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A. S. McNeilly and D. T. Baird

Summary. The effect of sustained high plasma levels of prolactin, induced by repeated 2-h i.v. injections of thyrotrophin-releasing hormone (TRH; 20 μg), on ovarian oestradiol secretion and plasma levels of LH and FSH was investigated during the preovulatory period in the ewe. Plasma levels of progesterone declined at the same rate after prostaglandin-induced luteal regression in control and TRH-treated ewes. However, TRH treatment resulted in a significant increase in plasma levels of LH and FSH compared to controls from 12 h after luteal regression until 5 to 6 h before the start of the preovulatory surge of LH. In spite of this, and a similar increase in pulse frequency of LH in control and TRH-treated ewes, ovarian oestradiol secretion was significantly suppressed in TRH-treated ewes compared to that in control ewes. The preovulatory surge of LH and FSH, the second FSH peak and subsequent luteal function in terms of plasma levels of progesterone were not significantly different between control and TRH-treated ewes.

These results show that TRH treatment, presumably by maintaining elevated plasma levels of prolactin, results in suppression of oestradiol secretion by a direct effect on the ovary in the ewe.

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P. W. Howie and A. S. McNeilly

An adequate inter-birth interval is of great importance because it enables a mother to recover her physical and emotional strength between pregnancies and confers upon the child advantages of better health and development (Morley, 1977). Breast feeding is the naturally evolved method of ensuring an adequate inter-birth interval but the contraceptive effect of breast feeding has been largely dismissed in developed countries on the grounds that it is an unreliable method of family planning for individual mothers (Kamal et al., 1969). The use of artificial contraception varies very widely in different countries, being much higher in developed nations than in the poorer developing countries (Text-fig. 1). Where the use of artificial contraceptives is low, breast feeding is the most widely used method of birth spacing and assumes major demographic importance. It is important, therefore, that the factors controlling lactational infertility should be understood so that guidelines can be developed which

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G. A. Lincoln, C. E. Lincoln and A. S. McNeilly

Summary. Seasonal cycles in testicular activity in rams were monitored in groups of wild (mouflon), feral (Soay) and domesticated breeds of sheep (Shetland, Blackface, Herdwick, Norfolk, Wiltshire, Portland and Merino) living outdoors near Edinburgh (56°N). The changes in the blood plasma concentrations of FSH, inhibin and testosterone, and the diameter of the testis were measured every half calendar month from 1 to 3 years of age. There were significant differences between breeds in the magnitude and timing of the seasonal reproductive cycle. In the mouflon rams, the seasonal changes were very pronounced with a 6–15-fold increase in the plasma concentrations of FSH, inhibin and testosterone from summer to autumn, and a late peak in testicular diameter in October. In the Soay rams and most of the domesticated breeds, the seasonal increase in the reproductive hormones occurred 1–2 months earlier with the peak in testicular size in September or October. In the two southern breeds (Portland and Merino), the early onset of testicular activity was more extreme with the seasonal maximum in August. In cross-bred rams, produced by mating Soay ewes (highly seasonal breed) with Portland or Merino rams (less seasonal breeds), there was a seasonal reproductive cycle that was intermediate compared to that of the parents. A comparison between all 11 breeds showed a significant correlation between the timing of the seasonal cycle in plasma FSH concentration and testicular diameter (time of peak FSH vs testis, r = 0·95).

The overall results in the rams are consistent with a primary role of FSH in dictating the seasonal cycle in testicular size and the secretion of inhibin. The earlier seasonal onset in the testicular cycle in the southern breeds of domesticated sheep, and the differences from the wild type, are taken to represent the effects of genetic selection for a longer mating season.

Keywords: seasonal breeding; pituitary gland; testis; inhibin; genetic variation; sheep

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A. S. McNeilly, J. A. Jonassen and H. M. Fraser

Summary. Active immunization of 6 Damline ewes against LHRH during seasonal anoestrus resulted in an inhibition of ovarian cyclicity throughout 2 subsequent breeding seasons. This was associated with a significant suppression of plasma LH and FSH concentrations but no significant effect on prolactin. The ovaries of LHRH-immunized ewes 30 months after primary immunization contained no follicles > 2·5 mm in diameter and a greater proportion of follicles between 1 and 2 mm were atretic than in control ewes (N = 8). In-vitro production of testosterone and androstenedione were similar in follicles 1–2 mm in both control and LHRH-immunized ewes (N = 6) and all had little or no ability to secrete oestradiol. However, basal and hCG-stimulated progesterone secretion was suppressed in the follicles from LHRH-immunized ewes. These results show that follicular development beyond 2·5 mm in the ewe is dependent on adequate stimulation by both LH and FSH.

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K. M. Henderson, A. S. McNeilly and I. A. Swanston

Summary. The concentrations of LH, FSH and prolactin, and oestradiol-17β, androstenedione, testosterone and progesterone were measured in follicular fluid from small, medium and large bovine follicles. As follicle size increased, there was a significant increase in median fluid concentrations of prolactin (2-fold) and oestradiol-17β (14-fold) and a significant decrease in concentrations of LH (to 73%), androstenedione (to 30%) and testosterone (to 10%). There was no relationship between follicle size and fluid concentrations of FSH or progesterone, or between fluid concentrations of FSH and the relative concentrations of androgen and oestradiol-17β. As follicle size increased there was a significant increase in the proportion of follicles in which follicular fluid concentrations of oestradiol-17β exceeded those of androgen. There was a significant relationship between follicular fluid concentrations of prolactin and progesterone; as fluid prolactin concentrations increased, the maximum concentration of progesterone observed decreased.

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G. A. Lincoln, A. S. McNeilly and C. L. Cameron

Summary. Six adult Soay rams were preconditioned to an artificial lighting regimen of alternating 4-month periods of long (16L:8D) and short days (8L:16D) for at least 10 months before blood samples were collected at hourly intervals for 24 h at various times. The abrupt change from long to short days resulted in a progressive decrease in plasma levels of prolactin, while that from short to long days had the reverse effect; the first response to the light changes was rapid, beginning within 6 days. During the periods of high secretion there was a 24-h cycle in plasma prolactin concentrations, with a peak in both the early dark and early light phases of each day. Changes in the relative magnitude of these peaks were observed in relation to the long-term alteration in prolactin secretion. Plasma levels of FSH were also measured and a close inverse relationship between gonadotrophin and prolactin secretion was observed.

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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


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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J. Brooks, W. J. Struthers and A. S. McNeilly

Gonadotrophin and inhibin concentrations were measured in anoestrous ewes after acute treatment with either saline, ovine follicular fluid (oFF), GnRH antagonist, or oFF and GnRH antagonist in combination. The increase in mean LH concentrations observed in ewes treated with oFF alone, was not seen in either of the groups treated with GnRH antagonist, in which LH pulsatility was completely inhibited. This result suggests that the LH rebound that follows follicular fluid treatment is GnRH dependent. Blockade of GnRH had no effect on the suppression of FSH seen after follicular fluid injection, indicating that this component of FSH secretion is independent of short-term GnRH input. After this initial suppression, a rebound release of FSH was seen in the group treated with oFF alone. The addition of GnRH antagonist appeared to decrease the rebound, suggesting that this rebound release of FSH may have a GnRH-dependent component. Inhibin concentrations in both oFF-treated groups increased after oFF injection and then declined to pretreatment values. However, a second rise in inhibin concentration, concomitant with the FSH rebound in ewes receiving oFF alone, was seen in the group treated with oFF and GnRH antagonist. As this rise in endogenous inhibin concentration could also act to suppress the rebound release of FSH, it cannot be conclusively proved from this study that GnRH input is required for the generation of the rebound release of FSH after treatment with oFF.

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H. M. Fraser, I. J. Clarke and A. S. McNeilly

Summary. Endogenous LH-RH in ewes was inhibited by active immunization or by injection of LH-RH antiserum. Plasma levels of LH and FSH were elevated in 3 ovariectomized control ewes but low in 3 LH-RH immunized ovariectomized ewes. Oestradiol benzoate (50 μg i.m.) caused a marked rise in LH concentrations in control ewes but not in the immunized ewes. In the immunized ewes the low plasma levels of FSH decreased even further 8–36 h after injection of oestrogen, indicating a direct inhibitory action of the steroid on the pituitary. Both groups responded to the oestrogen injection by a rise in plasma levels of prolactin and by exhibiting normal oestrous behaviour.

When the control ewes were again challenged with oestradiol benzoate and, after 10 h, given an i.v. injection of 75 ml antiserum to LH-RH, the LH surge was abolished in one animal and reduced in another. These experiments indicate that the continued presence of LH-RH is necessary for the occurrence of the oestrogen-induced LH surge in the ewe.

Administration of a stimulatory analogue of LH-RH released LH and FSH in control and immunized ewes but the responsiveness to further injections at intervals of 3 h decreased, particularly for FSH.