The paper presents an update of our 1993 model of ovarian follicular development in ruminants, based on knowledge gained from the past 15 years of research. The model addresses the sequence of events from follicular formation in fetal life, through the successive waves of follicular growth and atresia, culminating with the emergence of ovulatory follicles during reproductive cycles. The original concept of five developmental classes of follicles, defined primarily by their responses to gonadotrophins, is retained: primordial, committed, gonadotrophin-responsive, gonadotrophin-dependent and ovulatory follicles. The updated model has more extensive integration of the morphological, molecular and cellular events during folliculogenesis with systemic events in the whole animal. It also incorporates knowledge on factors that influence oocyte quality and the critical roles of the oocyte in regulating follicular development and ovulation rate. The original hypothetical mechanisms determining ovulation rate are retained but with some refinements; the enhanced viability of gonadotrophin-dependent follicles and increases in the number of gonadotrophin-responsive follicles by increases in the throughput of follicles to this stage of growth. Finally, we reexamine how these two mechanisms, which are thought not to be mutually exclusive, appear to account for most of the known genetic and environmental effects on ovulation rate.

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This study tested whether the effects of a short period of nutritional supplementation given to ewes during the luteal phase on follicle development and ovulation rate is associated with an increase in circulating concentrations of FSH, glucose or metabolic hormones. Oestrus was synchronised with two prostaglandin injections given 9 days apart and the supplement consisted of corn grain and soybean meal. Corriedale ewes with low body condition were randomly assigned to 2 groups: the control group (C; n 5 10) received a maintenance diet while the short-term supplemented group (STS; n 5 10) received double the maintenance diet over days 9 to 14 of the oestrous cycle (day 0 5 ovulation). Ovaries were examined daily by ultrasound and blood was sampled three times a day during the inter-ovulatory interval for measuring reproductive and metabolic hormones. On days 9, 11 and 14 of the oestrous cycle, half of the ewes from each group (n 5 5) were bled intensively to determine the concentrations of glucose, insulin, IGF-I and leptin. Plasma FSH, progesterone, oestradiol and androstenedione concentrations were similar among groups. Dietary supplementation increased plasma insulin concentrations from the first to the sixth day of supplementation and increased glucose concentrations on the third day, compared with control ewes. Plasma leptin concentrations were higher in STS ewes from the second to the fifth day of supplementation. The pattern of IGF-I concentrations was similar among groups. In STS ewes, the nutritional treatment prolonged the lifespan of the last non-ovulatory follicle, so fewer follicular waves developed during the cycle. In STS ewes, increased concentrations of glucose, insulin and leptin one day before ovulatory wave emergence were associated with increased numbers of follicles growing from 2 to 3 mm and with stimulation of the dominant follicle to grow for a longer period. We suggest that the mechanism by which short-term nutritional supplementation affects follicle development does not involve an increase in FSH concentrations, but may involve responses to increased concentrations of glucose, insulin and leptin, acting directly at the ovarian level. This effect is acute, since concentrations of all three substances decrease after reaching peak values on the third day of supplementation. The status of follicle development at the time of maximum concentrations of glucose and metabolic hormones may be one of the factors that determines whether ovulation rate increases or not.

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In the present study, it was investigated whether undernutrition affected the binding capacity, immunoreactivity and mRNA expression for uterine oestrogen and progesterone receptors (ER and PR, respectively) in sheep, as well as whether the responses were associated with changes in plasma concentrations of progesterone (P4), oestradiol (E2), glucose, fatty acids, insulin, leptin and insulin-like growth factor (IGF)-I during the oestrous cycle. Twenty ewes were fed either 1.5 (C) or 0.5 (L) times their maintenance requirements and were killed on Day 5 or 14 of the cycle (Day 0 = oestrus). Compared with Group C, Group L had higher concentrations of non-esterified fatty acids and lower concentrations of insulin, leptin and IGF-I. Group L also had higher plasma concentrations of P4 during the final days of the luteal phase. At oestrus in both treatment groups, there were peaks in the concentrations of glucose, insulin and IGF-I. For ER and PR, transcript expression, binding capacity and immunoreactivity were higher on Day 5 than on Day 14 of the cycle. The binding capacities for ER and PR were lower in Group L than in Group C on Day 5. Group C showed more immunoreactive staining for ER than did Group L in two of five cell types, whereas no effect of treatment was observed for PR immunoreactivity. There was more PR mRNA in the uterine horn contralateral to the corpus luteum in Group C than in Group L ewes. We conclude that undernutrition impairs steroid receptor expression and binding capacity. This may alter the uterine environment and help explain the reductions in embryo survival.

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We have developed an experimental model in which groups of ewes are simultaneously experiencing the first ovarian follicular wave of their oestrous cycle. We used this 'first-wave model' in a 2!2 factorial experiment (ten ewes per group) to study the effect of body condition (BC) and a short-term supplement on follicular dynamics and ovulation rate. The 'first-wave' was established by giving ewes three injections of prostaglandin (PG), 7 days apart. The 6-day supplement (lupin grain) began 2 days after the second PG injection and continued until the third. Follicles were studied by ultrasound, and blood was sampled to measure glucose and hormones. The supplement increased (P!0.01) the concentrations of glucose, insulin and leptin, decreased FSH concentrations (P!0.01) and tended to increase oestradiol concentrations (PZ0.06). The supplement tended to increase the number of 3 mm follicles (PZ0.06). Compared with low-BC ewes, high-BC ewes had more follicular waves (P!0.05), higher concentrations of insulin, leptin and IGF1 (P!0.05) and tended to have higher FSH concentrations (PZ0.09). Leptin and insulin concentrations remained high until the end of supplementation in high-BC ewes, whereas they decreased after the third day of supplementation in low-BC ewes. In conclusion, high concentrations of metabolic hormones in fat ewes are associated with the development of more follicular waves. When a supplement is superimposed on this situation, changes in glucose and metabolic hormones allow more follicles to be selected to ovulate.

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