Timing of puberty and synchronization of seasonal rhythms by simulated natural photoperiods in female Siberian hamsters

The timing of puberty is a critical life history trait of short-lived species; spring-born individuals mature rapidly and breed in the season of birth, whereas young born in mid- to late summer delay puberty until the next spring. The cues that govern the transition from rapid to delayed maturation in natural populations remain unknown. To identify ecologically relevant photoperiod cues that control timing of puberty, we monitored nine cohorts of female Siberian hamsters (Phodopus sungorus) born every 2 wk from 4 wk before to 12 wk after the summer solstice in a simulated natural photoperiod (SNP). Hamsters born by the summer solstice underwent rapid somatic growth and achieved puberty that summer; among females born 2-4 wk after the solstice, some delayed puberty by many weeks, whereas others manifested early puberty. Hamsters born 6 or more weeks after the solstice generally delayed puberty until the following spring. The transition from accelerated to delayed pubertal development in the SNP occurred at day lengths that induce early puberty when presented as static photoperiods. Despite differences in timing of birth and timing of puberty, fall and subsequent spring seasonal events occurred at similar calendar dates in all cohorts. We found no evidence that prenatal photoperiod history influenced postnatal development of female hamsters. Considered together with a parallel study on males, the present findings point to sex differences in responsiveness to natural photoperiod variations. In both sexes, incrementally changing photoperiods exert a strong organizing effect on seasonal rhythms.     

Differential effects of multiple short day lengths on body weights of gonadectomized siberian hamsters

Siberian hamsters (Phodopus sungorus) maintained under simulated natural photoperiods exhibit marked reductions in body weight as day lengths decrease in summer and fall. This experiment assessed whether the component of the seasonal body weight rhythm that is independent of gonadal hormones exhibits a graded dependence on decreasing day lengths or whether the entire program of weight loss is triggered by the crossing of a single critical day length in late summer. Male hamsters born into a photoperiod with 13 h light and 11 h dark (i.e., 13L : 11D) were castrated and transferred to simulated natural photoperiod for early April at 40 degrees N latitude. At the summer solstice (15L : 9D) some hamsters remained on that photoperiod whereas others experienced gradual decreases in day length. Three additional groups were moved to static photoperiods when day lengths had subsequently declined to 13L : 11D, 11L : 13D, or 9L : 15D, respectively. Day lengths decreasing to 13L : 11D were sufficient to suppress body weight but were less inhibitory than further decreases in day length. Hamsters identified as photononresponsive on the basis of daily activity rhythms increased body weight monotonically. These results establish that steroid-independent modulation of body weight depends on photoperiod in a graded fashion.     

A plastic interval timer synchronizes pubertal development of summer- and fall-born hamsters

Summer and fall decreases in day length induce reproductive regression in adult hamsters and delay reproductive maturation of their young. The following year pubertal development is triggered by an interval timer (IT) that renders animals refractory to inhibitory short day lengths after approximately 25 wk. Timing of gonadal and somatic development was examined among offspring born to Siberian hamsters in early-August vs. late-September day lengths. Pubertal maturation was delayed in both groups until late winter. Gonadal growth occurred at significantly later ages among August- vs. September-born males as did late-winter spurts in ponderal growth of both sexes. Timing of reproductive and somatic development depended on postnatal rather than prenatal photoperiod exposure and was unrelated to the circadian entrainment status of dams. When developmental patterns were assessed in relation to time of year, group differences were largely eliminated. Because the IT triggers these developmental events, its duration must be plastic. This plasticity facilitates a relative synchronization or entrainment of developmental milestones in hamsters born into different late-summer/early-fall photoperiods.     

Photoperiodic influences on ultradian rhythms of male siberian hamsters

Seasonal changes in mammalian physiology and behavior are proximately controlled by the annual variation in day length. Long summer and short winter day lengths markedly alter the amplitude of endogenous circadian rhythms and may affect ultradian oscillations, but the threshold photoperiods for inducing these changes are not known. We assessed the effects of short and intermediate day lengths and changes in reproductive physiology on circadian and ultradian rhythms of locomotor activity in Siberian hamsters. Males were maintained in a long photoperiod from birth (15 h light/day; 15 L) and transferred in adulthood to 1 of 7 experimental photoperiods ranging from 14 L to 9 L. Decreases in circadian rhythm (CR) robustness, mesor and amplitude were evident in photoperiods ≤14 L, as were delays in the timing of CR acrophase and expansion of nocturnal activity duration. Nocturnal ultradian rhythms (URs) were comparably prevalent in all day lengths, but 15 L markedly inhibited the expression of light-phase URs. The period (τ'), amplitude and complexity of URs increased in day lengths ≤13 L. Among hamsters that failed to undergo gonadal regression in short day lengths (nonresponders), τ' of the dark-phase UR was longer than in photoresponsive hamsters; in 13 L the incidence and amplitude of light-phase URs were greater in hamsters that did not undergo testicular regression. Day lengths as long as 14 L were sufficient to trigger changes in the waveform of CRs without affecting UR waveform. The transition from a long- to a short-day ultradian phenotype occurred for most UR components at day lengths of 12 L-13 L, thereby establishing different thresholds for CR and UR responses to day length. At the UR-threshold photoperiod of 13 L, differences in gonadal status were largely without effect on most UR parameters.     

European hamsters (Cricetus cricetus) show a transient phase of insensitivity to long photoperiods after gonadal regression

Annual rhythms of body weight and reproduction in the European hamster (Cricetus cricetus) are the result of an interaction between seasonal changes in day length (photoperiod) and seasonal changes in the responsiveness of animals to these photoperiods. The present study demonstrates that under natural conditions European hamsters are not able to perceive long photoperiods (i.e., a 16L:8D cycle) before mid-November. This is an important difference to other hamster species, in which regrowth of the gonads can be stimulated by exposure to long photoperiods at any stage of gonadal regression. The experiments also demonstrate the existence of an annual phase of sensitivity to long photoperiods that starts around mid-November and extends until March/April. During this phase of sensitivity, exposure to a long photoperiod (16L:8D) induced gonadal regrowth within 3 wk. Additional experiments with an accelerated photoperiodic lighting regimen indicated that a photoperiod of approximately 13 h is necessary to stimulate gonadal regrowth. Under natural light conditions in Stuttgart (48.46 degrees N), a photoperiod of 13 h is reached by the beginning of April, which fits well with the finding that the majority of animals kept under a natural light:dark cycle had well-developed gonads by the end of April. Nevertheless, these animals showed a rather variable timing of gonadal regrowth, ranging from early January to late April. This is most likely the result of two processes: first, an endogenous mechanism (photorefractoriness) that induces gonadal recrudescence without any photoperiodic information while the animals are still in their hibernation burrows, and second, a direct stimulatory effect of long photoperiods.     

Interval timer control of puberty in photoinhibited Siberian hamsters

Puberty, which is markedly delayed in male Siberian hamsters (Phodopus sungorus) born into short day lengths, is controlled by an interval timer regulated by the duration of nocturnal melatonin secretion. Properties of the interval timer were assessed by perturbing normal patterns of melatonin secretion in males gestated and maintained thereafter in 1 of 2 short day lengths, 10 h light/day (10 L) or 12L. Melatonin secretion of short-day hamsters was suppressed by constant light treatment or modified by daily injection of propranolol to mimic nocturnal melatonin durations typical of long-day hamsters. Constant light treatment during weeks 3 to 5 induced early incomplete gonadal growth in 12L but not 10 L hamsters but did not affect late onset of gonadal development indicative of puberty in either photoperiod. Propranolol treatment during postnatal weeks 3 to 5 induced transient growth of the testes and ultimately delayed the timing of puberty by 3 weeks. Similar treatments between weeks 5 and 7 or on alternate weeks for 24 weeks did not affect the interval timer. The first 2 weeks after weaning may constitute a critical period during which the interval timer is highly responsive to photoperiod. Alternatively, the hamsters' photoperiodic history rather than age or developmental stage may be the critical variable. The interpolation of long-day melatonin signals at the time of weaning does not appear to reset the interval timer to its zero position but may reduce timer responsiveness to long-day melatonin signals several weeks later.     

Exogenous T3 mimics long day lengths in Siberian hamsters

Siberian hamsters (Phodopus sungorus) exhibit seasonal cycles of reproduction driven by changes in day length. Day length is encoded endogenously by the duration of nocturnal melatonin (Mel) secretion from the pineal gland. Short-duration Mel signals stimulate reproduction and long-duration signals inhibit reproduction. The mechanism by which Mel signals are decoded at the level of neural target tissues remains uncharacterized. In Siberian hamsters, exposure to short day lengths or injections of Mel in long days results in a decrease in hypothalamic expression of type 2 iodothyronine deiodinase (Dio2) mRNA. Dio2 catalyzes the conversion of the thyroid hormone thyroxine to triiodothyronine (T3). Thus exposure to short and long day lengths should decrease and increase hypothalamic T3 concentrations, respectively. We tested the hypothesis that exogenous T3 administered to short-day hamsters would mimic exposure to long day lengths with respect to gonadal stimulation. Hamsters gestated and raised in short day lengths that exhibited photoinhibition of the testes were given daily subutaneous injections of T3 or saline vehicle for 4 wk beginning at week 12 of life. The results indicate that exogenous T3 induced gonadal growth in short-day hamsters and delayed spontaneous gonadal development by an interval equal to the number of weeks during which T3 was administered. T3 injections delayed gonadal regression if given coincident with the transfer of hamsters from long to short day lengths. These results suggest that T3 mimics long day exposure in Siberian hamsters and may serve as an intermediate step between the Mel rhythm and the reproductive response.     

Perinatal photoperiod organizes adult immune responses in Siberian hamsters (Phodopus sungorus)

Individuals of many nontropical rodent species display reproductive, immunological, and somatic responses to day length. In general, short day (SD) lengths inhibit reproduction and enhance immune function in the laboratory when all other conditions are held constant. Most studies to date have focused on seasonal variation in immune function in adulthood. However, perinatal photoperiods also communicate critical day length information and serve to establish a developmental trajectory appropriate for the time of year. Nontropical rodents born early in the breeding season undergo rapid reproductive development, presumably to promote mating success during their first reproductive season. Rodents born late in the breeding season suspend somatic growth and puberty until the following vernal breeding season. We tested the hypothesis that perinatal day lengths have similar enduring effects on the immune system of rodents. Siberian hamsters (Phodopus sungorus) were maintained prenatally and until weaning (21 days) in either SDs (8 h light:16 h dark) or long days (LD) (16 h light:8 h dark), then they were weaned into either the opposite photoperiod or maintained in their natal photoperiod, forming four groups (LD-LD, LD-SD, SD-LD, and SD-SD). After 8-wk in these conditions, cell-mediated immune activity was compared among groups. SD-SD hamsters of both sexes enhanced immune function relative to all other groups. The reproductive effects of perinatal photoperiod were not evident by the end of the experiment; circulating testosterone and cortisol sampled at the end of the experiment reflected the postweaning, but not the perinatal photoperiod. This experiment demonstrates long-lasting organizational effects of perinatal photoperiod on the rodent immune system and indicates that photoperiod-induced changes in the immune system are dissociable from changes in the reproductive system.     

Maternal photoperiodic history affects offspring development in Syrian hamsters

During the first 7 weeks of postnatal life, short day lengths inhibit the onset of puberty in many photoperiodic rodents, but not in Syrian hamsters. In this species, timing of puberty and fecundity are independent of the early postnatal photoperiod. Gestational day length affects postnatal reproductive development in several rodents; its role in Syrian hamsters has not been assessed. We tested the hypothesis that cumulative effects of pre- and postnatal short day lengths would restrain gonadal development in male Syrian hamsters. Males with prenatal short day exposure were generated by dams transferred to short day lengths 6 weeks, 3 weeks, and 0 weeks prior to mating. Additional groups were gestated in long day lengths and transferred to short days at birth, at 4 weeks of age, or not transferred (control hamsters). In pups of dams exposed to short day treatment throughout gestation, decreased testis growth was apparent by 3 weeks and persisted through 9 weeks of age, at which time maximum testis size was attained. A subset of males (14%), whose dams had been in short days for 3 to 6 weeks prior to mating displayed pronounced delays in testicular development, similar to those of other photoperiodic rodents. This treatment also increased the percentage of male offspring that underwent little or no gonadal regression postnatally (39%). By 19 weeks of age, males housed in short days completed spontaneous gonadal development. After prolonged long day treatment to break refractoriness, hamsters that initially were classified as nonregressors underwent testicular regression in response to a 2nd sequence of short day lengths. The combined action of prenatal and early postnatal short day lengths diminishes testicular growth of prepubertal Syrian hamsters no later than the 3rd week of postnatal life, albeit to a lesser extent than in other photoperiodic rodents.     

The timing of neuroendocrine sexual maturity in the male lamb by photoperiod

In spring-born female lambs, the long days of summer, followed by their gradual decrease, provide the seasonal cue necessary to time puberty to early autumn (approximately 30 wk of age). Male lambs begin spermatogenesis during mid-summer, some 20 wk before puberty occurs in females. Unlike young female lambs, male lambs attain puberty at the same age under a variety of photoperiodic manipulations, raising the possibility that sexual maturation in males is not affected by photoperiod. We have reinvestigated the role of photoperiod on puberty in the male lamb, using a more precise indicator of reproductive activation--the decreased sensitivity of the hypothalamo-pituitary axis to inhibitory steroid feedback leading to increased LH secretion. To test whether photoperiod can influence the onset of neuroendocrine sexual maturation in male lambs, this study compared the timing of the decrease in sensitivity to inhibitory steroid feedback in two groups of males under opposite photoperiodic conditions. Eight males were reared indoors from 2 wk of age under conditions simulating the natural increasing and decreasing day lengths around the summer solstice; an additional 7 males were exposed to a reversed simulated natural photoperiod in which the changes in day length were amplified and accelerated relative to outdoor conditions. Both groups of lambs were castrated and received s.c. implants of Silastic estradiol capsules to provide a constant steroid feedback signal. The timing of reduction in sensitivity to estradiol negative feedback, measured as a sustained increase in circulating of LH above 1.0 ng/ml, was used to define neuroendocrine sexual maturity.(ABSTRACT TRUNCATED AT 250 WORDS)     

Functional maturation of the gonads of Turkish hamsters under various photoperiods

The golden hamster, Mesocricetus auratus, is the only photoperiodic rodent to date that has been shown to fail to respond to inhibitory (i.e., short, less than 12.5 h/day) photoperiods until after pubertal onset. In other photoperiodic hamsters, mice, and voles, short photoperiods greatly retard gonadal maturation. The Turkish hamster, Mesocricetus brandti, is a photoperiodic rodent that as an adult is reproductively competent only on photoperiods of 15-17 h of light per day; photoperiods of less than 15 or greater than 17 h of light promote gonadal regression. In this report we addressed two questions: a) are prepubertal M. brandti photoperiodic, and b) if so, is gonadal maturation enhanced or suppressed by exposure to photoperiods of greater than 17 h of light per day? Turkish hamsters were raised on photoperiods of 12, 16, 20, or 24 (= LL) h of light per day. Testicular growth was retarded for 16 wk by 12L:12D. Very long days, 20L:4D, or LL did not retard testicular development. In females, pubertal onset, as indicated by first vaginal estrus, was delayed in young raised on 12L:12D and in 2 of 18 and 4 of 19 young raised on 20L:4D and LL, respectively. These results demonstrate that prepubertal Turkish hamsters are photoperiodic, but respond differently from adults to photoperiods greater than 17 h of light per day.     

Photorefractoriness and energy availability interact to permit facultative timing of spring breeding

In seasonally breeding mammals, vernal reproductive development is not directly triggered by increases in day length, rather, an endogenous program of photorefractoriness to short winter days initiates spontaneous development in advance of spring. The transition to the reproductive phenotype is energetically demanding. How food availability in late winter and early spring impacts the onset and expression of photorefractoriness is not known. In this study, male Siberian hamsters were born into a simulated natural photoperiod, and at the winter solstice, they were subjected to a restricted feeding protocol in which a daily food ration was provided in an amount equal to ad libitum (AL) intake during the weeks preceding the solstice. Over the next several months, AL-fed control hamsters exhibited spontaneous recrudescence or spontaneous development. In contrast, vernal reproductive development was abolished in most food-rationed hamsters. In food-rationed hamsters that did exhibit recrudescence, conspicuous delays in the onset of gonadal development and decreases in the magnitude of growth were evident. In all hamsters, the termination of food rationing triggered rapid gonadal development. The data indicate that late winter/early spring increases in environmental food availability are required for the normal manifestation of photorefractoriness-induced reproductive development and suggest that a function of photorefractoriness may be merely to disinhibit the reproductive axis from photoperiodic suppression. Vernal gonadal development or recrudescence appears to be strongly affected by proximate energy availability.     

Seasonal regulation of reproduction: altered role of melatonin under naturalistic conditions in hamsters

The seasonal reproductive cycle of photoperiodic rodents is conceptualized as a series of discrete melatonin-dependent neuroendocrine transitions. Least understood is the springtime restoration of responsiveness to winter-like melatonin signals (breaking of refractoriness) that enables animals to once again respond appropriately to winter photoperiods the following year. This has been posited to require many weeks of long days based on studies employing static photoperiods instead of the annual pattern of continually changing photoperiods under which these mechanisms evolved. Maintaining Siberian hamsters under simulated natural photoperiods, we demonstrate that winter refractoriness is broken within six weeks after the spring equinox. We then test whether a history of natural photoperiod exposure can eliminate the requirement for long-day melatonin signalling. Hamsters pinealectomized at the spring equinox and challenged 10 weeks later with winter melatonin infusions exhibited gonadal regression, indicating that refractoriness was broken. A photostimulatory effect on body weight is first observed in the last four weeks of winter. Thus, the seasonal transition to the summer photosensitive phenotype is triggered prior to the equinox without exposure to long days and is thereafter melatonin-independent. Distinctions between photoperiodic and circannual seasonal organization erode with the incorporation in the laboratory of ecologically relevant day length conditions.     

Amplitude modulation of the nightly melatonin rise in the neonatal lamb and the subsequent timing of puberty

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)     

Short day lengths delay development of the SNB neuromuscular system in the Siberian hamster,Phodopus sungorus

The Siberian hamster, Phodopus sungorus, breeds seasonally. In the laboratory, the seasonal breeding can be controlled by photoperiod, which affects the durations of nightly melatonin secretions. Winterlike short day lengths induce gonadal regression in adult animals, and pups born and maintained in short days undergo gonadal development much later than animals born into long days. The spinal nucleus of the bulbocavernosus (SNB) and its target muscles, the bulbocavernosus (BC) and levator ani (LA), comprise a sexually dimorphic, androgen-sensitive neuromuscular system involved in male reproduction. The SNB neuromuscular system was studied in male Siberian hamsters maintained from conception in short-day (8:16 h light/dark cycle) versus long-day (16:8 h light/dark cycle) conditions. At 40–47 days of age, development of three components of the SNB neuromuscular system were all significantly delayed in hamsters raised in the short photoperiod: BC/LA muscle weight, the size of SNB motoneuronal somata, and the area of the neuromuscular junctions at the BC/LA muscles of short-day hamsters were each significantly reduced relative to those of long-day counterparts. Thus, development of the SNB reproductive system is delayed under short day lengths in this species. © 1998 John Wiley & Sons, Inc. J Neurobiol 35: 355–360, 1998     

Gonadal hormone-dependent and -independent regulation of immune function by photoperiod in Siberian hamsters

Siberian hamsters (Phodopus sungorus) exhibit changes in reproductive and immune function in response to seasonal variations in day length. Exposure to short days induces gonadal regression and inhibits testosterone secretion. In parallel, short days enhance immune function: increasing leukocyte numbers and attenuating cytokine and behavioral responses to infection. We examined whether photoperiodic changes in leukocyte phenotypes and sickness behaviors are dependent on concurrent photoperiodic changes in gonadal function. Male hamsters were gonadectomized or sham-gonadectomized and either exposed to short days (9 h light/day; SD) or kept in their natal long-day (15 h light/day; LD) photoperiod for 10-13 wk. Blood samples were obtained for leukocyte enumeration, and hamsters were challenged with bacterial LPS, which induced behavioral (anorexia, reductions in nest building) and somatic (weight loss) sickness responses. Among gonad-intact hamsters, exposure to SD increased total and CD62L+ lymphocytes and CD3+ T lymphocytes in blood and significantly attenuated LPS-induced sickness responses. Independent of photoperiod, castration alone increased total and CD62L+ lymphocyte and CD3+ T lymphocyte numbers and attenuated somatic and anorexic sickness responses. Among castrated hamsters, SD exposure increased lymphocyte numbers and suppressed sickness behaviors. In castrated hamsters, the magnitude of most immunological effects of SD were diminished relative to those evident in gonad-intact hamsters. The SD phenotype in several measures of immunity can be instated via elimination of gonadal hormones alone; however, photoperiodic effects on immune function persist even in castrated hamsters. Thus, photoperiod affects the immune system and neural-immune interactions underlying sickness behaviors via gonadal hormone-dependent and -independent mechanisms.     

Short day lengths delay reproductive aging

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.     

Transient effects of photoperiod on reproduction in the Mexican bean beetle

ABSTRACT. Mexican bean beetles (Coccinellidae: Epilachna varivestis (Mulsant)) have an adult diapause and show a graded response to photoperiods intermediate between those that lead to the fastest gonadal maturation and those that lead to diapause. The rate of maturation is reduced by a few to several days at day lengths corresponding to dates after the summer solstice, and such delays in reproduction may be of ecological and evolutionary importance. Multinomial probit analysis is demonstrated as a method for studying more than two classes of response to a photoperiodic regime.     

Temperature dependence of gonadal regression in Syrian hamsters exposed to short day lengths

We sought to determine whether ambient temperature (T(a)) affects gonadal function by altering the rate at which circadian rhythms entrain to short day lengths. Syrian hamsters were housed in cages where they received 14 h of light per day ("long days," 14L) at 22 degrees C. Hamsters were then transferred to cages to receive 10 h of light per day ("short days," 10L) and kept at 5, 22, or 28 degrees C or were maintained in 14L at 22 degrees C. Body mass and estimated testis volume as well as duration of nocturnal locomotor activity (alpha), previously established as a reliable indicator of the duration of nocturnal melatonin secretion, were determined over the course of 24 wk. Testicular regression in short days was accelerated by 4 wk at 5 degrees C and delayed by 3 wk at 28 degrees C relative to 22 degrees C. The interval between alpha-expansion and initiation of testicular regression was markedly affected by T(a) with delays of 0, 3, and 6 wk at 5, 22, and 28 degrees C, respectively. All hamsters held at 5 and 22 degrees C underwent testicular regression, but 25% of those maintained at 28 degrees C failed to do so. We suggest that T(a) modulates testicular regression primarily by affecting responsiveness of neuroendocrine target tissues to long melatonin signals.     

Photoperiod and melatonin influence seasonal gonadal cycles in the grasshopper mouse (Onychomys leucogaster)

Development of the reproductive apparatus was delayed in grasshopper mice maintained from birth in short photoperiods (10 h light/day). The inhibitory effects of short photoperiods on sexual maturation eventually waned and mice in 10L:14D became reproductively active. Adult mice transferred from long (14 h light/day) to short photoperiods underwent testicular regression after 10 weeks and complete gonadal redevelopment after 30 weeks. A similar phenomenon was observed in adult female mice; oestrous cycles ceased within 3 weeks and resumed after 13 weeks in the short photoperiod. The regressive effects of short photoperiods on the male reproductive system were mimicked by daily injections of melatonin administered to mice housed in 14L:10D. Responsiveness of the female reproductive system to melatonin was reduced among photorefractory as compared to photosensitive mice. We suggest that the initial rate of sexual maturation and the timing of seasonal breeding in adult mice are regulated by photoperiod; effects of short daylengths on the neuroendocrine-reproductive axis appear to be mediated by the pineal gland.