Factors affecting folliculogenesis in ruminants

This review addresses the reasons for the lack of progress in the control of superovulation and highlights the importance of understanding the mechanisms underlying follicular development. The present inability to provide large numbers of viable embryos from selected females still restricts genetic improvement, whilst variability in ovarian response to hormones limit the present capacity for increasing reproductive efficiency.Females are born with a large store of eggs which rapidly declines as puberty approaches. If these oocytes are normal then there is scope for increasing the reproductive potential of selected females. Oocytes must reach a certain size before they can complete all stages of development and the final changes that occur late in follicular development. It is likely that oocytes that do not produce specific factors at precise stages of development will not be viable. Hence, it is important to characterize oocyte secreted factors since there are potential indicators of oocyte quality.The mechanisms that determine ovulation rate have still not been fully elucidated. Indeed follicular atresia, the process whereby follicles regress, is still not known. A better understanding of these processes should prove pivotal for the synchronization of follicular growth, for more precise oestrous synchronization and improved superovulatory response.Nutrition can influence a whole range of reproductive parameters however, the pathways through which nutrition acts have not been fully elucidated. Metabolic hormones, particularly insulin and IGFs, appear to interact with gonadotrophins at the level of the gonads. Certainly gonadotropins provide the primary drive for the growth of follicles in the later stages of development and both insulin and IGF-1, possibly IGF-2, synergize with gonadotrophins to stimulate cell proliferation and hormone production. More research is required to determine the effects of other growth factors and their interaction with gonadotropins.There is evidence, particularly from studies with rodents, that steroids can also modulate follicular growth and development, although information is very limited for ruminants. There may be a rôle for oestrogens in synchronizing follicular waves, to aid in oestrous synchronization regimes and for removing the dominant follicle to achieve improved superovulatory responses. However more information is required to determine whether these are feasible approaches.Heritability for litter size is higher in sheep than in cattle. Exogenous gonadotropins are a commercially ineffective means of inducing twinning in sheep and cattle. Although there are differences in circulating gonadotropin concentrations, the mechanism(s) responsible for the high ovulation appear to reside essentially within the ovaries. The locus of the Booroola gene, a major gene for ovulation rate, has been established but not specifically identified. However sheep possessing major genes do provide extremely valuable models for investigating the mechanisms controlling ovulation rate, including a direct contrast to mono-ovulatory species such as cattle.In conclusion, the relationship between oocyte quality, in both healthy follicles and those follicles destined for atresia, must be resolved before the future potential for increasing embryo yield can be predicted. In addition, a greater understanding of the factors affecting folliculogenesis in ruminants should ensure that the full benefits ensuing from the precise control of ovarian function are achieved. The improved use of artificial insemination and embryo transfer that would ensue from a greater understanding of the processes of folliculo genesis, coupled with the new technologies of genome and linkage mapping, should ensure a more rapid rate of genetic gain.