摘要
生发泡期卵母细胞染色质的一个共同特点是:随卵泡生长,弥散在整个细胞核区域中的染色质逐渐发生凝集,依物种不同,或在核仁周围凝集成环或形成其它凝集结构。先前有关卵母细胞染色质的研究主要集中在不同构型与减数分裂及发育能力的关系方面,对参与染色质凝集的分子及信号通路还知之不多。最近的研究表明后成性修饰,包括DNA甲基化及组蛋白修饰在调控染色质结构及基因表达过程中发挥了重要作用。本文以小鼠卵母细胞为实验模型,利用免疫荧光技术及相关特异性抑制剂,系统研究了DNA甲基化、组蛋白磷酸化和乙酰化在卵母细胞GV期染色质及MII期染色体凝集过程中的作用。结果发现:
1、MPF及MAPK抑制剂处理后,卵母细胞GV期染色质凝集正常发生,表明MPF与MAPK均不参与染色质凝集
2、卵母细胞GV期染色质与MII期染色体凝集均伴随DNA甲基化水平升高,但甲基化特异性抑制剂5-aza-cytidine处理后,染色质与染色体凝集仍然发生,表明DNA甲基化不是卵母细胞染色质及染色体凝集所必需
3、卵母细胞GV期染色质与MII期染色体凝集均伴随组蛋白磷酸化水平升高,但磷酸化特异性抑制剂ZM447439处理后,染色质与染色体凝集仍然发生,表明组蛋白磷酸化不是卵母细胞染色质及染色体凝集所必需
4、卵母细胞GV期染色质凝集伴随着组蛋白乙酰化水平升高,但HDACs抑制剂TSA处理后,GV染色质不再继续凝集但乙酰化水平仍然升高,表明HDACs参与GV期染色质凝集,但并非通过组蛋白乙酰化/去乙酰化途径
5、卵母细胞MII期染色体凝集伴随组蛋白去乙酰化,TSA处理后,组蛋白保持乙酰化状态,但染色体凝集正常发生,表明组蛋白去乙酰化不是染色体凝集所必需
6、GV期及MII期卵母细胞中,HATs/HDACs保持动态平衡,协同调节组蛋白的乙酰化/去乙酰化状态
受核胞质制备是核移植过程中的重要环节,作为受核胞质的卵母细胞必须除去自身遗传物质,此过程通常称之为去核。去核不完全往往会使克隆胚染色体出现多倍性或非整倍性现象,直接导致克隆胚胎发育失败。化学辅助去核是指利用药物处理卵母细胞使之形成含核物质的突起,再通过显微操作去除突起而去核。这种去核方式因其操作简便、对卵母细胞发育能力损伤较小而得到广泛应用。目前所用化学辅助去核药物一般为微管抑制剂,通过破坏纺锤体形成进而扰乱染色体分离,从而实现辅助去核。咖啡因和MG132通常用于升高MPF活性,尚未见将这两种药物应用于卵母细胞辅助去核的报道。本文首先研究了成熟培养液中添加半胱胺和胱氨酸对山羊卵母细胞孤雌胚胎及核移植胚胎发育的促进作用,然后利用此优化的成熟培养系统,首次将咖啡因和MG132这两种非微管抑制剂应用于山羊卵母细胞的化学辅助去核,结果发现:
1、成熟培养液中添加100μM半胱胺和200μM胱氨酸可促进山羊孤雌胚胎的体外发育
2、成熟培养液中添加100μM半胱胺和200μM胱氨酸可促进山羊体细胞核移植胚胎的体外发育
3、1mM的Caffeine和5μM的MG132处理30分钟可诱导80%山羊MII期卵母细胞产生胞质突起
4、与Demecolcine相比,Caffeine和MG132处理后不破坏山羊MII期卵母细胞的纺锤体结构
5、Caffeine和MG132处理诱导胞质突起后不损害山羊孤雌胚胎的体外发育能力
6、与盲吸去核相比,Caffeine和MG132诱导突起后去核率较高,并能提高核移植胚胎的体外发育能力
7、Caffeine处理诱导产生的胞质突起的持续时间长于MG132处理,与Demecolcine处理相当
8、Caffeine辅助去核的体细胞核移植胚胎可发育到期
A common feature in the configuration of germinal vesicle (GV) chromatin in most species was that diffuse chromatin condenses into perinucleolar rings or other corresponding condensed configurations with or without the perinucleolar rings, depending on species. Previous investigations focused on the correlation between different chromatin configuration and the acquisition of meiotic as well as developmental competence of oocytes, little was known concerning the cellular and signaling pathways regulating chromatin condensation.Recent studies have revealed that epigenetic modifications,such as DNA methylation and histone modifications,played important roles in the regulation of chromatin structure and gene expression. In the present study, using immunofluorescence and distinct specific inhibitors, we conducted a comprehensive analysis of the DNA methylation,histone phosphorylation and acetylation during chromatin and chromosome condensation in mouse oocytes. Results as follows:
1.GV oocytes treated with inhibitors of MPF and MAPK underwent chromatin condensation, suggested that chromatin condensation was not correlated with MPF and MAPK activities.
2. GV chromatin and MII chromosome condensation were associated with increased levels of DNA methylation, however, condensations remained occurred after treating oocytes with 5-aza-cytidine, a specific inhibitor of DNA methylation. These results indicated that DNA methylation was not required for chromatin and chromosome condensation.
3. GV chromatin and MII chromosome condensation were associated with increased levels of histone phosphorylation, however, condensations remained occurred after treating oocytes with ZM447439, a specific inhibitor of histone phosphorylation. These results indicated that histone phosphorylation was not required for chromatin and chromosome condensation.
4. GV chromatin condensation was associated with increased levels of histone acetylation, however,chromatin condensation did not occur while levels of acetylation remained increased after treatment with TSA,a inhibitor of HDACs. These evidences provided for a role of HDACs in the control of GV chromatin condensation,but not take place via histone acetylation/deacetylation-dependent mechanism.
5. MII chromosome condensation was associated with histone deacetylation, chromatin condensation remained continued while levels of acetylation increased after treatment with TSA. These results indicated that histone deacetylation was not required for chromosome condensation.
6. Dynamic equilibrium between HATs and HDACs activitie were presented in GV and MII oocytes, by which, to regulate the acetylation/deacetylation state. The preparation of recipient cytoplasm was one of the key steps of somatic cell nuclear transfer,the removal of chromosomes from recipient oocytes was named enucleation. Improper measures used in enucleation resulted in problems as aneuploidy or polyploidy with subsequent detrimental effects on development.Chemically assisted enucleation was an approach of choice, oocytes were exposed to the drug, and as a result of treatment, a cortical protrusion formed on the surface of the oocyte containing all the oocyte chromosomes, this protrusion can be easily removed by micromanipulation techniques.Chemically assisted enucleation seemed to be a very attractive procedure that could simplify conventional enucleation methods with a minimal decrease of the cytoplasmic volume and without reducing the viability of the resulting cytoplast. Chemically assisted enucleation previously reported mainly focused on anti-microtubule drugs.Exposured to these drugs impaired spindle rotation, altered chromosome migration, and thereby, resulted in the generation of enucleated oocytes.
Caffeine and MG132 were used to increase MPF activity, however,no study about these drugs has been reported in chemically assisted enucleation procedure. In the present study, effects of supplementation of cysteamine and cystine on the developmental capacity for parthenogenetic and SCNT embryos were investigated, and then, using the improved system of IVM, the protocol for caffeine and MG132 assisted enucleation of goat oocytes was first applied. Results as follows:
1. Simultaneous supplementation of 100μM cysteamine and 200μM cystine to maturation medium improved parthenogenetic blastulation of goat oocytes.
2. Simultaneous supplementation of 100μM cysteamine and 200μM cystine to maturation medium improved SCNT blastulation of goat oocytes.
3. A 30-min treatment with 1mM caffeine or 5μM MG132-induced cytoplasmic protrusions in over 80% of the MII oocytes.
4. Treatment with caffeine or MG132 did not impair the structure of oocyte spindle compared with demecolcine.
5. Treatment with caffeine or MG132 did not compromise the capacity of parthenogenetic development.
6. Rates of enucleation and blastocyst formation were significantly higher after caffeine or MG132-assisted than after blind aspiration enucleation.
7. After withdraw exposured to drugs, caffeine had longer persistence of the cytoplasmic protrusions than MG132, and equivalent to demecolcine.and so,caffeine was more suitable to application.
8. Production of live cloned goat from SCNT embryos using caffeine-enucleated cytoplasts demonstrated the potential of this new agent for nuclear transfer practice.
引文
Abeydeera LR,WangW-H, Cantley TC, Rieke A & Day BN. Coculture with follicular shell pieces can enhance the developmental competence of pig oocytes after in vitro fertilization: relevance to intracellular glutathione. Biol Reprod.1998, 58:213–218.
Adams RR, Maiato H, Earnshaw WC, Carmena M. 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 2001; 153:865–880.
Adenot, P.G., Szollosi, M.S., Chesne, P., et al. (1997). In vivo aging of oocyte influences the behavior of nuclei transferred to enucleated rabbit oocytes. Mol. Reprod. Dev. 46, 325–336.
Ajiro, K., Yasuda, H. and Tsuji, H. (1996). Vanadate triggers the transition from chromosome condensation to decondensation in a mitotic mutant (tsTM13) inactivation of p34cdc2/H1 kinase and dephosphorylation of mitosis-specific histone H3. Eur. J. Biochem. 241, 923-930.
Akiyama T., Kim J. M., Nagata M., Aoki F. Regulation of Histone Acetylation During Meiotic Maturation in Mouse Oocytes Mol. Reprod. Dev. 2004;69:222–227. Allemain GL. Deciphering the MAPK kinase pathway. Progress in Growth Factor Res 1994, 5: 291-334
Anderiesz C, Fong CY, Bongso A, Trounson AO. Regulation of human and mouse oocyte maturation in vitro with 6-dimethylaminopurine. Hum Reprod 2000; 15:379–388.
Assou S, Anahory T, Pantesco V, Le Carrour T, Pellestor F, Klein B, Reyftmann L, Dechaud H, De Vos J, Hamamah S. The human cumulus–oocyte complex gene-expression profile. Hum Reprod 2006;21:1705–1719.
Baguisi A,Behboodi E,Melican DT,Pollock JS,Destrempes MM,Cammuso C,Williams JL,Nims SD,Porter CA,Midura P,Palacios MJ,Ayres SL,Denniston RS et al., 1999.Production of goats by somatic cell nuclear transfer.Nat Biotechnol,17:456-61
Baguisi, A., and Overstrom, E.W. (2000). Induced enucleation in nuclear transfer procedures to produce cloned animals. Theriogenology 54, 209 (abstract).
Balasubramanyam, K., Altaf, M., Varier, R. A., Swaminathan, V., Ravindran, A., Sadhale, P. P., and Kundu, T. K. Polyisoprenylated Benzophenone, Garcinol, a Natural Histone Acetyltransferase Inhibitor, Represses Chromatin Transcription and Alters Global Gene Expression. (2004) J. Biol. Chem. 279, 33716-33726
Bannister, A.J., Zegerman, P., Partridge, J.F., Miska, E.A., Thomas, J.O., Allshire, R.C., Kouzarides, T., 2001. Selective recognition of methylated lysine 9 on histone H3 by the HP1 chromo domain. Nature 410, 120–124.
Beux GL, Richard FJ, Sirard MA. 2003. Effect of cycloheximide, 6-DMAP, roscovitine and butyrolactone I on resumption of meiosis in porcine oocytes. Theriogenology 60:1049-1058.
Bordignon V, Smith LC. Telophase enucleation: an improved method to prepare recipient cytoplasts for use in bovine nuclear transfer. Mol Reprod Dev 1998; 49: 29-36.
Bornslager EA, Mattel P, Schultz RM 1986. Involvement of cAMP-dependent protein kinase and protein phosphorylation in regulation of mouse oocyte maturation. Dev Biol,114:453-462
Borsuk E, Milik E. Fully grown mouse oocyte contains transcription inhibiting activity which acts through histone deacetylation. Mol Reprod Dev, 2005, 71(4): 509-515.
Bourc’his D, Xu GL, Lin CS, Bollman B & Bestor TH 2001 Dnmt3L and the establishment of maternal genomic imprints. Science 294 2536–2539.
Brownell JE,Allis CD.Special HATs for special occasions:linking histon acetylation to chromatin assembly and gene activation.Curr Opin Genet Dev,1996,6(2):176-184.
Bui HT, Hwang KC, Kim JH, VAN Thuan,Wakayama T,Miyano T. Effect of Vanadate on the Chromatin Configuration in Pig GV-oocytes.J Reprod Dev 2009;55:367-372
Bui HT, Van Thuan N, Kishigami S, Wakayama S, Hikichi T, Ohta H, Mizutani E, Yamaoka E,Wakayama T, Miyano T. Regulation of chromatin and chromosome morphology by histone H3 modifications in pig oocytes. Reproduction 2007;133:371–382.
Bui HT, Yamaoka E, Miyano T. Involvement of histone H3 (Ser10) phosphorylation in chromosome condensation without Cdc2 kinase and mitogen-activated protein kinase activation in pig oocytes. Biol Reprod 2004; 70:1843–1851
Campbell KHS.1999.Nuclear equivalence,nuclear transfer,and the cell cycle. Cloning,1:3-15. Campbell, K.H.S., McWhir, J., Ritchie, W.A., et al. (1996). Sheep cloned by nuclear transfer from a cultured cell line. Nature 380, 64–66.
Carmena M, Earnshaw WC. The cellular geography of aurora kinases. Nat Rev Mol Cell Biol 2003;4:842–854.
Castro A, Bernis C, Vigneron S, Labbe′J, Lorca T. The anaphase- promoting complex: a key factor in the regulation of cellcycle. Oncogene 2005, 24: 314-325. Chen HW, Tini M, Evans RM.HATs on and beyond chromatin.Current opinion in cell biology,2001,13:218–224
Chen T , Zhang YL, Jiang, Y., Liu, S.Z., Schatten, H., Chen, D.Y. and Sun, Q.Y. (2004) The DNA methylation events in normal and cloned rabbit embryos. FEBS Lett., 578, 69–72. Cheong H, Takahashi Y, Kanagawa H. Birth of mice after transplantation of early cell-cycle-stage embryonic nuclei into enucleated oocytes. Biol Reprod 1993; 48: 958-963.
Chesne P,Adenot PG,Viglietta C,Baratte M,Boulanger L,Renard JP.2002 Cloned rabbits produced by nuclear transfer from adult somatic cells.Nat.Biotechnol.20:366-369 Cheung, P., Allis, C.D., Sassone-Corsi, P., 2000. Signaling to chromatin through histone modifications. Cell 103, 263–271.
Cibelli JB,Stice SL,Golouke PJ,Kane JJ,Jerry J.Cloned transgenic calves produced from nonqyienscent fetal fibroblasts.Science,1998,280:1256-1258
Combelles CM, Cekleniak NA, Racowsky C, Albertini DF. Assessment of nuclear and cytoplasmic maturation in in-vitro matured human oocytes. Hum Reprod 2002;17:1006–1016.
Costa-Borges N, Paramio MT, Calderón G, SantalóJ, Ibá?ez E. Antimitotic treatments for chemically assisted oocyte enucleation in nuclear transfer procedures. Cloning Stem Cells. 2009 Mar;11(1):153-66.
Davis R. The mitogen-activated protein kinase signal transduction pathway. J Biol Chem 1993, 268:14553-14556
De Azevedo WF, Leclerc S, Meijer L, Havlicek L, Stmad M, Kim SH. Inhibition ofcyclin-dependent kinases by purine analogues: crystal structure of human cdk2 complexed with roscovitine. Eur J Biochem 1997;243:518–26.
De La Fuente R, Viveiros MM, Burns KH, Adashi EY, Matzuk MM, Eppig JJ. Major chromatin remodeling in the germinal vesicle (GV) of mammalian oocytes is dispensable for global transcriptional silencing but required for centromeric heterochromatin function. Dev Biol 2004; 275:447–458
de Matos DG & Furnus CC 2000 The importance of having high glutathione level after bovine in vitro maturation on embryo development: effect of b-mercaptoethanol, cysteine, and cystine. Theriogenology 53 761–771.
de Matos DG, Furnus CC & Moses DF 1997 Glutathione synthesis during in vitro maturation of bovine oocytes: role of cumulus cells. Biology of Reproduction 57 1420–1425.
de Matos DG, Furnus CC, Moses DF & Baldassarre H 1995 Effect of cysteamine on glutathione level and developmental capacity of bovine oocyte matured in vitro. Molecular Reproduction and Development 42 432–436.
de Matos DG, Gasparrini B, Pasqualini SR & Thompson JG 2002 Effect of glutathione synthesis stimulation during in vitro maturation of ovine oocytes on embryo development and intracellular peroxide content. Theriogenology 57 1443–1451.
de Matos DG, Nogueira D, Cortvrind R, Herrea C, Adriaenssens T, Pasqualini RS et al. 2003 Capacity of adult and prepubertal mouse oocytes to undergo embryo development in the presence of cysteamine. Molecular Reproduction and Development 64 214–218.
de Ruijter AJ, van Gennip AH, Caron HN, Kemp S, van Kuilenburg AB. Histone deacetylases (HDACs): characterization of the classical HDAC family. Biochem J 2003; 370:737–749.
de Vantery Arrighi C, Campana A & Schorderet-Slatkine S 2000 A role for the MEK-MAPK pathway in okadaic acid-induced meiotic resumption of incompetent growing mouse oocytes. Biology of Reproduction 63 658–665.
Debey P, Szollosi MS, Szollosi D, Vautier D, Girousse A, Besombes D. Competent mouse oocytes isolated from antral follicles exhibit different chromatin organization and follow different maturation dynamics. Mol Reprod Dev 1993;36:59–74.
Dinnyes, A., Dai, Y., Jiang, S., et al. (2000). High developmental rates of vitrified bovine oocytes following parthenogenetic activation, in vitro fertilization, and somatic cell nuclear transfer. Biol. Reprod. 63, 513–518.
Dominko, T., Chan, A., Simerly, C., et al. (2000). Dynamic imaging of the metaphase II spindle and maternal chromosomesin bovine oocytes: implications for enucleation efficiency verification, avoidance of parthenogenesis, and successful embryogenesis. Biol. Reprod. 62, 150–154.
Downs SM 1993. Purine control of mouse oocyte maturation: evidence that nonmetabolized hypoxanthine maintains meiotic arrest. Mol Reprod Dev 35(1): 82-94
Downs SM, Hunzicker-Dunn M 1995. Differential regulation of oocyte maturation and cumulus expansion in the mouse oocyte-cumulus cell complex by site-selective analogs of cyclic adenosine monophosphate. Dev Biol, 172(1): 72-85
Elsheikh, A.S., Takahashi, Y., Hishinuma, M., et al. (1997). Developmental ability of mouse late 2-cell stage blastomeres fused to chemically enucleated oocytes in vitro. J. Vet. Med. Sci. 59, 107–113.
Elsheikh, A.S., Takahashi, Y., Kanagawa, H., et al. (1998). Functional enucleation of mouse metaphase II oocytes with etoposide. Jpn. J. Vet. Res. 45, 217–220.
Endo T, Kano K, Naito K. Nuclear histone deacetylases are not required for global histone deacetylation during meiotic maturation in porcine oocytes. Biol Reprod.2008;78:1073-80.
Endo T, Naito K, Aoki F, Kume S, Tojo H. Changes in histone modifications during in vitro maturation of porcine oocytes. Mol Reprod Dev 2005; 71:123–128.
Endo T, Naito K, Kume S, Nishimura Y, Kashima K, Tojo H. Activities of maturation-promoting factor (MPF) and mitogen-activated protein kinase (MAPK) are not required for the global histone deacetylation observed after germinal vesicle breakdown (GVBD) in porcine oocytes. Reproduction 2006; 131:439–447.
Fair T,Hulshof SCJ,Hyttel P,Greve T,Boland M.1997.Nucleus ultrastructure and transcriptional activity of bovine oocytes in preantral and early antral follicles.Mol Reprod Dev 46:208–215
Fair T,Hyttel P,Greve T and Boland M.1996.Nucleolus sructure and transcriptional activity in relation to oocyte diameter in cattle.Mol Reprod Dev 43:503-512
Fang G, Yu H, Kirschner MW. Control of mitotic transitions by the anaphase-promoting complex. Philos. Trans. R. Soc. Lond. B Biol. Sci. 1999, 354: 1583-1590.
Faraone-Mennella, M.R., 2005. Chromatin architecture and functions: the role (s) of poly(ADP-RIBOSE) polymerase and poly(ADPribosyl)ation of nuclear proteins. Biochem. Cell. Biol. 83, 396–404.
Feng Q & Zhang Y 2001 The MeCP1 complex represses transcription through preferential binding, remodeling, and deacetylating methylated nucleosomes. Genes and Development 15 827–832.
Fischle, W., Wang, Y., Allis, C., 2003. Binary switches and modification cassettes in histone biology and beyond. Nature 425, 475–479.
Forsberg, E.J., Strelchenko, N.S., Augenstein, M.L., et al. (2002). Production of cloned cattle from in vitro systems. Biol. Reprod. 67, 327–333.
Fujita N, Takebayashi S, Okumura K, Kudo S, Chiba T, Saya H & Nakao M 1999 Methylation-mediated transcriptional silencing in euchromatin by methyl-CpG binding protein MBD1 isoforms. Molecular and Cellular Biology 19 6415–6426.
Fujita N, Watanabe S, Ichimura T, Tsuruzoe S, Shinkai Y, Tachibana M, Chiba T & Nakao M 2003 Methyl-CpG binding domain 1 (MBD1) interacts with the Suv39h1-HP1
heterochromatic complex for DNA methylation-based transcriptional repression. Journal of Biological Chemistry 278 24132–24138.
Fulka, J., Jr., and Moor, R.M. (1993). Noninvasive chemical enucleation of mouse oocytes. Mol. Reprod. Dev. 34, 427–430.
Funahashi H, Cantley TC, Stumpf TT, Terlouw SL & Day B 1994 Use of lowsalt culture medium with elevated oocyte glutathione levels and enhanced male pronuclear formation after in vitro fertilization. Biology of Reproduction 51 633–639.
Furnus CC, de Matos DG & Moses DF 1998 Cumulus expansions during in vitro maturation of bovine oocytes: relationship level and its role on subsequent embryo development. Molecular Reproduction and Development 51 76–83.
Gal FL, Gall L and Smedt VD. 1992. Changes in protein synthesis pattern during in vitromaturation of goat oocytes. Mol Reprod Dev 32:1-8.
Galli C., Lagutina I., Crotti G., Colleoni S., Turini P., Ponderato N., Duchi R., Lazzari G. Pregnancy: a cloned horse born to its dam twin. Nature 2003;424:635.
Gasparrini B, Boccia L, Marchandise J, Palo RD, George F, Donnay I & Zicarelli L 2006 Enrichment of in vitro maturation medium for buffalo (Bubalus bubalis) oocytes with thiol compounds: effects of cystine on glutathione synthesis and embryo development. Theriogenology 65 275–287.
Gasparrini B, Neglia G, Palo RD, Campanile G & Zicarelli L 2000 Effect of cysteamine during in vitro maturation on buffalo embryo development. Theriogenology 54 1537–1542.
Giet R, Glover DM. Drosophila aurora B kinase is required for histone H3 phosphorylation and condensin recruitment during chromosome condensation and to organize the central spindle during cytokinesis. J Cell Biol 2001; 152:669–682.
Glotzer, M., Murray, A.W., Kirschner, M.W. (1991). Cyclin is degraded by the ubiquitin pathway. Nature 349(6305), 132-138.
Golding MC & Westhusin ME 2003 Analysis of DNA (cytosine 5) methyltransferase mRNA sequence and expression in bovine preimplantation embryos, fetal and adult tissues. Gene Expression Patterns 3 551–558.
Gomez E, Rodriguez A, Goyache F, Diez C, Royo LJ, Moreira PN, Caamano JN, Moran E, Gutierrez-Adan A. 2004. Retinoid-Dependent mRNA expression and Poly-(A) contents in bovine oocytes meiotically arrested and/or matured in vitro. Mol Reprod Dev 69:101-108.
Goto H, Yasui Y, Nigg EA, Inagaki M. Aurora-B phosphorylates histone H3 at serine 28 with regard to the mitotic chromosome condensation. Genes Cells 2002; 7:11–17.
Gray SG, Ekstrom TJ. The human histone deacetylase family. Exp Cell Res 2001; 262:75–83.
Greising, T., and Jonas, L. (1999). The influence of enucleation on the ultrastructure of in vitro matured and enucleated cattle oocytes. Theriogenology 52, 303–312.
Grunstein, M., 1997. Histone acetylation in chromatin structure and transcription. Nature 389, 349–352.
Gu L, Wang Q, Wang CM, Hong Y, Sun SG, Yang SY, Wang JG, Hou Y, Sun QY, Liu WQ. Distribution and expression of phosphorylated histone H3 during porcine oocyte maturation. Mol Reprod Dev. 2008; 75:143-149.
Guthrie HD, Garrett WM. Changes in porcine oocyte germinal vesicle development as follicles approach preovulatory maturity. Theriogenology 2000;54:389–399.
Guy J, Hendrich B, Holmes M, Martin JE & Bird A 2001 A mouse Mecp2-null mutation causes neurological symptoms that mimic Rett syndrome. Nature Genetics 27 322–326.
Hagemann LJ,Beaumont SE,Berg M,Donnison MJ,Ledgard A,Peterson AJ, Schurmann A,Tervit HR.1999.Development Curing Single IVP of Bovine Oocytes From Dissected Follicles:Interactive Effects of Estrous Cycle Stage,Follicle Size and Atresia.Mol Reprod Dev 53:451-458
Han D, Lan GC, Wu YG, Han ZB, Wang HL, Tan JH. Factors affecting the etticiency and reversibility of roscovitine(ROS) block on the meiotic resumption of goat oocytes. Mol Reprod Dev. 2006,73:238-246.
Hanks SK, Quinn AM, Hunter T. The protein kinase family: Conserved features and deducedphylogeny of the catalytic domains. Science 1988, 241:42-52
Hardarson Th, Lundin K and Hamberger L. The position of the metaphase II cannot be predicted by the location of the first polar body in the human oocyte. Human Reprod 2000; 15: 1372-1376.
Hashimoto S, Minami N, Takakura R, Imai H. 2002. Bovine immature oocytes acquire development competence during meiotic arrest in vitro. Biol Reprod 66:1696-1701.
Hata K, Okano M, Lei H & Li E 2002 Dnmt3L cooperates with the Dnmt3 family of de novo DNA methyltransferases to establish maternal imprints in mice. Development 129 1983–1993.
Hayes, E., Galea, S., Verkuylen, A., et al. (2001). Nuclear transfer of adult and genetically modified fetal cells of the rat. Physiol. Genomics 27, 193–204.
Hendrich B & Bird A 1998 Identification and characterization of a family of mammalian methyl-CpG binding proteins. Molecular and Cellular Biology 18 6538–6547.
Hendrich B, Guy J, Ramsahoye B, Wilson VA & Bird A 2001 Closely related proteins MBD2 and MBD3 play distinctive but interacting roles in mouse development. Genes and Development 15 710–723.
Hendrich B, Hardeland U, Ng HH, Jiricny J & Bird A 1999 The thymine glycosylase MBD4 can bind to the product of deamination at methylated CpG sites. Nature 401 301–304.
Henery, C.C., Chesne, P., Lebourhis, D., et al. (1994). Developmental ability of bovine embryos after nuclear transfer based on the nuclear source: in vivo versus in vivo. Theriogenology 42, 695–702.
Hermann A, Schmitt S & Jeltsch A 2003 The human Dnmt2 has residual DNA-(cytosine-C5) methyltransferase activity. Journal of Biological Chemistry 278 31717–31721.
Hinrichs K, Choi YH, Love LB, Varner DD, Love CC, Walckenaer BE. Chromatin configuration within the germinal vesicle of horse oocytes: changes post mortem and relationship to meiotic and developmental competence. Biol Reprod 2005;72:1142–1150.
Hinrichs K, Love CC, Choi YH, Varner DD, Wiggins CN, Reinoehl C. 2002. Suppression of meiosis by inhibitors of m-phase proteins in horse oocytes with low meiotic competence. Zygote 10(1):37-45.
Hinrichs K, Schmidt AL. Meiotic competence in horse oocytes: interactions among chromatin configuration, follicle size, cumulus morphology, and season. Biol Reprod 2000;62:1402–1408.
Hinrichs K,Schmidt AL,Friedman PP,Selgrath JP,Martin MG.1993.In vitro maturation of horse oocytes:Characterization of chromatin configuration using fluorescence microscopy.Biol Reprod 48:363–370.
Homa ST 1988. Effects of cyclic AMP on the spontaneous meiotic maturation of cumulus-free bovine oocytes cultured in chemical defined medium. J Exp Zool, 248: 222-231
Howell CY, Bestor TH, Ding F, Latham KE, Mertineit C, Trasler JM & Chaillet JR 2001 Genomic imprinting disrupted by a maternal effect mutation in the Dnmt1 gene. Cell 104 829–838.
Hsu J, Sun Z, Li X, Reuben M, Tatchell K, Bishop D, Grushcow J, Brame C, Caldwell J, Hunt D et al. Mitotic phosporylation of histone H3 Is governed by Ipl1/aurora kinase andGLC7/PP1 phosphatase in budding yeast and nematodes. Cell 2000;102:279–291.
Hsu JY, Sun ZW, Li X, Reuben M, Tatchell K, Bishop DK, Grushcow JM, Brame CJ, Caldwell JA, Hunt DF, Lin R, Smith MM, Allis CD. Mitotic phosphorylation of histone H3 is governed by Ipl1/aurora kinase and Glc7/PP1 phosphatase in budding yeast and nematodes. Cell 2000; 102:279–291.
Hu HM, Chuang CK, Lee MJ, Tseng TC, Tang TK. Genomic organization, expression, and chromosome localization of a third aurora-related kinase gene, Aie1. DNA Cell Biol 2000;19:679–688.
Humblot P,Holm P,Lonergan P,Wrenzycki C,Lequarre AS,Joly CG Herrmann D,Lopes A,Rizos D,Niemann H,Callesen H.2005.Effect of Stage of Follicular Growth During Superovulation On Developmental Competence of Bovine Oocytes.Theriogenology.63:1149-1166
Hyttel P,Callesen H,Greve T.1986a,Ultrastructural Features of Preovulatory Oocyte Maturation in Superovulated Cattle J.Reprod.Fert 76:645-656
Hyttel P,Xu KP,Smith S and Greve T 1986b.Ultrastructural of In-Vitro Oocyte Maturation in Cattle.J Reprod Fert.78:615-625
Iguchi-Ariga SM & Schaffner W 1989 CpG methylation of the cAMPresponsive enhancer/promoter sequence TGACGTCA abolishes specific factor binding as well as transcriptional activation. Genes and Development 3 612–619.
Inoue S, and Salmon, ED.(1995). Force generation by microtubule assembly/disassembly in mitosis and related movements. Mol. Biol. Cell 6, 1619–1640.
Jelinkova L and Kubelka M. Neither Aurora B Activity nor Histone H3 Phosphorylation Is Essential for Chromosome Condensation During Meiotic Maturation of Porcine Oocytes. Biol Reprod 2006; 74, 905–912.
Jelinkova L, Kubelka M, Motlik J, Guerrier P. Chromatin condensation and histone H1 kinase activity during growth and maturation of rabbit oocytes. Mol Reprod Dev 1994; 37: 210-215.
Jenuwein, T., Allis, C.D., 2001. Translating the histone code. Science 293, 1074–1080. Jessus C, Rime H, Haccard O, Van Lint J, Goris J, Merlevede W, et al. Tyrosine phosphorylation of p34cdc2 and p42 during meiotic maturation of Xenopus oocyte. Antagonistic action of okadaic acid and 6-DMAP. Development 1991;111:813–20.
Jones PL, Veenstra GJ, Wade PA, Vermaak D, Kass SU, Landsberger N, Strouboulis J & Wolffe AP 1998 Methylated DNA and MeCP2 recruit histone deacetylase to repress transcription. Nature Genetics 19 187–191.
Josefsberg LB, Galiani D, Dantes A, Amsterdam A, Dekel N. The proteasome is involved in the first metaphase-to-anaphase transition of meiosis in rat oocytes. Biol Reprod 2000; 62:1270-1277.
JU JYH-C, Tsay C, Chi-Wai Ruan. 2003. Alterations and reversibility in the chromatin, cytoskeleton and development of pig oocytes treated with roscovitine. Mol Reprod Dev 64:482-491.
Kageyama S, Liu H, Kaneko N, Ooga M, Nagata M, Aoki F. Alterations in epigenetic modifications during oocyte growth in mice. Reproduction 2007;133:85–94.
Kaitna S, Pasierbek P, Jantsch M, Loidl J, Glotzer M. The aurora B kinase AIR-2 regulates kinetochores during mitosis and is required for separation of homologous Chromosomesduring meiosis. Curr Biol 2002;12:798–812.
Kalous J, Kubelka M, Rimkevicova Z, Guerrier P, Motlik J. Okadaic acid accelerates germinal vesicle breakdown and overcomes cycloheximide- and 6-dimethylaminopurine block in cattle. Dev Biol 1993;157:448–54.
Kaneda M, Okano M, Hata K, Sado T, Tsujimoto N, Li E & Sasaki H 2004 Essential role for de novo DNA methyltransferase Dnmt3a in paternal and maternal imprinting. Nature 429 900–903.
Kato Y., Tani T., Sotomaru Y., Kurokawa K., Kato J., Doguchi H., Yasue H., Tsunoda Y. Eight calves cloned from somatic cells of a single adult. Science 1998;282:2095-2098.
Katoh, K., Hammar, K., Smith, P.J.S., et al. (1999). Arrangement of radical actin bundles in the growth cone of Aplysia bag cell neurons shows the immediate past history of filopodial behavior. PNAS 96, 7928–7931.
Kawakami, M., Tani, T., Yabuuchi, A., et al. (2003). Effect of demecolcine and nocodazole on the efficiency of chemically assisted removal of chromosomes and the developmental potential of nuclear transferred porcine oocytes. Cloning Stem Cells 5, 379–387.
Keefe, D., Tran, P., Pellegrini, C., et al. (1997). Polarized light microscopy and digital image processing identify a multilaminar structure of the hamster zona pellucida. Hum. Reprod. 12, 1250–1252.
Kikuchi K, Naito K, Noguchi J, Shimada A, Kaneko H, Yamashita M, Aoki F, Tojo H, Toyoda Y. Maturation/M-phase promoting factor: a regulator of aging in porcine oocytes. Biol Reprod. 2000; 63: 715-722.
Kim JM, Liu H, Tazaki M, Nagata M, Aoki F. Changes in histone acetylation during mouse oocyte meiosis. J Cell Biol 2003; 162:37–46.
Kimura M, Matsuda Y, Yoshioka T, Okano Y. Cell cycle-dependent expression and centrosome localization of a third human aurora/Ipl1-related protein kinase, AIK3. J Biol Chem 1999;274:7334–7340.
Kitagawa M, Okabe T, Ogino H, Matsumoto H, Suzuki-Takahashi I, Kokubo T, Higashi H, Saitoh S, Taya Y, Yasuda H, et al. 1993. Butyrolactone I, a selective inhibitor of cdk2 and cdc2 kinase. Oncogene 8 (9) :2425-2432.
Kovo M, Kandli-Cohen M, Ben-Haim M, Galiani D, Carr DW and Dekel N.An active protein kinase A (PKA) is involved in meiotic arrest of rat growing oocytes. Reproduction (2006) 132 33–43.
Krischek C, Meinecke B. 2001. Roscovitine, a specific inhibitor of cyclin-dependent protein kinases, reversibly inhibits chromatin condensation during in vitro maturation of porcine oocytes. Zygote 9 (November) :309-316.
Kruhlak, M.J., M.J. Hendzel, W. Fischle, N.R. Bertos, S. Hameed, X.-J. Yang, E. Verdin, and D.P. Bazett-Jones. 2001. Regulation of global acetylation in mitosis through loss of histone acetyltransferases and deacetylases from chromatin. J. Biol. Chem. 276:38307–38319.
Kubelka M, Anger M, Kalous J, Schultz R, Motlik J. 2002. Chromosome condensation in pig oocytes:lack of a requirment for either cdc2 kinase or MAP kinase activity. Mol Reprod Dev 63:110-118.
Kubelka M, Motlik J, Schultz RM, Pavlok A. Butyrolactone I reversibly inhibits meiotic maturation of bovine oocytes, without influencing chromosome condensation activity.Biol Reprod 2000; 62:292–302.
Kubelka M, Rimkevicova Z, Guerrier P, Motlik J. Inhibition of protein synthesis affects histone H1 kinase, but not chromosome condensation activity, during the first meiotic division of pig oocytes. Mol Reprod Dev 1995; 41:63–69.
Kubota, C., Yamakuchi, H., Todoroki, J., et al. (2000). Six cloned calves produced from adult fibroblast cells after long-term culture. PNAS 97, 990–995.
Kubota, C., Yang, X., Dinnyés, A., et al. (1998). In vitro and in vivo survival of frozen thawed bovine oocytes after IVF, nuclear transfer and parthenogenetic activation. Mol. Reprod. Dev. 51, 281–286.
Kuo MH and David Allis C..Roles of histone acetyltransferases and deacetylases in gene regulation BioEssays 1998;20:615–626,
Kurosaka S, Nagao Y, Minami N, Yamada M, Imai H. Dependence of DNA synthesis and in vitro development of bovine NT embryos on the stage of the cell cycle of donor cells and recipient cytoplasts. Biol Reprod 2002; 67: 643-647.
Lan, G.C., Wu, Y.G., Han, D., et al. (2008). Demecolcine-assisted enucleation of goat oocytes: protocol optimization, mechanism investigation, and application to improve the developmental potential of cloned embryos. Cloning Stem Cells 10, 189–202.
Lee BC., Kim MK., Jang G., Oh HJ., Yuda F., Kim H.J., Hossein M.S., Kim J.J., Kang S.K., Schatten G., Hwang W.S. Dogs cloned from adult somatic cells. Nature 2005;436:641.
Lee HS, Yin XJ, Jin YX, Kim NH, Cho SG, Bae IH, Kong IK. Germinal vesicle chromatin configuration and meiotic competence is related to the oocyte source in canine. Anim Reprod Sci 2008;103:336–347.
Lefevre B,Gougeon A,Nome F,Testart J.1989.In vivo changes in oocyte germinal vesicle related to follicular quality and size at mid-follicular phase during stimulated cycles in the Cynomolgus monkey.Reprod Nutr Dev 29:523–532.
Leis JK, Kaplan NO. An acid phosphatase in the plasma membranes of human astrocytoma showing marked specificity towards phosphotyrosine protein. Proc Natl Acad Sci USA 1982; 79:6507–6511.
Li E., Beard, C., and Jaenisch, R. (1993). Role for DNA methylation in genomic imprinting. Nature 366: 362–365.
Li E., Bestor, T. H., and Jaenisch, R. (1992). Targeted mutation of the DNA methyltransferase gene results in embryonic lethality. Cell 69: 915–926.
Li GP, Bunch TD, White KL, Aston KI, Meerdo LN, Pate BJ, Sessions BR. Development, chromosomal composition, and cell allocation of bovine cloned blastocyst derived from chemically assisted enucleation and cultured in conditioned media. Mol Reprod Dev 2004a; 68: 189-197.
Li GP, Chen DY, Lian L, et al. (2001). Viable rabbits derived from reconstructed oocytes by germinal vesicle transfer after intracytoplasmic sperm injection (ICSI). Mol. Reprod. Dev. 58, 180–185.
Li GP, Meng QG, Wei P, Yu YS, Chang ZL and Tan JH. 2001. Effect of cycloheximide (CHX) on in vitro maturation of porcine oocytes. Acta Zoologial Sinica 47 (6) :684-690.
Li GP, White KL, Bunch TD. Review of enucleation methods and procedures used in animal cloning: state of the art. Cloning Stem Cells 2004; 6(1): 5-13.
Li X, Sakashita G, Matsuzaki H, Sugimoto K, Kimura K, Hanaoka F, Taniguchi H, FurukawaK, Urano T. Direct association with inner centromere protein (INCENP) activates the novel chromosomal passenger protein, Aurora-C. J Biol Chem 2004;279:47201–47211.
Li Z., Sun X., Chen J., Liu X., Wisely S.M., Zhou Q., Renard J.P., Leno G.H., Engelhardt J.F. Cloned ferrets produced by somatic cell nuclear transfer. Dev Biol 2006b;293:439-448.
Lincoln AJ, Wickramasinghe D, Stein P , Schultz RM, Palko ME, De Miguel MP, Tessarollo L & Donovan PJ. Cdc25b phosphatase is required for resumption of meiosis during oocyte maturation. nature genetics 2002;30:446-449.
Liu K, Wang YF, Cantemir C & Muller MT 2003 Endogenous assays of DNA methyltransferases: evidence for differential activities of DNMT1, DNMT2, and DNMT3 in mammalian cells in vivo. Molecular and Cellular Biology 23 2709–2719.
Liu L, Oldenbourg R, Trimarchi JR, Keefe DL. A reliable, noninvasive technique for spindle imaging and enucleation of mammalian oocytes Nat Biotechnol 2000a; 18: 223-225.
Liu L, Trimarchi JR, Oldenbourg R, Keefe DL. Increased birefringence in the meiotic spindle provides a new marker for the onset of activation in living oocytes. Biol Reprod 2000b; 63:251-258.
Liu L.2001.Cloning efficiency and differentiation.Nat Biotech,19:406
Liu Y,Sui HS,Wang HL,Yuan JH,Luo MJ,Xia P,Tan JH.2006.Germinal Vesicle Chromatin Configurations of Bovine Oocytes.Microsc Res Tech 69:799-807
Liu, J.L., Sung, L.Y., Barber, M., et al. (2002). Hypertonic medium treatment for localization of nuclear material in bovine metaphase II oocytes. Biol. Reprod. 66, 1342–1349.
Liu, J.L., Wang, M.K., Sun, Q.Y., et al. (2000). Effect of telophase enucleation on bovine somatic nuclear transfer. Theriogenology 54, 989–998.
Loi P, Ptak G, Barboni B, Fulka J, Cappai P, Clinton M. Genetic rescue of an endangered mammal by cross-species nuclear transfer using post-mortem somatic cells, Nat Biol 2001; 19:962-964.
Lonergan P, Dinnyes A, Fair T, Yang X, Boland M. 2000. Bovine oocyte and embryo development following meiotic inhibition with butyrolactone I. Mol Reprod Dev 57 (2) :204-209.
Lonergan P, Fair T, Khatir H, Cesaroni G, Mermillod. 1998. Effect of protein-synthesis inhibition before or during in vitro maturation on subsequent development of bovine oocytes. Theriogenology 50:417-431.
Lonergan P, Khatir H, Carolan C, Mermillod P. 1997. Bovine blastocysts production in vitro after inhibition of oocyte meiotic resumption for 24 hr. J Reprod Fertil 109:355-365.
Lonergan P,Monaghan P,Rizos D,Boland MP,Gordon I.1994.Effect of Follicle Size On Bovine Oocyte Quality and Developmental Competence Following Maturation,Fertilization,and Culture In Vitro.Mol Reprod Dev 37:48-53
Lonergan P,Rizos D,2003,7(6).Gutierrez-Adan A,Fair T,Boland MP.Effect of Culture Environment on Embryo Quality and Gene Expression-Experience From Animal Studies.Reprod Biomed Online.657-663
Marchal R, Tomanek M, Terqui M, Mermillod P. 2001. Effect of cell cycle dependent kinases inhibitor on nuclear and cytoplasmic maturation of porcine oocytes. Mol Reprod Dev 60:65-73.
Masui Y and markert CL. Cytoplasmic centrol of nuclear behavior during meiotic mutition of frog oocytes. J Exp Zool, 1971, 177:129-146.
Mattson BA,Albertini DF.1990.Oogenesis:Chromatin and microtubule dynamics during meiotic prophase.Mol Reprod Dev 25:374-383
McGrath, J., and Solter, D. (1983). Nuclear transplantation in the mouse embryo by microsurgery and cell fusion. Science 220, 1300–1302.
Meijer L, Borgne A, Mulner O, Chong JPJ, Blow JJ, Inagaki N, Inagaki M, Delcros JG, Moulinoux JP. 1997. Biochemical and cellular effects of roscovitine, a potent and selective inhibitor of the cyclin-dependent kinases cdc2 and cdk5. Eur J biochim 243:527-536.
Meijer L, Kim SH. 1997. Chemical inhibitors of cycline-dependent kinases. Meth Enzymol 283:113-128.
Meister A 1983 Selective modification of glutathione metabolism. Science 220 472–477. Meng, L., Rutledge, J., Zhu, Y., et al. (1996). Role of germinal vesicle on protein synthesis in rat oocyte during in vitro maturation. Mol. Reprod. Dev. 43, 228–235.
Mermillod P, Tomanek M, Marchal R, Meijer L. 2000. High developmental competence of cattle oocytes maintained at the germinal vesicle stage for 24 hours in culture by specific inhibition of MPF kinase activity. Mol Reprod Dev 55:89-95.
Mertineit C, Yoder JA, Taketo T, Laird DW, Trasler JM, Bestor TH. 1998. Sex-specific exons control DNA methyltransferase in mammalian germ cells. Development 125:889–897.
Miao Y, Ma S, Liu X, Miao D, Chang Z, Luo M, Tan J. Fate of the first polar bodies in mouse oocytes. Mol Reprod Dev 2004; 69: 66-76.
Mitalipov, S.M., White, K.L., Farrar, V.R., et al. (1999). Development of nuclear transfer and parthenogenetic rabbit embryos activated with inositol 1,4,5-trisphosphate. Biol. Reprod. 60, 821–827.
Miyara F,Migne C,Dumont-Hassan M,Meur AL,Cohen-Bacrie P,Aubriot FX,Glissant A,Nathan C,Douard S,Stanovici A,Debey P.2003. Chromatin configuration and transcriptional control in human and mouse oocytes.Mol Reprod Dev 64:458–470.
Miyara, F., Han, Z., Gao, S., et al. (2006). Non-equivalence of embryonic and somatic cell nuclei affecting spindle composition in clones. Dev. Biol. 289, 206–217.
Morgan DO, Fisher RP, Espinoza FH, Farrell A, Nourse J, Chamberlin H, Jin P. Control of eukaryotic cell cycle progression by phosphorylation of cyclin-dependent kinases. Cancer J Sci Am 1998, 4(Suppl 1): S77-S83.
Morgan DO. Cyclin-dependent kinases: engines, clocks, and microprocessors. Annu Rev Cell Dev Biol 1997, 13: 261-291.
Morgan DO. Principles of CDK regulation. Nature 1995; 374: 131-134. Morgan HD, Santos F, Green K, Dean W, Reik W. Epigenetic reprogramming in mammals. Hum Mol Genet 2005; 14:R47–R58.
Motlik J, Pavlok A, Kubelka M, Kalous J, Kalab P. 1998. Interplay between CDC2 kinase MAP kinase pathway during maturation of mammalian oocytes. Theriogenology 49:461-469.
Motlik J,Fulka J.1976.Breakdown of the germinal vesicle in pig oocytes in vivo and in vitro.J Exp Zool 198:155–162.
Motlik J,Koefoed-Johnsen HH,Fulka J.1978.Breakdown of the germinal vesicle in bovine oocytes cultivated in vitro.J Exp Zool 205:377–383.
Murnion ME, Adams RR, Callister DM, Allis CD, Earnshaw WC, Swedlow JR.Chromatin-associated protein phosphatase 1 regulates aurora-B and histone H3 phosphorylation. J Biol Chem 2001; 276: 26656–26665.
Nagai T, Ebihara M, Onishi A, Kubo M. Germinal vesicle stages in pig follicular oocytes collected by different methods. J Reprod Dev 1997; 43:339–343.
Nan X, Campoy FJ & Bird A 1997 MeCP2 is a transcriptional repressor with abundant binding sites in genomic chromatin. Cell 88 471–481.
Nan X, Ng HH, Johnson CA, Laherty CD, Turner BM, Eisenman RN & Bird A 1998 Transcriptional repression by the methyl-CpG-binding protein MeCP2 involves a histone deacetylase complex. Nature 393 386–389.
Ne′ant I, Guerrier P. 6-Dimethylaminopurine blocks starfish oocyte maturation by inhibiting a relevant protein kinase activity. Exp Cell Res 1988;176:68–79.
Ng HH, Jeppesen P & Bird A 2000 Active repression of methylated genes by the chromosomal protein MBD1. Molecular and Cellular Biology 20 1394–1406.
Ng HH, Zhang Y, Hendrich B, Johnson CA, Turner BM, Erdjument-Bromage H, Tempst P, Reinberg D & Bird A 1999 MBD2 is a transcriptional repressor belonging to the MeCP1 histone deacetylase complex. Nature Genetics 23 58–61.
Nigg E. Mitotic kinases as regulators of cell division and it’s checkpoints. Nat Rev Mol Cell Biol 2001;2:21–32.
Nogueira D,Cortvrindt R,De Matos DG,Vanhoutte L,Smitz J.2003.Effect of phosphodiesterase type 3 inhibitor on developmental competence of immature mouse oocytes in vitro.Biol Reprod 69:2045–2052.
Nour MS, Takahashi Y. Preparation of young preactivated oocytes with high enucleation efficiency for bovine nuclear transfer. Theriogenology 1999; 51: 661-666.
Ola SI,Wang Q,Ai JS,Yin S,Liang CG,Chen DY,Sun QY.2007.Meiotic competence and acetylation pattern of UV light classified mouse antral oocytes after meiotic arrest with isobutylmethylxanthine.Mol Reprod Dev 74:591–599
Onishi A., Iwamoto M., Akita T., Mikawa S., Takeda K., Awata T., Hanada H., Perry A.C. Pig cloning by microinjection of fetal fibroblast nuclei. Science 2000;289:1188-1190.
Parfenov V,Potchukalina G,Dudina L,Kostyuchek D,Gruzova M.1989. Human antral follicles:Oocyte nucleus and the karyosphere formation (electron microscopic and autoradiographic data).Gamete Res 22:219–231.
Pavlok A,Lucas-Hahn A,Niemann H.1992.Fertilization and DevelopmentalCompetence of Bovine Oocytes Derived From Different Categories of Antral Follicles.Mol Reprod Dev 31:63-67
Perreault SD, Barbee RR & Slott VL 1988 Importance of glutathione in the acquisition and maintenance of sperm nuclear decondensing activity in maturing hamster oocytes. Developmental Biology 125 181–186.
Peterson, C.L., Laniel, M.A., 2004. Histones and histone modifications. Curr. Biol. 14, R546–R551.
Peura TT and Trounson AO. Recycling bovine embryosfor nuclear transfer. Reprod Fertil Dev 1998; 10: 627-632.
Peura, T.T., Lewis, I.M., and Trounson, A.O. (1998a). The effect of recipient oocyte volume on nuclear transfer in cattle. Mol. Reprod. Dev. 50, 185–191.
Poleiaeva IA,Chen SH,Vaught TD,Page RL,Ball S.Cloned pigs produced by nuclear transferfrom adult somatic cell.Nature,2000,407:86-90
Ponderato N, Lagutina I, Crotti G, Turini P, Galli C, Lazzari G. 2001. Bovine oocytes treated prior to in vitro maturation with a combination of Butyrolactone I and Roscovitine at low doses maintain a normal developmental capacity. Mol Reprod Dev 60:579-585.
Prather, R.S., Sims, M.M., and First, N.L. (1989). Nuclear transplantation in early pig embryos. Biol. Reprod. 41, 414–418.
Prokhortchouk A, Hendrich B, Jorgensen H, Ruzov A, Wilm M, Georgiev G, Bird A & Prokhortchouk E 2001 The p120 catenin partner Kaiso is a DNA methylation dependent transcriptional repressor. Genes and Development 15 1613–1618.
Ray LB, SturgillTW. Insulin-stimulatedmicrotuble-associated protein kinase is phosphorylated on tyrosine and threonine in vivo[J]. Proc Natl Acad Sci USA,1988,85: 3 753–3 757.
Reik W, Dean W & Walter J 2001 Epigenetic reprogramming in mammalian development. Science 293 1089–1093.
Rime H, Neant I, Guerrier P, Ozon R. 6-Dimethylaminopurine (6-DMAP), a reversible inhibitor of the transition to metaphase during the firstmeiotic cell division of the mouse oocyte. Dev Biol 1989;133:169–79.
Rizos D,Fair T, Papadopoulos S, Boland MP, Lonergan P.2002.Developmental, Qualitative, and Ultrastructural Differences Between Ovine and Bovine Embryos Produced In Vivo or In Vitro. Mol Reprod Dev 62:320-327
Robertson KD & Wolffe AP 2000 DNA methylation in health and disease. Nature Reviews. Genetics 1 11–19.
Rodriguez-Gonzalez E, Lopez-Bejar M, Mertens MJ & Paramio MT 2003 Effects on in vitro embryo development and intracellular glutathione content of the presence of thiol compounds during maturation of prepubertal goat oocytes. Molecular Reproduction and Development 65 446–453.
Rogers E, Bishop JD, Waddle JA, Schumacher JM, Lin R. The aurora kinase AIR-2 functions in the release of chromosome cohesion in Caenorhabditis elegans meiosis. J Cell Biol 2002;157:219–229.
Roth SY, Allis CD. 1992. Chromatin condensation: does histone H1 dephosphorylation play a role? Trends Biochem Sci 17:93–98.
Rouleau, M., Aubin, R.A., Poirier, G.G., 2004. Poly(ADP-ribosyl)ated chromatin domains: access granted. J. Cell Sci. 117, 815–825.
Russo V, Martelli A, Berardinelli P, Di Giacinto O, Bernabo` N, Fantasia D, Mattioli M, Barboni B. Modifications in chromatin morphology and organization during sheep oogenesis. Microsc Res Tech 2007;70: 733–744.
Ruzov A, Dunican DS, Prokhortchouk A, Pennings S, Stancheva I, Prokhortchouk E & Meehan RR 2004 Kaiso is a genome-wide repressor of transcription that is essential for amphibian development. Development 131 6185–6194.
Ryan J, Llinas AJ, White DA, Turner BM, Sommerville J. Maternal histone deacetylase is accumulated in the nuclei of Xenopus oocytes as protein complexes with potential enzyme activity. J Cell Sci 1999; 112(pt 14): 2441–2452.
Saeki K, Nagao Y, Kishi M, Nagai M, Iritani A. 1998. Timing of completion of the first meiotic division in bovine oocytes after maintenance of meiotic arrest withcycloheximide and their subsequent development. J Vet Med Sci 60 (4) :523-526. Sarma, K., Reinberg, D., 2005. Histone variants meet their match. Nat. Rev., Mol. Cell Biol. 6, 139–149.
Sasai K, Katayama H, Stenoien DL, Fujii S, Honda R, Kimura M, Okano Y, Tatsuka M, Suzuki F, Nigg EA et al. Aurora-C kinase is a novel chromosomal passenger protein that can complement Aurora-B kinase function in mitotic cells. Cell Motil Cytoskeleton 2004;59:249–263.
Schoevers EJ, Bevers1 MM, Roelen BAJ and Colenbrander B. 2005. Nuclear and cytoplasmic maturation of sow oocytes are not synchronized by specific meiotic inhibition with roscocitine during in vitro maturation. Theriogenology 63:1111-1130.
Schramm RD, Tennier MT, Boatman DE, Bavister BD. Chromatin configurations and meiotic competence of oocytes are related to follicular diameter in nonstimulated rhesus monkeys. Biol Reprod 1993; 48:349–356.
Shi W, Dirim F, Wolf, E., Zakhartchenko, V. and Haaf, T. (2004) Methylation reprogramming and chromosomal aneuploidy in in vivo fertilized and cloned rabbit preimplantation embryos. Biol. Reprod. 71, 340–347.
Shin T,Kraemer D,Pryor J,Liu L,Rugila J,Howe L,Buck S,Murphy K,Lyons L,Westhusin M. 2002.A cat cloned by nuclear transplantation.Nature;415(6874):859
Silva CP, Kommineni K, Oldenbourg R, Keefe DL. The first polar body does not predict accurately the location of the metaphase II meiotic spindle in mammalian oocytes. Fertil Steril 1999;71: 719-721.
Simerly, C. Dominko, T., Navara, C., et al. (2003). Molecular correlates of primate nuclear transfer failures. Science 300, 297.
Sirard MA, Blondin P.1996.Oocyte Maturation and IVF in Cattle.Anim Reprod Sci 42:417-426
Smith, L.C. (1993). Membrane and intracellular effects of ultraviolet irradiation with Hoechst 33342 on bovine secondary oocytes matured in vitro. J. Reprod. Fertil. 99, 39–44. Smith, L.C., and Wilmut, I. (1990). Factors affecting the viability of nuclear transplantation embryos. Theriogenology 33, 83–98.
Spinaci M, Seren E & Mattioli M 2004 Maternal chromatin remodeling during maturation and after fertilization in mouse oocytes. Molecular Reproduction and Development 69 215–221.
Stice, S.L., Keefer, C.L., and Matthews, L. (1994). Bovine nuclear transfer embryos: oocytes activation prior to blastomere fusion. Mol. Reprod. Dev. 38, 61–68.
Sui HS,Liu Y,Miao DQ,Yuan JH,Qiao TW,Luo MJ,Tan JH.2005. Configurations of germinal vesicle(GV)chromatin in the goat differ from those of other species.Mol Reprod Dev 71:227–236.
Sun XS,Liu Y,Yue KZ,Ma SF,Tan JH.2004.Changes in germinal vesicle (GV)chromatin configurations during growth and maturation of porcine oocytes.Mol Reprod Dev 69:228–234.
Sutovsky P & Schatten G 1997 Depletion of glutathione during bovine oocyte maturation reversibly blocks the decondensation of the male pronucleus and pronuclear apposition during fertilization. Biology of Reproduction 56 1503–1512.
Swain JE, Ding J, Brautigan DL, Villa-Moruzzi E, Smith GD. Proper chromatin condensationand maintenance of histone H3 phosphorylation during mouse oocyte meiosis requires protein phosphatase activity. Biol Reprod 2007;76:628–638.
Swain JE, Ding J, Wu J, Smith GD.Regulation of spindle and chromatin dynamics during early and late stages of oocyte maturation by aurora kinases. Mol Hum Reprod 2008; 14: 291 - 299.
Swales AKE and Spears N.Genomic imprinting and reproduction. Reproduction (2005) 130 389–399
Swarup G, Vincer Speeg KJ, Cohen G, Garbers DL. Phosphotyrosylprotein phosphatase of TCRC-2 cells. J Biol Chem 1982; 257:7298– 7301.
Tan JH, Wang HL, Sun XS, Liu Y, Sui HS, Zhang J Chromatin configurations in the germinal vesicle of mammalian oocytes.Mol Hum Reprod. 2009, 15:1-9.
Tang LY, Reddy MN, Rasheva V, Lee TL, Lin MJ, Hung MS & Shen CK 2003 The eukaryotic DNMT2 genes encode a new class of cytosine- 5 DNA methyltransferases. Journal of Biological Chemistry 278 33613–33616.
Tani T, Kato Y, Tsunoda Y. Aberrant spindle assembly checkpoint in bovine somatic cell nuclear transfer oocytes. Front Biosci. 2007 Jan 1;12:2693-705.
Tani T, Kato Y, Tsunoda Y. Direct exposure of chromosomes to nonactivated ovum cytoplasm is effective for bovine somatic cell nucleus reprogramming. Biol Reprod 2001; 64:324-330.
Tani, T., Shimada, H., Kato, Y., et al. (2006). Demecolcine-assisted enucleation for bovine cloning. Cloning Stem Cells 8, 61–66.
Tatemoto H and Terada T.Involvement of cumulus cells stimulated by FSH in chromatin condensation and the activation of maturation-promoting factor in bovine oocytes. Theriogenology, 1998; 49: 1007-20.
Tatemoto H, Terada T. Involvement of cyclic AMP-dependent protein kinase in chromatin condensation before germinal vesicle breakdown in bovine oocytes. Anim Reprod Sci 1996; 44: 99-109.
Tatham, B.G., Dowsing, A.T., and Trounson, A.O. (1995). Enucleation by centrifugation of in vitro–matured bovine oocytes for use in nuclear transfer. Biol. Reprod. 53, 1088–1094.
Tatham, B.G., Sathananthan, A.H., Dharmawardena, V., et al. (1996). Centrifugation of bovine oocytes for nuclear micromanipulation and sperm microinjection. Hum. Reprod. 11, 1499–1503.
Taunton J, Hassig CA & Schreiber SL 1996 A mammalian histone deacetylase related to the yeast transcriptional regulator Rpd3p. Science 272 408–411. Tseng TC, Chen SH, Hsu YP, Tang TK. Protein kinase profile of sperm and eggs: cloning and characterization of two novel testis-specific protein kinases (AIE1, AIE2) related to yeast and fly chromosome segregation regulators. DNA Cell Biol 1998;17:823–833.
Tsuji H, Matsudo Y, Tsuji S, Hanaoka F, Hyodo M & Hori T. (1990) Isolation of temperature-sensitive CHO-K1 cell mutants exhibiting chromosomal instability and reduced DNA synthesis at nonpermissive temperature. Somat Cell Mol Genet. 16, 461 - 476.
Tsuji H, Matsudo YK, Ajiro K, Yasuda H, Hanaoka F, Hayashi A, Utsumi S, Ohba Y & Hori T. (1992) A temperature sensitive CHO-Kl cell mutant (tsTM13) defective in chromosome decondensation and spindle deconstruction in M phase.Exp Cell Res. 198, 201-213.
Tsunoda Y,Kato Y.The recent progress on nuclear transfer in mammalian.Zool Sci,2000,17:1177-1184.
Turner, B.M., 2000. Histone acetylation and an epigenetic code. BioEssays 22, 836–845.
Vajta G, Bartels P, Joubert J, de la Rey M, Treadwell R, Callesen H. Production of a healthy calf by somatic cell nuclear transfer without micromanipulators and carbon dioxide incubators using the Handmade Cloning (HMC) and the Submarine Incubation System (SIS). Theriogenology 2004; 62(8): 1465-72.
Vajta, G., Lewis, I.M., Trounson, A.O., et al. (2003). Handmade somatic cell cloning in cattle: analysis of factors contributing to high efficiency in vitro. Biol. Reprod. 68, 571–578.
Vajta, G., Maddox-Hyttel, P., Skou, C.T., et al. (2005). Highly efficient and reliable chemically assisted enucleation method for handmade cloning in cattle. Reprod. Fertil. Dev. 17, 791–797.
Van Thuan, N., Wakayama, S., Kishigami, S., et al. (2006). Donor centrosome regulation of initial spindle formation in mouse somatic cell nuclear transfer: roles of gamma-tubulin and nuclear mitotic apparatus protein 1. Biol. Reprod. 74, 777–787
Vanhoutte L De Sutter P, Nogueira D, Gerris J, Dhont M and Van der Elst J. Nuclear and cytoplasmic maturation of in vitro matured human oocytes after temporary nuclear arrest by phosphodiesterase 3-inhibitor.Hum Rerod 2007;22:1239-1246
Velilla, E., Lopez-Bejar, M., Rodriguez-Gonzalez, E., et al. (2002). Effect of Hoechst 33342 staining on developmental competence of prepubertal goat oocytes. Zygote 10, 201–208.
Verlhac MH, De Pennhart H, Maro B, Cobb MH, Clarke HJ. 1993. MAP kinase becomes stably activated at metaphase and is associated with microtubule-organizing centers during meiotic maturation of mouse oocytes. Dev Biol 158:330–340.
Verlhac MH, Kubiak JZ, Clarke HJ, Maro B. 1994. Microtubule and chromatin behavior follow MAP kinase activity but not MPF activity during meiosis in mouse oocytes. Development 120:1017–1025.
VerlhacMH,et al.Mos activates MAPkinase in mouse oocytes through two opposite pathways[J].The EMBO,2000,19:6 065.
Wakayama T,Perry AC,Zuccotti M,Johnson KR,Yanagimachi R.1998.Full-term development of mice from enucleated oocytes injected with cmulus cell nuclei.Nature 394:369-374
Wakayama, T., and Yanagimachi, R. (1998). Fertilisability and developmental ability of mouse oocytes with reduced amounts of cytoplasm. Zygote 6, 341–346.
Wang HL, Sui HS, Liu Y, Miao DQ, Lu JH, Liang B, Tan JH.Dynamic changes of germinal vesicle chromatin configuration and transcriptional activity during maturation of rabbit follicles. Fertility and Sterility.2009, 91: 1589-1594.
Wang Q, Wang CM, Ai JS, Xiong B, Yin S, Hou Y, Chen DY, Schatten H, Sun QY. Histone phosphorylation and pericentromeric histone modifications in oocyte meiosis. Cell Cycle 2006;5:1974–1982.
Wang Q, Yin S, Ai JS, Liang CG, Hou Y, Chen DY, Schatten H, Sun QY. Histone deacetylation is required for orderly meiosis. Cell Cycle 2006; 5: 766–774.
Wang, M.K., Liu, J.L., Li, G.P., et al. (2001). Sucrose pretreatment for enucleation: an efficient and non-damage method for removing the spindle of the mouse MII oocyte. Mol. Reprod. Dev. 58, 432–436.
Westhusin ME,Long CR,Shin T,Hill JR,Looney CR,Pryor JH,Piedrahita JA.2001 Cloning toreproduce desired genotypes.Theriogenology;55:35-49
Westhusin, M.E., Levanduski, M.J., Scarborough, R., et al. (1992). Viable embryos and normal calves after nuclear transfer into Hoechst stained enucleated demi-oocytes of cows. J. Reprod. Fertil. 95, 475–480.
Wickramasinghe D,Ebert KM,Albertini DF.1991.Meiotic Competence Acquisition is Associated With the Appearance of M-Phase Characteristics in Growing Mouse Oocytes.Dev Biol 143:162-172
Wilmut I,Schmieke AE,McWhir J,Kind AJ,Campbell KHS.1997.Viable offspring derived from fetal and adult mammalian cells.Nature,385:810-813
Woods GL, White KL, Vanderwall DK, Li GP, Aston KI, Bunch TD, Meerdo LN, Pate BJ. A mule cloned from fetal cells by nuclear transfer. Science 2003; 301: 1063.
Wu GM, Sun QY, Mao J, Lai LX, McCauley T, Park KW, Prather R, Didion B, Day B. 2002. High developmental competence of pig oocytes after meiotic inhibition with a specific M-phase promoting factor kinase inhibitor,butyrolactone I. Biol Reprod 67:170-177.
Yan X, Cao L, Li Q, Wu Y, Zhang H, Saiyin H, Liu X, Zhang X, Shi Q, Yu L. Aurora C is directly associated with Survivin and required for cytokinesis. Genes Cells 2005;10:617–626.
Yang, X., Zhang, L., Kovacs, A., et al. (1990). Potential of hypertonic medium treatment for embryo micromanipulation: II. Assessment of nuclear transplantation methodology, isolation, subzona insertion, and electrofusion of blastomeres to intact or functionally enucleated oocytes in rabbits. Mol. Reprod. Dev. 27, 118–129.
Yao LJ, Sun QY. Characterization of aurora-a in porcine oocytes and early embryos implies its functional roles in the regulation of meiotic maturation, fertilization and cleavage. Zygote 2005;13:23–30.
Yao LJ, Zhong ZS, Zhang LS, Chen DY, Schatten H, Sun QY. Aurora-A is a critical regulator of microtubule assembly and nuclear activity in mouse oocytes, fertilized eggs, and early embryos. Biol Reprod 2004;70: 1392–1399.
Yin, X.J., Tani, T., Yonemura, I., et al. (2002). Production of cloned pigs from adult somatic cells by chemically assisted removal of maternal chromosomes. Biol. Reprod. 67, 442–446.
Yoshida M, Ishigaki K, Nagai T, Chikyu M & Pursel VG 1993 Glutathione concentration during maturation and after fertilization in pig oocytes: relevance to the ability of oocytes to form male pronucleus. Biology of Reproduction 49 89–94.
Yoshida N, Brahmajosyula M, Shoji S, Amanai M, Perry AC. Epigenetic discrimination by mouse metaphase II oocytes mediates asymmetric chromatin remodeling independently of meiotic exit. Dev Biol 2007; 301:464–477.
Yoshida, M., M. Kijima, M. Akita, and T. Beppu. 1990. Potent and specific inhibition of mammalian histone deacetylase both in vivo and in vitro by trichostatin A. J. Biol. Chem. 265:17174–17179.
Yu F, Thiesen J & Stratling WH 2000 Histone deacetylase-independent transcriptional repression by methyl-CpG-binding protein 2. Nucleic Acids Research 28 2201–2206.
Zachariae W, Nasmyth K. Whose end is destruction: cell division and the anaphase-promoting complex. Genes Dev 1999, 13: 2039-2058.
Zhang, Y., 2003. Transcriptional regulation by histone ubiquitination and deubiquitination.Genes Dev. 17, 2733–2740.
Zhao X, Ueba T, Christie BR, Barkho B, McConnell MJ, Nakashima K, Lein ES, Eadie BD, Willhoite AR, Muotri AR, Summers RG, Chun J, Lee KF & Gage FH 2003 Mice lacking methyl-CpG binding protein 1 have deficits in adult neurogenesis and hippocampal function. PNAS 100 6777–6782.
Zhou P, Wu YG, Li Q, Lan GC,Wang G, Gao D, et al.The interactions between cysteamine, cystine and cumulus cells increase the intracellular glutathione level and developmental capacity of goat cumulus-denuded oocytes. Reproduction 2008;135:605–11.
Zhou Q, Renard JP,Friec GL,Brochard V,Beaujean N,Cherifi Y,Fraichard A,Cozzi J.2003 Generation of fertile cloned rats by regulating oocyte activation. Science, 302:1179
Zuccotti M, Piccinelli A, Giorgi Rossi P, Garagna S, Redi CA. Chromatin organization during mouse oocyte growth. Mol Reprod Dev 1995;41:479–485.