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B. Malpaux and F. J. Karsch

Summary. This study tested the hypothesis that short days can prolong the breeding season of the ewe when reproductive activity is initiated by an endogenous process, as opposed to when it is driven by exposure to short days. Suffolk ewes were ovariectomized and treated with Silastic capsules containing oestradiol; reproductive activity was monitored from serum concentrations of LH. In this model, a rise in LH is indicative of onset of the breeding season and the duration of the elevation in LH is an indicator of length of reproductive activity. The ewes were subjected to 6-month alternations between long and short photoperiods such that the LH rise began during exposure to the inhibitory long photoperiod (i.e. it resulted from an endogenous process). When short days were provided soon after such a spontaneous onset of the LH rise, the duration of reproductive activity was greater than that observed when ewes were maintained in long days throughout the LH elevation (194 ± 10 vs 155 ± 15 days; P < 0·02). Since the transition from anoestrus to the breeding season in Suffolk ewes maintained outdoors does not require a decrease in daylength and appears to be generated by an endogenous process, our results support the hypothesis that shortening photoperiod sustains the natural breeding season which begins in early autumn.

Keywords: seasonal reproduction; photoperiod; photorefractoriness; LH; breeding season; sheep

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J. E. Robinson and F. J. Karsch

Summary. The reproductive neuroendocrine response of Suffolk ewes to the direction of daylength change was determined in animals which were ovariectomized and treated with constant release capsules of oestradiol. Two groups of animals were initially exposed to 16 or 10 h light/day for 74 days. On day zero of the study, when one group of ewes was reproductively stimulated (elevated LH concentrations) and the other reproductively inhibited (undetectable LH concentrations), half the animals from each group were transferred to an intermediate daylength of 13 h light/day. The remaining ewes were maintained on their respective solstice photoperiods to control for photorefractoriness. LH concentrations rose in animals experiencing a 3 h decrease in daylength from 16L:8D to 13L:11D while LH concentrations fell to undetectable values in those that experienced a 3 h increase in daylength from 10L:14D to 13L:1 1D. The photoperiodic response of the Suffolk ewe, therefore, depends on her daylength history. Such a result could be explained if the 24-h secretory pattern of melatonin secretion, known to transduce photoperiodic information to the reproductive axis, was influenced by the direction of change of daylength. Hourly samples for melatonin were collected for 24h 17 days before and three times after transfer to 13L:1 ID. The melatonin secretory profile always conformed to daylength. Therefore, the mechanism by which the same photoperiod can produce opposite neuroendocrine responses must lie downstream from the pineal gland in the processing of the melatonin signal.

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N. L. Wayne, B. Malpaux and F. J. Karsch

Summary. Female Suffolk sheep were pinealectomized around the vernal equinox to eliminate the major environmental input to the reproductive system (photoperiod) and then either isolated from, or maintained with, pineal-intact gonad-intact sheep. The ewes were ovariectomized and treated with constant-release oestradiol implants and reproductive state was monitored by measuring serum LH concentrations. Pinealectomized ewes that were isolated from the normal flock showed a 2½-month delay in onset of the seasonal rise in LH values compared with that of pineal-intact controls (18 November vs 5 September). On the other hand, pinealectomized ewes that were maintained with the flock showed an onset of the seasonal rise in LH that was not delayed. These results suggest a timekeeping role for social cues for timing onset of the breeding season in an animal that normally relies on photoperiodic signals for temporal regulation of the seasonal reproductive cycle.

Keywords: photoperiod; pineal gland; seasonal reproduction; sheep; social cues

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J. F. ROCHE, F. J. KARSCH, D. L. FOSTER and P. J. DZIUK

The ovary exerts a dominant influence on the secretion of gonadotrophins from the pituitary gland (Nalbandov, 1964). Ovariectomy of ewes caused an increase in serum LH and FSH (Arendarčik & Maraček, 1968; Roche, Foster, Karsch & Dziuk, 1970). The purpose of this experiment was to determine which compartments of the ovary (follicles, CL or stroma) normally maintain the basal level of LH in serum during the luteal phase of the oestrous cycle and throughout anoestrus (Roche, Foster, Karsch, Cook & Dziuk, 1970; Denamur, 1972).

Twenty Grade Columbia ewes of 4 to 8 years of age were penned outdoors with `painted' vasectomized rams. Ewes were checked twice daily for marks and the day of oestrus was designated Day 0 of the oestrous cycle. The ovaries were exposed at

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D. O'Callaghan, F. J. Karsch, M. P. Boland, J. P. Hanrahan and J. F. Roche

Summary. Photoperiod may regulate seasonal reproduction either by providing the primary driving force for the reproductive transitions or by synchronizing an endogenous reproductive rhythm. This study evaluated whether breed differences in timing of the reproductive seasons of Finnish Landrace (Finn) and Galway ewes are due to differences in photoperiodic drive of the reproductive transitions or to differences in photoperiodic synchronization of the endogenous rhythm of reproductive activity. The importance of decreasing photoperiod after the summer solstice in determining the onset and duration of the breeding season was tested by housing ewes from the summer solstice in either a simulated natural photoperiod or a fixed summer-solstice photoperiod (18 h light:6 h dark; summer-solstice hold). Onset of the breeding season within each breed did not differ between these photoperiodic treatments, but Galway ewes began and ended their breeding season earlier than Finn ewes. The duration of the breeding season was shorter in Galway ewes on summer-solstice hold than on simulated natural photoperiod; duration did not differ between photoperiodic treatments in Finn ewes. The requirement for increasing photoperiod after the winter solstice for initiation of anoestrus was tested by exposing ewes from the winter solstice to either a simulated natural photoperiod or a winter-solstice hold photoperiod (8·5 h light:15·5 h dark). Onset of anoestrus within each breed did not differ between these photoperiodic treatments, but the time of this transition differed between breeds. These observations suggest that genetic differences in timing of the breeding season in Galway and Finn ewes do not reflect differences in the extent to which photoperiod drives the reproductive transitions, because neither breed requires shortening days to enter the breeding season or lengthening days to end it at appropriate times. These findings are consistent with the hypothesis that photoperiod synchronizes an endogenous rhythm of reproductive activity in both breeds and that genetic differences in timing of the breeding season reflect differences in photoperiodic synchronization of this rhythm.

Keywords: breeding season; sheep; photoperiod

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B. Malpaux, J. E. Robinson, M. B. Brown and F. J. Karsch

Summary. Three groups of ovariectomized Suffolk ewes bearing s.c. Silastic implants of oestradiol were subjected to a 90-day priming treatment of an inhibitory long photoperiod (16 h light/day; 16L:8D). On Day 0 of the experiment, they were moved to stimulatory photoperiods. One control group was transferred to 12L:12D and a second control group was transferred to 8L:16D; both groups remained in those photoperiods to determine the timing of reproductive induction and refractoriness. The experimental group was transferred to 12L:12D on Day 0 and then to 8L:16D on Day 55 to determine whether the further reduction in daylength could delay the development of refractoriness. Reproductive neuroendocrine condition was monitored by serum concentrations of LH and FSH. Both gonadotrophins remained elevated for a longer period of time in the experimental group receiving the second reduction in daylength than in either control group, indicating that the second photoperiodic drop delayed the onset of photorefractoriness. Measurement of 24-h patterns of circulating melatonin suggests that the prolonged stimulation of reproductive neuroendocrine activity in the experimental group resulted from a lengthening of the nocturnal melatonin rise. These findings indicate that refractoriness to an inductive photoperiod can be temporarily overcome by exposure to a shorter daylength, and that the change in duration of the nocturnal increase in melatonin secretion is important in photoperiodic signalling. Thus, in natural conditions, the decreasing autumnal daylength, and the resulting expansion of the nocturnal elevation in melatonin secretion, may be utilized to produce a breeding season of normal duration.

Keywords: seasonal reproduction; photorefractoriness; melatonin; oestradiol negative feedback; photoperiodic history; sheep

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B. Cook, F. J. Karsch, Jean W. Graber and A. V. Nalbandov

In many mammalian species, pregnancy triggers a luteotrophic mechanism that causes the corpora lutea (CL) to persist for a period longer than the lifespan of the cyclic CL. In marsupials, however, the life span of CL is not changed by pregnancy. In Didelphis marsupialis, for example, the CL reach maximum size on Day 3 after ovulation, begin to regress on Day 7 and are almost fully regressed by Day 13 when parturition occurs (Hartman, 1923). The next ovulation takes place about Day 28 regardless of whether pregnancy has intervened, provided no young are suckling (Hartman, 1923). Unlike the effects in sheep and some other species (Anderson, Bowerman & Melampy, 1963), hysterectomy also fails to prolong the life of CL in the opossum (Hartman, 1925) and there appears to be no known way in which the life of the didelphine CL can be extended. Because luteotrophic mechanisms induced by pregnancy and luteolytic mechanisms depending on the uterus seem to be absent in the opossum, the direct action of oestrogen on the lifespan of the opossum CL was investigated. Hoffmann (1960) suggested, as a consequence of his studies in women, that oestrogen acted directly on CL to induce regression, and a direct effect of oestrogen has been demonstrated for the CL of sheep (Cook, Karsch, Foster & Nalbandov, 1974) and rhesus monkeys (Karsch & Sutton, 1976).

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D. L. Foster, K. D. Ryan, R. L. Goodman, S. J. Legan, F. J. Karsch and S. M. Yellon

Summary. Intact female lambs were chronically treated with low levels of oestradiol by Silastic implant from 20 weeks of age. Reproductive cycles were initiated in only 33% of these lambs (3 of 9) compared to 80% of untreated females (11 of 14) by 45 weeks when the study was terminated. Moreover, in the 3 oestradiol-treated lambs which began cycles, the age at first oestrus was delayed 3 weeks (37 ± 1 weeks of age vs 34 ± 1 weeks of age for untreated controls). Retardation of the pubertal process was not due to absence of the pubertal rise in circulating LH. At about 32 weeks of age, chronic oestradiol treatment was no longer able to suppress tonic LH secretion and serum LH increased in intact, oestradiol-treated lambs. These results indicate that a maturational decrease in responsiveness to oestradiol inhibition of tonic LH secretion can be demonstrated in the intact female, as in the ovariectomized female. However, chronic oestradiol suppression of prepubertal LH secretion also delays onset of reproductive cycles. This finding raises the possibility that low tonic LH secretion, presumably in the form of slow pulses, is necessary for development or maintenance of ovarian function before puberty. In the absence of LH during the last part of sexual maturation, the ability of the ovary to respond to the high frequency LH pulses during the pubertal gonadotrophin rise may be delayed.