Termination of gonadal refractoriness in Turkish hamsters, Mesocricetus brandti

The Turkish hamster (Mesocricetus brandti) is a photoperiodic species. In this investigation, we characterized the photoperiodic requirements for termination of gonadal refractoriness, defined as the inability of the animal to respond to short-day treatment with gonadal regression. Paired testes weights were reduced to less than 20% of their original weight by 10 wk of 12L:12D treatment. This was followed by spontaneous testicular recrudescence (completed by Week 25 of 12L:12D treatment), the overt indication of refractoriness to short photoperiods. Next, the period of long-day exposure sufficient for termination of refractoriness was determined. Refractory males were exposed to 16L:8D for 5 to 20 wk. Ten weeks of 16L:8D treatment was enough for the animals to regain the sensitivity to a second challenge of 12L:12D treatment. Fifteen weeks of 20L:4D or 16L:8D terminated refractoriness in female Turkish hamsters; 20L:4D therefore was not interpreted as a short day by refractory hamsters. This was unexpected because in photosensitive animals this photoperiod acts like a short day, causing gonadal regression. These results suggest that Turkish hamsters are similar to Syrian hamsters in that both species require two or more months of long days in summer to recover sensitivity to the short days of the following fall.     

Gonadotrophin secretion and pituitary responsiveness to LHRH in castrated and intact male rabbits exposed to different photoperiods

Adult male wild rabbits were exposed to at least 16 weeks of 16L:8 D before experiments began. Plasma LH and FSH concentrations increased significantly (P less than 0.001) when rabbits were castrated in 16L:8D but declined when rabbits were transferred to 8L:16D. Concentrations had returned to normal for castrated rabbits in 16L:8D by 74 days after the start of the 8L:16D treatment. Treatment of intact male rabbits with an injection of LHRH before and after transfer to short daylengths caused a transient increase in plasma LH which lasted 50-80 min and this produced a concomitant rise in plasma testosterone. The daylength change had no effect on this response even though testicular size declined after the transfer to short daylengths. Rabbits moulted in response to exposure to 8L:16D. This suggests that hypothalamic activity responds to photoperiod and that changes in pituitary responsiveness to LHRH and steroid negative feedback are unimportant.     

Photoperiodic control of the termination of reproduction in Japanese quail (Coturnix coturnix japonica)

Typically, birds come into breeding in the spring as a response to long days, and end reproduction some weeks later by becoming refractory to those long days. The refractory state is subsequently dissipated by the short days of autumn and winter, so producing once again a bird that can respond to long days. Bird species differ in the extent to which refractoriness is developed; the present experiments took advantage of the relative, rather than the absolute, refractoriness in quail to measure quantitatively the dissipation process. Quail were made refractory by exposure to long days, then transferred to short days and at various times thereafter photostimulated with longer daylengths, the degree of photoresponsiveness being assessed by measuring changes in luteinizing hormone (LH) secretion or cloacal gland size or both. The most clearcut results came from using 13L:11D as the test stimulus to measure photoresponsiveness, and this indicated, in both intact and castrated quail, no response to 13L:11D after one week of short days, a minor response after two weeks, a strong response after three weeks and a full response after five weeks. Thus refractoriness appears to be dissipated gradually under short days, and not in an all-or-none fashion. Confirmation of this conclusion came from experiments in which refractory quail were moved to short days and after one or two weeks transferred to a range of long daylengths. After one week of short days no responses were obtained to 13L:11D or 14L:10D and moderate responses only to 16L:8D, but after two weeks of short days the magnitudes of all the responses were increased.(ABSTRACT TRUNCATED AT 250 WORDS)     

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

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

Long-day inhibition of reproduction and circadian photogonadosensitivity in Zembra Island wild rabbits (Oryctolagus cuniculus).

We investigated the effects of photoperiod on testicular activity in wild rabbits (Oryctolagus cuniculus) captured on Zembra Island (North Tunisia) and maintained in experimental photoperiodic conditions. Sexually inactive animals were subjected to alternate 3-mo periods of short days (8L:16D) and long days (16L:8D) for 1 yr. Testicular activity increased significantly and then decreased to levels equivalent to or lower than those measured during sexual quiescence after 1 mo of 8L:16D or 16L:8D, respectively. Eight groups of sexually active animals were also exposed to 8L:16D for 60 days. The light phase was divided into two photofractions (7.5 and 0.5 h). The short photofraction interrupted the dark phase 9.5-18.5 h after the beginning of the main photofraction. Testicular activity was inhibited if the short photofraction interrupted the dark phase 12.5 h or more after the beginning of the main photofraction. These results clearly confirm that photoperiod affects reproduction in this species: Short days stimulate reproduction, whereas long days inhibit it. The asymmetric pattern of skeleton photoperiods used demonstrated the existence of a circadian rhythm for photogonadosensitivity, with the photosensitive phase beginning 12.5 h after dawn. In this species, photoperiod length controls both the beginning and the end of the reproductive period. These results differ from those obtained with continental populations of wild rabbits, in which reproduction is inhibited by short day length. This difference may reflect genetic drift linked to the geographic isolation of this population, which is known to have been present on this small island for more than 2000 yr.     

Sustained reproductive responses in Djungarian hamsters (Phodopus sungorus) exposed to a single long day

Male and female Djungarian hamsters maintained from birth in a short photoperiod (8 h light per day; 8L:16D) showed substantial testicular and uterine growth in response to a single long photoperiod or a 15-min light pulse that interrupted the 16-h dark period at 18 days of age. These light regimens resulted in heavier testes and uteri at 30 and 35 days of age when compared with those of control animals. Similar results were obtained in hamsters maintained from birth to Day 18 in a long photoperiod (16L:8D), given a single longer day (20L:4D) or constant light on Day 18 and then transferred to a short photoperiod (8L:16D) on Day 19. At 35 days of age animals that received extended light treatment on Day 18 had significantly more developed reproductive structures than did control hamsters. The marked effects of brief light treatment in producing long-term changes in the reproductive axis provide a convenient mammalian model system in which to study neuroendocrine events that underlie photoperiodism.     

Evidence that the onset of the breeding season in the ewe may be independent of decreasing plasma prolactin concentrations

Ten ewes of each of two breeds, Dorset Horn (long breeding season) and Welsh Mountain (short breeding season), were given subcutaneous oestradiol-17 beta implants and then ovariectomized. Another 10 ewes of each breed were left intact. On 3 May 1982, all the ewes were housed in an artificial photoperiod of 16L:8D. After 4 weeks, half of the ewes of each breed and physiological state were abruptly exposed to a short-day (8L:16D) photoperiod while the others remained in long days (16L:8D). The time of onset of the breeding season was significantly (P less than 0.05) advanced in ewes switched to short days (12 August +/- 10 days) compared to those maintained in long days (4 September +/- 14 days). Dorset Horn ewes began to cycle (20 July +/- 7 days) significantly (P less than 0.001) earlier than Welsh Mountain ewes (19 September +/- 6 days). Disparities in the time of onset of cyclic activity in ewes of different breeds and daylength groups were echoed in disparities in the time at which plasma LH and FSH concentrations rose in oestrogen-implanted, ovariectomized ewes of the same light treatment group. Prolactin concentrations showed an immediate decrease in ewes switched to short days, but remained elevated in long-day ewes. Since the breeding season started in the presence of high prolactin concentrations in long-day ewes, it seems unlikely that prolactin is an important factor determining the timing of the onset of cyclic activity.     

Photoperiodic stimulation of pubertal development in male deer mice: involvement of the circadian system

Pubertal development in prairie deer mice (Peromyscus maniculatus bairdii) is accelerated by exposure of juveniles to a long-day photoperiod, and, conversely, retarded by exposure to short days. The purpose of the present study was to evaluate the possible involvement of the circadian system in the photoperiodic regulation of puberty. Weanling males, previously housed on a short-day light cycle of 6L:18D, were subjected to a "resonance" protocol in which they received one of the following light cycles: 6L:18D, 6L:30D, 6L:42D, 6L:54D, or 16L:8D. Post-weaning exposure to cycles of 16L:8D, 6L:30D, and 6L:54D stimulated reproductive organ growth as measured at 6 weeks of age. Exposure to cycles of 6L:18D and 6L:42D failed to stimulate reproductive development. These data support the hypothesis that young male deer mice use a circadian rhythm of responsiveness to light to measure photoperiodic time and, consequently, regulate pubertal development.     

Social cues attenuate photoresponsiveness of the male reproductive system in Siberian hamsters (Phodopus sungorus)

Transfer of adult Siberian hamsters (Phodopus sungorus) from long (16 h light and 8 h dark, 16L:8D) to short (8L:16D) daily photoperiods induces an involution of the gonads and a cessation of reproductive behavior 8 to 10 weeks later. However, when male and female long-day hamsters were paired on transfer to short photoperiods, the males' gonads did not undergo the typical short-day response. Similarly, when male long-day hamsters were paired with refractory females (i.e., females housed in short photoperiods for at least 28 weeks so that they became unresponsive to short photoperiods), the response of the males' reproductive system to short photoperiods also was attenuated. Thus, social cues can override or delay the effects of photoperiod on the testes of this species. These results suggest that the inhibitory effects of long durations of melatonin secretion (in response to short photoperiods) on the male hypothalamic-pituitary-gonadal axis may be attenuated by social cues such as contact with the opposite sex.     

The effect of duration and time of food availability on the photoperiodic response in the male house sparrow, Passer domesticus

We investigated the effects of food availability on the seasonal testicular growth in the photoperiodic house sparrow (Passer domesticus). Two experiments were performed, each lasting 4 weeks. In experiment 1, sparrows were exposed to natural (NDL; group 1), short (8L:16D; group 2) and long (16L:8D; groups 3-5) day lengths with access to food ad libitum (groups 1-3) or for 10 h (zeitgeber time (zt) 0-10, group 4; zt 0 is the time of light onset) or for 8 h (zt 8-16, group 5). Testes recrudesced under long, but not short and natural, day lengths, and the recrudescence under long days was influenced by the duration of food availability. In experiment 2, the sparrows were exposed to short (8L:16D, group 1) and long (16L:8D, groups 2-5) day lengths with access to food ad libitum (for groups 1 and 2) or for 6 h (for groups 3-5) at different times during the 16 h light period (group 3- zt 0-6, group 4- zt 5-11, group 5- zt 10-16). As the expected, the testes recrudesced only under long lengths, but the photoinduction was variable among the 4 groups. The testes grew to full size in groups 2 and 3 that received food either ad libitum or for 6 h at zt 0-6, but to sub-maximal size in the groups that received 6 h food either at zt 5-11 (group 4) or at zt 10-16 (group 5). Altogether, these results support the idea that the photoperiodic regulation of reproduction in a seasonally breeding species is influenced both by the duration and the time of food availability.     

Photoperiodic response of diapause induction in the pine caterpillar, Dendrolimus punctatus

The pine caterpillar, Dendrolimus punctatus (Walker) (Lepidoptera: Lasiocampidae), is a multivoltine pest of pine trees in China, overwintering as larvae. Winter diapause was induced by short day length. The critical night length was about 10 h 40 min at 25, 28, and 31 °C in the field, showing a temperature‐compensated diapause induction. Transfer experiments from a short night (L16:D8) to a long night (L12:D12) or vice versa at different times after hatching showed that sensitivity to day length was restricted to the first 14 days; the required day number for a 50% response at 25 °C was about 3.5 days for short nights but 7.5 days for long nights, indicating that short nights are photoperiodically more effective. When four successive short nights (L16:D8) were used to interrupt the long‐night regime (L12:D12) at different development stages and vice versa, the results showed that the highest sensitivity to photoperiod occurred on the 4th?8th day, corresponding to the second larval instar. Experiments of alternating short‐night (L16:D8) and long‐night (L12:D12) cycles during the larval period showed that the information of short nights as well as long nights could be accumulated. By rearing the larvae under conditions other than 24‐h light–dark cycles, we clearly showed that the dark period (scotophase) played a major role in the determination of diapause. The Nanda‐Hamner and Bünsow experiments failed to reveal rhythmic fluctuations with a period of about 24 h in the occurrence of diapause. Therefore, the photoperiodic clock in D. punctatus is an hourglass timer or a damped circadian oscillator.     

Effect of photoperiod length on body mass and testicular growth in the house sparrow (Passer domesticus) and brahminy myna (Sturnus pagodarum)

Two experiments studied the relative effects on body mass and testicular growth of stimulatory photoperiods applied simultaneously to two photosensitive species, the house sparrow (Passer domesticus) and brahminy myna (Sturnus pagodarum). Experiment 1 on the house sparrow consisted of two parts. In experiment 1A, beginning on 24 March 2002, short day pretreated sparrows were exposed for 12 weeks to 13L: 11D (13 h light: 11 h darkness), 20L: 4D and NDL (control). Experiment 1B was similar to 1A except that it used sparrows that were not treated with short days. This experiment was repeated at three different times in the year. Beginning on 29 December 2002 (for 24 weeks), 26 March 2003 (for 12 weeks) and 16 August 2003 (for 8 weeks), sparrows captured from the wild and acclimated to captive condition for 1 week were exposed to 13L: 11D and 20L: 4D. Each time, a group was maintained in NDL and served as the control. Experiment 2 was performed on myna and used an identical protocol. Beginning on 24 March 2002, myna that were captured from the wild and acclimated to captivity conditions were exposed for 16 weeks to 13L: 11D and 20L: 4D; a group was maintained in NDL and served as the control. There was photostimulation and subsequent regression of the testes on all day lengths except in the August group of experiment 1B. The effect on body mass was variable. Interestingly, however, the response to 20L:4D was relatively smaller as compared to 13L:11D. Taken together, these results confirm that the two species use photoperiods in control of their reproductive cycle, and tend to indicate that exposure to unnatural long photoperiods may in fact be unfavorable and could compromise gonadal growth and development.     

Altered light conditions during spring: effects on timing of migration and reproduction in migratory redheaded bunting (Emberiza bruniceps)

Photoperiod is the most consistent environmental cue, and therefore, any change in anticipated light environment may affect subsequent response under long days. To test this hypothesis, photosensitive migratory redheaded buntings (Emberiza bruniceps) were exposed to altered photoperiods for 4 weeks [short days (SP, 8L:16D, group 1; control), short days alternated with constant dim light (SP/LL_dim; group 2), constant dim light (LL_dim; group 3) and constant bright light (LL_bright; group 4)] before being transferred to long days (LP; 16L:8D) for 15 weeks. Group differences in long-day-induced responses were observed. The onset of migratory restlessness (Zugunruhe) was earliest in LL_bright but persisted for maximum period in LL_dim group. The LL_dim group attained peak testis size significantly delayed and had more food consumption under long days. The results suggest that the altered photoperiodic exposure during photosensitive stage affects the seasonal phenotypes such as migration and reproduction in migratory buntings.     

Regulation of seasonality in the migratory male blackheaded bunting (Emberiza melanocephala)

The present study was carried out on a Palearctic-Indian migratory species, the blackheaded bunting (Emberiza melanocephala), to understand the importance of photoperiodism and circannual rhythms in determining seasonality in changes in body mass and testis size in birds. An initial experiment determined the effects of duration and intensity of light on photoperiodic induction. The birds were exposed to different photoperiods (hours of light:hours of darkness; 11.5L:12.5D, 12L:12D, 12.5L:11.5D and 13L:11D) at the same (approximately 450 lux) light intensity, and to 13L:11D at different light intensities (50-, 100-, 400-, 800- and 1000-lux). The induction and subsequent regression of photoperiodic responses were dependent upon duration and intensity of the light period until these reached threshold. A second experiment investigated if an endogenous seasonal rhythm underlies photoperiodism in buntings. Birds maintained since February on a 8L: 16D photoperiod (a non-inductive short day length invariably used to ensure photosensitivity in photoperiodic species) were subjected periodically to 16L:8D (a long day length), one group every month from mid-March to mid-August. The magnitude of long day response in body mass and testes decreased as the duration of the short days progressed, but testicular response was restored in birds that were exposed to long days in July and August. The birds exposed simultaneously to short, long, and natural day lengths for 32 weeks underwent an induction-regression cycle under long days and natural day lengths, but not under short days in which a decrease in body mass occurred after about 20 weeks. The last experiment examined the importance of latitudinal migration on photoperiodism, by comparing the response to long days of three groups which included birds from populations those were held in the outdoor aviary for 1 or 2 years at 27 degrees N and those immediately arrived from their breeding grounds (approximately 40 degrees N). There was no difference in the photoperiodic induction among the three groups, indicating that neither experience to changing photoperiods during a migratory journey, nor to long photoperiods at breeding grounds, were critical for a subsequent response (initiation-termination-reinitiation) cycle. Taken together, these findings suggest that (1) the blackheaded bunting has its own endogenous timing program, which is regulated by the photoperiod, and (2) the photoperiodic programs of bunting are flexible enough to accommodate variations in the amplitude of environmental cycles. Thus, it appears that photoperiodism has evolved independently of the evolution of migration in this species.     

Photoperiodic variations of somatic and germ cell populations in the Soay ram testis

Seasonal changes in the Leydig, Sertoli and stem cells (undifferentiated A0 and cyclic A1) spermatogonia and the daily spermatid production were analysed in the testes of adult Soay rams exposed to short days (8L:16D) or long days (16L:8D) for 12 weeks. The total numbers of Leydig, Sertoli and stem cells (A0 + A1) were not affected by the treatments, but the size of the Leydig and the Sertoli cells, the efficiency of spermatogenesis (i.e. the number of male gametes produced by an A1 spermatogonium) and the daily sperm production were all significantly reduced in the rams exposed to long days. There was a positive correlation between the concentration of FSH and testosterone and many of the cytological changes consistent with a causal role for these hormones in mediating the effects of photoperiod on the testicular function in the ram.     

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

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

光照和温度对根田鼠褐色脂肪组织产热能力的影响

测定了根田鼠在室内驯化2周后褐色脂肪组织（BAT）的蛋白含量、线粒体蛋白含量和细胞色素C氧化酶活性的变化。动物进行如下4种处理:23℃,16L:8D;23℃,8L：16D;5℃,16L：8D和5℃,8L:16D.。结果表明,短光照可刺激根田鼠的BAT线粒体细胞色素C氧化酶的活性增加;低温驯化可导致根田鼠的BAT线粒体蛋白含量增加;短光照加低温可进一步促进酶的活性增加。结合个体水平的产热特点,说明根田鼠在产热能力的季节驯化过程中,光周期是一主要的季节调节信号,环境温度进一步增强光照的作用,环境温度和光周期共同作用以诱导其产热能力等方面的生理调节。研究结果支持了环境温度和光周期共同作用以调节动物产热变化的学说。     

Seasonal modulation of diurnal food consumption in Indian songbirds

Seasonal changes in daily food consumption have a direct bearing with energy requirement of bird that is in turn associated with life history stage of birds. We compared seasonal changes in daily food intake in adult male migratory redheaded bunting (Emberiza bruniceps) that over winters in Indian subcontinent with those in non-migratory blackheaded munia to reiterate the same. We also compared daily food eating pattern (DFEP) in wintering blackheaded and redheaded buntings, closely related Emberizidae finches to establish circadian nature of feeding behavior and how it varied at species level. The birds were held under short days (8L:16D; 8 h of light and 16 h of darkness) and two hourly food consumption was measured to profile their DFEP. Further, we extended the study to establish how the circadian pattern of food consumption varied depending on birds’ physiological state and effect of photoperiod in adult male redheaded buntings. Redheaded buntings DFEP and locomotor activity were compared in pre-migratory months of February (spring) and September (autumn). The results suggest that September (photorefractory) birds exhibit clear bimodality in their feeding behavior as compared to (photosensitive) birds in February. Another experiment compared bird’s DFEP held under short (8L:16D) and long (16L:8D) days for 5 weeks and suggested that under long days, prolonged hours of photophase render adaptive advantage to birds for positive energy budgeting. The present study clearly establishes the circadian nature of feeding behavior and that it modulates over seasons. The bimodal i.e. morning and evening peaks of food consumption suggest morning–evening food entrainable oscillators, however this needs to be investigated with mechanistic approach in future studies.     

Effect of intranasal irrigation with zinc sulphate on male-induced oestrus in the wild mouse, Mus musculus domesticus

Adult males of the subtropical house sparrow (Passer domesticus) were subjected to continuous darkness (DD) or short days (8L:16D) after photostimulation (15L:9D) for 30 days. One group was retained on the long day (15L:9D) schedule. Another two groups of photosensitive birds were placed under DD or 8L:16D. The photostimulated birds under DD or 8L:16D showed testicular regression and the rate of regression was similar in both the treatments. The photosensitive birds, on the other hand, did not show any initiation in testicular growth and development in either of the treatments. Long day birds (15L:9D) maintained enlarged testes without regression (the onset of photorefractoriness) during the treatment period. These results conform with the external coincidence model, but could also be explained by a modified internal coincidence model.     

Short days induce premature reproductive maturation in juvenile starlings, Sturnus vulgaris

Castration of juvenile and photorefractory adult starlings caused no immediate increase in circulating concentrations of LH. In castrated juveniles and adults exposed to natural changes in daylength, plasma LH increased between mid-October and mid-November, although the increase was more rapid in adults. In castrated photorefractory adults, plasma LH increased 3-5 weeks after transfer to artificial short days (8L:16D). In castrated juvenile starlings plasma LH increased 4-6 weeks after transfer to 8L:16D, irrespective of the age of the birds. Birds as young as 17 weeks had high LH concentrations. These results suggest that the reproductive system of juvenile starlings is in the same state as that of photorefractory adults, and therefore the activation of the hypothalamo-pituitary axis for the first time in juveniles is analagous to the termination of photorefractoriness in adults.