Stimulation of reproductive activity in anovulatory Alpine goats exposed to bucks treated only with artificially long days

Two experiments were conducted in a subtropical latitude to determine the response of Alpine male goats to a treatment with artificially long days (experiment 1), and the response of anovulatory lactating Alpine does exposed to males treated only with artificially long days (experiment 2). In experiment 1, one group of males was kept under natural photoperiod (n = 4) while another was exposed to 2.5 months of artificially long days (16 h of light/day) from 1 December (n = 4). Plasma testosterone concentrations were determined weekly. Intensity of odor of males was determined every 2 weeks. Sexual behavior of bucks was observed during 3 days about 90 days after the end of the long day treatment. A treatment-by-time interaction was detected for testosterone secretion (P < 0.001). In control males, low plasma concentrations of testosterone were observed from March to June. In contrast, in long-day treated males, high levels of testosterone were observed from March to June (P < 0.05). A treatment-by-time interaction was detected for the intensity of male odor (P < 0.01). The male odor was stronger in long-day treated bucks than in untreated ones from March to June (P < 0.05). The number of ano-genital sniffing, nudging and flehmen was greater in long-day treated males than in untreated ones when exposed to anestrous does (P < 0.05). In experiment 2, one group of males was left under natural photoperiod variations (n = 5) and the other (n = 5) was submitted to the same photoperiodic treatment described in experiment 1. On 3 May, three untreated and three long-day treated males were put in contact with anestrous Alpine does left under natural photoperiod. Fertility was higher in does exposed to light-treated males (36/45, 80%) than those in contact with untreated ones (3/45, 7%; P < 0.05). Prolificacy was similar (P > 0.05) in does exposed to treated (1.8 ± 0.1) and untreated males (1.7 ± 0.3). These results indicate that the sexual activity of Alpine male goats raised in subtropical latitudes can be induced using only artificially long days and that such males are effective in stimulating reproductive activity in anovulatory females in late spring.     

Effects of photoperiod on pituitary gonadotropin levels in masu salmon

We examined the effects of photoperiod on pituitary levels of two types of gonadotropin (GTH), GTH I and GTH II, in masu salmon Oncorhynchus masou to study their mechanism of synthesis. In Experiment 1, the effects of long or short photoperiod combined with castration were examined using 8-month-old precocious males. Castration was carried out in early August and then the fish were reared under a short (8L16D) or long (16L8D) photoperiod for 60 days. In Experiment 2, the effects of photoperiod combined with testosterone treatment were examined using 12-month-old immature females. Silastic tubes containing testosterone (500 microg /fish) or vehicle were implanted intra-peritoneally in early October. Fish were reared under 16L8D for 60 days, and then half of the fish were transferred to 8L16D, while the remaining fish were kept under 16L8D until Day 90. In Experiment 1, GTH I contents were higher under 16L8D than under 8L16D in the castrated group on Day 30. Moreover, GTH I contents were higher in the castrated group than the control group under 16L8D on Day 30. GTH II contents increased with testicular maturation in the control groups, whereas they remained at low levels in the castrated groups regardless of photoperiodic treatment. In Experiment 2, GTH I contents did not change remarkably in all the groups, while GTH II contents were remarkably increased by testosterone treatment regardless of photoperiodic treatment. These results indicate that the synthesis of GTH I and GTH II are differently regulated by photoperiod and testosterone in masu salmon.     

The response of luteinizing hormone secretion to photoperiod is modified by the level of nutrition in female Mediterranean goats

This paper reports the influence of nutrition on the photoperiodic control of luteinizing hormone (LH) secretion in female Mediterranean goats (i.e., goats from the Mediterranean area in general). Ovariectomized, oestradiol-treated goats were subjected to two consecutive intervals of 3 months of long days followed by 3 months of short days (group LDSD, N=20), or vice versa (group SDLD, N=20). The LDSD and SDLD does were also randomly assigned to one of two nutrition groups that received either 1.1 (H group, N=10) or 0.7 (L group, N=10) times their maintenance requirements. Live weight and body condition score were determined weekly and LH concentrations twice per week. To establish the pulsatility of secretion of LH, three periods of intensive sampling were undertaken. Melatonin was determined after a period of 45 short or long days. All photoperiod/nutrition groups showed large variations in LH concentrations according to photoperiod, with nutrition having a significant effect (P<0.001). The mean time between the shift from long to short days and the stimulation of LH secretion, and between the shift from short to long days and the inhibition of LH secretion, was different in each nutrition group (at least P<0.05). No differences were seen in the frequency of LH pulses between the nutrition groups, but differences between sampling periods were observed (P<0.001). Melatonin secretion was not affected by food supply. These results confirm: (1) that Mediterranean female goats are sensitive to photoperiod, (2) that this environmental cue may control the timing of pituitary activity under natural conditions, and (3) suggest that nutrition plays an important role in the effect of photoperiod on LH secretion.     

Continuous light after a long-day treatment is equivalent to melatonin implants to stimulate testosterone secretion in Alpine male goats

In rams, artificial long days followed by continuous light stimulate testosterone secretion during the non-breeding season. The objective of this study was to determine whether artificial long days followed by continuous light could stimulate testosterone secretion in Alpine bucks as well as in those exposed to long days followed by a melatonin treatment. All bucks were kept in shaded open pens. Control males were exposed to natural photoperiod conditions (n=5). Males of the two experimental groups were exposed to 2.5 months of long days from 1 December (n=5 each). On 16 February, one group of males was exposed to 24 h of light per day until 30 June; the other group was exposed to natural variations of photoperiod and received two s.c. melatonin implants. Testicular weight was determined every 2 weeks, and the plasma testosterone concentrations once a week. In the control and the two photoperiodic-treated groups, a treatment×time interaction was detected for testicular weight and plasma testosterone concentrations (P<0.001). In control bucks, testicular weight increased from January and peaked in June, whereas in both photoperiodic-treated groups, this variable increased from January, but peaked in April, when the values were higher than in controls (P<0.05). In the control group, plasma testosterone concentrations remained low from January to June, whereas in both photoperiodic-treated groups, this variable remained low from January to March; thereafter, these levels increased in both photoperiodic-treated groups, and were higher than controls in April and May (P<0.05). We conclude that continuous light after a long-day treatment stimulate testosterone secretion in Alpine male goats during the non-breeding season as well as the long days followed by a melatonin treatment. Therefore, continuous light could replace the implants of melatonin.     

Reproductive photoresponsiveness in Aethomys ineptus and A. namaquensis (Rodentia: Muridae) from southern Africa

Male Tete veld rats Aethomys ineptus and Namaqua rock mice A. namaquensis were independently subjected to a long photoperiod of 16L:8D and a short photoperiod of 8L:16D for 3 months in order to assess the effect of photoperiod on testicular function. Testicular parameters examined included testicular mass relative to body mass, testicular volume, seminiferous tubule diameters and changes in circulating plasma testosterone concentration. On a long day (LD) length, both species exhibited significantly higher testicular mass relative to body mass and testicular volume, and larger seminiferous tubule diameters. Circulating plasma testosterone concentrations in A. ineptus were not significantly different between the two photoperiodic treatments, but A. namaquensis showed significantly higher testosterone concentration in males on a LD length than on a short day (SD) length. This suggests that both A. ineptus and A. namaquensis are photoperiodically responsive to LDs.     

Photoperiodic regulation of annual testicular events in the Indian major carp Catla catla

The importance of photoperiods in the regulation of annual testicular events in the carp Catla catla was evaluated by subjecting them to either long (16 h light : 8 h dark) or short (8 h light : 16 h dark) photoperiods for 30 days during the preparatory, prespawning, spawning and postspawning phases of an annual gonadal cycle. In each reproductive phase, testicular responsiveness to subjected photoperiods was determined by comparing the gonadal status in corresponding groups of control or natural photoperiodic fish. The values of testicular weight, gametogenic index, as well as testicular activity of two steroidogenic enzymes (Δ⁵3β‐, and 17β‐hydroxysteroid dehydrogenase), and the serum titre of testosterone were considered as the indices of functional status of the testis in the fish concerned. During the prespawning phase, exposure of fish to a daily long photoperiod schedule resulted in precocious maturation of testis, while retardation of testicular growth was noted under the influences of short photoperiod. However, none of the employed photo‐schedules could influence the gametogenic and steroidogenic functions of the testis in the remaining part of the gonadal cycle. Collectively, the present study provides evidence for the first time that in the case of a commercially important carp, Catla catla, artificial, long photoperiods may be used for advanced testicular maturation, while reductions in maturation‐associated growth and deterioration in flesh quality may be avoided by submitting the fish to shorter day lengths during the prespawning phase of the reproductive cycle.     

The role of nutrition in the regulation of luteinizing hormone secretion by the opioidergic, dopaminergic, and serotonergic systems in female Mediterranean goats

This study examined which neural mechanism (opioid, dopaminergic, or serotonergic system) is involved in the regulation of luteinizing hormone (LH) secretion, with and without nutritional modulation, at different times of the photoperiodic cycle. Goats were randomly distributed into two experimental groups that received either 1.1 (high group; n = 18) or 0.7 (low group; n = 18) times the nutritional maintenance requirements. The goats were exposed to alternations of 3 mo of long days and 3 mo of short days. Plasma LH concentrations were measured twice a week. The effects of intravenous injections of naloxone (endogenous opioid receptor antagonist), pimozide (dopaminergic(2) receptor antagonist), and cyproheptadine (serotonin 5-hydroxytryptamine(2) receptor antagonist) on LH secretion were assessed during challenges in three different photoperiodic situations: the onset of LH stimulation by short days (OnsetSD), the onset of LH inhibition by long days (OnsetLD), and during the LH inhibition by long days (LateLD). The role of the different neural systems was clearly modified by the level of nutrition. In the low-nutrition group, only naloxone increased LH concentrations during onsetLD (P < 0.05). However, in the high-nutrition group, naloxone increased the concentration and pulsatility of LH (P < 0.05) in onsetSD and onsetLD. Pimozide increased LH concentration and pulsatility (P < 0.05) in onsetLD and LH concentration in lateLD (P < 0.001). Finally, cyproheptadine significantly increased LH concentration at all three times (P < 0.001). These results provide evidence that all three systems are involved in the inhibition of LH release in onsetLD, and that the opioid and serotonin mechanisms are involved during the onsetSD that were enhanced by a high plane of nutrition.     

Differential response of type 2 deiodinase gene expression to photoperiod between photoperiodic Fischer 344 and nonphotoperiodic Wistar rats

The molecular basis of seasonal or nonseasonal breeding remains unknown. Although laboratory rats are generally regarded as photoperiod-insensitive species, the testicular weight of the Fischer 344 (F344) strain responds to photoperiod. Recently, it was clarified that photoperiodic regulation of type 2 iodothyronine deiodinase (Dio2) in the mediobasal hypothalamus (MBH) is critical in photoperiodic gonadal regulation. Strain-dependent differences in photoperiod sensitivity may now provide the opportunity to address the regulatory mechanism of seasonality by studying Dio2 expression. Therefore, in the present study, we examined the effect of photoperiod on Dio2 expression in photoperiod-sensitive F344 and photoperiod-insensitive Wistar rats. A statistically significant difference was observed between short and long days in terms of testicular weight and Dio2 expression in the F344 strain, while no difference was observed in the Wistar strain. These results suggest that differential responses of the Dio2 gene to photoperiod may determine the strain-dependent differences in photoperiod sensitivity in laboratory rats.     

Refractoriness to short and long days determines the end and onset of the breeding season in subtropical goats

The objective was to determine whether refractoriness to short and long days were involved in the end and onset of the breeding season, respectively, in goats adapted to subtropical latitudes. Ovariectomized does given a subcutaneous implant constantly releasing estradiol-l7 β (OVX+E) were used in two experiments. Plasma LH concentrations were determined twice weekly. In Experiment 1, the control group remained in an open-shed pen (natural day length and ambient temperature). Two experimental groups were placed in light-proof buildings (with natural temperature variations). One group was exposed to natural simulated increasing days (winter to spring), whereas the other was exposed to a winter solstice photoperiod (10 h of light) from December 21 to April 28. In Experiment 2, the control group remained under natural day length and ambient temperature. One experimental group was exposed to natural simulated decreasing days (summer to autumn), whereas the other group was exposed to a summer solstice photoperiod (14 h of light) from June 21 to October 20. In Experiment 1, the breeding season was not prolonged in does maintained in the winter solstice day length. Mean dates of decrease in LH secretion (end of the breeding season) did not differ significantly between does exposed to natural (February 3 ± 5 d) or natural simulated photoperiod (January 26 ± 14 d) and those exposed to constant short days of winter solstice (February 4 ± 10 d). In Experiment 2, the onset of the breeding season was not delayed in does maintained in the summer solstice day length. Mean dates of increase in LH secretion (onset of the breeding season) did not differ significantly between does exposed to natural (September 7 ± 8 d) or natural simulated photoperiod (September 18 ± 10 d) and those exposed to constant long days photoperiod of summer solstice (September 24 ± 4 d). In goats adapted to a subtropical environment, we concluded that: 1) the end of breeding season was due to refractoriness to short days, and not the inhibitory effect of increasing day length; and 2) the onset of the breeding season was due to refractoriness to long days, and not a stimulatory effect of decreasing day length.     

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.     

Artificial long days in addition to exogenous melatonin and daily contact with bucks stimulate the ovarian and oestrous activity in Mediterranean goat females

One experiment was conducted to determine whether the treatment with artificial long days and exogenous melatonin can induce reproductive activity during spring (seasonal anoestrus) in Mediterranean goats that are in daily contact with bucks and whether this treatment causes a variation in the reactivation of the reproductive activity in the normal breeding season. The experiment started on 4 November 2005 and finished on 27 October 2006. Thirty-four adult and barren does were used, distributed into two groups balanced according to their live weight (LW) and body condition score (BCS). Seventeen females were exposed to long days (16 h of light/day) from 14 November 2005 to 20 February 2006. On 20 February, they received one s.c. melatonin implant (LD-M group) and were exposed to natural photoperiodic changes in an open shed. The other females during the experiment were placed in an open shed under natural photoperiod and remained as the control group (C group). The C and LD-M groups were keeping in contact with males during the whole experiment. During the experiment, the LW, BCS and plasma progesterone concentrations were measured weekly, oestrous activity was tested daily using entire aproned bucks, and ovulation rate was evaluated by laparoscopy 7 days after positive identification of the oestrus. A clear treatment-time interaction was observed for plasma progesterone concentrations (P < 0.001), with a period of high progesterone concentrations during the natural seasonal anoestrus in the LD-M group. Although 94.1% of females in the LD-M group presented ovarian activity during this period, no female in group C did. Resumption of ovarian activity in the subsequent natural breeding season was 2 weeks later in the LD-M group in comparison with group C (P < 0.05). We can conclude that in Mediterranean goat breeding systems, when females are in daily contact with bucks, the treatment with 3 months of long days and melatonin implant at the end of the light photoperiodic treatment can induce ovarian and oestrous activity during the seasonal anoestrus. Finally, this treatment causes a short delay in the subsequent reactivation of ovarian activity in the natural breeding season.     

Reciprocal interaction between seasonal testis and thyroid activity in Zembra Island wild rabbits (Oryctolagus cuniculus): effects of castration, thyroidectomy, temperature, and photoperiod

The reproductive cycle of wild rabbits (Oryctolagus cuniculus) living in Zembra Island (North Tunisia) is dependent on an external factor, the photoperiod: the gonads are inhibited by long days and stimulated by short days or melatonin implants. Here we studied the role of an internal factor, thyroid hormones and the possible thyroid-gonadal interrelationships, in animals captured on Zembra Island and maintained in natural conditions of photoperiod and temperature. We determined the seasonal profile of the thyroid and testis cycles and investigated the effects of castration and thyroidectomy on the seasonal testosterone and thyroxine cycles. Plasma thyroxine and testosterone levels followed similar, parallel seasonal patterns, with a peak in autumn (October) and low values from January to August. In thyroidectomized animals, plasma testosterone levels, although significantly higher than those in controls (P < 0.001), remained low throughout the 13 mo of the experiment, and no testicular reactivation was observed in the fall. In castrated animals, despite the increase in thyroxine concentration in the 3 mo following castration (P < 0.01), plasma thyroxine levels remained low during the 2 yr of the study. We then investigated the combined effects of long days (16L:8D) and moderately high temperature (25 degrees C) on these two endocrine axes. In constant gonado-inhibiting conditions (16L:8D), whether the temperature was kept constantly high or allowed to fluctuate naturally, no reactivation of the thyroid and testicular axes was observed in the fall. In control animals, the peaks of testosterone and thyroxine concentrations observed in September were larger (P < 0.001) than those in animals subjected to the same natural photoperiod conditions but with constantly high temperature. The lower level of autumnal testis stimulation (P < 0.001) in animals maintained in conditions of constant high temperature (25 degrees C) may be attributed to the low thyroxine levels induced by high temperature. These results clearly confirm that the thyroid and testicular cycles display similar seasonal variations and show that the thyroid and gonadal axes are strictly interdependent. This study provides the first demonstration, for a given species, that the seasonal reactivation of gonad activity is controlled by the thyroid, and thyroid activity is controlled by the gonads.     

Changes in the testicular binding of luteinizing hormone and plasma testosterone concentrations in the Djungarian hamster subjected to different photoperiods and temperatures and effects of long-term testosterone treatment on the binding

The effects of artificial photoperiod, temperature, and long-term testosterone treatment on testicular luteinizing hormone (LH) binding were studied in adult male Djungarian hamsters. In hamsters transferred to long-day (LD; 16 hr light, 8 hr dark) photoperiod 8 weeks after adaptation in short-day (SD; 8 hr light, 16 hr dark) photoperiod of 25 degrees C, testicular growth was associated with an increase in the total LH binding per two testes and a decrease in LH binding per unit testicular weight. Plasma testosterone levels reached a peak 47 days after transfer to LD and tended to decrease thereafter, while the testes continued growing. In contrast, when hamsters reared under LD conditions at 25 degrees C for 12 weeks were transferred to SD, testicular regression was associated with a decrease in plasma testosterone and the total LH binding per two testes and an increase in LH binding per unit testicular weight. A significant decrease in LH binding per unit weight compared to SD controls was observed in those hamsters exposed to SD with continuous testosterone treatment. The testosterone treatment tended to induce decrease in the total LH binding. Scatchard plot analyses of the binding suggested that changes in LH binding were due to changes in the number of binding sites. When sexually mature male hamsters were subjected for 8 weeks to two different ambient temperatures (7 degrees C and 25 degrees C) and photoperiods (LD and SD), the difference between the two temperature groups was statistically not significant regarding the weights of testes, epididymides, and prostates; plasma testosterone levels; and LH binding in either LD or SD group. These results suggest that photoperiod is a more important environmental factor than temperature for the regulation of testicular activity and LH receptors and that testosterone reduces the number of LH receptors per unit testicular weight in adult male Djungarian hamsters.     

Effects of melatonin feeding on smoltification in masu salmon (Oncorhynchus masou)

Influences of photoperiod on plasma melatonin profiles and effects of melatonin administration on long-day-induced smoltification in masu salmon (Oncorhynchus masou) were investigated in order to reveal the roles of melatonin in the regulation of smoltification in salmonids. Under light-dark (LD) cycles, plasma melatonin levels exhibited daily variation, with higher values during the dark phase than during the light phase. The duration of nocturnal elevation under short photoperiod (LD 8:16) was longer than that under long photoperiod (LD 16:8). Melatonin feeding (0.01, 0.1 and 1 mg/kg body weight) elevated plasma levels of melatonin in a dose-dependent manner for at least 7 h but not for 24 h. When masu salmon reared under short photoperiod were exposed to long photoperiod (LD 16:8) and fed melatonin (1 mg/kg body weight) 7 hours before the onset of darkness, a significantly smaller proportion of smolts appeared in the melatonin-fed group after 32 days than in the control group. However, after 59 days of the treatment, there was no difference in the proportion of smolts between the control and melatonin-treated groups. Thus, melatonin feeding mimicked the effects of short photoperiod, which delays but does not completely suppress smoltification. These results indicate that the day length is transduced into changes in the duration of nocturnal elevation in plasma melatonin levels, and that artificial modification of the plasma melatonin pattern possibly delays the physiological processes of smoltification induced by long-day photoperiodic treatment.     

Circadian participation in the photoregulation of testis activity and prolactin secretion in the mink, a short-day breeder

It has been demonstrated that an endogenous mechanism is involved in photoperiodic time measurement in the mink, a short-day-breeding mannal. A study of testicular activity (testicular volume, plasma testosterone concentration) and plasma prolactin level was carried out in sexually resting minks (the experiment began in November). Groups of minks were kept in the natural photoperiod or subjected to different resonance light-dark (LD) cycles (LD 4:8, LD 4:20, LD 4:32, LD 4:44); an additional group of animals was reared in an ahemeral photoperiod (LD 4:16). A rapid increase of testicular activity was observed in control animals or those kept in LD 4:20 (T 24) and LD 4:44 (T 48). In the other groups of animals, those kept in LD 4:8 (T 12), LD 4:32 (T 36), and LD 4:16 (T 20), testicular function remained at rest. Prolactin secretion was, in contrast, stimulated in the groups kept in LD 4:8 (T 12). LD 4:32 (T 36), and LD 4:16 (T 20), and remained low in the groups kept in LD 4:20 (T 24) and LD 4:44 (T 48). These results show that the effects of the different photoperiodic regimens do not depend on the duration of the photophase, but rather on the period of the LD cycles. The LD cycles that allow an increase of testicular function are those that are inhibitory to reproduction in birds and long-day-breeding mammals. To explain these results, it is suggested that in the mink exposure to light during the circadian photosensitive phase induces inhibition of testicular activity and stimulation of prolactin secretion. To explain the opposite effects of a single photoperiod on testicular function and secretion of prolactin, the hypothesis has been advanced that, in the mink, long days might simultaneously inhibit hypothalamic luteinizing-hormone-releasing hormone (LH-RH) activity and prolactin-inhibiting factor (PIF) activity.     

Abolition of the seasonal release of luteinizing hormone and testosterone in Alpine male goats (Capra hircus) by short photoperiodic cycles.

This study was performed to determine whether rapid alternation between long and short days abolished seasonal variations in the activity of the hypothalamo-pituitary-testis axis observed normally in Alpine and Saanen male goats during the year. Three groups of 6 males were used: group 1 remained in open sheds under the natural annual change in daylength from 16 h of light (long day) to 8 h of light (short day). Group 2 was exposed to 1 month of long days alternated with 1 month of short days; and group 3 to 2 months of long days alternated with 2 months of short days. In group 1, blood samples were taken in December, February and June; in groups 2 and 3, samples were obtained once during short and long days for the melatonin assay. For luteinizing hormone and testosterone determinations monthly samples from group 1 were obtained from September to August while, in groups 2 and 3, blood samples were taken on 4 occasions during long and short days. Weekly blood samples were taken from all groups during the whole of the experiment to measure prolactin and testosterone concentrations. Melatonin profiles indicated that secretion by the pineal gland of male goats from the treated groups adapted to rapid changes in daylength: duration of nocturnal secretion was close to that of the dark period. Treated goats were also able to transduce this signal adequately and always responded to long days by increasing their prolactin concentration (mean +/- s.e.m.; group 2: 62.4 +/- 6.8 ng/ml; group 3: 102.3 +/- 15.7 ng/ml) and to short days with a decrease in prolactin concentrations (35.0 +/- 3.6 and 46.1 +/- 9.5 ng/ml, respectively). In the treated groups, luteinizing hormone pulse frequency varied with day length. In group 2, it was higher in long days (1.1 +/- 0.3 pulses in 8 hours) than in short days (0.7 +/- 0.3) while, in group 3, this frequency was higher in short days (1.9 +/- 0.3) than in long days (0.5 +/- 0.2). Testosterone secretion also varied with daylength; in group 2, the testosterone concentrations were maximum during long days (5.8 +/- 1.4 ng/ml) while in group 3 the maximum testosterone concentrations occurred during short days (6.4 +/- 1.2 ng/ml). These results lead to the conclusion that rapid alternation of long and short days either attenuated (group 3) or prevented (group 2) seasonal changes in the activity of the hypothalamo-pituitary axis.(ABSTRACT TRUNCATED AT 400 WORDS)     

Circadian and photoperiodic time measurement in male Syrian hamsters following lesions of the melatonin-binding sites of the paraventricular thalamus.

Autoradiographic studies using [125I]iodomelatonin in several species, including the Syrian hamster, have revealed that the rostral region of the anterior paraventricular nucleus of the thalamus (aPVT) contains a very high density of binding sites for melatonin. In two studies, small or large bilateral electrolytic lesions of the aPVT were made in adult male hamsters maintained on long days (LD 16:8). The hamsters were then transferred to short days (LD 8:16) to test whether testicular regression could occur in response to a decrease in photoperiod. Serum prolactin concentrations were measured as a second photoperiodic response. All unoperated control hamsters showed the typical short-day photoperiodic response: A decrease in serum luteinizing hormone (LH) and prolactin concentrations and testicular regression all occurred within 6 weeks in short days, followed by the development of scotorefractoriness. Lesions of the aPVT did not significantly affect the rate or the degree of the short-day-induced decline in serum levels of LH or prolactin, nor the pattern of testicular regression and the subsequent expression of refractoriness. To enable us to determine whether the aPVT might be involved in the entrainment or the expression of circadian rhythms, locomotor activity was monitored continuously in lesioned and control groups in Experiment 2, prior to and following the switch to short days. The reduction in photoperiod (involving an 8-hr advance in the time of lights-off and an 8-hr extension of the dark phase) caused a decompression of the nocturnal activity bout of control animals, so that after 2 weeks in short days, activity onset had also advanced to regain its phase relationship to the timing of lights-off. A similar pattern of reentrainment was observed in lesioned animals, and no differences were observed between treatment groups in the rate of entrainment and decompression. In addition, both intact controls and animals bearing large bilateral lesions of the aPVT exhibited robust free-running circadian rhythms of locomotor activity when held under constant dim red light. In summary, the integrity of the aPVT is not necessary for the seasonal response of the reproductive axis and prolactin secretion to photoperiod, nor for photic entrainment of activity rhythms, in the Syrian hamster.     

Effect of constant-release implants of melatonin on seasonal cycles in reproduction, prolactin secretion and moulting in rams

Seasonal cycles in the size of the testes, blood plasma concentration of testosterone, FSH and prolactin, intensity of the sexual skin flush, timing of rutting behaviour and moulting of the body coat were recorded in Soay rams after s.c. implantation of melatonin contained in a Silastic envelope which increased the circulating blood levels of melatonin to 200-600 pg/ml for many months. Two groups of 8 adult rams were held under alternating periods of short days (8L:16D) and long days (16L:8D) to drive the seasonal cycles and the treatments with melatonin were initiated during the long or short days, and one group of 8 ram lambs was kept out of doors and given implants during the long days of summer (4 melatonin-implanted and 4 control (empty implants) rams per group). The treatments demonstrated that melatonin implants during exposure to long days resulted in a rapid 'switch on' of reproductive redevelopment similar to that produced by exposure to short days melatonin implants prevented the rams from showing the normal responses to changes in the prevailing photoperiod rendering them nonphotoperiodic; and long-term cyclic changes in testicular activity, prolactin secretion and other characteristics occurred in the melatonin-implanted rams; the pattern was similar to that previously observed in rams exposed to prolonged periods of short days. The overall results are consistent with the view that melatonin is the physiological hormone that relays the effects of changing photoperiod on reproduction and other seasonal features, and that continuous exogenous melatonin from an implant interferes with the normal 'signal' and produces an over-riding short-day response.     

Male reproductive condition is the limiting factor of efficiency in the male effect during seasonal anestrus in female goats

Two experiments were conducted to determine whether the failure of males to induce sexual activity in goats during seasonal anestrus is due to unresponsiveness of females to male stimulus or insufficient stimulation from males. In the first study, one group of males (sexually inactive, SI; n = 4) was kept under natural photoperiod while the other (sexually active, SA; n = 4) was subjected to 2.5 mo of long days (16L:8D) and received 2 s.c. implants of melatonin. Two mo later, 2 different flocks of anovulatory goats previously separated from bucks were exposed to either SI (n = 34) or SA (n = 40) bucks. Progesterone assays and estrous behavior were used to determine ovarian and behavioral responses of the females to teasing. Of the goats exposed to SI males, only 2 ovulated, and none showed estrous behavior during the 35 days of the study. In contrast, all females (40 of 40) in contact with SA males ovulated and showed at least one estrous behavior during the first 11 days following male introduction (P < 0.001). Overall, 38 of 40 females stimulated with SA bucks were diagnosed pregnant at Day 35, according to progesterone assay (versus 0 in SI-treated group: P < 0.001). To control for a possible difference of responsiveness between flocks, the experiment was repeated 1 yr later using a single flock of goats divided into 2 groups. Again, over the first 14 days, 1 of 33 goats showed estrous behavior in the SI-treated group versus 27 of 33 in the SA-treated group (P < 0.001). Therefore, treating bucks with long days and melatonin increased their teasing capacity to induce sexual activity in females during anestrus. These results indicate that the absence of response to teasing at this time of the year is not due to female unresponsiveness, but to insufficient stimulation from the male.     

Importance of retinal photoreceptors to the photoperiodic control of seasonal breeding in the ewe

Two experiments were performed to determine whether the eyes are necessary for photoperiodic control of reproduction in ewes. In the first, intact and estradiol-treated ovariectomized (OVX + E) ewes were housed in each of 2 photoperiod-controlled rooms with a vasectomized ram and subjected to 90-day alternations between long and short days. Prior to blinding, long days initiated anestrus in intact ewes and a suppression of serum luteinizing hormone (LH) levels in OVX + E ewes; short days caused onset of estrous cycles and an increase in LH levels in the intact and OVX + E ewes, respectively. After 1.5 years of such photoperiodic control, all ewes were blinded by bilateral orbital enucleation. Photoperiodic control was lost following blinding, but circannual alternations between cyclicity and anestrus or high and low LH levels, were maintained in most ewes for the remaining 2.5 years of the study. In one group of OVX + E ewes, serum LH levels remained synchronized to the 90-day shifts in photoperiod for about 1 year after blinding. Once the sighted ram was removed from the room, however, the 90-day rhythm in LH disappeared and a circannual pattern of LH became evident, suggesting that blind ewes may receive photoperiodic information from a sighted ram. This possibility was supported by the results of the second experiment in which 12 additional OVX + E ewes were blinded and exposed to 90 long days and 90 short days in the absence of a sighted ram. In these ewes, serum LH levels were not controlled by the changes in photoperiod. These results are consistent with the following conclusions: 1) the eyes are necessary for perception of photoperiod in the ewe and 2) ewes have an endogenous circannual rhythm of reproduction and/or they can be controlled by other environmental signals in the absence of photoperiodic input. Further, the results lead to the hypothesis that blind ewes can receive photoperiodic information indirectly from a sighted ram.