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The dorsomedial nucleus (DMN) of the hypothalamus, the only site within the mediobasal hypothalamus of Syrian hamsters that both binds melatonin and has abundant concentrations of androgen receptors, has been proposed as a target tissue for induction of seasonal changes in brain sensitivity to steroid negative feedback. We tested whether DMN ablation, which does not interfere with pineal gland secretion of melatonin in short day lengths, prevents testicular regression by altering sensitivity to steroid negative feedback. Hamsters with DMN lesions, unlike control hamsters, failed to undergo testicular regression after transfer from a long (14 h light/day) to a short day length (8 h light/day); however, increased negative-feedback inhibition of follicle-stimulating hormone by testosterone was not compromised by ablation of the DMN, indicating that this tissue is not an essential mediator of seasonal changes in feedback sensitivity. We propose a redundant neural network comprised of multiple structures, each of which contributes to neuroendocrine mechanisms, that determines the effect of short days on gonadal function.

Long days stimulate and short days (SDs) inhibit the reproductive axis of photoperiodic rodents. In long-day Turkish hamsters, unlike most other rodents, elimination of pineal melatonin secretion by constant light or pinealectomy initiates a cycle of gonadal involution and recrudescence outwardly similar to that induced by short days. The present study assessed whether short days and constant light induce the seasonal reproductive cycle via common or different interval timing mechanisms. Male hamsters that had undergone gonadal involution in SDs for 8 or 14 weeks were treated with LL for 14 and 8 weeks, respectively. If SDs and LL act via independent mechanisms, then gonadal quiescence of SD-regressed males, which normally lasts 10 weeks, might be extended by LL treatment; alternatively, if SDs and LL act on the same timer, or the timer cannot be retriggered, then LL will not extend the duration of reproductive quiescence. Neither of these outcomes materialized. Instead, male hamsters exposed to LL while reproductively quiescent exhibited accelerated gonadal recrudescence. Extended LL treatment did not restore responsiveness to SDs in photorefractory hamsters. In Turkish hamsters, photoperiodic history determines whether constant light inhibits or stimulates the hypothalamic-pituitary-testicular axis., (© 2011 Sage Publications)

Photoperiodism research has relied on static day lengths and abrupt transitions between long and short days to characterize the signals that drive seasonal rhythms. To identify ecologically relevant critical day lengths and to test the extent to which naturally changing day lengths synchronize important developmental events, we monitored nine cohorts of male 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). SNP hamsters born from 4 wk before to 2 wk after the solstice underwent rapid somatic and gonadal growth; among those born 4-6 wk after the solstice, some delayed puberty by many weeks, whereas others manifested early puberty. Hamsters born eight or more weeks after the solstice failed to undergo early testicular development. The transition to delayed development occurred at long day lengths, which induce early puberty when presented as static photoperiods. The first animals to delay puberty may do so predominantly on the basis of postnatal decreases in day length, whereas in later cohorts, a comparison of postnatal day length to gestational day length may contribute to arrested development. Despite differences in timing of birth and timing of puberty, autumn gonadal regression and spring gonadal and somatic growth occurred at similar calendar dates in all cohorts. Incrementally changing photoperiods exert a strong organizing effect on seasonal rhythms by providing hamsters with a richer source of environmental timing cues than are available in simple static day lengths.

In mammals, removal of one testis results in compensatory testicular hypertrophy (CTH) of the remaining gonad. Although CTH is ubiquitous among juveniles of many species, laboratory rats, laboratory mice, and humans unilaterally castrated in adulthood fail to display CTH. We documented CTH in pre- and postpubertally hemi-castrated Syrian and Siberian hamsters and tested whether day length affects CTH in juvenile and adult Siberian hamsters. Robust CTH was evident in long-day hemi-castrates of both species and was preceded by increased serum FSH concentrations in juvenile Siberian hamsters. In sharp contrast, CTH was undetectable in short-day hemi-castrated Siberian hamsters for several months and only made its appearance with the development of neuroendocrine refractoriness to short day lengths; serum FSH concentrations of juveniles also did not increase above sham-castrate values until the onset of refractoriness. Long-day hemi-castrated Siberian hamsters with hypertrophied testes underwent complete gonadal regression after transfer to short days, albeit at a reduced rate for the first 3 weeks of treatment. Blood testosterone concentrations of adult hamsters did not differ between long-day hemicastrates and sham-castrates 9-12 weeks after surgery. We conclude that CTH is suppressed by short day lengths in Siberian hamsters at all ages and stages of reproductive development; in short day lengths, but not long day lengths, the remaining testis produces sufficient negative feedback inhibition to restrain FSH hypersecretion and prevent CTH.

We assessed the influence of perinatal melatonin on reproductive development and adult responsiveness to melatonin. Testicular growth in an intermediate day length (14 : 10 h light/dark cycle) was substantially reduced in Siberian hamsters gestated by pinealectomised compared to pineal-intact females; gonadal development was normalised in offspring of pinealectomised dams that were pinealectomised at 3-4 days of age. Hamsters deprived of melatonin only during gestation, or both pre- and postnatally, underwent testicular involution during treatment with melatonin in adulthood. Photoperiodic histories acquired prenatally did not endure as long as those acquired by adult hamsters. Hamsters first exposed to melatonin in adulthood were not more proficient in acquiring photoperiodic histories than were normal males. These findings indicate that pre- versus postnatal differences in melatonin signal duration determine rates of testicular development. Exposure to melatonin perinatally does not appear to organise the neuroendocrine substrate that mediates effects of day length and melatonin on the gonads of adult hamsters.

Seasonal variation in prepubertal penile growth has not previously been studied. The present study assessed the influence of daylength and androgens on penile development in the Siberian hamster (Phodopus sungorus). Adult penile masses were achieved at 18 and 8 weeks of age in hamsters maintained from birth under short (10 h light:14 h dark) versus long (14 h light:10 h dark) daylengths, respectively. Insulin-like growth factor I concentrations, previously implicated in penile growth, did not differ between hamsters maintained in short versus long daylengths. Gonadectomized juvenile males maintained in short and long daylengths and administered testosterone attained adult penile masses well in advance of untreated gonad-intact males maintained in short daylengths. Hamsters from both photoperiods, castrated as juveniles and first treated with testosterone in adulthood, also achieved adult penile masses. The photoinhibited gonad is insufficient to promote penile growth, and prepubertal gonadal secretions during short daylengths are not necessary for eventual penile development. Among young born near the end of the mating season, onset of neuroendocrine refractoriness to short daylengths at about 100 days of age and subsequent gonadal development induces growth in all reproductive tissues. Timing of puberty and increased androgen secretion controlled by daylength are the primary determinants of postnatal penile growth, which may also be affected by prenatal and early postnatal organizational actions of androgens.

Siberian hamsters transferred from a long (16 h light/day [16 L]) to an intermediate (13.5 L) day length (DL) undergo testicular regression within 2 months followed approximately 2 months later by "spontaneous" testicular recrudescence. Recovery of gonadal function after prolonged exposure to intermediate DLs is thought to reflect development of neuroendocrine refractoriness to intermediate-duration melatonin signals. The authors tested the alternative hypothesis that testicular recrudescence in 13.5 L occurs when the "memory" for the 16-L photoperiod fades and hamsters can no longer compare the 13.5-L to the prior 16-L day length. Adult hamsters transferred from 16 L to 13.5 L that underwent testicular involution were either maintained continuously in 13.5 L for 41 weeks or given a supplementary 2-week treatment of 16 L before being returned to 13.5 L. The supplementary treatment was administered either after hamsters had been in 13.5 L for 10 weeks and had involuted testes, or after 24 weeks, when the gonads had undergone recrudescence. The authors found that 16 L treatment administered at week 10 delayed final gonadal recrudescence by approximately 12 weeks; similar 16-L treatment at week 24 induced a second gonadal regression when animals were returned to 13.5 L. The most parsimonious hypothesis to account for these findings is that gonadal recrudescence in intermediate DLs reflects fading of the "memory" for prior long DLs rather than induction of refractoriness to melatonin signals generated in intermediate DLs.

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.

We investigated the impact of frequency and pattern of melatonin signals on reproductive development in Siberian hamsters. Juvenile males gestated in short day lengths and housed in constant illumination to suppress melatonin secretion were infused with melatonin for 5 h either once or twice per day for 20 days. Melatonin infusions at either frequency produced equivalent increases in testes and body weights that exceeded those of animals infused with saline but were indistinguishable from those of hamsters transferred to long day lengths. The reproductive system appears to be maximally stimulated by a single short melatonin signal each day. Other animals kept from birth in a short photoperiod were treated 6 h after onset of darkness with the beta-adrenergic receptor antagonist DL-propranolol to shorten melatonin secretion on the night of injection but not on subsequent nights. This permitted interpolation of short nightly melatonin signals of 4-5 h duration against a background of long melatonin signals of 10-12 h duration on other nights. Treatment regimes that maintained a 1:1 ratio of short to long melatonin signals for 8 wk stimulated reproductive development; a 1:2 signal ratio, in each of three different patterns, was uniformly ineffective. The number of successive short melatonin signals had little influence on the interval across which successive melatonin signals were summated to influence photoperiodic traits. The neuroendocrine axis appears more responsive to short melatonin signal frequency than pattern for development of the summer phenotype.

Long summer days unequivocally stimulate, and short winter days inhibit reproduction in Siberian hamsters. By contrast, intermediate-duration day lengths (12.5-14 h long) either accelerate reproductive development or initiate regression of the reproductive apparatus. Which of these outcomes transpires depends on an animal's photoperiodic history, suggesting that hamsters must encode a representation of prior photoperiods. The duration of nocturnal melatonin secretion is the endocrine representation of day length, but nothing is known about how long it takes to establish photoperiodic histories or how long they endure. Hamsters exposed for 2 or more weeks to long summer day lengths acquired a long-day photoperiodic history that determined subsequent reproductive responses to intermediate-duration day lengths and melatonin signals. The memory for long-day lengths persisted in pinealectomized hamsters for 6.5 weeks, faded significantly after 13 weeks, and was functionally absent after 20 weeks. These findings indicate that hamsters are influenced only by relatively recent day lengths and melatonin signals and ignore earlier ones that might cause them to misinterpret the salience of current day lengths.

Short day lengths induce testicular regression in seasonally breeding Syrian hamsters. To test whether the ventromedial hypothalamus is necessary to maintain reproductive quiescence once testicular regression has been achieved, photoregressed male hamsters were subjected to lesions of the ventromedial hypothalamus (VMHx), pinealectomy (Pinx), or sham operation (Sham). VMHx hamsters underwent accelerated gonadal recrudescence compared to Pinx and Sham hamsters. Recovery of prolactin concentrations (PRL) to values characteristic of long-day hamsters was hastened in the VMHx animals compared to Sham hamsters. Concentrations of follicle stimulating hormone (FSH) increased prematurely in both the VMHx and Pinx animals, beginning a few weeks after surgery. By the time the gonads had undergone recrudescence and the hamsters were refractory to melatonin, PRL and FSH concentrations had returned to baseline long-day values in all groups; there was no evidence of hypersecretion of either hormone in any of the animals with lesions. Melatonin concentrations of VMHx hamsters did not differ from those of sham-operated animals, but because only a single determination was made, it remains possible that VMH damage altered the duration of nightly melatonin secretion. An intact VMH appears to be essential for the continued maintenance of reproductive suppression induced by exposure to short day lengths; these and earlier findings suggest that the VMH-dorsomedial hypothalamic complex mediates regression of the reproductive apparatus during decreasing day lengths of late summer and early autumn and also is necessary to sustain regression during the winter months.

The impact of photoperiodic history was studied in male hamsters kept from birth to 6 wk of age (Week 0) in long-day (LD; 16 h light per day) or short-day (SD; 10 h light per day) photoperiods. At Week 0, hamsters were transferred to constant light and were s.c. infused daily with melatonin (MEL; 4 ng/h) until Week 12. Groups of SD hamsters received 10-h MEL infusions or MEL infusions of durations that decreased linearly by 1-2 min/day from 10 h at Week 0 to 7.5 h at Week 12. LD hamsters were infused for 5 h/day or in patterns that decreased from 10 to 7.5 h or increased from 5 to 7.5 h during Weeks 0-12. MEL signals of decreasing duration produced opposite effects on the gonads of LD and SD hamsters: decreasing MEL durations induced complete gonadal regression in LD hamsters (testis weight = 42 +/- 3 mg) but stimulated testis growth in SD hamsters (testis weight = 276 +/- 57 mg; p < 0.05). MEL infusions that were of shorter but increasing duration induced gonadal regression in LD hamsters. The present experiments indicate that MEL signals of a given sequence and duration can produce either gonadal growth or regression depending on the hamster's photoperiodic history. The pattern of change in MEL secretion, as well as absolute duration of the MEL signal, is important for implementing seasonal cycles of reproduction.

In seasonally breeding rodent species, a fraction of the population is unresponsive to short day lengths (DL) and remains reproductively competent during winter. We previously observed that incidence of nonresponsiveness to short days was affected by photoperiodic history. Here we tested whether exposure to long DL (18 h light/day; 18L) renders animals unresponsive to short DL (10L). Hamsters, maintained from birth in 10L, were transferred at week 6 to 18L or 14L. Ten weeks later (week 16), groups were transferred to 10L for 10 wk. All hamsters maintained in short DL from birth had undeveloped testes at week 6. At week 26, however, 92% of hamsters previously kept in 18L failed to undergo complete gonadal regression in 10L, compared with only 10% of hamsters previously in 14L. Entrainment of locomotor activity in 10L in nonresponsive hamsters resembled that typically observed under long DL. Exposure to 18L may induce nonresponsiveness by altering interactions of component circadian oscillators that mediate gonadal regression in short DL.

Adult male Siberian hamsters transferred from 16 h light/day (16L) to 14 h light/day (14L) underwent gonadal regression and recrudescence over the course of 24 wk; the duration of reproductive quiescence was shorter by 10 wk than in hamsters transferred from 16L to 10L. A decrease in day length (DL) at week 12 from 14L to 10L greatly extended the duration of testicular involution, whereas transfer at week 12 from 10L to 8L was without effect. Hamsters that had manifested gonadal regression and recrudescence in 14L immediately initiated a second regression when challenged with 10L. Intermediate DLs induce gonadal regression and recrudescence without rendering hamsters refractory to shorter DLs. Intermediate DLs sufficient to initiate gonadal regression may be too long to trigger the interval timer that eventually induces refractoriness to shorter DLs and may be successively interpreted as short and long days, respectively, by Siberian hamsters.

Studies of rodent photoperiodism almost without exception have employed fixed day lengths (DLs) and abrupt transitions from long to short DLs. Because the natural progression of changes in DL carries predictive information and may have physiological consequences, we determined seasonal fluctuations in testis size and body weight in Siberian hamsters maintained in a year-long pattern of increasing and decreasing DLs. A cycle of gonadal development, regression, and recrudescence, and corresponding changes in body weight, were observed in hamsters maintained in separate simulated natural photoperiods (SNPs) in which DLs neither fell below nor exceeded the putative critical DL of 13 h. Gradually decreasing DLs as long as 15.3 h induced gonadal regression, and DLs as short as 12.3 h supported ponderal growth, depending on the hamster's prior photoperiodic history. DLs experienced by hamsters during development influenced the adult incidence of responsiveness to short DLs. Photorefractoriness to short DLs occurred earlier in hamsters kept in static 10-h than in 12-h DLs. Increasing DLs in winter had little impact on the rate of gonadal recrudescence and weight gain. These data extend earlier investigations in showing the photoperiodic history determines gonadal responses over a broad range of DLs and influences gonadal responsiveness to short DLs and the triggering of the interval timer underlying recrudescence.

Photoregressed Siberian hamsters exposed at 19 days of age to constant light for 24 h manifested increased testicular development 16 days later. Hamsters that sustained lesions of the suprachiasmatic nucleus (SCN) 3 days after the light pulse had significantly heavier gonads than did sham-operated animals or hamsters pinealectomized at 19 days of age. Enhanced reproductive development after SCN ablation is not due solely to lesion-induced elimination of melatonin secretion. SCN neural activity beginning 3 days after the light pulse is not necessary for light-induced gonadal growth.

In Siberian hamsters, long and short durations of nightly melatonin secretion are associated with testicular regression and development, respectively; surprisingly, the absence of a nightly melatonin signal induced by pinealectomy is less effective than a short (4-6 h) melatonin duration for promoting gonadal development. We compared the patterns of serum FSH and prolactin concentrations underlying pinealectomy-induced and photostimulated testicular growth in juvenile hamsters. Pinealectomy in photoinhibited hamsters on Day 18 produced significant gonadal development compared to that in short-day controls by Day 23. By Day 28, however, testes weights of pinealectomized hamsters were significantly lower than those of intact hamsters transferred to a long photoperiod. Pinealectomy was associated with significant elevations in serum FSH and prolactin concentrations over baseline by Day 21, but peak values were attenuated compared with those induced by exposure to long days. FSH titers of pinealectomized animals were significantly lower than those of photostimulated animals between Days 21 and 30 but were significantly greater on Days 35 and 40; prolactin values were comparable in pinealectomized and photostimulated hamsters between Days 21 and 25 but were significantly lower in the former group thereafter. Blunted elevation of FSH and prolactin secretion may underlie the slower rates of gonadal maturation observed in pinealectomized, juvenile hamsters. The neuroendocrine systems of pinealectomized hamsters appear to be in a state intermediate between those characteristic of hamsters kept in long and in short day lengths.

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.

Pinealectomy completely prevented gonadal regression as well as reduction in body weight and white adipose tissue content of the gonadal and retroperitoneal fat deposits in male meadow voles transferred from long to short day lengths. Pineal influences on pelage characteristics depended on which parameter was assessed. For instance, the increase in guard hair length observed in short-day control voles was blocked by pinealectomy; however, a similar increase in underhair length was unaffected by removal of the pineal gland. Photoperiod-dependent changes in fat deposition, testicular activity, and guard hair length presumably rely on altered pineal secretory activity to transduce the effects of day length on the neuroendocrine axis; however, mechanisms independent of pineal activity may be capable of mediating photoperiodic control of underhair growth.

After 15 wk in a short photoperiod (10 h of light/day), adult male voles weighed 25% less and consumed 33% less food than did voles in a long photoperiod (14 h light/day). Neither body weight nor food intake differed among long- and short-day castrated voles. After 19 wk, castrated long-day voles weighed less than did intact animals. Voles reduced their body weight during the first 15 wk in the short photoperiod and increased their body mass during the succeeding 15 wk. Body mass of short-day voles was positively correlated with combined testes weight. Voles in the short photoperiod collected less nesting material than did their long-day counterparts at week 31. Pelage characteristics at week 32 were not affected by castration or by photoperiod. We conclude that the decreased body mass of male voles during the late summer, fall, and early winter reflects a decrease in circulating levels of testicular hormones; gonadal recrudescence and increased circulating androgen levels are in part responsible for the subsequent seasonal increase in body weight. Part of the increased body mass in late winter in field populations or in laboratory voles after 15 wk in the short photoperiod is independent of the resumption of gonadal activity. Fluctuations in neural and endocrine processes, rather than availability of food, are the principal factors underlying seasonal changes in energy balance.

The gonads of hamsters maintained in short photoperiods first undergo regression and then spontaneous recrudescence. The latter is indicative of refractoriness of the neuroendocrine axis to short day lengths. We tested the hypothesis that the absence of long-day stimulation contributes to the development of refractoriness to short photoperiods. Male Syrian hamsters were maintained for 26 weeks in a short-day photoperiod (8 hr light/day). Testicular regression was confirmed in all animals at week 10; thereafter different groups of hamsters were stimulated with one long day (16 hr light) every 10, 15, 20 or 30 days until week 26. Control groups were maintained continuously in long or short photoperiods, respectively. The rate of gonadal recrudescence of short-day hamsters was not affected by intermittent stimulation with long photoperiods. After 26 weeks, testes in all short-day groups had undergone recrudescence. Refractoriness to short day lengths is not counteracted by long photoperiods. The role of the pineal gland in the development of refractoriness is considered.

The testes of hamsters exposed to short days (10 hr of light per day) regress within 13 weeks. Administration of 7.5% deuterium oxide to hamsters lengthens the period of free running circadian activity rhythms by 2.2% and prevents testicular regression during short-day exposure. This consistent with predictions derived from an external coincidence model for photoperiodic time measurement: Deuterium oxide changes phase relationships between the light-dark cycle and the circadian system, the hamster's daily photosensitive phase is stimulated with light during short days, and the testes remain large. Conservation of the period of circadian rhythms within narrow limits has adaptive significance for hamster photoperiodism and for the occurrence and phasing of the annual reproductive cycle.

Onset of breeding activity tends to occur later in the year for deer mice from progressively higher elevations. We tested the hypothesis that the critical day length for development and maintenance of the reproductive system is longer for mice from higher elevations. Offspring of deer mice trapped at elevations of 1219 and 3048 m were raised from birth to 55 days of age in photo-periods that provided 10, 11, 12, 13 or 16 h light/day. Short day lengths (less than 13 h light/day) retarded maturation of the reproductive apparatus of juveniles from both elevations. The critical photoperiod for development of the reproductive apparatus was 1 h shorter for mice derived from stock trapped at 3048 m as compared to 1219 m. The gonads of adult mice from each elevation were unresponsive to photoperiod and remained functional in all day lengths tested (10-13 h light/day). Photoperiod does not appear to be a proximate factor underlying the altitudinal gradient in reproduction of male deer mice.

White-footed mice were maintained in short or long photoperiods from birth to 60 days of age (10 h vs. 14 h of light per day). Testes weights and spermatogenesis were substantially reduced in short daylengths. Pinealectomy at 5-7 days of age eliminated the suppressive effect of photoperiod on the reproductive system. However, testicular development was not retarded in intact males kept from 25 to 60 days of age in short daylengths. Exposure to short daylengths prior to 25 days of age contributes to photoperiodic inhibition of testicular development. Removal of the pineal gland did not consistently affect gonadal maturation in long photoperiods. The pineal gland transduces the effects of short daylengths on reproductive development. Some effects of long daylengths on the neuroendocrine axis of white-footed mice may also be mediated by the pineal gland.

Laboratory rats traditionally are classified as nonphotoperiodic because variations in daylength have little or no effect on their gonadal function. After olfactory bulbectomy, however, rats show clear evidence of photoperiodic regulation of the gonads. The present study demonstrates, by means of resonance experiments, that the testicular response to daylength in rats is mediated by a circadian photoperiodic time measurement system similar to that of photoperiodic rodents. Olfactory-bulbectomized rats were maintained in fixed photoperiods in which a 6 h light period was coupled with dark periods of 18, 30, 42, or 54 h (6L:18D, 6L:30D, etc.); a fifth group was maintained in a 45L:10D photoperiod. Rats from the 6L:30D, 6L:54D and 14L:10D photoperiods had testes and seminal vesicle weights, plasma testosterone titers and spermatogenesis indices indicative of functional reproductive status. Rats exposed to the 6L:18D and 6L:42D photoperiods had reduced testicular and seminal vesicle weights, lower testosterone levels and reduced spermatogenesis. We hypothesize that photo- and nonphotoperiodic rodent species use similar mechanisms for distinguishing long from short photoperiods, but differ in the extent to which discrimination of short daylengths is transduced into altered gonadal activity.

Plasma luteinizing hormone (LH) levels were determined at monthly intervals in intact and orchidectomized ground squirrels maintained in a constant 14L:10D photoperiod at a temperature of 23 +/- 2 degrees C. Animals were orchidectomized or sham operated at different ages and at different stages of the annual reproductive cycle. LH levels were elevated in orchidectomized as compared to intact males within 1 month, and often within 48 h, after orchidectomies performed either before or after the normal breeding season. LH levels tended to remain chronically elevated in long-term orchidectomized males. Although negative feedback regulation of LH by the testes was evident at most stages of the reproductive cycle, effects of orchidectomy on LH were most pronounced during the nonbreeding season when plasma LH titers of intact males were at their annual nadirs, and less pronounced or absent during the breeding season when levels were normally elevated in intact males. For most of the year male gonadal hormones exert negative feedback on LH release. We hypothesize that under the influence of a circannual timing mechanism, sensitivity to steroid feedback is reduced as the breeding season approaches and LH secretion increases despite the attendant rise in plasma androgen levels. These results contrast with data for female squirrels in which there is a seasonally restricted and characteristically brief elevation of plasma LH after ovariectomy; in females, but not in males, onset and termination of LH secretion appear to be regulated by a circannual clock that functions independently of gonadal secretions.

Testis size and spermatogenesis were monitored serially in individual golden-mantled ground squirrels before, during, and after the hibernation season. During hibernation, animals spent 81% of days in torpor at body temperatures of 3-4 degrees C. Torpor bouts of 6 days duration were interspersed with brief arousals from torpor during which animals were normothermic. In the 5 mo between December (when animals entered hibernation) and April (when torpor was spontaneously terminated), the estimated mass of testes increased gradually from 500 to 1100 mg, but spermatogenesis did not advance beyond pachytene spermatocytes, which were present before hibernation began. In contrast, during the month after torpor was terminated, testes increased rapidly to 3500 mg and after 31 days, spermatozoa were found in the epididymides. We suggest that the limited testis growth that occurred during the hibernation season was restricted to intervals during which squirrels were aroused from torpor. The major portion of gonadal growth and spermatogenesis in the laboratory, and presumably in the field, occurs after ground squirrels have regained the normothermic state. Since males are reproductively mature when first trapped in spring, these findings suggest that males are normothermic for several weeks before they emerge from their hibernacula in the spring.

The 4-5-mo hibernation season of golden-mantled ground squirrels consists of extended torpor bouts interspersed with brief, periodic intervals of normothermic arousal. Plasma levels of testosterone (T), luteinizing hormone (LH), and follicle-stimulating hormone (FSH) and degree of scrotal pigmentation were measured in torpid and aroused male ground squirrels throughout a season of hibernation and in active animals after the termination of torpor. T was basal in torpid animals; beginning 3 weeks before torpor ended, T was elevated in normothermic males during the first half of periodic arousals but returned to basal levels before animals reentered torpor. After the last (terminal) arousal from torpor, T levels were moderately elevated for 4 wk and maximal for the next 6 wk before they returned to basal values. LH patterns were similar to those of T; however, levels of T and LH were positively correlated only in aroused or posthibernation males. FSH levels remained constant and low during most of the heterothermic season but increased in several torpid males within 3 days of terminal arousal. FSH levels peaked 2 wk after the end of heterothermy. Scrotal pigmentation developed over the first 4 wk after terminal arousal. Maturation of reproductive function occurs during the 4 wk after termination of heterothermy, but elevated levels of T during arousals and variable levels of FSH in the last days of torpor suggest that activation or increased sensitivity of the hypothalamic-pituitary-gonadal axis is important in the termination of heterothermy in ground squirrels.

Gonadectomy did not eliminate circannual cycles of body mass and food intake in male pallid bats maintained for up to 30 months in short or long photoperiods at a constant temperature of 23 +/- 2 degrees C. Generation and expression of circannual rhythms is independent of the annual testicular cycle. Initial peak body and testicular masses were recorded within one month of each other but the peak in food intake occurred approximately eight months after animals attained peaks in body mass. Increased metabolic efficiency appears to act independently of changes in food intake to effect seasonal increases in body mass of pallid bats.