Reproductive photorefractoriness in rams and accompanying changes in the patterns of melatonin and prolactin secretion

Exposure of rams to alternating 16-week cycles of long and short days (16L:8D and 8L:16D) results in periods of testicular regression followed by testicular redevelopment, and there is an inverse relationship between the blood levels of prolactin and testis activity. In this study, two groups of rams were held under long or short day lengths for a period of 94 weeks. When held under either long or short days for more than 16 weeks, the animals showed spontaneous changes in gonadal activity and in the secretion of prolactin, both of which were no longer correlated with the prevailing photoperiod, i.e., they became photorefractory. The photorefractoriness was characterized by cyclical changes in testis function which were independent of day length. The period of these spontaneous cycles was similar during both treatment regimens (long days: 40.9 +/- 1.5 weeks; short days: 38.1 +/- 0.33 weeks), suggesting the presence of an endogenous pacemaker for the reproductive system. The changes in blood prolactin levels during photorefractoriness were no longer correlated with testis activity, and though cyclical, the period lengths differed under the two regimens (long days: 31.8 +/- 1.4 weeks; short days: 48.6 +/- 0.3 weeks). The rates of change in testis function and prolactin secretion were slower during the refractory state than during the photosensitive state. Upon switching the rams to a different photoperiod after the 94 weeks of exposure to fixed day lengths, the rams showed relatively rapid testicular and prolactin responses. Photoperiodic information appears to be relayed to the endocrine system through the daily pattern of melatonin secretion by the pineal. We measured the daily blood levels of melatonin on several occasions during phases of photosensitivity and photorefractoriness in the same group of rams. During the first 21 weeks under both lighting treatments, the rams showed synchronized daily patterns in their blood levels of melatonin, with elevated levels occurring mainly during the daily period of darkness. Similar synchronized daily rhythms were also seen when the rams were switched to a different photoperiod following 94 weeks of exposure to either long or short days. Between Weeks 21 and 94, the daily rhythms of melatonin did not occur consistently in all rams; often, the patterns differed markedly between individual rams held under the same day length and peak levels of melatonin were not always confined to periods of darkness.(ABSTRACT TRUNCATED AT 400 WORDS)     

Photoperiod entrainment of testosterone, luteinizing hormone, follicle-stimulating hormone, and prolactin cycles in rams in relation to testis size and semen quality

Adult rams were exposed to photoperiod treatments over 2 years to study the influence of light regimes on pituitary-testicular activity and semen quality. Initially, all rams (12 per group) were exposed to 3 months of long days (16L:8D). Group 1 was then exposed to a regime of continuous short days (8L:16D) and Groups 2, 3, and 4 were exposed to 4 months of short days alternated with 1, 2, or 4 months, respectively, of long days. Every 2 weeks, serum hormone levels and scrotal circumference were determined and semen quality was evaluated. Regular cycles in pituitary and testicular activities corresponding to the period of the lighting regime resulted in Groups 2, 3, and 4, but not in Group 1. In general, the change from long days to short days induced increases in follicle-stimulating hormone (FSH), luteinizing hormone (LH), and testosterone levels, scrotal size and sperm numbers and a decrease in prolactin. The reverse occurred after subsequent exposure to long days. After 4 months of long days, testicular regression was complete, but when long-day exposure was reduced, less regression occurred. With continuous exposure to short days, FSH and testosterone remained above basal levels, prolactin levels were depressed, scrotal size remained near the maximum, and elevated numbers of motile sperm were sustained.     

What photoperiodic signal is provided by a continuous-release melatonin implant?

The purpose of this study was to evaluate whether the insertion of a continuous-release melatonin implant into ewes provides a short-day photoperiodic signal or acts as a functional pinealectomy (provides no specific photoperiodic signal but renders ewes incapable of responding to changes in photoperiod). Ewes primed with 60 long days (18L:6D) during the spring were moved to intermediate day length (13L:11D) for 66 days and then given one of five treatments: 1) short-day control, second drop in photoperiod to 8L:16D; 2) intermediate-photoperiod control, kept on 13L:11D; 3) pinealectomy and kept on 13L:11D; 4) melatonin implant and kept on 13L:11D; 5) melatonin implant and moved to 8L:16D. Mean number of estrous cycles per group and total duration of reproductive activity were determined. Ewes in all groups began to exhibit estrous cycles after the initial reduction in photoperiod. The number of estrous cycles and duration of reproductive activity differed among groups. The number of estrous cycles and duration of reproductive activity was extended in ewes receiving the second drop in photoperiod compared to that of the intermediate-photoperiod controls. Pinealectomized ewes had a number of estrous cycles and duration of reproductive activity similar to those of ewes maintained on the intermediate photoperiod. Melatonin implants increased the number of estrous cycles and prolonged reproductive activity in ewes maintained on the intermediate photoperiod; melatonin implants did not prevent the extension of reproductive activity in ewes receiving the second photoperiodic drop to the short daylength.(ABSTRACT TRUNCATED AT 250 WORDS)     

Influence of melatonin on the testicular regression induced by subcutaneous testosterone pellets in male rats kept in long or short photoperiod

Daily afternoon injections of 25 micrograms melatonin for 12 weeks had no effect on testicular weights of male rats kept in long photoperiod (14L:10D); similarly, exposure of rats to short photoperiod (2L:22D) had no effect on gonadal weight. However, rats maintained in a long or short photoperiod and implanted every 2 weeks with a 15 mm Silastic pellet containing testosterone showed a significant reduction in testicular weight; this effect was more pronounced in rats exposed to a short photoperiod. Melatonin injections in testosterone-treated rats in a long photoperiod exacerbated the inhibitory effects of testosterone alone. Subcutaneous 2-weekly implants of a beeswax pellet containing 1 mg melatonin reversed the effects of the melatonin injections on relative testicular weights but not those due to short photoperiod exposure. Testosterone implants significantly reduced pituitary LH values in long and short photoperiod-exposed animals, more particularly in those exposed to short photoperiod. Melatonin injections alone or in combination with melatonin pellets did not further exaggerate the depression in pituitary LH due to testosterone alone in long photoperiod-exposed animals; similarly melatonin pellets did not reverse the depression in pituitary LH observed. No significant differences in plasma prolactin concentrations or in thyroxine concentrations or free thyroxine index were observed after any combination of treatments. We therefore suggest that the effects observed with short photoperiod may be due to melatonin.     

Role of the pituitary gland in the development of photorefractoriness and generation of long-term changes in prolactin secretion in rams

Hypothalamo-pituitary disconnected Soay rams were exposed to two photoperiodic treatments: 1) constant long days (16L:8D) for 48 wk after pretreatment under short days (LD group), and 2) constant short days (8L:16D) for 48 wk after pretreatment under long days (SD group). In the LD group, plasma prolactin (PRL) concentrations increased from 0 to 8 wk (maximum: 143.3 +/- 8.4 microg/l; 8.8 +/- 1. 2 wk), decreased from 9 to 34 wk (minimum: 15.6 +/- 1.6 microg/l; 34. 5 +/- 1.5 wk), and finally increased again under the constant conditions, with a similar cyclical pattern for all individuals. In the SD group, PRL concentrations showed an inverse pattern (minimum: 8.6 +/- 2.6 microg/l; 17.1 +/- 2.0 wk; maximum: 46.4 +/- 5.5 microg/l; 30.2 +/- 3.2 wk), with more variability. Plasma concentrations of FSH were basal in both groups. The duration of the daily nocturnal melatonin peak (measured at 10, 24, and 44 wk) remained close to 8 h under long days (high-fidelity melatonin signal) but decreased significantly (13.8 h to 9.3 h) under short days (low-fidelity melatonin signal). The results support the conclusion that the melatonin signal encoding photoperiod acts within the pituitary gland to induce both acute (inductive) and chronic (refractory) effects photoperiod on PRL secretion.     

Sexual maturation of ewes raised without ram exposure in a controlled lighting environment

Finn x Dorset ewe lambs (n = 70) born in the spring (March 28 to April 6) from two successive lambing seasons were evaluated for age at first ovulation in the absence of mature rams. Ewe lambs were born in a controlled, short light (8L:16D) photoperiod or in ambient light (13L:11D). At about 10 to 11 wk of age, ewe lambs were allocated to a short (8L:16D) or long (16L:8D) light environment. Plasma progesterone (P(4)) concentrations were measured as an index of first ovulation. First exposure of ewes to sexually mature rams was in November. Most ewe lambs (77%) ovulated before ram exposure. More lambs (P < 0.025) born in ambient light and raised in short light reached puberty with typical cycles of plasma progesterone compared to other treatments. Long days tended to retard the onset of puberty. Although pregnancy rate did not differ across light treatments, more ewes became pregnant from the ambient-light born and short-light raised treatment. Photoperiod is an important factor affecting the onset of sexual maturation and genesis of normal luteal progesterone secretion in the ewe lamb.     

The effects of a sudden decrease or increase in daylength on prolactin secretion in the ram

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.     

Profiles of luteinizing hormone, follicle-stimulating hormone, testosterone and prolactin in rams of diverse breeds: effects of contrasting short (8L:16D) and long (16L:8D) photoperiods

Mature rams of Polled Dorset, Finnish Landrace, Rambouillet and Suffolk breeding were maintained in a temperature-controlled environment and exposed to two consecutive cycles of short (8L:16D) followed by long (16L:8D) days. Serum hormone concentrations were determined in weekly samples and in 24-h profiles characterized at the end of each lighting schedule (i.e., 12, 24, 36 and 48 weeks). In all four breeds, the pituitary-testicular axis was more active during short days as compared with long days and the magnitudes of changes in serum luteinizing hormone (LH), follicle-stimulating hormone (FSH) and testosterone concentrations were greater for the two most seasonal breeds, Finnish Landrace and Suffolks. In comparison to other breeds, Finnish Landrace rams had significantly (P less than 0.05) higher mean LH levels, showed the greatest number of LH peaks/24 h, and had the highest mean testosterone levels at the end of both periods of short days, while Rambouillet rams had significantly (P less than 0.05) lower testosterone. Rambouillets also showed the smallest changes in pulsatile LH and testosterone secretion and displayed the least number of LH peaks/24 h following short days. Serum FSH levels were significantly (P less than 0.05) higher in Finnish Landrace and Suffolk rams than in Polled Dorsets and Rambouillets after 12 weeks of short days. Breed differences in serum LH, FSH and testosterone were not apparent following long days. Prolactin levels in Rambouillet rams were significantly (P less than 0.05) lower than in the other breeds following both periods of long days. These results indicate that breed differences exist in mature rams with regard to hormone secretory profiles. Breed differences in serum gonadotropin and testosterone are only apparent during short days when the hypothalamo-pituitary-testicular axis in rams is considered most active. Likewise, breed differences in prolactin are noticeable only during long days when secretion of this hormone is enhanced. Breed differences in LH, FSH and testosterone secretion in rams during short days might be related to seasonality of mating and/or fecundity of breed types.     

A comparison of the efficiency of melatonin treatments in advancing oestrus in ewes

Early oestrous cycles were induced in adult, maiden, 18-month-old Suffolk-cross ewes, maintained from birth in natural photoperiod by the following treatments applied from mid-June: subcutaneous implantation of melatonin (1 g) in Silastic packets, daily, oral, melatonin administration (3 mg/ewe) at 15:30 h, an artificial photoperiod of 8L:16D (lights on 07:30 h). Ovarian cycles began 5-10 weeks before those of control ewes maintained in a natural photoperiod. In contrast, the onset of ovarian cycles in ewes given s.c. implants of melatonin (1 g) in April, and a further group in May, was highly variable, and not significantly different from that of the control ewes. Plasma melatonin profiles in sheep with implants showed a night-time rise super-imposed on a constant level, which was itself within the physiological night-time range. Implant-derived melatonin declined with time but remained at or above physiological night-time levels for at least 3 1/2 months. These results indicate that melatonin implants in June, but not in April or May, advance onset of oestrus in the non-lactating, adult ewe. The effects of melatonin implants in June on onset of ovarian cycles were indistinguishable from those of melatonin feeding or artificial short photoperiod initiated at this time of year.     

Effects of photoperiod, melatonin implants and castration on molting and on plasma thyroxine, testosterone and prolactin levels in the European badger (Meles meles)

1. The seasonal molt, which lasts six months in the badger, begins in mid-July and ends at the beginning of winter. It occurs under natural long-day conditions, following the seasonal drop in plasma testosterone levels, concomitant with high levels of thyroxine and prolactin. 2. To examine the role of the different factors involved (day length, prolactin, thyroxine, testosterone), different groups of badgers, divided into subgroups of castrated or intact animals, were subjected to the influence of long days (20L: 4D), short days (4L:20D) or the effect of subcutaneous melatonin implants. 3. In all cases, castration resulted in a significantly earlier onset of molting 1-3 months, depending on the group, regardless of the experimental conditions (20L:4D, 4L:20D, melatonin). 4. However, molting started earliest in animals subjected to long days, irrespective of whether they were castrated or intact. 5. In the melatonin-implanted badgers, molting started either early (castrated animals), or late or not at all (intact animals). 6. Lastly, in castrated badgers subjected to experimental photoperiods (short days or long days) or melatonin implants, the period of molting was shortened from 6 months (intact outdoor animals) to 4 months. 7. The advance in shedding was always related to an early drop in testosterone (or an absence of testosterone in the castrated animals) and to a higher or earlier increase in thyroxine levels.(ABSTRACT TRUNCATED AT 250 WORDS)     

Endogenous circannual rhythms and photorefractoriness of testis activity, moult and prolactin concentrations in mink (Mustela vison).

Mink are seasonal photosensitive breeders; testis activity is triggered when days have less than 10 h light. Increasing and decreasing plasma concentrations of prolactin induce the spring and autumn moults. In a 5 year experiment, males were maintained under short days (8 h light:16 h dark) at 13 degrees C or long days (16 h light:8 h dark) at 21 degrees C, winter and summer conditions, respectively. Under winter and summer conditions, circannual cycles of prolactin secretion and moulting were observed at intervals of about 11 months. Recurrence of testis cycles was not evident. In a second experiment, males were maintained under an 8 h light:16 h dark cycle from the winter solstice or under 10 h light:14 h dark, 12 h light:12 h dark or 14 h light:10 h dark cycles from 10 February. Under 8 h light:16 h dark cycle, testis regression was slightly later than under natural conditions, indicating photorefractoriness. However, mink remained sensitive to light: the longer the photoperiod, the faster the testis regression. In a third experiment, males were transferred under 8 h light:16 h dark or 16 h light:8 h dark from 15 May (group 1), 12 June (group 2) or 4 July (group 3); males submitted to long days received melatonin capsules on the day of transfer. Increasing concentrations of follicle-stimulating hormone (FSH) and luteinizing hormone (LH) and testis volume were shown by half the males in group 2 and nearly all the males in group 3; the constant release of melatonin from implants was more efficient than short days; but in the three groups, prolactin concentrations decreased in the few days after short-day or melatonin treatment. Overall, the results demonstrate endogenous circannual rhythms of prolactin secretion, body weight and moulting. Although a refractory period to short days was observed, the annual cycle of testis activity totally relies on the annual changes in daylength.     

Effects of constant darkness and constant light on circadian organization and reproductive responses in the ram

The relationship between circadian rhythms in the blood plasma concentrations of melatonin and rhythms in locomotor activity was studied in adult male sheep (Soay rams) exposed to 16-week periods of short days (8 hr of light and 16 hr of darkness; LD 8:16) or long days (LD 16:8) followed by 16-week periods of constant darkness (dim red light; DD) or constant light (LL). Under both LD 8:16 and LD 16:8, there was a clearly defined 24-hr rhythm in plasma concentrations of melatonin, with high levels throughout the dark phase. Periodogram analysis revealed a 24-hr rhythm in locomotor activity under LD 8:16 and LD 16:8. The main bouts of activity occurred during the light phase. A change from LD 8:16 to LD 16:8 resulted in a decrease in the duration of elevated melatonin secretion (melatonin peak) and an increase in the duration of activity corresponding to the changes in the ratio of light to darkness. In all rams, a significant circadian rhythm of activity persisted over the first 2 weeks following transfer from an entraining photoperiod to DD, with a mean period of 23.77 hr. However, the activity rhythms subsequently became disorganized, as did the 24-hr melatonin rhythms. The introduction of a 1-hr light pulse every 24 hr (LD 1:23) for 2 weeks after 8 weeks under DD reinduced a rhythm in both melatonin secretion and activity: the end of the 1-hr light period acted as the dusk signal, producing a normal temporal association of the two rhythms. Under LL, the 24-hr melatonin rhythms were disrupted, though several rams still showed periods of elevated melatonin secretion. Significant activity rhythms were either absent or a weak component occurred with a period of 24 hr. The introduction of a 1-hr dark period every 24 hr for 2 weeks after 8 weeks under LL (LD 23:1) failed to induce or entrain rhythms in either of the parameters. The occurrence of 24-hr activity rhythm in some rams under LL may indicate nonphotoperiodic entrainment signals in our experimental facility. Reproductive responses to the changes in photoperiod were also monitored. After pretreatment with LD 8:16, the rams were sexually active; exposure to LD 16:8, DD, or LL resulted in a decline in all measures of reproductive function. The decline was slower under DD than LD 16:8 or LL.(ABSTRACT TRUNCATED AT 400 WORDS)     

LH and testosterone responses to GnRH in red deer (Cervus elaphus) stags kept in a manipulated photoperiod

Six red deer stags from age 4 months were kept in a light-proof room under an artificial photoperiod consisting of 5.5 cycles of alternate 2-month periods of 16 h light and 8 h dark (16L:8D) and 8L:16D. At 2 or 3 weekly intervals from 10 months of age through 4 cycles, the stags were anaesthetized with xylazine and challenged i.v. with 10 micrograms GnRH. Blood samples were withdrawn immediately before and 10 and 60 min after injection. LH and testosterone concentrations were measured in all samples by RIA. Antler status was recorded daily. Peak LH values on each sampling day occurred in the sample taken 10 min after GnRH stimulation while peak testosterone occurred in the sample taken at 60 min. There were 4 cycles of LH and testosterone secretion accompanied by 4 antler cycles in the stags. The highest LH responses were detected during short days (8L:16D), and the highest testosterone responses were detected around the time of the change from short to long days. The responses of both hormones were lowest at the end of periods of long days or the beginning of short days. The increased pituitary LH response to GnRH was evident 4 weeks after the change to short days which are stimulatory for gonadal development. Antler casting occurred at the end of long days and cleaning at the end of short days. It is considered that antler cycles were due to the ability of the stags to vary release of LH and testosterone in response to changes in the artificial photoperiod.     

Test of ML23 as an antagonist to the effects of melatonin in the ram

In Exp. 1, four groups of 8 yearling Soay rams were housed under long days (16L:8D) to induce reproductive quiescence and were treated daily for 12 weeks with: (I) vehicle (2 or 4 ml 50% ethanol/water), (II) ML23 (2 mg), (III) melatonin (2 mg) and (IV) melatonin and ML23 (2 mg of each). All treatments were given orally in the mid-light phase. In the rams receiving melatonin (Group III) there was an earlier increase in the plasma concentrations of FSH and testosterone and regrowth of the testes compared to the controls (time to maximum testicular diameter: 10.0 +/- 0.5 and 15.3 +/- 1.2 weeks). These differences were reversed after the end of the 12-week treatments when rapid testicular regression occurred in melatonin-treated rams but not in the controls. In the group receiving ML23 and melatonin (Group IV), there was early reactivation and regression of the reproductive axis as in the melatonin group (testis max. 9.9 +/- 0.7 and 10.0 +/- 0.5 weeks) while in the group receiving ML23 alone (Group II) there was a slower redevelopment and regression as in the controls (testis max. 15.7 +/- 1.1 and 15.3 +/- 1.2 weeks). The comparison between the 4 groups in the changes in the blood concentrations of prolactin, voluntary food intake and total body weight also indicated that the treatment with ML23 failed to modify the effect of melatonin (combined treatment vs melatonin) or the effect of the long day photoperiod (ML23 vs vehicle).(ABSTRACT TRUNCATED AT 250 WORDS)     

Endogenous circannual cycles of ovarian activity and changes in prolactin and melatonin secretion in wild and domestic female sheep maintained under a long-day photoperiod

The present study examines the ovulatory activity of wild and domesticated ewes subjected to either a constant photoperiod of long days (16L:8D) or natural changes in daily photoperiod for 16 mo. The aim was to determine whether an endogenous reproductive rhythm controls seasonal reproductive activity in these sheep, and how the photoperiod might affect this. The effects of long-day photoperiods on long-term changes in prolactin and melatonin secretion were also evaluated. The two species showed changes in reproductive activity under the constant photoperiod conditions, suggesting the existence of an endogenous rhythm of reproduction. This rhythm was differently expressed in the two types of ewe (P < 0.05), with the domestic animals exhibiting much greater sensitivity to the effects of long days. A circannual rhythm of plasma prolactin concentration was also seen in both species and under both photoperiod conditions, although in both species the amplitude was always lower in the long-day animals (P < 0.01). The duration of the nocturnal melatonin plasma concentrations reflected the duration of darkness in both species and treatments. The peak melatonin concentration did not differ between seasons either under natural or long-day photoperiods.     

Prolactin cycles in sheep under constant photoperiod: evidence that photorefractoriness develops within the pituitary gland independently of the prolactin output signal

The present study investigated photorefractoriness in the prolactin (PRL) axis in hypothalamopituitary-disconnected (HPD) sheep exposed to prolonged long days. In experiment 1, HPD Soay rams transferred from short (8L:16D) to long (16L:8D) days for 48 wk to induce a cycle of activation, decline (photorefractoriness), and reactivation in PRL secretion were treated chronically with bromocriptine (dopamine-receptor agonist) or vehicle from the onset of photorefractoriness. Bromocriptine (0.01-0.04 mg kg-1 day-1; 12-24 wk of long days) blocked PRL release and caused a rebound response after the treatment, but it had no effect on the long-term PRL cycle (posttreatment PRL minimum, mean +/- SEM, 35.3 +/- 0.6 and 37.0 +/- 0.4 wk for bromocriptine and control groups, respectively; not significant). In experiment 2, HPD rams were treated with sulpiride (dopamine-receptor antagonist) during photorefractoriness. Sulpiride (0.6 mg/kg twice daily; 22-30 wk of long days) induced a marginal increase in blood PRL concentrations, but again, it had no effect on the long-term PRL cycle (PRL minimum, 37.9 +/- 0.4 and 37.6 +/- 0.9 wk for sulpiride and control groups, respectively; not significant). The 24-h blood melatonin profile consistently reflected the long-day photoperiod throughout, and blood FSH concentrations were minimal, confirming the effectiveness of the HPD surgery. The results support the conclusion that photorefractoriness is regulated at the level of the pituitary gland independently of the PRL output signal.     

Effect of photoperiod and melatonin on testis development and regression in wild European rabbits (Oryctolagus cuniculus)

Testis size in male wild rabbits (Oryctolagus cuniculus) kept in an enclosure in Cambridgeshire, U.K. was maximal during May and June and minimal between October and December. Regression occurred after the summer solstice and recrudescence occurred after the winter solstice. Rabbits kept in long days for 25 wk showed no sign of spontaneous testis regression. Hence, testis regression during the summer is probably not due to development of refractoriness to long days. Testis regression occurred in rabbits transferred from long (16L:8D) to short (8L:16D) days. Within 8 wk of the transfer spontaneous regrowth of the testes occurred and the rabbits moulted, and after 16 wk the testes had recovered to their size before the transfer. Subcutaneous implants of melatonin given to rabbits in long days mimicked the effect of a transfer to short days by causing testis regression followed by recrudescence. Moult occurred in rabbits immediately after short day- or melatonin-induced testis regression. The study demonstrates that seasonal reproduction in male wild rabbits in Britain is largely controlled by changes in photoperiod and that this is probably mediated via the pineal gland.     

Pineal and ovarian response to 22- and 24-h days in the ewe

Melatonin secretion in ewes was entrained by 22-h light-dark cycles whether of long (16L:6D) or short (6L:16D) photoperiod. In photoperiods of 6L:16D, a phase-delay of melatonin secretion was evident, leading to a dark-phase duration shorter than that found in 8L:16D. Early onset of estrus was induced in anestrous ewes kept in 8L:16D, but not 6L:16D, from 22 July compared to controls in natural light. In photoperiods of 16L:6D, the melatonin profile corresponded precisely to the dark phase. Early offset of estrus was induced in estrous ewes kept in both 18L:6D and 16L:6D from 18 December compared to controls in natural light. Thus, when the duration of melatonin secretion was appropriate to the long photoperiod (16L:6D), but with a constantly changing phase position, a long-day reproductive response was found. Activity-rest cycles were not entrained by 16L:6D; thus the synchronization of melatonin and activity-rest cycles does not appear to be essential for the induction of a long-day reproductive response. These results support the hypothesis that the duration, not the circadian-phase position, of melatonin is critical to the induction of photoperiodic effects.     

Geographic differences for delay of sexual maturation in Peromyscus leucopus: effects of photoperiod, pinealectomy, and melatonin

Effects of short-day photoperiod, pinealectomy, and melatonin on sexual maturation were tested in Peromyscus leucopus from either Connecticut (CT) or Georgia (GA). Laboratory reared-stocks from CT and GA were exposed to short daylength (photoperiod) from birth or 25 days of age. At 12 wk of age, delay in sexual maturation was indicated in most CT mice by decreased testis length, combined testes weight, and seminal vesicle weight. Conversely, GA animals did not delay sexual maturation when exposed to short-day photoperiod from either birth or 25 days of age. These results indicate that responses to short daylengths differ for juvenile CT and GA populations. In a second experiment, pinealectomized or sham-operated CT males were exposed to short-day (9L:15D) or long-day (16L:8D) photoperiod from birth. Pinealectomy blocked the effect of short daylength on reproduction. Therefore, the pineal must be involved in the delay of sexual maturation observed for short-day CT mice. The effects of melatonin, a pineal gland hormone, were tested with chronic s.c. implants or daily injections. In CT mice given either melatonin implants or afternoon injections, sexual maturation was delayed. GA mice were insensitive to all melatonin treatments. Further, no differences in circadian organization (phase angle, duration of activity, period under constant dark) between GA and CT animals were apparent. Collectively, these studies indicate that melatonin is involved in the mechanism responsible for delay of sexual maturation in CT mice. Short-day insensitivity of GA Peromyscus leucopus probably results from a deficiency in the melatonin effector pathway and is not due to a disruption of circadian organization.     

Effects of adrenalectomy on photoperiod-induced changes in release of luteinizing hormone and prolactin in ovariectomized ewes

Finnish Landrace x Southdown ewes were ovariectomized (OVX) and subjected to daily photoperiods of 16L:8D (Group I) or 8L:16D (Group II) for 84 days. Ewes were then either adrenalectomized (ADX) (N = 5 for Group I; N = 4 for Group II) or sham ADX (N = 6 for Groups I + II). After surgery, ewes in Group I were subjected to 8L:16D for 91 days and 16L:8D for 91 days whereas ewes in Group II were exposed to 16L:8D for 91 days and 8L:16D for 91 days. Oestradiol implants were inserted into all ewes on Day 148. Sequential blood samples were taken at 28, 56, 91, 119, 147 and 168 days after surgery to determine secretory profiles of LH and prolactin. Photoperiod did not influence LH release in Group I in the absence of oestradiol. Although photoperiod influenced frequency and amplitude of LH pulses in Group II before oestradiol treatment, adrenalectomy did not prevent these changes in patterns of LH release. However, in Group II the increase in LH pulse amplitude during exposure to long days was greater (P less than 0.01) in adrenalectomized ewes than in sham-operated ewes. Mean concentrations of LH increased in ADX ewes on Days 91 (P = 0.07) and 119 (P less than 0.05). Adrenalectomy failed to influence photoperiod-induced changes in mean concentrations of LH, amplitude of LH pulses and frequency of LH pulses in the presence of oestradiol. Concentrations of prolactin were influenced by photoperiod. In Groups I and II concentrations of prolactin increased (P less than 0.01) after adrenalectomy, but the magnitude of this effect decreased over time.(ABSTRACT TRUNCATED AT 250 WORDS)