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1.
The pineal gland, through its nocturnal melatonin secretion, mediates the effects of inhibitory (long) and stimulatory (short) photoperiod on reproduction in female sheep. Earlier studies revealed that duration of the nighttime melatonin rise is important in determining the inhibitory effect of day length on reproduction in the ewe. The present study tested whether the duration is also important in mediating the inductive response to short days. Pinealectomized ewes, housed under long days, received a short-day melatonin infusion (16-h duration) for 90 days. Reproductive status was monitored from the response to estradiol negative feedback of luteinizing hormone (LH) secretion. This short-day melatonin pattern led to unambiguous reproductive induction, despite the exposure to inhibitory long days. The increase in serum LH was comparable, in terms of latency and magnitude, to that in pinealectomized controls receiving the same short-day melatonin pattern under short days, and in pineal-intact controls transferred from long to short days. Since the reproductive status conformed to the length of time that melatonin was elevated each day rather than to photoperiod, these results support the conclusion that duration of the nighttime melatonin rise mediates the reproductive response of the ewe to an inductive photoperiod. In all, the melatonin rhythm is considered an integral component of the physiologic mechanism measuring day length; through duration of its nocturnal secretion, melatonin encodes both inhibitory and stimulatory photoperiods.  相似文献   

2.
This study examined whether or not the reproductive response of female sheep to photoperiod varies with seasonal reproductive state. The specific objective was to test the hypothesis that the reproductive response to a long-day pattern of melatonin varies with the reproductive state of the ewe. The response examined was the synchronization of reproductive neuroendocrine induction (rise in serum luteinizing hormone, or LH) following nocturnal infusion of melatonin into pinealectomized ewes for 35 consecutive nights. This infusion restored a pattern of circulating melatonin similar to that in pineal-intact ewes maintained in a long photoperiod (LD 16:8). The ewes had been pinealectomized and without melatonin replacement for 16-25 months prior to the study. They were in differing reproductive states at the start of the infusion, as their endogenous reproductive rhythm had become desynchronized among individuals and with respect to time of year. Noninfused pinealectomized ewes served as controls. Regardless of the reproductive state at the start of the 35-day infusion of the long-day pattern of melatonin, all treated ewes exhibited the same reproductive neuroendocrine response after the infusion was ended. This consisted of a synchronized rise in LH some 6-8 weeks after the infusion was terminated, the maintenance of a high level of serum LH for some 15 weeks, and a subsequent precipitous fall in LH to a very low level. These results provide evidence that a long-day pattern of melatonin can synchronize reproductive neuroendocrine induction in the ewe, regardless of reproductive condition, and thus do not support the hypothesis that this response differs with seasonal reproductive state.  相似文献   

3.
Experiments were conducted to examine whether the refractoriness of the Suffolk ewe to the reproductive effects of day length is associated with a deficit in the generation of the circadian rhythm of melatonin secretion or in the postpineal processing of this photoperiodic message. Using serum luteinizing hormone (LH) concentrations in ovariectomized ewes bearing constant-release estradiol implants as a marker of reproductive induction, ewes with intact pineal glands were found to become unresponsive to fixed artificial photoperiods that initially had been either inductive (short days) or inhibitory (long days). The loss of the photoperiodic response was not associated with notable changes in the 24-h secretory pattern of melatonin, which remained characteristically low throughout the day and rose at night. In pinealectomized ewes, nightly infusion of a stimulatory pattern of melatonin (simulating that seen on short days) initially provoked reproductive induction; this response then lessened over much the same time course that pineal intact ewes became refractory to short days. These results support the hypothesis that photorefractoriness reflects a deficit in the postpineal processing of the photoperiodic message. Further, in view of recent evidence that photorefractoriness normally leads to both onset and cessation of the breeding season in Suffolk ewes maintained outdoors, these findings suggest that the loss of response to the melatonin signal contributes to at least one of these reproductive transitions, the cessation of the breeding season, under natural environmental conditions.  相似文献   

4.
The main objective of the study was to test the hypothesis that the phase of melatonin release with respect to the light-dark cycle mediates the effects of photoperiod on the reproductive response of the ewe. To test the phase hypothesis, we eliminated endogenous melatonin secretion by pinealectomy and then restored physiological levels of serum melatonin with rises of the same duration but at different phases of the light-dark cycle (either at night or in the middle of the day). Serum melatonin patterns were determined by radioimmunoassay in samples taken hourly for 24 h. The reproductive state was monitored by measuring serum luteinizing hormone (LH) in ovariectomized ewes treated with constant-release estradiol implants. Infusion of a long-day pattern of melatonin was equally effective in maintaining reproductive suppression when given during the night or the middle of the day. LH remained low for approximately 150 days and then rose as ewes became refractory to the inhibitory melatonin signal. These results do not support the phase hypothesis. Rather, they are consistent with the hypothesis that the duration of the nocturnal secretion of melatonin codes for day length.  相似文献   

5.
Although many species display endogenous circannual rhythms of biological activity that are synchronized by day length, the specific photoperiodic requirements for synchronizing such rhythms are not established for any species. We tested the hypothesis that the circannual reproductive rhythm of sheep can be synchronized by exposure to just one or two discrete blocks of photoperiodic information each year. Ewes were pinealectomized to prevent their ability to transduce photoperiodic information into altered reproductive neuroendocrine activity. During the 53/4 yr following pinealectomy, specific photoperiodic signals were restored for discrete periods of time via replacement of 24-h patterns of melatonin, the pineal hormone that transmits photic information to the reproductive neuroendocrine axis. The ewes were kept in a 12-mo photoycycle that alternated between short (8L:16D) and long (16L:8D) days every 6 mo and that was 6 mo out of phase with the geophysical year. Pineal-intact control ewes exhibited synchronous annual reproductive cycles. Noninfused pinealectomized control ewes did not exhibit synchronous cycles. Pinealectomized ewes infused with alternating 70-day blocks of short- and long-day patterns of melatonin every 6 mo for the first 21/2 yr of the experiment exhibited synchronous annual reproductive cycles that were 6 mo out of phase with those of ewes maintained outdoors. This synchrony persisted when the frequency of the melatonin treatment was reduced to just one 70-day block of a long-day pattern of melatonin each 365 days. Cycle period was 368 +/- 3 days; standard deviation of the date of onset of reproductive induction averaged only 3 days. Our study provides the first direct evidence that a single block of photoperiodic information a year can synchronize a circannual rhythm.  相似文献   

6.
The 24-h pattern of melatonin secretion was evaluated in Suffolk ewes during prolonged exposure to an inductive photoperiod to assess whether altered secretion of melatonin could account for the eventual loss of response to stimulatory photoperiod (photorefractoriness). Secretory patterns of melatonin were determined approximately every two weeks in samples obtained hourly for 24-48 h. Sampling was begun one week before the switch from inhibitory (long) to inductive (short) photoperiod and continued for 150 days, by which time all but one of the ewes were unresponsive to that stimulatory day length. Melatonin was measured in two different radioimmunoassay systems. Reproductive state was monitored by luteinizing hormone secretion in ovariectomized ewes bearing constant-release estradiol implants. No evidence for disruption of the melatonin pattern was observed on any occasion. The duration and the phase of the melatonin elevation relative to the light/dark cycle did not vary with time of exposure to short days. These findings indicate that refractoriness of the Suffolk ewe to an inductive photoperiod is not caused by an inappropriate secretory pattern of melatonin.  相似文献   

7.
The objective of this work was to evaluate the role of short photoperiod in timing the onset and duration of reproductive activity in ewes. The perception of photoperiod was disrupted by pinealectomy following transfer from long (17L:7D) to short (8.5L:15.5D) photoperiod and the subsequent reproductive response was monitored. Ovariectomized ewes given Silastic implants containing estradiol-17 beta were exposed to long days until Day 0 (May 24) and then were allocated to the following groups (n = 5-6/group): Group 1) short-day control--moved to short days; Groups 2 to 5) pinealectomy after 0, 30, 60, or 90 short days, respectively; Group 6) long-day hold--kept on long days; Group 7) long days after 60 short days--moved to short days on Day 0 and returned to long days on Day 60. Six ewes kept outdoors served as additional controls. Reproductive neuroendocrine activity was assessed from plasma LH concentrations, high values being indicative of the breeding season and low values indicative of anestrus. Time of reproductive neuroendocrine activity onset (LH rise) did not differ among animals in the 7 groups kept indoors, but was advanced (p less than 0.05) relative to that of ewes outdoors. In contrast, duration of the LH elevation differed among ewes in groups kept indoors.(ABSTRACT TRUNCATED AT 250 WORDS)  相似文献   

8.
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.  相似文献   

9.
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.  相似文献   

10.
Summary A recent study has shown that olfactory bulbectomy (BX) will prevent reproductive regression associated with short photoperiod in male golden hamsters. The results of experiments reported in this paper show that bulbectomized hamsters on long or short photoperiod still show a large nocturnal elevation in pineal melatonin production and that BX inhibits the reproductive regression induced by exogenous melatonin in pinealectomized hamsters. The data therefore indicate that BX does not inhibit short photoperiod induced testicular regression by altering melatonin secretion.  相似文献   

11.
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.  相似文献   

12.
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.  相似文献   

13.
Short day lengths delay reproductive aging   总被引:1,自引:0,他引:1  
Caloric restriction and hormone treatment delay reproductive senescence in female mammals, but a natural model of decelerated reproductive aging does not presently exist. In addition to describing such a model, this study shows that an abiotic signal (photoperiod) can induce physiological changes that slow senescence. Relative to animals born in April, rodents born in September delay their first reproductive effort by up to 7 mo, at which age reduced fertility is expected. We tested the hypothesis that the shorter day lengths experienced by late-born Siberian hamsters ameliorate the reproductive decline associated with advancing age. Short-day females (10L:14D) achieved puberty at a much later age than long-day animals (14L:10D) and had twice as many ovarian primordial follicles. At 10 mo of age, 86% of females previously maintained in short day lengths produced litters, compared with 58% of their long day counterparts. Changes in pineal gland production of melatonin appear to mediate the effects of day length on reproductive aging; only 30% of pinealectomized females housed in short days produced litters. Exposure to short days induces substantial decreases in voluntary food intake and body mass, reduced ovarian estradiol secretion, and enhanced production of melatonin. One or more of these changes may account for the protective effect of short day lengths on female reproduction. In delaying reproductive senescence, the decrease in day length after the summer solstice is of presumed adaptive significance for offspring born late in the breeding season that first breed at an advanced chronological age.  相似文献   

14.
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.  相似文献   

15.
Reproduction is a complex and energy demanding function. When internal and external conditions might impair reproductive success (negative energy balance, stress, harsh season) reproductive activity has to be repressed. Recent evidence suggests that these inhibitory mechanisms operate on Kiss1-expressing neurons, which were recently shown to be implicated in the regulation of GnRH release. Hamsters are seasonal rodents which are sexually active in long photoperiod and quiescent in short photoperiod. The photoperiodic information is transmitted to the reproductive system by melatonin, a pineal hormone whose secretion is adjusted to night length. The photoperiodic variation in circulating melatonin has been shown to synchronize reproductive activity with seasons, but the mechanisms involved in this effect of melatonin were so far unknown. Recently we have observed that Kiss1 mRNA level in the arcuate nucleus of the Syrian hamster is lower in short photoperiod, when animals are sexually quiescent. Notably, intracerebroventricular infusion of Kiss1 gene product, kisspeptin, in hamsters kept in short photoperiod is able to override the inhibitory photoperiod and to reactivate sexual activity. The inhibition of Kiss1 expression in short photoperiod is driven by melatonin because pinealectomy prevents decrease in Kiss1 mRNA level in short photoperiod and melatonin injection in long photoperiod down regulates Kiss1 expression. Whether melatonin acts directly on arcuate Kiss1 expressing neurons or mediates its action via interneurons is the subject of the current investigations.  相似文献   

16.
In this study, the authors asked whether pinealectomy or temporary exposure to a stimulatory photoperiod affects the timing of spontaneous testicular recrudescence in adult Siberian hamsters chronically exposed to short days (9:15 light:dark). In Experiment 1, hamsters were pinealectomized after 6, 9, or 12 weeks in short days. Pinealectomy after 9 or 12 weeks did not affect the timing of spontaneous gonadal growth (27.7 +/- 1.9 and 25.4 +/- 1.3 weeks, respectively) compared to sham-operated controls (28.6 +/- 0.9 weeks). Enlarged testes occurred earlier in animals that were pinealectomized after 6 weeks in short days (21.8 +/- 2.1 weeks). In Experiment 2, adult hamsters were exposed to short days for 9 weeks, transferred to long days (16:8 light:dark) for 4 weeks, and then returned to short days for 23 additional weeks. Although long-day interruption caused gonadal growth in 15 out of 19 hamsters, the temporary long-day exposure did not affect the timing of spontaneous gonadal growth following return to short days (28.2 +/- 0.9 weeks) in 10 of the 15, relative to the timing observed in control hamsters continuously maintained in short days (28.2 +/- 1.1 weeks). Four out of 19 hamsters did not show gonadal growth following long-day exposure. Spontaneous gonadal growth in these hamsters (28.0 +/- 1.4 weeks) also occurred at the same time as controls. The remaining 5 hamsters exhibited enlarged testes following long-day exposure (12.0 +/- 0.0 weeks) but were refractory to the second short-day exposure. All hamsters exhibited entrainment of wheel-running activity following the change in photoperiod. A final group of 13 animals were pinealectomized before long-day transfer. They exhibited gonadal growth (at 17.2 +/- 0.8 weeks) but failed to regress a second time when returned to short days. The timing of gonadal growth in these animals was delayed relative to the sham-operated hamsters temporarily transferred to long days (Experiment 2) but accelerated relative to the hamsters pinealectomized at 9 weeks, which remained continuously in short days (Experiment 1). The results of both experiments suggest that a pineal-independent process mediates the timing of spontaneous gonadal growth in Siberian hamsters chronically exposed to a short-day photoperiod.  相似文献   

17.
The effects of pinealectomy on pituitary and plasma gonadotropin levels and gonadal development in female goldfish exposed to various photoperiod-temperature regimes during different seasons were examined. Pinealectomy during autumn had no effect on either pituitary or plasma hormone levels or gonadal development. When goldfish are pinealectomized in spring and exposed to long photoperiod conditions, the ovaries regress and plasma gonadotropin levels are significantly depressed compared to sham operated animals. Sham operated goldfish exposed to short photoperiod conditions in spring had regressing ovaries whereas pinealectomized animals under this regime either spawned or had ovaries in the late vitellogenic phase. Plasma gonadotropin titres in the pinealectomized group were significantly lower than those of sham operated animals. The pineal can be either stimulatory or inhibitory to gonadal development depending on the photoperiod regime to which the animals are exposed. The pineal apparently influences gonadal activity by modulating gonadotropin secretion. A diurnal variation in plasma gonadotropin levels was also observed in both sham operated and pinealectomized goldfish exposed to a long photoperiod warm-temperature regime in spring.  相似文献   

18.
Long-term effects of subcutaneous melatonin implants on intrahypothalamic LHRH content and on pulsatile luteinizing hormone release have been investigated in ganglionectomized male mink. Animals were submitted to bilateral removal of the superior cervical ganglion in mid-April. A preliminary study revealed that plasma LH concentrations remain at a basal level throughout the year following ganglionectomy. In a second experiment, one month after ganglionectomy and transfer from the natural photoperiod environment to short daylengths (LD 4:20), melatonin pellets were subcutaneously implanted to overcome deafferentation of the pineal. Progressive effects of treatment were studied 7 days, 15 days, and one, two and three months after insertion of the melatonin implants. The intra-hypothalamic LHRH content in ganglionectomized mink was at a basal level similar to that observed during seasonally sexual quiescence, or after exposure to inhibitory long days (LD 20:4). A significant and transient elevation in LHRH content was observed already after fifteen days, and also one month after insertion of melatonin implants. This resulted in mean values similar to those observed during the breeding season, or after exposure to stimulatory short days (LD 4:20). A decrease in hypothalamic LHRH content started after two months. No pattern of pulsatile LH secretion was recorded in ganglionectomized untreated mink. A significant increase in all parameters of pulsatile LH secretion was observed fifteen days after the elevation of LHRH content induced by melatonin treatment, and maximum values were reached after two months. Pituitary activity tended to decrease after three months, characterized in particular by a significant decrease in the mean frequency of LH pulses. In addition, the increase in pulsatile characteristics of LH release occurred two months before the peripheral renewal of testicular activity. Apparently, the reproductive endocrine function in ganglionectomized mink treated with melatonin implants is restored more rapidly at the hypothalamic level than at the pituitary or testicular levels.  相似文献   

19.
Long-term effects of subcutaneous melatonin implants on intrahypothalamic LHRH content and on pulsatile luteinizing hormone release have been investigated in ganglionectomized male mink. Animals were submitted to bilateral removal of the superior cervical ganglion in mid-April. A preliminary study revealed that plasma LH concentrations remain at a basal level throughout the year following ganglionectomy. In a second experiment, one month after ganglionectomy and transfer from the natural photoperiod environment to short daylengths (LD 4:20), melatonin pellets were subcutaneously implanted to overcome deafferentation of the pineal. Progressive effects of treatment were studied 7 days, 15 days, and one, two and three months after insertion of the melatonin implants. The intra-hypothalamic LHRH content in ganglionectomized mink was at a basal level similar to that observed during seasonally sexual quiescence, or after exposure to inhibitory long days (LD 20:4). A significant and transient elevation in LHRH content was observed already after fifteen days, and also one month after insertion of melatonin implants. This resulted in mean values similar to those observed during the breeding season, or after exposure to stimulatory short days (LD 4:20). A decrease in hypothalamic LHRH content started after two months. No pattern of pulsatile LH secretion was recorded in ganglionectomized untreated mink. A significant increase in all parameters of pulsatile LH secretion was observed fifteen days after the elevation of LHRH content induced by melatonin treatment, and maximum values were reached after two months. Pituitary activity tended to decrease after three months, characterized in particular by a significant decrease in the mean frequency of LH pulses. In addition, the increase in pulsatile characteristics of LH release occurred two months before the peripheral renewal of testicular activity. Apparently, the reproductive endocrine function in ganglionectomized mink treated with melatonin implants is restored more rapidly at the hypothalamic level than at the pituitary or testicular levels.  相似文献   

20.
Spring-born female lambs require a decrease in day length for the normal timing of puberty the following autumn. If this decrease occurs early in postnatal life (i.e. 0-10 weeks), puberty is delayed. This study tested the hypothesis that failure of the neonatal lamb to respond to the critical long-day to short-day signal is due to inadequate nocturnal melatonin secretion. The approach was to artificially increase, to adult levels, the low nighttime rises of melatonin during the early postnatal period. Eight female lambs served as controls; they were raised on short days until 17 wk of age, and then exposed to 5 wk of long days, after which they were returned to short days. This alternating sequence of photoperiods during mid-development would be expected to induce normal puberty. Sixteen experimental females were exposed to the critical block of long days much earlier; they were placed in long days between 2 and 7 wk of age and in short days thereafter. Half (n = 8) received no further treatment. The other half (n = 8) were infused nightly with melatonin during the 8-h dark phase of the 5-wk, long-day photoperiod. This increased the amplitude of the natural nighttime melatonin rises 3- to 4-fold, well into the adult range.(ABSTRACT TRUNCATED AT 250 WORDS)  相似文献   

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