Variation in the timing of the reproductive season among breeds of sheep in relation to differences in photoperiodic synchronization of an endogenous rhythm.

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.     

Termination of the breeding season in the Suffolk ewe: involvement of an endogenous rhythm of reproduction

Two studies were performed to determine if the transition into anestrus in the Suffolk ewe results from the lack of a decrease in photoperiod, a signal suggested to be important in maintaining the breeding season, or from an obligatory turn-off that reflects the expression of an endogenous rhythm of reproduction. Shortly after the autumnal equinox, three groups of ovariectomized ewes bearing s.c. Silastic implants of estradiol were placed in different lighting environments. A control group was exposed to normal variations in day length until the winter solstice and held on that day length thereafter. The two experimental groups experienced either a continuously decreasing day length or a continuously increasing duration of elevated melatonin levels during each 24-h period until mid-March. Reproductive activity, assessed by circulating levels of luteinizing hormone (LH), was marginally extended (2-3 wk) in the experimental groups. In a second experiment, two groups of ovary-intact ewes were exposed to photoperiodic treatments shortly after the winter solstice. A control group was held on the winter solstice day length (10L:14D) until the end of the study in mid-March. The experimental group experienced a 3-h photoperiodic reduction to 7L:17D. It remained to determine if that abrupt decrease could maintain reproductive induction. Again, the cessation of reproductive induction was marginally delayed in the experimental group (2-3 wk). This marginal delay in the arrest of reproductive activity seen in both experiments indicates that the lack of decrease in day length around and after the winter solstice may play some role in timing the end of the breeding season. However, our inability to prevent or markedly delay the termination of reproductive activity leads to the conclusion that the primary reason for the transition into anestrus is an obligatory turn-off. This obligatory process may be the expression of an underlying endogenous rhythm of reproduction.     

Photoperiodic requirements for timing onset and duration of the breeding season of the ewe: Synchronization of an endogenous rhythm of reproduction

Summary A study was conducted to test the hypothesis that different portions of the annual photoperiodic cycle play different roles in timing the breeding season of the ewe, Ovis aries, an animal in which an endogenous rhythm generates the seasonal reproductive transitions. Adult female sheep were pinealectomized to disrupt transduction of photoperiodic cues at 4 times of the year (summer and winter solstices, vernal and autumnal equinoxes), and the effects on seasonal reproductive neuroendocrine activity were evaluated. Time of pinealectomy greatly influenced the subsequent seasonal reproductive cycle such that the following inferences are possible. Lengthening days between the winter and summer solstices synchronize reproductive onset to the appropriate time of year. The relatively long days around the summer solstice act to suppress reproductive activity and forestall the start of the breeding season until late summer/early autumn. The shortening days between the summer solstice and autumnal equinox maintain a normal intensity and duration of reproductive neuroendocrine induction during the impending breeding season. However, the shortening days between the autumnal equinox and winter solstice (i.e., after breeding season onset) do not appear to play a critical role in maintaining the breeding season of that year, but may provide important cues for timing the breeding season of the following year.Abbreviations LH luteinizing hormone Presented in preliminary form at the 21st Annual Meeting of the Society for the Study of Reproduction, 1988, Biol Reprod 38 (Suppl. 1): 184 (Abstract 408). This work was performed in partial fulfillment of the requirements for the degree of Ph.D. at The University of Michigan (to N.L.W.), and was funded by NIH-HD-18337 and NIH-HD-18258, T-32-HD-07048, the Institut National de la Recherche Agronomique (France) and the Office of the Vice President for Research of The University of Michigan.     

Seasonal variations in circadian rhythms coincide with a phase of sensitivity to short photoperiods in the European hamster

European hamsters (Cricetus cricetus) show pronounced seasonal changes in their physiology and behavior. The present study provides a detailed analysis of the temporal relationship between seasonal cycles of reproduction and body mass and seasonal changes of two circadian parameters, i.e., locomotor activity and 6-sulphatoxymelatonin (aMT6s) excretion, in individual animals kept under natural light conditions. Our results demonstrate a characteristic pattern of locomotor activity and aMT6s excretion observed around the summer solstice, i.e., from mid-May to mid-July. During this time, locomotor activity was characterized by a high level of activity and an early activity onset, while the nightly elevation of melatonin was reduced to baseline levels. These seasonal changes in aMT6s excretion and locomotor activity were only loosely related to changes in the reproductive status of the animals, but correlated well with a period of the annual cycle during which the animals were sensitive to short days. They may therefore reflect a specific state of the circadian pacemaker system within the SCN and can thus be a valuable tool to further characterize molecular and physiological mechanisms of photoperiodic time measurements in European hamsters.     

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.     

The interaction between photoperiod and nutrition and its effects on seasonal rhythms of reproduction in the ewe

In sheep, the seasonal patterns of reproductive activity are driven primarily by the annual photoperiodic cycle, but can also respond to other environmental factors, such as nutrition, yet little is known about the mechanisms underlying this interaction. This study was designed to define the interaction between photoperiodic and nutritional cues on seasonal patterns of ovarian activity, and to determine if there is a central interaction between these cues. Groups of Ile-de-France ewes were maintained in two nutritional states (restricted and well fed) under a simulated annual photoperiod of 8-16 h of light per day over two breeding seasons. At the end of the first breeding season, half of the animals of each group were ovariectomized (OVX) and fitted subcutaneously with estradiol implants. Low nutritional status shortened the season of ovarian activity, determined from the pattern of progesterone concentrations, by modifying the timing of seasonal transitions between periods of ovarian activity and anestrus. The same results were observed for the seasonal rhythm of neuroendocrine activity, assessed in the OVX ewes, from the pattern of luteinizing hormone concentrations. These results were then confirmed for neuroendocrine activity induced by a photoperiodic treatment. We conclude that nutrition centrally modulates the interpretation of photoperiod to affect seasonal reproductive transitions. The mechanisms of this interaction are discussed in the paper.     

Importance of changing photoperiod and melatonin secretory pattern in determining the length of the breeding season in the Suffolk ewe

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.     

Melatonin Does Not Modulate Testicular Responsiveness to Altered Photoperiods. A Study During Different Phases of the Annual Gonadal Cycle in Roseringed Parakeets (Psittacula krameri)

The roseringed parakeet has been shown to exhibit a variable testicular responsiveness to both altered photoperiodic regimens and to treatment with melatonin during different phases of the annual gonadal cycle. Adult male roseringed parakeets were held under either natural photoperiods (NP), or long photoperiods (LP; 16L 8D), or short photoperiods (SP; 8L 16D) for a total period of 90 days. From day 46 onward, half of the total birds in each group were administered with the vehicle of melatonin, and the other birds were injected daily in the afternoon with melatonin (25 µg/ 100 g body wt.) till the end of the experiment. An identical experimental schedule was followed during the four different (preparatory, progressive, pre-breeding, and breeding) phases of the annual testicular cycle. The testicular activities in various bird groups were evaluated by volumetric, gravimetric, histometric and karyometric measurements, and by quantitative histological studies. The findings revealed that exogenous melatonin may exert either a suppressive influence or none at all on the testicular functions in relation to the photoperiodic schedule as well as to the reproductive phase of the concerned bird, but in no case modulates gonadal responsiveness to artificially altered photoperiods.     

Melatonin Does Not Modulate Testicular Responsiveness to Altered Photoperiods. A Study During Different Phases of the Annual Gonadal Cycle in Roseringed Parakeets (Psittacula krameri)

The roseringed parakeet has been shown to exhibit a variable testicular responsiveness to both altered photoperiodic regimens and to treatment with melatonin during different phases of the annual gonadal cycle. Adult male roseringed parakeets were held under either natural photoperiods (NP), or long photoperiods (LP; 16L 8D), or short photoperiods (SP; 8L 16D) for a total period of 90 days. From day 46 onward, half of the total birds in each group were administered with the vehicle of melatonin, and the other birds were injected daily in the afternoon with melatonin (25 µg/ 100 g body wt.) till the end of the experiment. An identical experimental schedule was followed during the four different (preparatory, progressive, pre-breeding, and breeding) phases of the annual testicular cycle. The testicular activities in various bird groups were evaluated by volumetric, gravimetric, histometric and karyometric measurements, and by quantitative histological studies. The findings revealed that exogenous melatonin may exert either a suppressive influence or none at all on the testicular functions in relation to the photoperiodic schedule as well as to the reproductive phase of the concerned bird, but in no case modulates gonadal responsiveness to artificially altered photoperiods.     

Photoperiodic counter of diapause induction in Pseudopidorus fasciata (Lepidoptera: Zygaenidae)

Induction of larval diapause is a photoperiodically controlled event in the life history of the moth Pseudopidorus fasciata. In the present study, the photoperiodic counter of diapause induction has been systematically investigated. The required day number (RDN) for a 50% response was determined by transferring from a short night (LD 16:8) to a long night (LD 12:12) or vice versa at different times after hatching, The RND differed significantly between short- and long-night cycles at different temperatures. The RDN for long-night cycles at 20, 22, 25 and 28 degrees C was 11.5, 9.5, 7.5 and 8.5 days, respectively. The RDN for short-night cycles was 3 days at 22 degrees C and 5 days at 20 degrees C indicating that the effect of one short night was equivalent to the effect of 2-3 long nights effect. Night-interruption experiments of 24h photoperiods by a 1 h light pulse showed that the most crucial event for the photoperiodic time measurement in this moth was whether the length of pre-interruption (D(1)) or the post-interruption (D(2)) scotophases exceeded the critical night length (10.5 h). If D(1) or D(2) exceeded 10.5 h diapause was induced. The diapause-averting effect of a single short-night cycle (LD 16:8) against a background of long nights (LD 12:12) showed that the photoperiodic sensitivity was greatest during the first 7 days of the larval period and the highest sensitivity was on the fourth day. Both non-24 and 24 h light-dark cycle experiments revealed that the photoperiodic counter in P. fasciata is able to accumulate both long and short nights during the photosensitive period, but in different ways. The information from short-night cycles seems to be accumulated one by one in contrast to long-night cycles where six successive cycles were necessary for about 50% diapause induction and eight cycles for about 90% diapause. These results suggest the accumulation of long-night and short-night cycles may be based on different mechanisms.     

Amplitude of the plasma melatonin nycthemeral rhythms is not associated with the dates of onset and offset of the seasonal ovulatory activity in the Ile-de-France ewe

We investigated if absolute (nocturnal) or relative (nocturnal/diurnal ratio) plasma melatonin concentrations were associated with the seasonal ovulatory activity in Ile-de-France ewes. Ninety-six and 121 ewes in two different groups of the same flock were used to determine the potential existence of a relationship between melatonin concentrations at the summer and winter solstices, and the dates of onset and offset of the ovulatory activity, respectively. The dates of the first and last ovulations were estimated by assaying progesterone in plasma samples taken once weekly. Mean +/- SEM (1) plasma melatonin concentrations at the summer solstice and at the winter solstice were 302.4 +/- 19.4 and 412.0 +/- 18.7 pg x mL(-1), respectively, (2) date of onset of the breeding season 29 Jul. +/- 1.6 days, (3) date of offset of the breeding season 24 Jan. +/- 2.2 days. In spite of a large variability in the different traits studied here, the analyses of correlation and regression indicated that neither the absolute nor relative melatonin concentrations were significantly related with the dates of onset or offset of ovulatory activity. Therefore, we concluded that absolute or relative plasma melatonin concentrations are not linked to the seasonal breeding activity, in Ile-de-France ewes.     

PHASE ADVANCE AFTER ONE OR THREE SIMULATED DAWNS IN HUMANS

A specially designed apparatus that can simulate the waveform of the dawn or dusk signal at any latitude and any day of the year has been shown to phase shift the circadian pacemaker in rodents and primates at a fraction of the illuminance previously used. Until recently, it was considered that rather high illuminances or rather long exposure episodes to room light were necessary to phase shift human circadian rhythms. This experiment shows that, under controlled conditions of a modified constant routine protocol, a single dawn signal is sufficient to phase advance the timing of the onset of secretion of the pineal hormone melatonin. The significant phase advance of salivary melatonin of 20 minutes, which is enhanced to 34 minutes after three consecutive dawn signals, is small, but appears to be of sufficient magnitude to entrain the human circadian pacemaker, which has an endogenous period of about 24.2h. (Chronobiology International, 17(5), 659–668, 2000)     

Phase advance after one or three simulated dawns in humans

A specially designed apparatus that can simulate the waveform of the dawn or dusk signal at any latitude and any day of the year has been shown to phase shift the circadian pacemaker in rodents and primates at a fraction of the illuminance previously used. Until recently, it was considered that rather high illuminances or rather long exposure episodes to room light were necessary to phase shift human circadian rhythms. This experiment shows that, under controlled conditions of a modified constant routine protocol, a single dawn signal is sufficient to phase advance the timing of the onset of secretion of the pineal hormone melatonin. The significant phase advance of salivary melatonin of 20 minutes, which is enhanced to 34 minutes after three consecutive dawn signals, is small, but appears to be of sufficient magnitude to entrain the human circadian pacemaker, which has an endogenous period of about 24.2h. (Chronobiology International, 17(5), 659-668, 2000)     

Importance of photoperiodic signal quality to entrainment of the circannual reproductive rhythm of the ewe

An endogenous circannual rhythm drives the seasonal reproductive cycle of a broad spectrum of species. This rhythm is synchronized to the seasons (i.e., entrained) by photoperiod, which acts by regulating the circadian pattern of melatonin secretion from the pineal gland. Prior work has revealed that melatonin patterns secreted in spring/summer entrain the circannual rhythm of reproductive neuroendocrine activity in sheep, whereas secretions in winter do not. The goal of this study was to determine if inability of the winter-melatonin pattern to entrain the rhythm is due to the specific melatonin pattern secreted in winter or to the stage of the circannual rhythm at that time of year. Either a summer- or a winter-melatonin pattern was infused for 70 days into pinealectomized ewes, centered around the summer solstice, when an effective stimulus readily entrains the rhythm. The ewes were ovariectomized and treated with constant-release estradiol implants, and circannual cycles of reproductive neuroendocrine activity were monitored by serum LH concentrations. Only the summer-melatonin pattern entrained the circannual reproductive rhythm. The inability of the winter pattern to do so indicates that the mere presence of a circadian melatonin pattern, in itself, is insufficient for entrainment. Rather, the characteristics of the melatonin pattern, in particular a pattern that mimics the photoperiodic signals of summer, determines entrainment of the circannual rhythm of reproductive neuroendocrine activity in the ewe.     

Refractoriness to melatonin and short daylengths in early seasonal quiescence in the Bennett's wallaby (Macropus rufogriseus rufogriseus)

Adult female Bennett's wallabies were treated with reductions in daylength, melatonin implants or injections of melatonin 2 h before dusk in early or mid-seasonal reproductive quiescence. In early reproductive quiescence (5 weeks after the winter solstice) reproductive quiescence did not end in response to 3 or 5 h of reduced daylength or in response to injections (400 ng/kg) or implants (0.5 g in a Silastic rubber envelope) of melatonin. Reductions in daylength at this time of year did, however, result in an extension of the circadian pattern of melatonin secretion. In mid-reproductive quiescence (21 weeks after the winter solstice) treatment with a 5 h reduction in daylength, melatonin injections administered 2 h before dusk or melatonin implants did result in the termination of reproductive quiescence and reactivation of the quiescent corpus luteum within a period of 5 days. The results of these experiments indicate that, in early reproductive quiescence, the Bennett's wallaby is refractory to the influence of reduced daylength or melatonin, although capable of responding to such reduced days in terms of an increased duration of melatonin secretion. Bennett's wallabies therefore exhibit a refractoriness to short days similar to that of some seasonal eutherians although it remains to be established whether this refractory response is the cause of the transition to seasonal reproductive quiescence.     

Photoperiodic history and a changing melatonin pattern can determine the neuroendocrine response of the ewe to daylength

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:11D. 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 24 h 17 days before and three times after transfer to 13L:11D. 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.     

Effects of melatonin implants in pony mares. 2. Long-term effects

The effects of melatonin implant treatment over a 4 wk period at the summer solstice on the transition into and out of the following anovulatory season were evaluated in ovary-intact and ovariectomized mares. Melatonin implants tended to delay the timing of the final ovulation of the breeding season (P = 0.0797) in the ovary-intact mares. Although the decline in LH secretion associated with the end of the breeding season was parallel between treatments and ovarian statuses, the rate of LH secretion, as expressed by its mathematical accumulation, was lower in ovariectomized, melatonin-treated mares than in ovariectomized, control mares suggesting that melatonin administration advanced the offset of the breeding season in ovariectomized mares (P = 0.0001). The first ovulation of the subsequent breeding season was significantly delayed in the melatonin-treated mares as compared with that of control mares (P = 0.0031). During reproductive recrudescence, the time of the onset of the increase in LH secretion was similar among all 4 groups but the patterns of LH secretion were different for each treatment and ovarian status combination (P = 0.0112). Mares with melatonin implants had a slower rate of increase in LH secretion than control mares (P = 0.0001), and ovariectomized mares had a faster rate of LH increase than intact mares (P = 0.0001). These results suggest that melatonin implants during the summer solstice can alter the annual reproductive rhythm in mares and support the concept that endocrine patterns of reproductive recrudescence are not entirely independent of the ovary.     

Photoperiodic polyphenisms in rodents: neuroendocrine mechanisms, costs, and functions

Annual changes in daylength figure prominently in the generation of seasonal rhythms in reproduction, and a wide variety of mammals use ambient photoperiod as a proximate cue to time critical reproductive events. Nevertheless, within many reproductively photoperiodic mammalian species, there exist individuals--termed "photoperiod nonresponders"--that fail to adopt a seasonal breeding strategy and instead exhibit reproductive competence at a time of year when their conspecifics are reproductively quiescent. Photoperiod nonresponsiveness has been principally characterized by laboratory observations--over half of the species known to be reproductively photoperiodic contain a proportion of nonresponsive individuals. The study of nonresponders has generated basic insights regarding photic regulation of reproduction in mammals. The neuroendocrine mechanisms by which the short-day photoperiodic signal is degraded or lost in nonresponders varies between species: differences in features of the circadian pacemaker, which provides photoperiodic input to the reproductive neuroendocrine system, have been identified in hamsters; changes in the responsiveness of hypothalamic gonadotrophs to melatonin and as-yet-unspecified inhibitory signals have been implicated in voles and mice. Individuals that continue to breed when their conspecifics refrain might enjoy higher fitness under certain circumstances. Statements regarding the adaptive function of reproductive nonresponsiveness to photoperiod require additional information on the costs (metabolic and fitness) of sustaining reproductive function during the winter months and how these costs vary as a function of environmental conditions. Reproductive nonresponders thus continue to represent a challenge to theories that extol the adaptive function of seasonality. Several nonexclusive hypotheses are proposed to account for the maintenance of nonresponsive individuals in wild rodent populations.     

Molecular characterization of the long-day response in the Soay sheep, a seasonal mammal

In mammals, seasonal timekeeping depends on the generation of a nocturnal melatonin signal that reflects nightlength/daylength. To understand the mechanisms by which the melatonin signal is decoded, we studied the photoperiodic control of prolactin secretion in Soay sheep, which is mediated via melatonin responsive cells in the pars tuberalis of the pituitary. We demonstrate that the phases of peak expression of the clock genes Cryptochrome1 (Cry1), Period1 (Per1), and RevErbalpha respond acutely to altered melatonin secretion after a switch from short to long days. Cry1 is activated by melatonin onset, forming the dusk component of the molecular decoder, while Per1 expression at dawn reflects the offset of melatonin secretion. The Cry1-Per1 interval immediately adjusts to the melatonin signal on the first long day, and this is followed within 24 hr by an increase in prolactin secretion. The timing of peak RevErbalpha expression also responds to a switch to long days due to altered melatonin secretion but does not immediately reset to an entrained long-day state. These data suggest that effects of melatonin on clock gene expression are pivotal events in the neuroendocrine response and that pars tuberalis cells can act as molecular calendars, carrying a form of "photoperiodic memory."     

Refractoriness to inhibitory day lengths initiates the breeding season of the Suffolk ewe

Recent evidence indicates that the breeding season of the Suffolk ewe ends because of loss of response to a day length that was previously inductive. This condition of photorefractoriness could potentially also initiate reproduction, as is the case in several long-day breeding rodents. In this study we determined if ewes enter their breeding season because they become refractory to the long ambient photoperiods of late summer. On the summer solstice, 3 groups of ovariectomized ewes (n = 6) bearing s.c. Silastic implants of estradiol (OVX + E) were placed in different day length treatments: 1) natural photoperiod; 2) artificial photoperiod, stimulating natural day lengths; or 3) artificial photoperiod equivalent to that of the summer solstice (16.25L). Entry into the breeding season is associated with a striking (greater than 30-fold) increase in circulating levels of luteinizing hormone (LH). Timing of the onset of the breeding season was not delayed in ewes maintained on the summer solstice photoperiod; LH levels rose simultaneously in all groups. We conclude that ewes normally begin breeding not because they are actively driven to do so by decreasing or short days, but because they become refractory to prevailing long days. Because the pattern of circulating melatonin, which is known to transduce the photoperiodic message, remained entirely appropriate to day length, we believe that the mechanism responsible for photorefractoriness resides in the postpineal processing of the melatonin signal.