References

Albuz, F. K., Sasseville, M., Lane, M., Armstrong, D. T., Thompson, J. G., and Gilchrist, R. B. (2010). Simulated physiological oocyte maturation (SPOM): a novel in vitro-maturation system that substantially improves embryo yield and pregnancy outcomes. Hum. Reprod. 25, 2999–3011.
| CrossRef | CAS | PubMed |

Ali, A., and Sirard, M. A. (2002). Effect of the absence or presence of various protein supplements on further development of bovine oocytes during in vitro maturation. Biol. Reprod. 66, 901–905.
| CrossRef | CAS | PubMed |

Bagg, M. A., Nottle, M. B., Armstrong, D. T., and Grupen, C. G. (2009). Effect of follicle size and dibutyryl cAMP on the cAMP content and gap-junctional communication of porcine prepubertal cumulus–oocyte complexes during IVM. Reprod. Fertil. Dev. 21, 796–804.
| CrossRef | CAS | PubMed |

Banwell, K. M., Lane, M., Russell, D. L., Kind, K. L., and Thompson, J. G. (2007). Oxygen concentration during mouse oocyte in vitro maturation affects embryo and fetal development. Hum. Reprod. 22, 2768–2775.
| CrossRef | CAS | PubMed |

Bateman, H. L., Bond, J. B., Campbell, M., Barrie, M., Riggs, G., Snyder, B., and Swanson, W. F. (2009). Characterization of basal seminal traits and reproductive endocrine profiles in North American river otters and Asian small-clawed otters. Zoo Biol. 28, 107–126.
| CrossRef | CAS | PubMed |

Bilodeau, S., Fortier, M. A., and Sirard, M. A. (1993). Effect of adenylate cyclase stimulation on meiotic resumption and cyclic AMP content of zona-free and cumulus-enclosed bovine oocytes in vitro. J. Reprod. Fertil. 97, 5–11.
| CrossRef | CAS | PubMed |

Bilodeau-Goeseels, S. (2006). Effects of culture media and energy sources on the inhibition of nuclear maturation in bovine oocytes. Theriogenology 66, 297–306.
| CrossRef | PubMed |

Bornslaeger, E. A., and Schultz, R. M. (1985). Regulation of mouse oocyte maturation: effect of elevating cumulus cell cAMP on oocyte cAMP levels. Biol. Reprod. 33, 698–704.
| CrossRef | CAS | PubMed |

Chen, J., Chi, M. M., Moley, K. H., and Downs, S. M. (2009). cAMP pulsing of denuded mouse oocytes increases meiotic resumption via activation of AMP-activated protein kinase. Reproduction 138, 759–770.
| CrossRef | CAS | PubMed |

Conti, M., Andersen, C. B., Richard, F. J., Shitsukawa, K., and Tsafriri, A. (1998). Role of cyclic nucleotide phosphodiesterases in resumption of meiosis. Mol. Cell. Endocrinol. 145, 9–14.
| CrossRef | CAS | PubMed |

Dekel, N., Aberdam, E., and Sherizly, I. (1984). Spontaneous maturation in vitro of cumulus-enclosed rat oocytes is inhibited by forskolin. Biol. Reprod. 31, 244–250.
| CrossRef | CAS | PubMed |

Desmarais, J. A., Bordignon, V., Lopes, F. L., Smith, L. C., and Murphy, B. D. (2004). The escape of the mink embryo from obligate diapause. Biol. Reprod. 70, 662–670.
| CrossRef | CAS | PubMed |

Downs, S. M. (2011). Mouse versus rat: profound differences in meiotic regulation at the level of the isolated oocyte. Mol. Reprod. Dev. 78, 778–794.
| CrossRef | CAS | PubMed |

Downs, S. M., and Hunzicker-Dunn, M. (1995). Differential regulation of oocyte maturation and cumulus expansion in the mouse oocyte–cumulus cell complex by site-selective analogues of cyclic adenosine monophosphate. Dev. Biol. 172, 72–85.
| CrossRef | CAS | PubMed |

Downs, S. M., Daniel, S. A. J., and Eppig, J. J. (1988). Induction of maturation in cumulus cell-enclosed mouse oocytes by follicle-stimulating hormone and epidermal growth factor: evidence for a positive stimulus of somatic cell origin. J. Exp. Zool. 245, 86–96.
| CrossRef | CAS | PubMed |

Downs, S. M., Daniel, S. A. J., Bornslaeger, E. A., Hoppe, P. C., and Eppig, J. J. (1989). Maintenance of meiotic arrest in mouse oocytes by purines: modulation of cAMP levels and cAMP phosphodiesterase activity. Gamete Res. 23, 323–334.
| CrossRef | CAS | PubMed |

Downs, S. M., Humpherson, P. G., and Leese, H. J. (2002). Pyruvate utilization by mouse oocytes is influenced by meiotic status and the cumulus oophorous. Mol. Reprod. Dev. 62, 113–123.
| CrossRef | CAS | PubMed |

Edwards, R. G. (1965). Maturation in vitro of mouse, sheep, cow, pig, rhesus monkey and human ovarian oocytes. Nature 208, 349–351.
| CrossRef | CAS | PubMed |

Fan, H. Y., Li, M. Y., Tong, C., Chen, D. Y., Xia, G. L., Song, X. F., Schatten, H., and Sun, Q. Y. (2002). Inhibitory effects of cAMP and protein kinase C on meiotic maturation and MAP kinase phosphorylation in porcine oocytes. Mol. Reprod. Dev. 63, 480–487.
| CrossRef | CAS | PubMed |

Funahashi, H., and Day, B. N. (1993). Effects of duration of exposure to hormone supplements on cytoplasmic maturation of pig oocytes in vitro. J. Reprod. Fertil. 98, 179–185.
| CrossRef | CAS | PubMed |

Funahashi, H., Cantley, T. C., and Day, B. N. (1997). Synchronization of meiosis in porcine oocytes by exposure to dibutyryl cyclic adenosine monophosphate improves developmental competence following in vitro fertilization. Biol. Reprod. 57, 49–53.
| CrossRef | CAS | PubMed |

Gilchrist, R. B. (2011). Recent insights into oocyte–follicle cell interactions provide opportunities for the development of new approaches to in vitro maturation. Reprod. Fertil. Dev. 23, 23–31.
| CrossRef | PubMed |

Gilchrist, R. B., and Thompson, J. G. (2007). Oocyte maturation: emerging concepts and technologies to improve developmental potential in vitro. Theriogenology 67, 6–15.
| CrossRef | PubMed |

Godard, N. M., Pukazhenthi, B. S., Wildt, D. E., and Comizzoli, P. (2009). Paracrine factors from cumulus-enclosed oocytes ensure the successful maturation and fertilization in vitro of denuded oocytes in the cat model. Fertil. Steril. 91, 2051–2060.
| CrossRef | CAS | PubMed |

Herrick, J. R., Brad, A. M., and Krisher, R. L. (2006a). Chemical manipulation of glucose metabolism in porcine oocytes: effects on nuclear and cytoplasmic maturation in vitro. Reproduction 131, 289–298.
| CrossRef | CAS | PubMed |

Herrick, J. R., Lane, M., Gardner, D. K., Behboodi, E., Memili, E., Blash, S., Echelard, Y., and Krisher, R. L. (2006b). Metabolism, protein content and in vitro embryonic development of goat cumulus–oocyte complexes matured with physiological concentrations of glucose and l-lactate. Mol. Reprod. Dev. 73, 256–266.
| CrossRef | CAS | PubMed |

Herrick, J. R., Bond, J. B., Magarey, G. M., Bateman, H. L., Krisher, R. L., Dunford, S. A., and Swanson, W. F. (2007). Toward a feline optimised culture medium: impact of ions, carbohydrates, essential amino acids, vitamins and serum on development and metabolism of IVF-derived feline embryos relative to embryos grown in vivo. Biol. Reprod. 76, 858–870.
| CrossRef | CAS | PubMed |

Herrick, J. R., Campbell, M., Levens, G., Moore, T., Benson, K., D’Agostino, J., West, G., Okeson, D. M., Coke, R., Portacio, S. C., Leiske, K., Kreider, C., Polumbo, P. J., and Swanson, W. F. (2010). In vitro fertilization and sperm cryopreservation in the black-footed cat (Felis nigripes) and sand cat (Felis margarita). Biol. Reprod. 82, 552–562.
| CrossRef | CAS | PubMed |

Horner, K., Livera, G., Hinckley, M., Trinh, K., Storm, D., and Conti, M. (2003). Rodent oocytes express an active adenylyl cyclase required for meiotic arrest. Dev. Biol. 258, 385–396.
| CrossRef | CAS | PubMed |

Jensen, J. T., Schwinof, K. M., Zelinski-Wooten, M. B., Conti, M., DePaolo, L. V., and Stouffer, R. L. (2002). Phosphodiesterase 3 inhibitors selectively block the spontaneous resumption of meiosis by macaque oocytes in vitro. Hum. Reprod. 17, 2079–2084.
| CrossRef | CAS | PubMed |

Kawashima, I., Okazaki, T., Noma, N., Nishibori, M., Yamashita, Y., and Shimada, M. (2008). Sequential exposure of porcine cumulus cells to FSH and/or LH is critical for appropriate expression of steroidogenic and ovulation-related genes that impact oocyte maturation in vivo and in vitro. Reproduction 136, 9–21.
| CrossRef | CAS | PubMed |

Laforest, M. F., Pouliot, E., Guéguen, L., and Richard, F. J. (2005). Fundamental significance of specific phosphodiesterases in the control of spontaneous meiotic resumption in porcine oocytes. Mol. Reprod. Dev. 70, 361–372.
| CrossRef | CAS | PubMed |

Lane, M., Gardner, D. K., Hasler, M. J., and Hasler, J. F. (2003). Use of G1.2/G2.2 media for commercial bovine embryo culture: equivalent development and pregnancy rates compared to co-culture. Theriogenology 60, 407–419.
| CrossRef | PubMed |

Lastro, M., Collins, S., and Currie, W. B. (2006). Adenylyl cyclases in oocyte maturation: a characterization of AC isoforms in bovine cumulus cells. Mol. Reprod. Dev. 73, 1202–1210.
| CrossRef | CAS | PubMed |

Liang, C. G., Huo, L. J., Zhong, Z. S., Chen, D. Y., Schatten, H., and Sun, Q. Y. (2005). Cyclic adenosine 3′,5′-monophosphate-dependent activation of mitogen-activated protein kinase in cumulus cells is essential for germinal vesicle breakdown of porcine cumulus-enclosed oocytes. Endocrinology 146, 4437–4444.
| CrossRef | CAS | PubMed |

Littell, R. C., Milliken, G. A., Stroup, W. W., and Wolfinger, R. D. (1996). ‘SAS System for Mixed Models’. (SAS Institute Inc.: Cary, USA.)

Luciano, A. M., Modina, S., Vassena, R., Milanesi, E., Lauria, A., and Gandolfi, F. (2004). Role of intracellular cyclic adenosine 3′,5′-monophosphate concentration and oocyte–cumulus cells communications on the acquisition of the developmental competence during in vitro maturation of bovine oocyte. Biol. Reprod. 70, 465–472.
| CrossRef | CAS | PubMed |

Luvoni, G. C., Chigioni, S., Perego, L., Lodde, V., Modina, S., and Luciano, A. M. (2006). Effect of gonadotrophins during in vitro maturation of feline oocytes on oocyte–cumulus cells functional coupling and intracellular glutathione. Anim. Reprod. Sci. 96, 66–78.
| CrossRef | CAS | PubMed |

Mehlmann, L. M. (2005). Stops and starts in mammalian oocytes: recent advances in understanding the regulation of meiotic arrest and oocyte maturation. Reproduction 130, 791–799.
| CrossRef | CAS | PubMed |

Nogueira, D., Ron-El, R., Friedler, S., Schachter, M., Raziel, A., Cortvrindt, R., and Smitz, J. (2006). Meiotic arrest in vitro by phosphodiesterase 3-inhibitor enhances maturation capacity of human oocytes and allows subsequent embryonic development. Biol. Reprod. 74, 177–184.
| CrossRef | CAS | PubMed |

Norris, R. P., Ratzan, W. J., Freudzon, M., Mehlmann, L. M., Krall, J., Movsesian, M. A., Wang, H., Ke, H., Nikolaev, V. O., and Jaffe, L. A. (2009). Cyclic GMP from the surrounding somatic cells regulates cyclic AMP and meiosis in the mouse oocyte. Development 136, 1869–1878.
| CrossRef | CAS | PubMed |

Ozawa, M., Nagai, T., Somfai, T., Nakai, M., Maedomari, N., Fahrudin, M., Karja, N. W. K., Kaneko, H., Noguchi, J., Ohnuma, K., Yoshimi, N., Miyazaki, H., and Kikuchi, K. (2008). Comparison between effects of 3-isobutyl-1-methylxanthine and FSH on gap-junctional communication, LH-receptor expression and meiotic maturation of cumulus–oocyte complexes in pigs. Mol. Reprod. Dev. 75, 857–866.
| CrossRef | CAS | PubMed |

Papadopoulos, S., Rizos, D., Duffy, P., Wade, M., Quinn, K., Boland, M. P., and Lonergan, P. (2002). Embryo survival and recipient pregnancy rates after transfer of fresh or vitrified, in vivo- or in vitro-produced ovine blastocysts. Anim. Reprod. Sci. 74, 35–44.
| CrossRef | CAS | PubMed |

Pursel, V. G., Wall, R. J., Rexroad, C. E., Hammer, R. E., and Brinster, R. L. (1985). A rapid whole-mount staining procedure for nuclei of mammalian embryos. Theriogenology 24, 687–691.
| CrossRef |

Racowsky, C. (1984). Effect of forskolin on the spontaneous maturation and cyclic AMP content of rat oocyte–cumulus complexes. J. Reprod. Fertil. 72, 107–116.
| CrossRef | CAS | PubMed |

Racowsky, C. (1985a). Effect of forskolin on the spontaneous maturation and cyclic AMP content of hamster oocyte–cumulus complexes. J. Exp. Zool. 234, 87–96.
| CrossRef | CAS | PubMed |

Racowsky, C. (1985b). Effect of forskolin on maintenance of meiotic arrest and stimulation of cumulus expansion, progesterone and cyclic AMP production by pig oocyte–cumulus complexes. J. Reprod. Fertil. 74, 9–21.
| CrossRef | CAS | PubMed |

Richard, F. J., Tsafriri, A., and Conti, M. (2001). Role of phosphodiesterase type 3A in rat oocyte maturation. Biol. Reprod. 65, 1444–1451.
| CrossRef | CAS | PubMed |

Sanbuissho, A., Coskun, S., and Lin, Y. C. (1992). Role of cyclic adenosine monophosphate (cAMP) in vitro on bovine oocyte maturation. Theriogenology 38, 153–163.
| CrossRef | CAS | PubMed |

Sasseville, M., Albuz, F. K., Côté, N., Guillemette, C., Gilchrist, R. B., and Richard, F. J. (2009). Characterization of novel phosphodiesterases in the bovine ovarian follicle. Biol. Reprod. 81, 415–425.
| CrossRef | CAS | PubMed |

Sato, E., and Koide, S. S. (1984). Forskolin and mouse oocyte maturation in vitro. J. Exp. Zool. 230, 125–129.
| CrossRef | CAS | PubMed |

Schroeder, A. C., and Eppig, J. J. (1984). The developmental capacity of mouse oocytes that matured spontaneously in vitro is normal. Dev. Biol. 102, 493–497.
| CrossRef | CAS | PubMed |

Schultz, R. M., Montgomery, R. R., and Belanoff, J. R. (1983). Regulation of mouse oocyte meiotic maturation: implication of a decrease in oocyte cAMP and protein dephosphorylation in commitment to resume meiosis. Dev. Biol. 97, 264–273.
| CrossRef | CAS | PubMed |

Shimada, M., and Terada, T. (2002). Roles of cAMP in regulation of both MAP kinase and p34cdc2 kinase activity during meiotic progression, especially beyond the MI stage. Mol. Reprod. Dev. 62, 124–131.
| CrossRef | CAS | PubMed |

Shu, Y. M., Zeng, H. T., Ren, Z., Zhuang, G. L., Liang, X. Y., Shen, H. W., Yao, S. Z., Ke, P. Q., and Wang, N. N. (2008). Effects of cilostamide and forskolin on the meiotic resumption and embryonic development of immature human oocytes. Hum. Reprod. 23, 504–513.
| CrossRef | CAS | PubMed |

Sirard, M. A., and First, N. L. (1988). In vitro inhibition of oocyte nuclear maturation in the bovine. Biol. Reprod. 39, 229–234.
| CrossRef | CAS | PubMed |

Somfai, T., Kikuchi, K., Onishi, A., Iwamoto, M., Fuchimoto, D. I., Papp, A. B., Sato, E., and Nagai, T. (2003). Meiotic arrest maintained by cAMP during initiation of maturation enhances meiotic potential and developmental competence and reduces polyspermy of IVM/IVF porcine oocytes. Zygote 11, 199–206.
| CrossRef | CAS | PubMed |

Songsasen, N., and Wildt, D. E. (2007). Oocyte biology and challenges in developing in vitro maturation systems in the domestic dog. Anim. Reprod. Sci. 98, 2–22.
| CrossRef | CAS | PubMed |

Stokes, Y. M., Clark, A. R., and Thompson, J. G. (2008). Mathematical modelling of glucose supply toward successful in vitro maturation of mammalian oocytes. Tissue Eng. 14, 1539–1547.
| CrossRef | CAS |

Sutton-McDowall, M. L., Gilchrist, R. B., and Thompson, J. G. (2004). Cumulus expansion and glucose utilization by bovine cumulus–oocyte complexes during in vitro maturation: the influence of glucosamine and follicle-stimulating hormone. Reproduction 128, 313–319.
| CrossRef | CAS | PubMed |

Tang-Pedersen, M., Westergaard, L. G., Erb, K., and Mikkelsen, A. L. (2012). Combination of IVF and IVM in naturally cycling women. Reprod. Biomed. Online 24, 47–53.
| CrossRef | PubMed |

Thomas, R. E., Armstrong, D. T., and Gilchrist, R. B. (2002). Differential effects of specific phosphodiesterase isoenzyme inhibitors on bovine oocyte meiotic maturation. Dev. Biol. 244, 215–225.
| CrossRef | CAS | PubMed |

Thomas, R. E., Armstrong, D. T., and Gilchrist, R. B. (2004a). Bovine cumulus cell–oocyte gap junctional communication during in vitro maturation in response to manipulation of cell-specific cyclic adenosine 3′,5′-monophosphate levels. Biol. Reprod. 70, 548–556.
| CrossRef | CAS | PubMed |

Thomas, R. E., Thompson, J. G., Armstrong, D. T., and Gilchrist, R. B. (2004b). Effect of specific phosphodiesterase isoenzyme inhibitors during in vitro maturation of bovine oocytes on meiotic and developmental capacity. Biol. Reprod. 71, 1142–1149.
| CrossRef | CAS | PubMed |

Vanhoutte, L., Nogueira, D., Gerris, J., Dhont, M., and DeSutter, P. (2008). Effect of temporary nuclear arrest by phosphodiesterase 3-inhibitor on morphological and functional aspects of in vitro-matured mouse oocytes. Mol. Reprod. Dev. 75, 1021–1030.
| CrossRef | CAS | PubMed |

Wildt, D. E., Comizzoli, P., Pukazhenthi, B., and Songsasen, N. (2010). Lessons from biodiversity – the value of non-traditional species to advance reproductive science, conservation and human health. Mol. Reprod. Dev. 77, 397–409.
| CrossRef | CAS | PubMed |

Wood, T. C., Byers, A. P., Jennette, B. E., and Wildt, D. E. (1995). Influence of protein and hormone supplementation on in vitro maturation and fertilization of domestic cat eggs. J. Reprod. Fertil. 104, 315–323.
| CrossRef | CAS | PubMed |

Yoshioka, K., Suzuki, C., Itoh, S., Kikuchi, K., Iwamura, S., and Rodriguez-Martinez, H. (2003). Production of piglets derived from in vitro-produced blastocysts fertilized and cultured in chemically defined media: effects of theophylline, adenosine and cysteine during in vitro fertilization. Biol. Reprod. 69, 2092–2099.
| CrossRef | CAS | PubMed |

Zhang, M., Su, Y. Q., Sugiura, K., Xia, G., and Eppig, J. J. (2010). Granulosa cell ligand NPPC and its receptor NPR2 maintain meiotic arrest in mouse oocytes. Science 330, 366–369.
| CrossRef | CAS | PubMed |