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J. J. Bass, A. S. McNeilly and H. E. Moreton

Summary. In bulls there was no increase in plasma FSH and only a small increase in LH over the first 14 weeks of age. In steers (castrated) plasma LH and FSH were unchanged for the first 3 weeks but increased significantly at 7 and 14 weeks. After 100 μg Gn-RH, LH release in bulls was minimal until 7 and 14 weeks and there was no comparable rise for FSH. LH and FSH responded to Gn-RH throughout the trial in the steers. The neonatal calf testes selectively inhibited the release of FSH from the pituitary even when challenged with Gn-RH.

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A. S. McNeilly, Anna Glasier, Julie Jonassen and P. W. Howie

Introduction

There is no doubt that in breast-feeding women, suckling with its associated hyperprolactinaemia prevents the resumption of ovarian activity for prolonged periods (see McNeilly, 1979). The extent of this suppression varies greatly among species but in all for which there are adequate data it appears to depend critically upon the intensity of the suckling stimulus (Lamming, 1978). Our recent data from women show that in the pattern of suckling, frequency and duration, throughout the day, are both key factors in maintaining the elevation of basal levels of prolactin associated with lactation (McNeilly, Howie & Houston, 1980a; Howie & McNeilly, 1982). Suckling also releases large quantities of prolactin, maintaining a physiological hyperprolactinaemic state which is directly associated with the duration of lactational amenorrhoea (Delvoye, Badawi, Demaegd & Robyn, 1978; Duchen & McNeilly, 1980). The question remains, how does suckling suppress ovarian activity?

The levels of prolactin in blood during peak

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A. S. McNeilly, W. J. Crow and H. M. Fraser

Summary. Progesterone secretion has been observed to be episodic in the late luteal phase of the oestrous cycle of ewes and is apparently independent of luteinizing hormone (LH). This study investigated the effects of suppressing the pulsatile release of LH in the early or late luteal phase on the episodic secretion of progesterone. Six Scottish Blackface ewes were treated i.m. with 1 mg kg−1 body weight of a potent gonadotrophin-releasing hormone (GnRH) antagonist on either day 4 or day 11 of the luteal phase. Six ewes received saline at each time and acted as controls. Serial blood samples were collected at 10 or 15 min intervals between 0 and 8 h, 24 and 32 h, and 48 and 56 h after GnRH antagonist treatment and daily from oestrus (day 0) of the treatment cycle for 22 days. Oestrous behaviour was determined using a vasectomized ram present throughout the experiment. Progesterone secretion was episodic in both the early and late luteal phase with a frequency of between 1·6 and 3·2 pulses in 8 h. The GnRH antagonist abolished the pulsatile secretion and suppressed the basal concentrations of LH for at least 3 days after treatment. This suppression of LH, in either the early or late luteal phase, did not affect the episodic release of progesterone. Daily concentrations of progesterone in plasma showed a minimal reduction on days 11 to 14 after GnRH antagonist treatment on day 4, although this was significant (P < 0·05) only on days 11 and 13. There was no effect of treatment on day 11 on daily progesterone concentration, and the timing of luteolysis and the duration of corpus luteum function was unaffected by GnRH antagonist treatment on either day 4 or day 11. These results indicate that the episodic secretion of progesterone during the luteal phase of the oestrous cycle in ewes is independent of LH pulses and normal progesterone secretion by the corpus luteum can be maintained with minimal basal concentrations of LH.

Keywords: LH; progesterone; FSH; corpus luteum; GnRH antagonist; sheep

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M. Caillol, M. Mondain-Monval, M. Meunier and A. S. McNeilly

Summary. In the brown hare, fertile mating takes place from the beginning of December to September. Seasonal variations of basal concentrations of LH and FSH, and pituitary response to a monthly i.v. injection of LHRH were studied in intact control females and in females ovariectomized during the seasonal anoestrus (OVX1) or during the breeding season (OVX2). In intact females, both basal and LHRH-stimulated LH levels showed an annual variation, with minimal values during anoestrus. During the breeding season, the LH response to LHRH exhibited a biphasic pattern. In contrast, there was no clear seasonal variation in basal and LHRH-stimulated FSH concentrations. After ovariectomy during anoestrus, basal LH remained low for 2 months and began to increase in December. After ovariectomy during the breeding season, LH basal concentrations increased within a few days after the operation. Thereafter, LH values remained high in both groups of females until September, and decreased significantly as in intact females. The pattern of LH release after LHRH remained monophasic in the two groups of ovariectomized females. In OVX1 females, the LH response increased as early as October, was maximum from December to April and decreased progressively until October. In OVX2 females, the LH response decreased regularly after ovariectomy to a minimum in October. In the 2 groups of ovariectomized females, basal FSH concentrations and pituitary response to LHRH rose rapidly after ovariectomy and did not vary significantly thereafter. These results showed a direct central effect of season on the regulation of basal concentrations of LH, modulated by a negative feed-back of ovarian secretions during the breeding season. In intact hares, the enhanced LH response after LHRH during the breeding season was related to an acute positive effect of ovarian secretions. The regulation of FSH was less dependent on season and remained under a negative control of the ovary throughout the year.

Keywords: brown hare; LH; FSH; LHRH; ovariectomy; season

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J. K. Hodges, Cilla Henderson and A. S. McNeilly

Summary. Oestrone, oestradiol-17β and oestriol were measured in plasma samples from non-pregnant and pregnant African elephants shot in the wild. Enzymic hydrolysis of plasma showed that approximately 90 and 96% of the total (i.e. conjugated plus unconjugated) concentrations of oestrone and oestradiol-17β, respectively were represented by conjugated hormones. Unconjugated oestrogens remained low ( <50 pg/ml) in all samples, with no distinction between non-pregnant and pregnant animals. Levels of total oestrone during pregnancy varied between 160 and 594 pg/ml but were not significantly different from non-pregnant values. Total oestradiol-17β concentrations were significantly elevated during pregnancy (P <0·01) and, despite considerable individual variation (193–1428 pg/ml), were consistently higher than non-pregnant values after 6 months of gestation. The elevated levels of oestradiol-17β resulted in a reversal of the total oestradiol-17β:oestrone concentration ratio at about 6 months of pregnancy. Concentrations of total oestriol did not exceed 103 pg/ml. An indirect method of measurement indicated that oestradiol-17β sulphate was probably the most abundant circulating oestrogen during pregnancy in the African elephant.

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S. M. Rhind, P. J. Goddard, S. R. McMillen and A. S. McNeilly

Ovarian follicle development in response to FSH infusion was investigated in Scottish Blackface ewes with high and low body condition scores in which endogenous gonadotrophin secretion and follicle development to ≥ 2.5 mm diameter was suppressed using subcutaneous implants containing a GnRH agonist. In two experiments conducted during the normal breeding season, groups of 20 (Expt 1) and 15 (Expt 2) ewes were fed to achieve body condition scores ≥2·75 (high; H) or ≤1·75 (low; L). In both experiments GnRH agonist implants were inserted four weeks before FSH was infused for 72 h at 7 μg h−1 to group H animals or at 5 μg h−1 to group L animals; the infusion rates were designed to ensure similar circulating FSH concentrations in animals of both groups. In Expt 2, additional subcutaneous implants containing oestradiol were inserted 21 days after insertion of GnRH agonist implants and 7 days before the FSH infusion began. In both experiments, FSH infusion was associated with an increase in circulatory concentrations of LH (P < 0.01) and FSH (P < 0·001), but there was no difference with body condition in mean circulating gonadotrophin concentrations, the numbers of ovarian follicles ≥ 2.5 mm diameter, the proportion of these follicles that were oestrogenic or the mean rate of oestradiol secretion in vitro. It is concluded that differences in body condition of ewes do not affect the responsiveness of the ovary to FSH, in the presence or absence of oestradiol, as measured by the number, size and steroidogenic capacity of ovarian follicles present following FSH infusion.

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L. E. A. ROWSON, A. S. McNEILLY and C. A. O'BRIEN

While surgical transfer of bovine eggs presents little difficulty, non-surgical attempts at transfer by deposition into the uterus through the cervical canal have been singularly unsuccessful. The occurrence of uterine contractions can be recorded on a kymograph almost immediately after non-surgical manipulation of the uterus. Isolated successes have been reported, however, when the uterus was distended with carbon dioxide at the time of egg transfer (Rowson & Moor, 1966; Vincent, Mills & Rundell, 1969). It was found that both natural eggs and artificial eggs in the form of resin spheres of the same size and density were ejected from the uterus within a few hours of their non-surgical deposition within the horn. A possible reason for this rejection might be the

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J M Young, S Henderson, C Souza, H Ludlow, N Groome and A S McNeilly

Little is known about the role of activin B during folliculogenesis. This study investigated the expression levels of activin/inhibin subunits (βA, βB, and α), steroid enzyme, and gonadotrophin receptors in theca (TC) and granulosa cells (GC) by QPCR and activin A and B and inhibin A protein levels in follicular fluid (FF) of developing sheep follicles during estrus and anestrus. The effect of activin B on androgen production from primary TC cultures in vitro was also assessed. During folliculogenesis, in anestrus and estrus, FF activin B concentrations and thecal and GC activin βB mRNA levels decreased as follicle diameter increased from 1–3 to >6 mm regardless of estrogenic status. Estrogenic preovulatory follicles had reduced concentrations of FF activins B and A, and TC and GCs expressed higher levels of activin βA mRNA at 3–4 mm, and TCs more inhibin α mRNA at >4 mm stages of development compared with nonestrogenic follicles. Activin B decreased androstenedione production from primary TCs in vitro, an effect blocked by inhibin A. Thus, sheep follicles 1–3 mm in diameter contained high FF levels of activin B, which decreased as the follicle size increased, and, like activin A, suppressed thecal androgen production in vitro, an effect blocked by inhibin. Furthermore, the theca of large estrogenic follicles expressed high levels of inhibin α and activin βA mRNA suggesting local thecal derived inhibin A production. This would inhibit the negative effects of thecal activins B and A ensuring maximum androgen production for enhanced estradiol production by the preovulatory follicle(s).

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D. T. Baird, T. Bäckström, A. S. McNeilly, S. K. Smith and C. G. Wathen

Summary. To investigate the mechanism of suppression of follicular development during the luteal phase of the human menstrual cycle, the corpus luteum was enucleated surgically from 10 women at various times after ovulation. In the 24 h after CL enucleation there was an immediate and rapid fall in the concentration of oestradiol and progesterone and a temporary decline in the concentration of FSH and LH. Within 3 days, however, all 10 women showed evidence of renewed follicular activity as indicated by a progressive rise in the concentration of oestradiol. This rise was preceded by a rise in the concentration of FSH and LH, and ovulation, as indicated by a mid-cycle surge in LH and rise in the concentration of plasma progesterone, occurred 16–19 days after enucleation. There was no significant difference in the time to ovulation following enucleation at different times of the luteal phase. The post-operative follicular phase, measured from the time of enucleation, was 3 days longer than that observed pre-operatively from the first day of menstrual bleeding. In the follicular phase of post-operative cycles the concentration of FSH was higher and that of oestradiol lower than the corresponding values before surgery.

These results indicate that the absence of healthy antral follicles in the luteal phase of the cycle is due to the inhibitory effects of the corpus luteum. The fact that, after CL enucleation, emergence of the dominant follicle was always preceded by a rise in the concentration of FSH and LH suggests that suppression of gonadotrophins by ovarian steroids secreted by the corpus luteum is responsible for the inhibition of follicular development during the luteal phase of the cycle.

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F. G. L. Hartley, B. K. Follett, S. Harris, D. Hirst and A. S. McNeilly

The endocrine basis of reproductive failure in red fox vixens was examined over two breeding seasons in a total of 11 animals. Weekly blood samples were assayed for progesterone, prolactin, LH and cortisol. Vaginal smears taken every 2 days over the oestrous period indicated that all vixens had mated. Vixens that successfully gave birth to a litter of cubs demonstrated significantly higher plasma progesterone and prolactin concentrations but significantly lower cortisol concentrations than did females that had ovulated, but then failed to whelp. There were no significant differences in plasma LH concentrations. These data suggest that reproductive losses could result from lowered plasma progesterone concentrations, possibly resulting from inadequate luteotrophic support by prolactin. A stress-induced mechanism of reproductive failure is implicated and is discussed in relation to social suppression of reproduction.