation become compromised during postovulatory and in vitro oocyte aging; thus, increasing the probability of faulty chromosome segregation. Recent data have shown that the efficacies of the spindle assembly checkpoint and the chromosome cohesion proteins diminish as oocytes age postovulation and during in vitro culture. Such changes represent potential models for studying aneuploidy. Prior to describing the biochemical and cellular organelle changes found in aged oocytes and their effect on chromosome segregation, an overview of the molecular details surrounding chromosome segregation is presented.




postovulatory ageing; oocyte; aneuploidy; premature centromere separation


Adams RR, Maiato H, Earnshaw WC & Carmena M. 2001a. Essential roles of Drosophila inner centromere protein (INCENP) and aurora B in histone H3 phosphorylation, metaphase chromosome alignment, kinetochore disjunction, and chromosome segregation. J Cell Biol 153 865-880

Adams RR, Eckley DM, Vagnarelli P, Wheatley SP, Gerloff DL, Mackay AM, Svingen PA, Kaufman SH & Earnshaw WC 2001b Human INCENP colocalizes with the Aurora-B/AIRK kinase on chromosomes and is overexpressed in tumor cells. Chromosoma 110 65-74

Adler I-D 1990 Aneuploidy studies in mammals. In Mendelsohn ML, Albertini RJ, editors. Mutation and the Environment, Part B: Metabolism, Testing Methods, and Chromosomes, New York: Wiley-Liss, p 285-293

Adler I-D 1993 Synopsis of the in vivo results obtained with the 10 known or suspected aneugens tested in the CEC collaborative study. Mutat Res 287 131-137

Adler I-D, Schmid TE & Baumgartner A 2002 Induction of aneuploidy in male mouse germ cells detected by the sperm–FISH assay: a review of the present data base. Mutat Res 504 173-182

Agarwal R & Cohen-Fix O 2002 Mitotic regulation: the fine tuning of separase activity. Cell Cycle 1 255-257

Ahonen LJ, Kallio MJ, Daum JR, Bolton M, Manke IA, Yaffe MB, Stukenberg PT & Gorbsky GJ 2005 Polo-like kinase 1 creates the tension-sensing 3F3/2 phosphoepitope and modulates the association of spindle-checkpoint proteins at kinetochores. Curr Biol 15 1078-1089

Akiyama T, Nagata M & Aoki F 2006 Inadequate histone deactylation during oocyte meiosis causes aneuploidy and embryo death in mice. Proc Natl Acad Sci USA 103 7339-7344

Allen JW, Liang JC, Carrano AV & Preston RJ 1986 Review of literature on chemical-induced aneuploidy in mammalian male germ cells. Mutat Res 167 123-137

Alexandru G, Uhlmann F, Mechtler K, Poupart MA & Nasmyth K 2001 Phosphorylation of the cohesion subunit Scc1 by Polo/Cdc5 kinase regulates sister chromatid separation in yeast. Cell 105 459-472

Amor DJ, Kalitsis P, Sumer H & Choo KH 2004 Building the centromere: from foundation proteins to 3D organization. Trends Cell Biol 14 359-368

Anahory T, Andreo B, Regnier-Vigourouox G, Souile JP, Baudouin M, Demaille J & Pellestor F 2003 Sequential multiple probe fluorescence in-situ hybridization analysis of human oocytes and polar bodies by combining centromeric labeling and whole chromosome painting. Mol Hum Report 9 577-585

Anderiesz C, Ferraretti A, Magli C, Fiorentino A, Fortini D, Gianaroli L, Jones GM & Trounson AO 2000 Effect of recombinant human gonadotrophins on human, bovine and marine oocyte meiosis. Hum Report 15 1140-1148

Andreassen PR, Martina SN & Margolis RL 1996 Chemical induction of mitotic checkpoint override in mammalian cells results in aneuploidy following a transient metalloid state. Mutat Res 372 181-194

Angell RR 1991 Predivision in human oocytes at meiosis I: a mechanism for trisomy formation in man. Hum Genet 86 383-387

Angell RR 1994 Aneuploidy in older women; higher rates of aneuploidy in oocytes from older women. Hum Report 9 1199-1201

Angell RR, Xian J, Keith J, Ledger W & Baird DT 1994 First meiotic division abnormalities in human oocytes: mechanism of trisomy formation. Cytogenet Cell Genet 65 194-202

Arion D, Meier L, Brizuela L & Beach D 1988 Cdc2 is a component of the M phase-specific histone H1 kinase: evidence for identity with MPF. Cell 55 371-378

Austin CR 1967 Chromosome deterioration in aging eggs of the rabbit. Nature 213 1018-1019

Austin CR 1970 Ageing and reproduction: post-ovulatory deterioration of the egg. J Reprod Fert Suppl 12 39-53

Baker DJ, Jeganathan KB, Cameron JD, Thompson M, Juneja S, Kopecka A, Kumar R, Jenkins RB, de Groen PC, Roche P & van Deursen JM 2004 BubR1 insufficiency causes early onset of aging-associated phenotypes and infertility in mice. Nat Genet 36 744-749

Barnes FL, Collas P, Powell R, King WA, Westhusin M & Shepherd D 1993 Influence of recipient oocyte cell cycle stage on DNA synthesis, nuclear envelope breakdown, chromosome constitution, and development in nuclear transplant bovine embryos. Mol Rep Dev 36 33-41

Baudat F, Manova K, Yuen JP, Jasin M & Keeney S 2000 Chromosome synapsis defects and sexually dimorphic meiotic progression in mice lacking Spo11. Mol Cell 6 989-998

Beatrice MC, Stiers DL & Pfeiffer DR 1984 The role of glutathione in the retention of Ca2+ by liver mitochondria. J Biol Chem 259 1279-1287

Bernard P, Maure JF & Javerzat JP 2001 Fission yeast Bub1 is essential in setting up the meiotic pattern of chromosome segregation. Nat Cell Biol 3 522-526

Bettencourt-Dias M, Giet R, Sinka R, Mazumdar A, Lock WG, Balloux PJ, Safiropoulos PJ, Yamaguchi S, Winter S, Carthew RW, Cooper M, Jones D, Frenz L & Glover DM 2004 Genome-wide survey of protein kinases required for cell cycle progression. Nature 432 980-987

Bharadwal R & Yu H 2004 The spindle checkpoint, aneuploidy, and cancer. Oncogene 23 2016-2027

Biggins S & Walczak CE 2003 Captivating capture: how microtubules attach to kinetochores. Curr Biol 13 R449-R460

Blazak WF 1987 Incidence of aneuploidy in farm animals. In Vig BK, Sandberg AA, editors. Aneuploidy, Part A: Incidence and Etiology. New York: Alan R. Liss. p.103-116

Boerjan ML & deBoer P 1990 First cell cycle of zygotes of the mouse derived from oocytes aged postovulation in vivo and fertilized in vivo. Mol Rep Dev 25 155-163

Boernslaeger EA, Mattei P & Schultz RM 1986 Involvement of cAMP-dependent protein kinase and protein phosphorylation in regulation of mouse oocyte maturation. Dev Biol 114 453-462

Bolton MA, Lan W, Powers SE, McCleland ML, Kuang J & Stukenberg PT 2002 Aurora B kinase exists in a complex with survivin and INCENP and its kinase activity is stimulated by survivin binding and phosphorylation. Mol Biol Cell 13 3064-3077

Bond DJ & Chandley AC 1983 Aneuploidy, Oxford Monographs on Medical Genetics, No. 11, Oxford: Oxford University Press, p 1-198

Brunet S, Pahlavan G, Taylor S & Maro B 2003 Functionality of the spindle checkpoint during the first meiotic division of mammalian oocytes. Reproduction 126 443-450

Buonomo SB, Clyne RK, Fuchs J, Loidl J, Uhlmann F & Nasmyth K 2000 Disjunction of homologous chromosomes in meiosis I depends on proteolytic cleavage of the meiotic cohesion Rec8 by separin. Cell 103 387-398

Carvalho A, Carmena M, Sambade C, Earnshaw WC & Wheatley SP 2003 Survivin is required for stable checkpoint activation in taxol-treated HeLa cells. J Cell Sci 116 2987-2998

Cascio SM & Wassarman PM 1982 Program of early development in the mammal: post-transcriptional control of a class of proteins synthesized by mouse oocytes and early embryos. Dev Biol 89 397-408

Champoux JJ 2001 DNA topoisomerases: structure, function, and mechanism. Annu Rev Biochem 70 369-413

Chandley AC 1987 Aneuploidy: an overview, In Vig BK, Sandberg AA, editors. Aneuploidy: Part A: Incidence and Etiology. New York: Alan R. Liss. p 1-8

Chen RH, Shevehenko A, Mann M & Murray AW 1998 Spindle checkpoint protein Xmad1 recruits Xmad2 to unattached kinetochores. J Cell Biol 143 283-295

Choi T, Aoki F, Mori M, Yamashita M, Nagahama Y & Kohmoto K 1991 Activation of p34cdc2 protein kinase activity in meiotic and mitotic cell cycles in mouse oocytes and embryos. Development 113 789-795

Chung E & Chen RH 2002 Spindle checkpoint requires Mad1-bound and Mad1-free Mad2. Mol Biol Cell 13 1501-1511

Cimini D 2007 Detection and correction of merotelic kinetochore orientation by Aurora B, its partners. Cell Cycle 6 1558-1564

Cimini D 2008 Merotelic kinetochore orientation, aneuploidy, and cancer. Biochem Biophys Acta doi:10.1016/j.bbcan.2008.05.003

Ciosk R, Shirayama M, Shevchenko A, Tanaka T, Toth A, Shevchenko A & Nasmyth K 2000 Cohesin’s binding to chromosomes depends on a separate complex consisting of Scc2 and Scc4 proteins. Mol Cell 4 243-254

Clarke AS, Tang TT, Ooi DL & Orr-Weaver TL 2005 POLO kinase regulates the Drosophila centromere cohesion protein MEI-S332. Dev Cell 8 53-64

Clyne RK, Katis VL, Jessop L, Benjamin KR, Herskowitz I, Lichten M & Nasmyth K 2003 Polo-like kinase Cdc5 promotes chiasmata formation and cosegregation of sister chromatids at meiosis I. Nat Cell Biol 5 379-382

Cobb J, Reddy RK, Park C & Handel MA 1997 Analysis of expression and function of topoisomerase I and II during meiosis in male mice. Mol Rep Dev 46 489-498

Cohen P, Holmes CF & Tsukitani Y 1990 Okadaic acid: a new probe for the study of cellular regulation. Trends Biochem Sci 15 98-102

Cohen-Fix O, Peters JM, Kirschner MW & Koshland D 1996 Anaphase initiation in Saccharomyces cerevisiae is controlled by the APC-dependent degradation of the anaphase inhibitor Pds1p. Genes Dev 10 3081-3093

Collas P, Sullivan EJ & Barnes FL 1993 Histone H1 kinase activity in bovine oocytes following calcium stimulation. Mol Rep Dev 34 224-231

Colledge WH, Carlton MBL, Udy GB & Evans MJ 1994. Disruption of c-mos causes parthenogenetic development of unfertilized mouse eggs. Nature 370 65-68

Collins J & Crosignani PG 2005 Fertility and ageing. Hum Reprod Update 11 261-276

Coulam CB, Jeyendran RS, Fiddler M & Pergament E 2007. Discordance among blastomeres renders preimplantation genetic diagnosis for aneuploidy ineffective. J Assist Reprod Genet 24 37-41

Coux O, Tanaka K & Goldberg AL 1996 Structure and function of the 20S and 26S proteasomes. Annu Rev Biochem 65 801-847

Craig JM, Earnshaw WC & Vagnarelli P 1999 Mammalian centromeres: DNA sequence, protein composition, and role in cell cycle progression. Exptl Cell Res 246 249-262

Craig JM & Choo KHA 2005 Kiss and break up – a safe passage to anaphase in mitosis and meiosis. Chromosoma 114 252-262

Cukurcam S, Hegele-Hartung C & Eichenlaub-Ritter U 2003 Meiosis-activating sterol protects oocytes from precocious chromosome segregation. Hum Reprod 18 1908-1917

Cupisti S, Conn CM, Fragouli E, Whalley K, Mills JA, Faed MJ & Delhanty JD 2003 Sequential FISH analysis of oocytes and polar bodies reveals aneuploidy mechanisms. Prenat Diagn 23 663-668

Cuthbertson KSR & Cubbbold PH 1985 Phorbol ester and sperm activate mouse oocytes by inducing sustained oscillations in cell Ca2+. Nature 316 541-542

Dai Y, Fulka Jr. J & Moor R 2003a Checkpoint controls in mammalian oocytes. In Trounson AO, Gosden RG, editors. Biology and Pathology of the Oocyte. Cambridge UK: Cambridge University Press. p 120-132

Dai W, Huang X & Ruan Q 2003b Polo-like kinases in cell cycle checkpoint control. Frontiers Biosci 8 1128-1133

Dai W, Wang Q, Liu TY, Swamy M, Fang YQ, Xie SQ, Mahmood R, Yang YM, Xu M & Rao CV 2004 Slippage of mitotic arrest and enhanced tumor development in mice with BUBR1 haploinsufficiency. Cancer Res 15 440-445

Dailey T, Dale B, Cohen J & Nunné S 1996 Association between nondisjunction and maternal age in meiosis II oocytes. Am J Hum Genet 59 176-184

Dekel N 1988 Regulation of oocyte maturation: the role of cAMP. Ann NY Acad Sci 541 211-216

Dekel N 1996 Protein phosphorylation-dephosphorylation in the meiotic cell cycle of mammalian oocytes. Rev Reprod 1 82-88

Dekel N 2005 Cellular, biochemical and molecular mechanisms regulating oocyte maturation. Mol Cell Endocrinol 234 19-25

De Pennart H, Helene-Verlhac M, Cibert C, Santa Maria A & Maro B 1993 Okadaic acid induces spindle lengthening and disrupts the interaction of microtubules with the kinetochores in metaphase II-arrested mouse oocytes. Dev Biol 157 170-181

Dewar H, Tanaka K, Nasmyth K & Tanaka TU 2004 Tension between two kinetochores suffices for their bi-orientation on the mitotic spindle. Nature 428 93-97

Diaz H & Esponda P 2004 Postovulatory ageing induces structural changes in the mouse zona pellucida. J Submicrosc Cytol Pathol 36 211-217

Dinardo S, Voelkel KA & Sternglanz RL 1984 DNA topoisomerase mutant of S. cerevisiae: topoisomerase II is required for segregation of daughter molecules at the termination of DNA replication. Proc Natl Acad Sci USA 81 2616-2620

Ditchfield C, Johnson VL, Tighe A, Ellston R, Haworth C, Johnson T, Mortlock A, Keen N & Taylor SS 2003 Aurora B couples chromosome alignment with anaphase by targeting BubR1, Mad2, and Cenp-E to kinetochores. J Cell Biol 161 267-280

Dobles M, Liberal V, Scott ML, Benezra R & Sorger PK 2000 Chromosome missegregation and apoptosis in mice lacking the mitotic checkpoint protein Mad2. Cell 101 635-645

Dorée M, Le Peuch C & Morin N 1995 Onset of chromosome segregation at the metaphase to anaphase transition of the cell cycle. Prog Cell Cycle Res 1 309-318

Dorer RK, Zhong S, Tallarico JA, Wong WH, Mitchison TJ & Murray AW 2005 A small-molecule inhibitor of Mps1 blocks the spindle-checkpoint response to a lack of tension on mitotic chromosomes. Curr Biol 15 1070-1076

Downes CS, Mullinger AM & Johnson RT 1991 Inhibitors of DNA topoisomerase II prevent chromatid separation in mammalian cells but do not prevent exit from mitosis. Proc Natl Acad Sci USA 88 8895-8899

Downs SM, Daniel SAJ, Bornslaeger EA, Hoppe PC & Eppig JJ 1989 Maintenance of meiotic arrest in mouse oocytes by purines: modulation of cAMP levels and cAMP phosphodiesterase activity. Gamete Res 23 323-334

Draetta G & Beach D 1988 Activation of cdc2 protein kinase during mitosis in human cells: Cell cycle dependent phosphorylation and subunit rearrangement. Cell 54 17-26

Duesbery NS, Choi T, Brown KD, Wood KW, Resau J, Fukasawa K, Cleveland DW & Vande-Woude GF 1997 CENP-E is an essential kinetochore motor in maturing oocytes and is masked during mos-dependent, cell cycle arrest at metaphase II. Proc Natl Acad Sci USA 94 9165-9170

Dunphy WG & Kumagai A 1991 The cdc25 protein contains an intrinsic phosphatase activity. Cell 67 189-196

Earnshaw WC & Cooke CA 1991 Analysis of the distribution of the INCENPs throughout mitosis reveals the existence of a pathway of structural changes in the chromosomes during metaphase and early events in cleavage furrow formation. J Cell Sci 98 443-461

Eastmond DA, Schuler M & Rupa DS 1995 Advantages and limitations of using fluorescence in situ hybridization for the detection of aneuploidy in interphase human cells. Mutat Res 348 153-162

Edwards RG & Gates AH 1959 Timing of the stages of the maturation divisions, ovulation, fertilization and the first cleavage of eggs of adult mice treated with gonadotrophins. J Endocrin 18 292-304

Eichenlaub-Ritter U 1993 Studies on maternal age-related aneuploidy in mammalian oocytes and cell cycle control. In Sumner AT, Chandley AC, editors. Chromosomes Today, Vol. 11. London: Chapman & Hall. p 323-336

Eichenlaub-Ritter U 1996 Parental age-related aneuploidy in human germ cells and offspring: a story of past and present. Environ Mol Mutagen 28 211-236

Eichenlaub-Ritter U, Chandley AC & Gosden RG 1986 Alterations to the microtubular cytoskeleton and increased disorder of chromosome alignment in spontaneously ovulated mouse oocytes in vivo: an immunofluorescence study. Chromosoma 94 337-345

Eichenlaub-Ritter U, Stahl A & Luciani JM 1988 The microtubular cytoskeleton and chromosomes of unfertilized human oocyte aged in vitro. Hum Genet 80 259-264

Eichenlaub-Ritter U, Baart E, Yin H & Betzendahl I 1996 Mechanisms of spontaneous and chemically-induced aneuploidy in mammalian oogenesis: basis of sex-specific differences in response to aneugens and the necessity for further tests. Mutat Res 372 279-294

Eichenlaub-Ritter U 2003 Aneuploidy in aging oocytes and after toxic insult. In Trounson A, Gosden RG, editors. Biology and pathology of the oocyte; its role in fertility and reproductive medicine. Cambridge, UK: Cambridge University Press. p 220-257

Eijpe M, Heyting C, Gross B & Jessberger R 2000 Association of mammalian SMC1 and SMC3 proteins with meiotic chromosomes and synaptonemal complexes. J Cell Sci 113 673-682

Eijpe M, Offenberg H, Jessberger R, Revenkova E & Heyting C 2003 Meiotic cohesion REC8 marks the axial elements of rat synaptonemal complexes before cohesions SMC1β and SMC3. J Cell Biol 160 657-670

Ekins JG & Shaver EL 1975 Cytogenetics of postimplantation rabbit conceptuses following delayed fertilization. Teratology 13 57-64

Emery BR, Wilcox AL, Aoki VW, Peterson CM & Carrell DT 2005 In vitro oocyte maturation and subsequent delayed fertilization is associated with increased embryo aneuploidy. Fertil Steril 84 1027-1029

Exley GE, Tang C, McElhinny AS & Warner CM 1999 Expression of caspase and BCL-2 apoptotic family members in mouse preimplantation embryos. Biol Rep 61 231-239

Fan HY & Sun QY 2004 Involvement of mitogen-activated protein kinase cascade during oocyte maturation and fertilization in mammals. Biol Rep 70 535-547

Fang G 2002 Checkpoint protein BubR1 acts synergistically with Mad2 to inhibit anaphase-promoting complex. Mol Biol Cell 13 755-766

Fissore RA, Kurokawa M, Knott J, Zhang M & Smyth J 2002 Mechanisms underlying oocyte activation and postovulatory aging. Reproduction 124 745-754

Ford JH 1981 Nondisjunction. In Burgio GR, Fraccaro M, Tiepolo L, Wolf U, editor. Trisomy 21. Berlin: Springer. p 103-143

Fragouli E, Wells D, Doshi A, Gotts S, Harper JC & Delhanty JD 2006 Complete cytogenetic investigation of oocytes from a young cancer patient with the use of comparative genomic hybridization reveals meiotic errors. Prenat Diagn 26 71-76

Fraser LR 1979 Rate of fertilization in vitro and subsequent nuclear development as a function of the post-ovulatory age of the mouse egg. J Rep Fertil 55 153-160

Fulka Jr. J, Jung T & Moor RM 1992 The fall of biological maturation promoting factor (MPF) and histone H1 kinase activity during anaphase and telophase in mouse oocytes. Mol Rep Dev 32 378-382

Fulton BP & Whittingham DG 1978 Activation of mammalian oocytes by intracellular injection of calcium. Nature 273 149-151

Gabrielli BG, Roy LM & Maller JL 1993 Requirement for cdk2 in cytostatic factor-mediated metaphase II arrest. Science 259 1766-1769

Gautier J, Solomon MJ, Booher RN, Bazan JF & Kirschner ME 1991 cdc25 is a specific tyrosine phosphatase that directly activates p34cdc2. Cell 67 197-211

Geijsen N, Horoschak M, Kim K, Gribnau J, Eggan K & Daley GQ 2004 Derivation of embryonic germ cells and male gametes from embryonic stem cells. Nature 427 148-154

George MA, Pickering SJ, Braude PR & Johnson MH 1996 The distribution of α- and λ tubulin in fresh and aged human and mouse oocytes exposed to cryoprotectant. Mol Hum Rep 6 445-456

Gianfortoni JG & Gulyas BJ 1985 The effects of short-term incubation (aging) of mouse oocytes on in vitro fertilization, zona solubility, and embryonic development. Gamete Res 11 59-68

Giet R, Petretti C & Prignet C 2005 Aurora kinases, aneuploidy and cancer, a coincidence or a real link? Trends Cell Biol 15 241-250

Glickman MH & Ciechanover A 2002 The ubiquitin-proteasome proteolytic pathway: destruction for the sake of construction. Physiol Rev 82 373-428

Glover DM, Ohkura H & Tavares A 1998 Polo-like kinases: a team that plays throughout mitosis. Genes Dev 12 3777-3787

Goldberg AL 1995 Functions of the proteasome: the lysis at the end of the tunnel. Science 268 522- 523

Goldstein LS 1980 Mechanisms of chromosome orientation revealed by two meiotic mutants in Drosophila melanogaster. Chromosoma 78 79-111

Gorbsky GJ 1994 Cell cycle progression and chromosome segregation in mammalian cells cultured in the presence of the topoisomerase II inhibitors ICRF-187 [(+)-1,2-bis(3,5-dioxopiperazinyl-1-yl) propane; ADR-529] and ICRF-159 (Razoxane). Cancer Res 54 1042-1048

Gordo AC, Wu H, He CL & Fissore RA 2000 Injection of sperm cytosolic factor into mouse metaphase II oocyte induces different development fates according to the frequency of [Ca2+] oscillations and oocyte age. Biol Reprod 62 1370-1379

Gordo A, He CL, Smith S & Fissore RA 2001 Mitogen-activated protein kinase plays a significant role in metaphase II arrest, spindle morphology and maintenance of maturation promoting factor activity in bovine oocytes. Mol Rep Dev 59 106-114

Gordo AC, Rodrigues P, Kurokawa M, Jellerette T, Exley GE, Warner C & Fissore R 2002 Intracellular calcium oscillations signal apoptosis rather than activation in in vitro aged mouse eggs. Biol Reprod 66 1828-1837

Goto H, Kiyono T, Tomono Y, Kawajiri A, Urano T, Furukawa K, Nigg EA & Inagaki M 2006 Complex formation of Plk1 and INCENP required for metaphase-anaphase transition. Nat Cell Biol 8 180-187

Goulding SE & Earnshaw WC 2005 Shugoshin: a centromeric guardian senses tension. Bioessays 27 588-591

Gregan J, Rabitsch PK, Sakem B, Csutak O, Latypov V, Lehmann E, Kohli J & Nasmyth K 2005 Novel genes required for meiotic chromosome segregation are identified by a high-throughput knockout screen in fission yeast. Curr Biol 15 1663-1669

Haering CH, Lowe J, Hochwagen A & Nasmyth K 2002 Molecular architecture of SMC proteins and the yeast cohesion complex. Mol Cell 9 773-788

Haering CH & Nasmyth K 2003 Building and breaking bridges between sister chromatids. BioEssays 25 1178-1191

Hafez ESE 1993 Reproduction in Farm Animals, 6th edn. Philadelphia: Lea & Febiger. p 144-164

Hagstrom KA, Holmes VF, Cozzarelli NR & Meyer BJ 2002 C. elegans condensing promotes mitotic chromosome architecture, centromere organization, and sister chromatid segregation during mitosis. Genes Dev 16 729-742

Hagting A, den Elzen N, Vodermaier HC, Waizenegger IC, Peters J-M & Pines J 2002 Human securin proteolysis is controlled by the spindle checkpoint and reveals when the APC/C switches from activation by Cdc20 to Cdh1. J Cell Biol 157 1125-1137

Hall VJ, Compton D, Stojkovic P, Nesbitt M, Herbert M, Murdoch A & Stojkovic M 2007 Developmental competence of human in vitro aged oocytes as host cells for nuclear transfer. Hum Reprod 22 52-62

Handel MA, Cobb J & Eaker S 1999 What are the spermatocyte’s requirements for successful meiotic division? J Exp Zool 285 243-250

Handel MA & Sun F 2005 Regulation of meiotic cell divisions and determination of gamete quality: impact of reproductive toxins. Semin Reprod Med 23 213-221

Hanks S, Coleman K, Reid S, Plaja A, Firth H, Fitzpatrick D, Kidd A, Mehes K, Nash R, Robin N, Shannon N, Tolmie J, Swansbury J, Itthum A, Douglas J & Rahman N 2004 Constitutional aneuploidy and cancer predisposition caused by biallelic mutations in BUB1B. Nat Gen 36 1159-1161

Hansmann I 1983 Factors and mechanisms involved in nondisjunction and X-chromosome loss. In Sandberg AA, editor. Cytogenetics of the Mammalian X Chromosome, Part A: Progress and Topics in Cytogenetics, Vol. 3A. New York: Alan R. Liss. p 131-170

Hansmann I & Pabst B 1992 Nondisjunction by failures in the molecular control of oocyte maturation. Ann Anat 174 485-490

Hartman T, Stead K, Koshland D & Guacci V 2000 Pds5 is an essential chromosomal protein required for both sister chromatid cohesion and condensation in Saccharomyces cerevisiae. J Cell Biol 151 613-626

Hashimoto N, Watanabe N, Furuta Y, Tamemoto H, Sagata N, Yokoyama M, Okazaki K, Nagayoshi M, Takeda N, Ikawa Y & Aizawa S 1994 Parthenogenetic activation of oocytes in c-mos deficient mice. Nature 370 68-71

Hashimoto N 1996 Role of c-mos proto-oncogene product in the regulation of mouse oocyte maturation. Horm Res 46 11-14

Hassold T, Chiu D & Yamane JA 1984 Parental origin of autosomal trisomies. Ann Human Genet 48 129-144

Hassold TJ 1985 The origin of aneuploidy in humans. In Dellarco VL, Voytek PE, Hollaender A, editors. Aneuploidy: Etiology and

Mechanisms. New York: Plenum Press. p 103-115

Hassold T & Sherman S 1993 The origin of non-disjunction in humans. In Sumner AT, Chandley AC, editors. Chromosomes Today. London, UK: Chapman and Hall. p 313-322

Hassold T & Hunt PA 2001 To err (meiotically) is human: the genesis of human aneuploidy. Nat Rev Genet 2 280-291

Hauf S, Cole RW, LaTerra S, Zimmer C, Schnapp G, Walter R, Heckel A, van Meel J, Rieder CL & Peters JM 2003 The small molecule Hesperadin reveals a role for Aurora B in correcting kinetochore-microtubule attachment and in maintaining the spindle assembly checkpoint. J Cell Biol 161 281-294

Hauf S, Roitinger E, Koch B, Dittrich CM, Mechtler K & Peters JM 2005 Dissociation of cohesion during early mitosis depends on phosphorylation of SA2. PloS Biol 3 0419-0432

Hecht F & Hecht BK 1987 Aneuploidy in humans: dimensions, demography, and dangers of abnormal numbers of chromosomes. In Vig BK, Sandberg AA, editors. Aneuploidy, Part A: Incidence and Etiology. New York: Alan R. Liss. p 9-49

Herbert M, Levasseur M, Homer H, Yallop K, Murdoch A & McDougall A 2003 Homologue disjunction in mouse oocytes requires proteolysis of securin and cyclin B. Nat Cell Biol 5 1023-1025

Hiller SG 2001 Gonadotropic control of ovarian follicular growth and development. Mol Cell Endocrinol 179 39-46

Hodges CA, LeMaire-Adkins R & Hunt PA 2001 Coordinating the segregation of sister chromatids during the first meiotic division: evidence for sexual dimorphism. J Cell Sci 114 2417-2426

Hodges CA, Revenkova E, Jessberger R, Hassold T & Hunt PA 2005 SMC1β-deficient female mice provide evidence that cohesions are a missing link in age-related nondisjunction. Nat Genet 37 1351-1355

Holm C, Stearns T & Botstein D 1989 DNA topoisomerase II must act at mitosis to prevent nondisjunction and chromosome breakage. Mol Cell Biol 9 159-168

Homa ST, Carroll J & Swann K 1993 The role of calcium in mammalian oocyte maturation and egg activation. Hum Reprod 8 1274-1281

Homer HA, McDougall A, Levasseur M, Yallop K, Murdoch AP & Herbert M 2005a. Mad2 prevents aneuploidy and premature proteolysis of cyclin B and securing during meiosis I in mouse oocytes. Genes Dev 19 202-207

Homer HA, McDougall A, Levasseur M, Murdoch AP & Herbert M 2005b RNA interference in meiosis I human oocytes: towards and understanding of human aneuploidy. Mol Hum Rep 11 397-404

Hook EB 1985a The impact of aneuploidy upon public health: mortality and morbidity associated with human chromosome abnormalities. In Dellarco VL, Voytek PE, Hollaender A, editors. Aneuploidy: Etiology and Mechanisms. New York: Plenum Press. p 7-33

Hook EB 1985b Maternal age, paternal age, and human chromosome abnormality: nature, magnitude, etiology, and mechanisms of effects, In Dellarco VL, Voytek PE, Hollaender A, editors. Aneuploidy: Etiology and Mechanisms, New York: Plenum Press. p 117-132

Hoque MT & Ishikawa F 2001 Human chromatid cohesion component hRad21 is phosphorylated in M phase and associated with metaphase chromosomes. J Biol Chem 276 5059-5067

Hoque MT & Ishikawa F 2002 Cohesin defects lead to premature sister chromatid separation, kinetochore dysfunction, and spindle-assembly checkpoint activation. J Biol Chem 277 42306-42314

Howell BJ, Moree B, Farrar EM, Stewart S, Fang G & Salmon ED 2004 Spindle checkpoint protein dynamics at kinetochores in living cells. Curr Biol 14 953-964

Howlett SK 1986 A set of proteins showing cell cycle dependent modification in the early mouse embryo. Cell 45 387-396

Huang JC, Yan LY, Lei ZL, Miao YL, Shi LH, Yang JW, Wang Q, Ouyang YC, Sun QY & Chen DY 2007 Changes in histone acetylation during postovulatory aging of mouse oocytes. Biol Reprod 77 666-670

Hwang LH, Lau LF, Smith DL, Mistrot CA, Hardwick KG, Hwang ES, Amon A & Murray AW 1998. Budding yeast Cdc20: a target of the spindle checkpoint. Science 279 1041-1044

Hyman AA & Mitchison TJ 1991 Two different microtubule-based motor activities with opposite polarities in kinetochores. Nature 351 206-211

Igarashi H, Takahashi T, Takahashi E, Tezuka N, Nakahara K, Takahashi K & Kurachi H 2005 Aged mouse oocytes fail to readjust intracellular adenosine triphosphates at fertilization. Biol Reprod 72 1256-1261

Ikui AE, Yang CP, Matsumoto T & Horwitz S 2005 Low concentrations of taxol cause mitotic delay followed by premature dissociation of p55CDC from Mad2 and BubR1 and abrogation of the spindle checkpoint, leading to aneuploidy. Cell Cycle 4 1385-1388

Ishikawa H & Endo A 1995 Combined effects of maternal age and delayed fertilization on the frequency of chromosome anomalies in mice. Hum Reprod 10 883-886

Jallepalli PV, Waizenegger I, Bunz F, Langer S, Speicher MR, Peters JM, Kinzler KW, Vogelstein B & Lengauer C 2001 Securin is required for chromosomal stability in human cells. Cell 105 445-457

Jeffreys, CA, Burrage PS & Bickel SE 2003 A model system for increased meiotic nondisjunction in older oocytes. Curr Biol 13 498-503

Jesus C, Rime H, Haccard O, Van Lint J, Goris J, Merlevede W & Ozon R 1991 Tyrosine phosphorylation of p34cdc2 and p42 during meiotic maturation of Xenopus oocyte: antagonistic action of okadaic acid and 6-DMAP. Development 111 813-820

Johnson VL, Scott MI, Holt SV, Hussein D & Taylor SS 2004 Bub1 is required for kinetochore localization of BubR1, Cenp-E, Cenp-F and Mad2, and chromosome congression. J Cell Sci 117 1577-1589

Juberg RC 1983 Origin of chromosomal abnormalities: evidence for delayed fertilization in meiotic nondisjunction. Hum Genet 64 122-127

Juetten J & Bavister B 1983 Effects of egg aging on in vitro fertilization and first cleavage division in the hamster. Gamete Res 8 219-230

Kalitsis P, Earle E, Fowler KJ & Choo KH 2000 Bub3 gene disruption in mice reveals essential mitotic spindle checkpoint function during early embryogenesis. Genes Dev 14 2277-2282

Kallio M, Eriksson JE & Gorbsky GJ 2000 Differences in spindle association of the mitotic checkpoint protein Mad2 in mammalian spermatogenesis and oogenesis. Dev Biol 225 112-123

Kallio MJ, McCleland ML, Stukenberg PT & Gorbsky GJ 2002 Inhibition of aurora B kinase blocks chromosome segregation, overrides the spindle checkpoint, and perturbs microtubule dynamics in mitosis. Curr Biol 12 900-905

Kamieniecki RJ, Shanks RM & Dawson DS 2000 Slk19p is necessary to prevent separation of sister chromatids in meiosis I. Curr Biol 10 1182-1190

Karsenti E 1991 Mitotic spindle morphogenesis in animal cells. Semin Cell Biol 4 251-260

Katis VL, Galova M, Rabitsch KP & Nasmyth K 2004a Maintenance of cohesion at centromeres after meiosis I in budding yeast requires a kinetochore-associated protein related to MEI-S332. Curr Biol 14 560-572

Katis VL, Matos J, Mori S, Shirahige K, Zachariae W & Nasymth K 2004b Spo13 facilitates monopolin recruitment to kinetochores and regulates maintenance of centromeric cohesion during yeast meiosis. Curr Biol 14 2183-2196

Kaufman MH 1983 Early Mammalian Development: Parthenogenetic Studies. Cambridge: Cambridge Univ. Press. p 1-276

Kerrebrock AW, Miyazaki WY, Birnby D & Orr-Weaver TL 1992 The Drosophila mei-S332 gene promotes sister-chromatid cohesion in meiosis following kinetochore differentiation. Genetics 130 827-841

Kienitz A, Vogel C, Morales I, Muller R & Bastians H 2005 Partial downregulation of MAD1 causes spindle checkpoint inactivation and aneuploidy, but does not confer resistance towards taxol. Oncogene 24 4301-4310

Kikuchi K, Izaike Y, Noguchi J, Furukawa T, Daen FP, Naito K & Toyoda Y 1995 Decrease of histone H1 kinase activity in relation to parthenogenetic activation of pig follicular oocytes matured and aged in vitro. J Reprod Fert 105 325-330

Kikuchi K, Naito K, Noguchi J, Shimada A, Kaneko H, Yamashita M, Aoki F, Tojo H & Toyoda Y 2000 Maturation/M-phase promoting factor: a regulator of aging in porcine oocytes. Biol Rep 63 715-722

Kim NH, Moon SJ, Prather RS & Day BN 1996 Cytoskeletal alteration in aged porcine oocytes and parthenogenesis. Mol Rep Dev 43 513-518

Kirsch-Volders M, Cundari E & Verdoodt B 1998 Towards a unifying model for the metaphase/anaphase transition. Mutagenesis 13 321-335

Kitajima TS, Kawashima SA & Watanabe Y 2004 The conserved kinetochore protein shugoshin protects centromeric cohesion during meiosis. Nature 427 510-517

Kitajima TS, Sakuno T, Ishiguro K, Iemura S, Natsume t, Kawashima SA & Watanabe Y 2006. Shugoshin collaborates with protein phosphatase 2A to protect cohesin. Nature 441 46-52

Klein F, Mahr P, Galova M, Buonomo SB, Michaelis C, Nairz K & Nasmyth K 1999 A central role for cohesions in sister chromatid cohesion, formation of axial elements, and recombination during yeast meiosis. Cell 98 91-103

Kline D & Kline JT 1992 Repetitive calcium transients and the role of calcium exocytosis and cell cycle activation in the mouse egg. Dev Biol 149 80-89

Knowlton AL, Lan W & Stukenberg PT 2006 Aurora B is enriched at merotelic attachment sites, where it regulates MCAK. Curr Biol 16 1705-1710

Kosower NS & Kosower EM 1978 The glutathione status of cells. Int Rev Cytol 54 109-160

Kotani S, Tanaka H, Yasuda H & Todokoro K 1999 Regulation of APC by phosphorylation and regulatory factors. J Cell Biol 146 791-800

Kubiak JZ 1989 Mouse oocytes gradually develop the capacity for activation during the metaphase II arrest. Dev Biol 136 537-545

Kuliev A, Cieslak J, Ilkevitch Y & Verlinsky Y 2003 Chromosomal abnormalities in a series of 6733 human oocytes in preimplantation diagnosis for age-related aneuploidies. Rep Biomed Online 6 54-59

Lampson MA, Renduchitala K, Khodjakov A & Kapoor TM 2004 Correcting improper chromosome-spindle attachments during cell division. Nat Cell Biol 6 232-237

Lamson SH & Hook EB 1980 A simple function for maternal age-specific rates of Down syndrome in the 20-to-48 year age range and its biological implications. Am J Hum Genet 32 743-753

Lavoie BD, Hogan E & Koshland D 2002 In vivo dissection of the chromosome condensation machinery: reversibility of condensation distinguishes contributions of condensin and cohesion. J Cell Biol 156 805-815

Lee BH & Amon A 2003 Role of Polo-like kinase CDC5 in programming meiosis I chromosome segregation. Science 300 482-486

Lee BH, Kiburz BM & Amon A 2004 Spo13 maintains centromeric cohesion and kinetochore coorientation during meiosis I. Curr Biol 14 2168-2182

Lee J, Miyano T & Moor RM 2000 Localization of phosphorylated MAP kinase during transition from meiosis I to Meiosis II in pig oocytes. Zygote 8 119-125

Lee J, Iwai T, Yokota T & Yamashita M 2003 Temporally and spatially selective loss of Rec8 protein from meiotic chromosomes during mammalian meiosis. J Cell Sci 116 2781-2790

Lee J, Okada K, Ogushi S, Miyano T, Miyake M & Yamashita M 2006 Loss of Rec8 from chromosome arm and centromere region is required for homologous chromosome separation and sister chromatid separation, respectively, in mammalian oocytes. Cell Cycle 5 1448-1455

Lee JY & Orr-Weaver TL 2001 The molecular basis of sister-chromatid cohesion. Annu Rev Cell Dev Biol 17 753-777

Lee JY, Hayashi-Hagihara A & Orr-Weaver TL 2005 Roles and regulation of the Drosophila centromere cohesion protein MEI-332 family. Phil Trans R Soc B 360 543-552

Lens SMA & Medema RH 2003 The survivin/Aurora B complex: its role in coordinating tension and attachment. Cell Cycle 2 507- 510

Lewis WH & Wright ES 1935 On the early development of the mouse egg. Carnegie Inst Contrib Embryol 25 113-143

Li Y & Benezra R 1996 Identification of a human mitotic checkpoint gene hsMAD2. Science 274 246-248

Liang XW, Zhu JQ, Miao YL, Liu JH, Wei L, Lu SS, Hou Y, Schatten H, Lu KH & Sun QY 2008 Loss of methylation imprint of Snrpn in postovulatory aging mouse oocytes. Biochem Biophys Res Comm 371 16-21

Lim AS, Ho AT & Tsakod MF 1995 Chromosomes of oocytes failing in-vitro fertilization. Hum Rep 10 2570-2575

Liu L, Ju JC & Yang X 1998 Differential inactivation of maturation-promoting factor and mitogen-activated protein kinase following parthenogenetic activation of bovine oocytes. Biol Rep 59 537-545

London SN & Mailhes JB 2001 Tamoxifen-induced alterations in meiotic maturation and cytogenetic abnormalities in mouse oocytes and 1-cell zygotes. Zygote 9 97-104

Longo FJ 1981 Changes in the zona pellucida and plasmalemmae of aging mouse oocytes. Biol Rep 25 399-411

Lorca T, Cruzalegul FH, Fesquet D, Cavadore D, Mery JC, Means A & Doree M 1993 Calmodulin-dependent protein kinase II mediates inactivation of MPF and CSF upon fertilization of Xenopus eggs. Nature 366 270-273

Losada A, Yokochi T, Kobayashi R & Hirano T 2000 Identification and characterization of SA/Scc3p subunits in the Xenopus and human cohesion complexes. J Cell Biol 150 405-416

Losada A, Hirano M & Hirano T 2002. Cohesin release is required for sister chromatid resolution, but not for condensing-mediated compaction at the onset of mitosis. Genes Dev 16 3004-3016

Luo X, Fang G, Coldiron M, Lin Y, Yu H, Kirschner MW & Wagner G 2000 Structure of the Mad2 spindle assembly checkpoint protein and its interaction with Cdc20. Nat Struct Biol 7 224-229

Ma W, Zhang D, Hou Y, Li YH, Sun QY, Sun, XF & Wang WH 2005 Reduced expression of MAD2, BCL2, MAP kinase activity in pig oocytes after in vitro aging are associated with defects in sister chromatid segregation during meiosis II and embryo fragmentation after activation. Biol Rep 72 373-383

Magli C, Gianaroli L & Ferraretti AP 2001 Chromosomal abnormalities in embryos. Mol Cell Endocrinol 183 29-34

Magnaghi-Jaulin, Eot-Houllier G, Fulcrand G & Jaulin C 2007 Histone deacetylase inhibitors induce premature sister chromatid separation and override of the mitotic spindle assembly checkpoint. Cancer Res 67 6360-6367

Mailhes JB, Preston RJ & Lavappa KS 1986 Mammalian in vivo assays for aneuploidy in female germ cells Mutat Res 320 139-148

Mailhes JB 1987 Incidence of aneuploidy in rodents. In Vig BK, Sandberg AA, editors. Aneuploidy, Part A: Incidence and Etiology. New York: Alan R. Liss. p 67-101

Mailhes JB & Yuan ZP 1987 Differential sensitivity of mouse oocytes to colchicine-induced aneuploidy. Environ Mol Mutagen 10 183-188

Mailhes JB, Marchetti F & Aardema MJ 1993 Griseofulvin-induced aneuploidy and meiotic delay in mouse oocytes: effect of dose and harvest time. Mutat Res 300 155-163

Mailhes JB & Marchetti F 1994 Chemically-induced aneuploidy in mammalian oocytes. Mutat Res 320 87-111

Mailhes JB & Marchetti F 1994a. The influence of postovulatory ageing on the retardation of mouse oocyte maturation and chromosome segregation induced by vinblastine. Mutagenesis 9 541-545

Mailhes JB, Marchetti F, Phillips Jr. GL & Barnhill DR 1994 Preferential pericentric lesions and aneuploidy induced in mouse oocytes by the topoisomerase II inhibitor etoposide. Terat Carcinogen Mutagen 14 39-51

Mailhes JB 1995 Important biological variables that can influence the degree of chemical-induced aneuploidy in mammalian oocytes and zygotes. Mutat Res 339 155-176

Mailhes JB, Young D, Aardema MJ & London SN 1997a Thiabendazole-induced cytogenetic abnormalities in mouse oocytes. Environ Mol Mutagen 29 367-371

Mailhes JB, Young D & London SN 1997b 1,2-Propanediol-induced premature centromere separation in mouse oocytes and aneuploidy in one-cell zygotes. Biol Rep 57 92-98

Mailhes JB, Young D & London SN 1998 Postovulatory ageing of mouse oocytes in vivo and premature centromere separation and aneuploidy. Biol Rep 58 1206-1210

Mailhes JB, Carabatsos MJ, Young D, London SN, Bell M & Albertini DF 1999 Taxol-induced meiotic maturation delay, spindle defects, and aneuploidy in mouse oocytes and zygotes. Mutat Res 423 79-80

Mailhes JB, Hilliard C, Lowery M & London SN 2002 MG-132, an inhibitor of proteasomes and calpains, induced inhibition of oocyte maturation and aneuploidy in mouse oocytes. Cell & Chromosome 1 1-7

Mailhes JB, Hilliard, Fuseler JW & London SN 2003a Okadaic acid, an inhibitor of protein phosphatase 1 and 2A, induces premature separation of sister chromatids during meiosis I and aneuploidy in mouse oocytes in vitro. Chromosome Res 11 619-631

Mailhes JB, Hilliard C, Fuseler JW & London SN 2003b Vanadate, an inhibitor of tyrosine phosphatases, induced premature anaphase in oocytes and aneuploidy and polyploidy in mouse bone marrow cells. Mutat Res 538 101-107

Mailhes JB, Mastromatteo C & Fuseler JW 2004 Transient exposure to the Eg5 kinesin inhibitor monastrol leads to syntelic orientation of chromosomes and aneuploidy in mouse oocytes. Mutat Res 559 153-167

Marchetti F & Mailhes JB 1995 Variation of mouse oocyte sensitivity to griseofulvin-induced aneuploidy during the second meiotic division. Mutagenesis 10 113-121

Marchetti F, Bishop JB, Lowe X, Generoso WM, Hozier J & Wyrobek AJ 2001 Etoposide induces heritable chromosomal aberrations and aneuploidy during male meiosis in the mouse. Proc Natl Acad Sci USA 98 3952-3957

Marston JH & Chang MC 1964 The fertilizable life of ova and their morphology following delayed insemination in mature and immature mice. J Exp Zool 155 237-252

Marston AL & Amon A 2004 Meiosis: cell-cycle controls shuffle and deal. Nat Rev Mol Cell Biol 5 983-987

Marston AL, Tham WH, Shah H & Amon A 2004 A genome-wide screen identifies genes required for centromeric cohesion. Science 303 1367-1370

McConnell JM, Campbell L & Vincent C 1995 Capacity of mouse oocytes to become activated depends on completion of cytoplasmic but not nuclear meiotic maturation. Zygote 3 45-55

McGuinness BE, Hirota T, Kudo NR, Peters JM & Nasmyth K 2005 Shugoshin prevents dissociation of cohesin from centromeres during mitosis in vertebrate cells. PLoS Biol 3 e86

Meraldi P, Honda R & Nigg EA 2004. Aurora kinases link chromosome segregation and cell division to cancer susceptibility. Curr Opin Genet Dev 14 29-36

Michaelis C, Ciosk R & Nasmyth K 1997 Cohesins: chromosomal proteins that prevent premature separation of sister chromatids. Cell 91 35-45

Michel LS, Liberal V, Chatterjee A, Kirchwegger R, Pasche B, Gerald W, Dobles M, Lorger PK, Murty VVVS & Benerza R 2001 MAD2 haplo-insufficiency causes premature anaphase and chromosome instability in mammalian cells. Nature 409 355-359

Michel L, Diaz-Rodriguez E, Narayan G, Hernando E, Vundavalli M & Benezra R 2004 Complete loss of the tumor suppressor MAD2 causes premature cyclin B degradation and mitotic failure in human somatic cells. Proc Natl Acad Sci USA 101 4459-4464

Miller BM & Adler I-D 1992 Aneuploidy induction in mouse spermatocytes. Mutagenesis 7 69-76

Miyazaki WY & Orr-Weaver TL 1992 Sister-chromatid misbehavior in Drosophila ord mutants. Genetics 132 1047-1061

Miyazaki WY & Orr-Weaver TL 1994 Sister-chromatid cohesion in mitosis and meiosis. Ann Rev Genet 28 167-187

Moos J, Visconti PW, Moore GD, Schultz RM & Kopf GS 1995 Potential role of mitogen-activated protein kinase (MAP) in pronuclear envelope assembly and disassembly following fertilization of mouse eggs. Biol Reprod 53 692-699

Moses RM & Masui Y 1994 Enhancement of mouse egg activation by the kinase inhibitor, 6-dimethylainopurine (6-DMAP). J Exp Zool 270 211-218

Morita Y & Tilly JL 1999 Oocyte apoptosis: like sand through an hourglass. Dev Biol 213 1-17

Munne S 2002 Preimplantation genetic diagnosis of numerical and structural chromosome abnormalities. Reprod Biomed Online 4 183-196

Murray AW 1998 MAP kinases in meiosis. Cell 92 157-159

Musacchio A & Hardwick KG 2002 The spindle checkpoint: structural insights into dynamic signaling. Nat Rev Mol Cell Biol 3 731-741

Nagai T 1987 Parthenogenetic activation of cattle follicular oocytes in vitro with ethanol. Gamete Res 16 243-249

Nasmyth K, Peters JM & Uhlmann F 2000 Splitting the chromosome: Cutting the ties that bind sister chromatids. Science 288 1379-1384

Nasmyth K 2001 Disseminating the genome: joining, resolving, and separating sister chromatids during mitosis and meiosis. Annu Rev Genet 35 673-745

Nasmyth K 2002 Segregating sister genomes: the molecular biology of chromosome separation. Science 297 559-565

Nasmyth K & Schleiffer A 2004 From a single double helix to paired double helices and back. Phil Trans R Soc Lond B 359 99-108

Nasmyth K 2005 How do so few control so many? Cell 120 739-746

Nguyen HG, Chinnappan D, Urano T & Ravid K 2005 Mechanism of Aurora-B degradation and its dependency on intact KEN and A-boxes: identification of an aneuploidy-promoting property. Mol Cell Biol 25 4977-4992.

Nicolaidis P & Petersen MB 1998 Origin and mechanisms of non-disjunction in human autosomal trisomies. Human Reprod 13 313-319

Nicklas RB 1997 How cells get the right chromosomes. Science 275 632-637

O’Neill GT & Kaufman MH 1988 Influence of postovulatory aging on chromosome segregation during the second meiotic division in mouse oocytes: a parthenogenetic analysis. J Exptl Zool 248 125-131

Ono T, Fang Y, Spector DL & Hirano T 2004 Spatial and temporal regulation of condensins I and II in mitotic chromosome assembly in human cells. Mol Biol Cell 15 3296-3308

Orrenius S, Burkitt MJ, Kass GN, Dypbukt JM & Nicotera P 1992 Calcium ions and oxidative cell injury. Ann Neurol 32 S33-S42.

Pacchierotti F 1988 Chemically induced aneuploidy in germ cells of mouse. In Vig BK, Sandberg AA, editors. Aneuploidy, Part B: Induction and Test Systems, Progress and Topics in Cytogenetics, Vol. 7b. New York: Alan R. Liss. p 123-139

Pacchierotti F & Ranaldi R 2006 Mechanisms and risk of chemically induced aneuploidy in mammalian germ cells. Curr Pharm Des 12 1489-1504

Pacchierotti F, Adler I-D, Eichenlaub-Ritter U & Mailhes JB 2007 Gender effects on the incidence of aneuploidy in mammalian germ cells. Environ Res 104 46-69

Pahlavan G, Polanski Z, Kalab P, Golsteyn R, Nigg EA & Maro B 2000 Characterization of polo-like kinase 1 during meiotic maturation of the mouse oocyte. Dev Biol 220 392-400

Panizza S, Tanaka T, Hochwagen A, Eisenhaber F & Nasmyth K 2000 Pds5 cooperates with cohesion in maintaining sister chromatid cohesion. Curr Biol 10 1557-1564

Parisi S, McKay MJ, Molnar M, Thompson MA, van der Spek PJ, van Drunen-Schoenmaker E, Kanaar R, Lehmann E, Hoeijmakers JH & Kohli J 1999 Rec8p, a meiotic recombination and sister chromatid cohesion phosphoprotein of the Rad21p family conserved from fission yeast to humans. Mol Cell Biol 19 3515-3528

Parra M, Viera A, Gomez R, Page J, Benavente R, Santos JL, Rufas JS & Suja JA 2004 Involvement of the cohesion Rad21 and SCP3 in monopolar attachment of sister kinetochores during mouse meiosis I. J Cell Sci 117 1221-1234

Parry JM, Henderson L & Mckay JM 1995 Procedures for the detection of chemically induced aneuploidy: recommendations of a UK Environmental Mutagen Society working group. Mutagenesis 10 1-14

Pasierbek P, Jantsch M, Melcher M, Schleiffer A, Schweizer D & Loidl J 2001 A Caenorhaboditis elegans cohesion protein with functions in meiotic chromosome pairing and disjunction. Genes Dev 15 1349-1360.

Paules RS, Buccione R, Moschel RC, Vande Woude GF & Eppig JJ 1989 Mouse mos protooncogene product is present and functions during oogenesis. Proc Natl Acad Sci USA 86 5395-5399

Pellestor F 1991 Frequency and distribution of aneuploidy in human female gametes. Hum Genet 86 283-288

Pellestor F, Andreo B, Arnal F, Humeau C & Demaille J 2002 Mechanisms of non-disjunction in human female meiosis: the co-existence of two modes of malsegregation evidenced by the karyotyping of 1397 in-vitro unfertilized oocytes. Hum Rep 17 2134-2145

Pellestor F, Andreo, Arnal F, Humeau C & Demaille J 2003 Maternal aging and chromosome abnormalities: new data drawn from in vitro unfertilized aged oocytes. Hum Genet 112 195-203

Pellestor F, Anahory T & Hamamah S 2005 The chromosomal analysis of human oocytes. An overview of established procedures. Hum Rep Update 11 15-32

Pellestor F, Andreo B, Anahory T & Hammamah S 2006 The occurrence of aneuploidy in human: lessons from the cytogenetic studies of human oocytes. Eur J Med Genet 49 103-116

Perez GI, Tao XJ & Tilly JL 1999. Fragmentation and death (a.k.a. apoptosis) of ovulated oocytes. Mol Human Rep 5 414-420

Peter M, Castro A, Lorca T, Le Peuch C, Magnaghi-Jaulin Doree M & Labbe JC 2001 The APC is dispensable for first meiotic division in Xenopus oocytes. Nature Cell Biol 3 83-87

Peters JM 2002 The anaphase-promoting complex: proteolysis in mitosis and beyond. Mol Cell 9 931-943

Petronczki M, Siomos MF & Nasmyth K 2003 Un Ménage à Quatre: The molecular biology of chromosome segregation in meiosis. Cell 112 423-440

Pickering SJ, Johnson MH, Braude Pr & Houliston E 1988 Cytoskeletal organization in fresh, aged and spontaneously activated human oocytes. Hum Rep 3 978-989

Plachot M 2003 Genetic analysis of the oocyte - A review. Placenta 24 S66-S69

Plachot M, deGrouchy J, Junca AM, Mandelbaum J, Salat-Baroux J & Cohen J 1988. Chromosome analysis of human oocytes and embryos: does delayed fertilization increase chromosome imbalance? Hum Rep 3 125-127

Prawitt D, Brixel L, Spangenberg C. Eshkind L, Heck R, Oesch F, Zabel B & Bockamp E 2004 RNAi knock-down mice: an emerging technology for post-genomic functional genetics. Cytogenet Genome Res 105 412-421

Prieto I, Suja JA, Pezzi N, Kremer L, Martinez AC, Rufas JS & Barbero JL 2001 Mammalian STAG3 is a cohesion specific to sister chromatid arms in meiosis I. Nat Cell Biol 3 761-766

Prieto I, Tease C, Pezzi N, Buesa JM, Ortega S, Kremer L, Martinez A, Martinez AC, Hulten MA & Barbero JL 2004 Cohesion component dynamics during meiotic prophase I in mammalian oocytes. Chromosome Res 12 197-213

Prieto I, Pezzi N, Buesa JM, Kremer L, Barthelemy I, Carreiro C, Roncal F, Martinez A, Gomez L, Fernandez R, Martinez AC & Barbero JL 2002 STAG2 and Rad21 mammalian mitotic cohesions are implicated in meiosis. EMBO Rep 3 543-550

Rabitsch KP, Gregan J, Schleiffer A, Javerzat JP, Eisenhaber F & Nasmyth K 2004 Two fission yeast homologs of Drosophila Mei-S332 are required for chromosome segregation during meiosis I and II. Curr Biol 14 287-301

Racowsky C 1993 Somatic control of meiotic status in mammalian oocytes, In Haseltine FP, Heyner S, editors. Meiosis II, Contemporary Approaches to the Study of Meiosis. Wash. D.C.: Amer Assoc Adv Science Press. p 107-116

Rajagopalan H & Lengauer C 2004 Aneuploidy and cancer. Nature 432 338-341

Revenkova E, Eijpe M, Heyting C, Gross B & Jessberger R 2001 Novel meiosis-specific isoform of mammalian SMC1. Mol Cell Biol 21 6984-6998

Revenkova E & Jessberger R 2005 Keeping sister chromatids together: cohesins in meiosis. Reproduction 130 783-790

Rhodes DR, Yu J, Shanker K, Deshpande N, Varambally R, Ghosh D, Barrette T, Pandey A & Chinnaiyan AM 2004 Large-scale meta-analysis of cancer microarray data identifies common transcriptional profiles of neoplastic transformation and progression. Proc Natl Acad Sci USA 101 9309-9314

Riedel CG, Katis VL, Katou Y, Itoh T, Helmhart W, Galova M, Petronczki M, Gregan J, Cetin B, Mudrak I, Ogris E, Mechtler K, Pelletier L, Bucholz F, Shirahige K & Nasmyth K 2006 Protein phosphatase 2A protects centromeric sister chromatid cohesion during meiosis I. Nature 441 53-61

Rieder CL & Palazzo RE 1992 Colcemid and the mitotic cycle. J Cell Sci 102 387-392

Rieder CL & Salmon ED 1998 The vertebrate cell kinetochore and its role during mitosis. Trends Cell Biol 8 310-318

Rieder CL, Schultz A, Cole R & Sluder G 1994 Anaphase onset in vertebrate somatic cells is controlled by a checkpoint that monitors sister kinetochore attachment at the spindle. J Cell Biol 127 1301-1310

Rieder CL & Maiato H 2004 Stuck in division or passing through: what happens when cells cannot satisfy the spindle assembly checkpoint. Dev Cell 7 637-651

Rime H, Neant I, Guerrier P & Ozon R 1989 6-dimethylaminopurine (6-DMAP), a reversible inhibitor of the transition to metaphase during the first meiotic cell division of the mouse oocyte. Dev Biol 133 169-179

Rodman TC 1971 Chromatid disjunction in unfertilized ageing oocytes. Nature 233 191-193

Romanienko PJ & Camerini-Otero RD 2000 The mouse Spo11 gene is required for meiotic chromosome synapsis. Mol Cell 6 975-987

Rose D, Thomas W & Holm C 1990 Segregation of recombined chromosomes in meiosis requires DNA topoisomerase II. Cell 60 1009-1017

Rosenbusch B 2004 The incidence of aneuploidy in human oocytes assessed by conventional cytogenetic analysis. Hereditas 141 97-105

Russo A & Pacchierotti F 1988 Meiotic arrest and aneuploidy induced by vinblastine in mouse oocytes. Mutat Res 202 215-221

Sagata N 1996 Meiotic metaphase arrest in animal oocytes: its mechanisms and biological significance. Trends Cell Biol 6 22-28

Sagata N 1997 What does Mos do in oocytes and somatic cells? BioEssays 19 13-21

Saito H, Koike K, Saito T, Nohara M, Kawagoe S & Hiroi M 1993 Aging changes in the alignment of chromosomes after human chorionic gonadotropin stimulation may be a possible cause of decreased fertility in mice. Horm Res 39 28-31

Sakai N & Endo A 1988 Effects of delayed mating on preimplantation embryos in spontaneously ovulated mice. Gamete Res 19 381-385

Sakurada K, Ishikawa H & Endo A 1996 Cytogenetic effects of advanced maternal age and delayed fertilization on first-cleavage mouse embryos. Cytogen Cell Genet 72 46-49

Salah SN & Nasmyth K 2000 Destruction of the securing Pds1p occurs at the onset of anaphase during both meiotic divisions in yeast. Chromosoma 109 27-34

Salic A, Waters JC & Mitchison TJ 2004 Vertebrate shugoshin links sister centromere cohesion and kinetochore microtubule stabilization in mitosis. Cell 118 567-578

Salmon ED, Cimini D, Cameron LA & DeLuca JG 2005 Merotelic kinetochores in mammalian tissue cells. Phil Trans R Soc B 360 553-568

Schlecht U & Primig M 2003 Mining meiosis and gametogenesis with DNA microarrays. Reproduction 125 447-456

Schönthal S 1992 Okadaic acid-a valuable new tool for the study of signal transduction and cell cycle regulation? New Biol 4 16-21

Schultz RM 1988 Regulatory functions of protein phosphorylation in meiotic maturation of mouse oocytes in vitro. In Haseltine FP, First NL, editors. Meiotic Inhibition: Molecular Control of Meiosis. New York: Alan R Liss. p 137-151

Schultz RM, Montgomery RR & Belanoff JR 1983 Regulation of mouse oocyte maturation: implication of a decrease in oocyte cAMP and protein dephosphorylation in commitment to resume meiosis. Dev Biol 97 264-273

Schultz RM 1986 Molecular aspects of mammalian oocyte growth and maturation, in Rossant J, Pederson RA, editors. Experimental Approach to Mammalian Embryonic Development. Cambridge UK: Cambridge University Press. p 195-237

Schwartz DA & Schultz RM 1991 Stimulatory effect of okadaic acid, an inhibitor of protein phosphatases, on nuclear envelope breakdown and protein phosphorylation in mouse oocytes and one-cell zygotes. Dev Biol 145 119-127

Segers I, Adriaenssens T, Coucke W, Cortvrindt R & Smitz J 2008 Timing of nuclear maturation and postovulatory aging in oocytes of in vitro-grown mouse follicles with of without oil overlay. Biol Rep 78 859-868

Shah JV, Botvinick E, Bonday Z, Fumari F, Berns M & Cleveland DW 2004 Dynamics of centromere and kinetochore proteins; implications for checkpoint signaling and silencing. Curr Biol 14 942-952

Shamu CE & Murray AW 1992 Sister chromatid separation in frog extracts requires DNA topoisomerase II activity during anaphase. J Cell Biol 117 921-934

Shang C, Hazbun TR, Cheesman IM, Arand J, fields S, Drubin DG & Barnes G 2003 Kinetochore protein interactions and their regulation by Aurora kinase Ipl1p. Mol Biol Cell 14 3342-3355

Shaver EL & Carr DH 1967 Chromosome abnormalities in rabbit blastocysts following delayed fertilization. J Reprod Fert 14 415-420

Shonn MA, McCarroll R & Murray AW 2000 Requirement of the spindle checkpoint for proper chromosome segregation in budding yeast meiosis. Science 289 300-303

Simerly C, Balczon R, Brinkley BR & Schatten G 1990 Microinjected centromere [corrected] kinetochore antibodies interfere with chromosome movement in meiotic and mitotic mouse oocytes. J Cell Biol 111 1491-1504

Singh B & Arlinghaus RB 1997 Mos and the cell cycle. Prog Cell Cycle Res 3 251-259

Siomos MF, Badrinath A, Pasierbek P, Livingstone D, White J, Glotzer M & Nasmyth K 2001 Separase is required for chromosome segregation during meiosis I in Caenorhabditis elegans. Curr Biol 11 1825-1835

Sluder G 1979 Role of spindle microtubules in the control of cell cycle timing. J Cell Biol 80 674-691

Sluder G & Rieder CL 1993 The events and regulation of anaphase onset. In Vig BK, editor. Chromosome Segregation and Aneuploidy, NATO ASI Series, Vol. H72. Berlin: Springer-Verlag. p 211-224

Sluder G & McCollum D 2000 The mad ways of meiosis. Science 289 254-255

Smith AL & Lodge JR 1987 Interactions of aged gametes: in vitro fertilization using in vitro-aged sperm and in vivo-aged ova in the mouse. Gamete Res 16 47-56

Sobajima T, Aoki F & Kohomoto K 1993 Activation of mitogen-activated protein kinase during meiotic maturation in mouse oocytes. J Rep Fert 97 389-394

Sonada E, Matsusaka T, Morrison C, Vagnarelli P, Hoshi O, Ushiki T, Nojima K, Fukagawa T, Waizenegger IC, Peters JM, Earnshaw WC & Takeda S 2001 Scc1/Rad21/Mcd1 is required for sister chromatid cohesion and kinetochore function in vertebrate cells. Dev Cell 1 759-770

Stein P, Svoboda P & Schultz RM 2003 Transgenic RNAi in mouse oocytes: a simple and fast approach to study gene function. Dev Biol 256 187-193

Stern BM 2002 Mitosis: aurora gives chromosomes a healthy stretch. Curr Biol 12 R316-318

Steuerwald N, Cohen J, Herrera RJ, Sandalinas M & Brenner CA 2001 Association between spindle assembly checkpoint expression and maternal age in human oocytes. Mol Hum Reprod 7 49-55

Steuerwald N, Steuerwald MD & Mailhes JB 2005 Postovulatory aging of mouse oocytes leads to decreased MAD2 transcripts and increased frequencies of premature centromere separation and anaphase. Mol Hum Rep 11 623-630

Strausfeld U, Labbe JC, Fesquet D, Cavadore JC, Picard A, Sadhu K, Russell P & Doree M 1991 Dephosphorylation and activation of a p34cdc2 / cyclin B complex in vitro by human CDC25 protein. Nature 351 242-245

Sumara I, Vorlaufer E, Stukenberg PT, Kelm O, Redemann N, Nigg EA & Peters JM 2002 The dissociation of cohesion from chromosomes in prophase is regulated by Polo-like kinase. Mol Cell 9 515-525

Sumara I, Gimenez-Abian JF, Gerlich D, Hirota T, Kraft C, de la torre C, Ellenberg J & Peters JM 2004 Roles of polo-like kinase 1 in the assembly of functional mitotic spindles. Curr Biol 14 1712-1722

Sun FY, Schmid TE, Schmid E, Baumgartner A & Adler ID 2000 Trichlorfon induces spindle disturbances in V79 cells and aneuploidy in male mouse germ cells. Mutagenesis 15 17-24

Swain JE & Smith GD 2007 Reversible phosphorylation and regulation of mammalian oocyte meiotic chromatin remodeling and segregation. Soc Reprod Fert Suppl 63 343-358

Szollosi D 1971 Morphological changes in mouse eggs due to aging in the fallopian tube. Am J Anat 130 209-226

Szollosi D 1975 Mammalian eggs ageing in the fallopian tubes. In Blandeau RJ, editor. Ageing Gametes. Basel: Karger. p 98-121

Szollosi MS, Debey P, Szollosi D, Rime H & Vautier D 1991 Chromatin behavior under influence of puromycin and 6-DMAP at different stages of mouse oocyte maturation. Chromosoma 100 339-354

Szollosi MS, Kubiak JZ, Debey P, de Pennart H, Szollosi D & Maro B 1993 Inhibition of protein kinases by 6-dimethylaminopurine accelerates the transition to interphase in activated mouse oocytes. J Cell Sci 104 861-872

Takahashi T, Takahashi E, Igarashi H, Tezuka N & Kurachi H 2003 Impact of oxidative stress in aged mouse oocytes on calcium oscillations at fertilization. Mol Rep Dev 66 143-152

Takenaka K, Moriguchi T & Nishida E 1998 Activation of the protein kinase p38 in the spindle assembly checkpoint and mitotic arrest. Science 280 599-602

Tanaka T, Fuchs J, Loidl J & Nasmyth K 2000 Cohesin ensures bipolar attachment of microtubules to sister centromeres and resists their precocious separation. Nat Cell Biol 2 492-499

Tanaka TU, Rachidi N, Janke C, Pereira G, Galova M, Schiebel E, Stark MJ & Nasmyth K 2002 Evidence that the IpI1-Sli15 (Aurora kinase-INCENP) complex promotes chromosome bi-orientation by altering kinetochore-spindle pole connections. Cell 108 317-329

Tang Z, Sun Y, Harley SE, Zou H & Yu H 2004 Human Bub1 protects centromeric sister-chromatid cohesion through Shugoshin during mitosis. Proc Natl Acad Sci USA 101 18012-18017

Tang Z, Shu H, Qi W, Mahmood NA, Mumby MC & Yu H 2006 PP2A is required for centromeric location of Sgo1 and proper chromosome segregation. Dev Cell 10 575-585

Tateno H & Kamiguchi Y 2001 Meiotic stage-dependent induction of chromosome aberrations in Chinese hamster primary oocytes exposed to topoisomerase II inhibitor etoposide. Mutat Res 476 139-148

Tarin JJ, Vendrell FJ, Ten J, Blanes R, Van Blerkom J & Cano A 1996 The oxidizing agent tertiary butyl hydroperoxide induces disturbances in spindle organization, c-meiosis, and aneuploidy in mouse oocytes. Mol Hum Rep 2 895-901

Tatone C, Carbone MC, Gallo R, Monache SD, Di Cola M, Alesse E & Amicarelli F 2006 Age-associated changes in mouse oocytes during postovulatory in vitro culture: possible role for meiotic kinases and survival factor BCL2. Biol Rep 74 395-402

Taylor SS, Ha E & McKeon F 1998 The human homolog of Bub3 is required for kinetochore localization of Bub1 and a Mad3/Bub1-related protein kinase. J Cell Biol 142 1-11

Taylor SS, Hussein D, Wang Y, Elderkin S & Morrow CJ 2001 Kinetochore localization and phosphorylation of the mitotic checkpoint components Bub1 and BubR1 are differentially regulated by spindle events in human cells. J Cell Sci 114 4385-4395

Taylor SS, Maria I, Scott F & Holland AJ 2004 The spindle checkpoint: a quality control mechanism which ensures accurate chromosome segregation. Chromosome Res 12 599-616

Terret ME, Wassmann K, Waizenegger I, Maro B, Peters JM & Verlhac MH 2003 The meiosis I-to-meiosis II transition in mouse oocytes requires separase activity. Curr Biol 13 1797-1802

Thein KH, Kleylein-Sohn J, Nigg EA & Gruneberg U 2007 Astrin is required for the maintenance of sister chromatid cohesion and centrosome integrity. J Cell Biol 178 345-354

Tiveron C, Marchetti F, Bassani B & Pacchierotti F 1992 Griseofulvin-induced aneuploidy and meiotic delay in female mouse germ cells. I. Cytogenetic analysis of metaphase II oocytes. Mutat Res 266 143-150

Tombes RM, Semerly C, Borisy GG & Schatten G 1992. Meiosis, egg activation, and nuclear envelope breakdown are differentially reliant on Ca2+ independent in the mouse oocyte. J Cell Biol 117 799-811

Tomonaga T, Nagao K, Kawasaki Y, Furuya K, Murakami A, Morishita J, Yuasa Y, Sutani T, Kearsey SE, Uhlmann F, Nasmyth K & Yanagida M 2000 Genes Dev 14 2757-2770

Tong C, Fan HY, Lian l, Li SW, Chen DY, Schatten H & Sun QY 2002 Polo-like kinase 1 is a pivotal regulator of microtubule assembly during mouse oocyte meiotic maturation, fertilization, and early embryonic mitosis. Biol Rep 67 546-554

Toyooka Y, Tsunekawa N, Akasu R & Noce T 2003 Embryonic stem cells can form germ cells in vitro. Proc Natl Acad Sci USA 100 11457-11462

Toth A, Ciosk R, Uhlmann F, Galova M, Schleiffer A & Nasmyth K 1999 Yeast cohesin complex requires a conserved protein, Eco1p(Ctf7), to establish cohesion between sister chromatids during DNA replication. Genes Dev 13 320-333

Tsurumi C, Hoffmann S, Geley S, Graeser R & Polanski Z 2004 The spindle assembly checkpoint is not essential for CSF arrest of mouse oocytes. J Cell Biol 167 1037-1050

Uhlmann F, Lottspeich F & Nasmyth K 1999 Sister-chromatid separation at anaphase onset is promoted by cleavage of the cohesion subunit Scc1. Nature 400 37-42

Uhlmann F 2001 Chromosome cohesion and segregation in mitosis and meiosis. Curr Opin Cell Biol 13 754-761

Uhlmann F 2003a Chromosome cohesion and separation: from men to molecules. Curr Biol 13 R104- R114

Uhlmann F 2003b Separase regulation during mitosis. Biochem Soc Symp 70 243-251

Vagnarelli P & Earnshaw WC 2004 Chromosomal passengers: the four-dimensional regulation of mitotic events. Chromosoma 113 211-222

Vandre DD & Willis VL 1992 Inhibition of mitosis by okadaic acid: possible involvement of a protein phosphatase 2A in the transition from metaphase to anaphase. J Cell Sci 101 79-91

Van Heemst D & Heyting C 2000 Sister chromatid cohesion and recombination in meiosis. Chromosoma 109 10-26

Verlhac M.H., Kubiak JZ, Clarke HJ & Maro BH 1994 Microtubule and chromatin behavior follow MAP kinase activity but not MPF activity during meiosis in mouse oocytes. Development 120 1017-1025

Vialard F, Petit C, Bergere M, Gomes DM, Martel-Petit V, Lombroso R, Ville Y, Gerard H & Selva J 2006 Evidence of a high proportion of premature unbalanced separation of sister chromatids in the first polar bodies of women of advanced age. Hum Rep 21 1172-1178

Vickers AD 1969 Delayed fertilization and chromosomal anomalies in mouse embryos. J Rep Fert 20 69-76

Vigneron S, Prieto S, Bernis C, Labbe JC, Castro A & Lorca T 2004 Kinetochore localization of spindle checkpoint proteins: Who controls whom. Mol Biol Cell 15 4584-4596

Vincent C, Cheek TR & Johnson MH 1992 Cell cycle progression of parthenogenetically activated mouse oocytes to interphase is dependent on the level of internal calcium. J Cell Sci 103 389-396

Full Text: PDF


  • There are currently no refbacks.

Copyright © 2017 Lorem Ipsum Press