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Olfactory bulbectomy prevents the gonadal regression associated with short photoperiod in male golden hamsters.

In male rats, pre-pubertal olfactory bulbectomy (BX) unmasks photoperiodic responsiveness so that maintenance of BX animals on short photoperiod results in a pineal gland mediated gonadal regression (Nelson and Zucker, Neuroendocrinology, 32 (1981) 266). The objective of the present study was to determine the effect of BX in Syrian hamsters, a species where short photoperiod alone induces a pineal gland induced testicular regression. Pre-pubertal (25-days old) Syrian hamsters underwent either sham BX, BX, or pinealectomy (PX) plus BX. One half of each group was then placed on an LD 14:10 photoperiod and the other half on LD 6:18. Thirteen weeks later all animals were decapitated. In the animals on LD 14:10, BX resulted in a slight but highly significant increase in testes weight but had no effect on other parameters examined. Sham BX hamsters on LD 6:18 had regressed testes and reduced serum testosterone levels, but these effects of short photoperiod were reversed in BX hamsters, so that the testes and seminal vesicles remained large. In addition, serum LH and testosterone levels were elevated following BX of the animals on LD 6:18. These results indicate that the olfactory bulbs tonically inhibit the reproductive system even in male hamsters maintained on long photoperiod and that olfactory bulbectomy prevents gonadal regression in male Syrian hamsters on short photoperiod, an effect which is the opposite of results obtained in rats on short photoperiod. Elucidation of the neural pathways involved and the nature of the relationship between the olfactory bulb and the pineal gland or photoperiodism remains for further investigation.

Animals↗

Variations in the photoperiodic cloacal response of Japanese quail: association with testes weight and feather color.

The size of the cloacal gland was found to be a reliable indicator of testicular activity of Japanese quail. Six experiments were performed to examine the effects of alternating long and short photoperiod on the size of the cloacal gland of male Japanese quail. Three types of photoperiodic cloacal responses were distinguished. Type I birds became refractory to short photoperiods after they had experienced 5 weeks or more of short days. They maintained large cloacal glands under subsequent condition of alternating long and short photoperiod. Type II birds were intermediate types I and III birds did not become refractory to short photoperiods after experiencing 5 weeks or more of short days. The cloacal glands responded to conditions of alternating long and short photoperiods with increases or decreases in size. Feather color on the throat was found to correspond to the type of cloacal response. Type I birds had brick-red throat feathers. Type II birds had white feathers intermingled with brick-red feathers. Type III had white throat feathers. The percentages of types I, II, and III observed in the experimental population was 67, 18, and 15%, respectively. Type III birds were used to study the effects of blinding on the cloacal response to short photoperiod. Five out of eight blinded type III birds did not lose the responsiveness to short photoperiod. These results are consistent with the view that extraocular photoreceptors participate in the photoperiodic gonadal response of Japanese quail.

Aging↗

Effects of photoperiod and temperature on gonadal maturation, and plasma steroid and gonadotropin levels in a cyprinid fish, the honmoroko Gnathopogon caerulescens.

The honmoroko (Gnathopogon caerulescens), a small, spring-summer-breeding cyprinid fish, was exposed to various temperature and photoperiod regimes during different phases of its annual reproductive cycle. In autumn, a clear photoperiodism was seen, especially at warm temperatures (18-24 degrees). The gonadosomatic index, plasma gonadotropin (GtH), and gonadal steroids were higher in fish held at long photoperiod (15L/9D) than in those held at short photoperiod (12L/12D); the differences were greater as the temperature increased. In early spring, gonadal recrudescence progressed at temperatures above 7 degrees, regardless of photoperiod. Photoperiod did not affect plasma GtH and gonadal steroid levels at this time of the year. During its spawning season, a long photoperiod (15L/9D) was again required to maintain gonadal activity and high plasma GtH and gonadal steroid levels, especially at high temperature (25 degrees). In early summer, high temperatures (25 and 30 degrees) suppressed gonadal activity; however, gonadal steroid levels were not depressed, and plasma GtH levels actually increased. The data suggest that under natural conditions, high water temperatures in summer may act directly at the gonadal level, and is probably the factor which terminates the breeding season for this species. The results of these four experiments indicate that the honmoroko exhibits a photoperiodic response during the autumn month and its spawning season, and is dependent on the water temperature. The effects of photoperiod on gonadal maturation appear to be GtH mediated, whereas the suppressive effects of high temperature are not.

Animals↗

Neuroimmunology: modulation of the hamster immune system by photoperiod.

Groups of adult male Syrian hamsters were kept in a long photoperiod (LD 14:10) or a short photoperiod (LD 10:14). After 12 weeks, half of the animals in each light:dark cycle were immunized with an immunogenic amino acid polymer. Exposure to short photoperiod was associated with a significant reduction in testicular, accessory sex organ, splenic and brown fat weights. However, photoperiod length did not influence whole body, thymic, adrenal or kidney weights. Spleens of immunized animals in the long photoperiod were significantly heavier than those of unimmunized animals in the long photoperiod, and both were heavier than spleens from immunized or unimmuized animals in the short photoperiod. This reflected increased splenic lymphocyte and macrophage counts. However, there was no difference in antibody production between animals kept in different photoperiods. These results demonstrate that the daily photoperiod length affects both hamster reproductive competence as well as selected immune parameters (splenic weight and mononuclear cell hyperplasia) but does not alter antibody production.

Adipose Tissue, Brown↗

In vitro reprogramming of the photoperiodic clock in an insect brain-retrocerebral complex.

Pupal diapause in the tobacco hornworm Manduca sexta is a photoperiodically induced event expressed by the absence of release of the cerebral peptide prothoracicotropic hormone. A short-day photoperiod (light-dark, 12:12) during larval development programs the brain to forego the release of prothoracicotropic hormone in the pupal stage. Prothoracicotropic hormone release can occur in the pupae and diapause can be averted if developing larvae experience an intervening long-day photoperiod (light-dark, 16:8) for 3 days early in the last larval instar. Implanted brains from larvae that received such long-day stimulation can reverse the diapause commitment of short-day-reared diapause-destined larvae, thus overriding the diapause program of the recipient. A system was established that demonstrated that this photoperiodic reprogramming of the insect's brain occurs in vitro. Brain-retrocerebral complexes from day 1 instar larvae were cultured in vitro for 3 days under long-day conditions. The effect of this non-diapause photoperiod on the diapause program of these brain-retrocerebral complexes was assessed by assaying the effect of the brains from these cultures on the incidence of non-diapause development in diapause-destined larval hosts. Brains that received a long-day photoperiod reversed the pupal diapause program of the recipient larvae, whereas brains that received a short-day photoperiod were ineffective. Therefore, the larval brain-retrocerebral complex of this insect is capable of photoperiodic light reception in vitro, in that it can sense the number of light-dark cycles to which it is exposed, store this environmental input, and express the information 9 days later at another stage of development. These findings demonstrate that the photoperiodic clock of an animal can be reprogrammed by a light-dark cycle in vitro.

Animals↗

Timing of photorefractoriness in the European starling: significance of photoperiod early and late in the reproductive cycle.

In European starlings, as in many other birds inhabiting higher latitudes, gonads develop in response to the increasing daylengths in early spring. Later in the year, however, the hypothalamo-pituitary-gonadal axis becomes refractory to the previously stimulatory long photoperiods and the gonads regress in summer. The present study addresses the question of when during the gonadal growth phase photorefractoriness is determined. A 13-h photoperiod induces testicular development and subsequent testicular regression associated with refractoriness in male starlings. An 11-h photoperiod, in contrast, induces only testicular development, and photorefractoriness never develops. When starlings were transferred to an 11-h photoperiod, either 12 or 25 days following exposure to a 13-h photoperiod, their testes developed to full size, but remained large to the end of the experiment, i.e. refractoriness did not develop. The same was even true of most birds in a third group that were transferred to an 11-h photoperiod after 46 days of the 13-h photoperiod, when gonads had developed to near maximal size. These data show that, in contrast to some other species of passerine birds, the onset of photorefractoriness does not become fixed before the testes have undergone considerable development, and that the photoperiodic conditions experienced at the end of the testicular growth phase are still effective in determining the precise time of onset of photorefractoriness. It is suggested that this peculiarity of the starling is related to the fact that its gonadal development begins rather early in spring and, hence, under much shorter photoperiods than the other species studied.

Animals↗

Genetic Regulation of Development in Sorghum bicolor (X. Greatly Attenuated Photoperiod Sensitivity in a Phytochrome-Deficient Sorghum Possessing a Biological Clock but Lacking a Red Light-High Irradiance Response).

The role of a light-stable, 123-kD phytochrome in the biological clock, in photoperiodic flowering and shoot growth in extended photoperiods, and in the red light-high irradiance response was studied in Sorghum bicolor using a phytochrome-deficient mutant, 58M (ma3R ma3R), and a near-isogenic wild-type cultivar, 100M (Ma3 Ma3). Since chlorophyll a/b-binding protein mRNA and ribulose bisphosphate carboxylase small subunit mRNA cycled in a circadian fashion in both 58M and 100M grown in constant light, the 123-kD phytochrome absent from 58M does not appear necessary for expression or entrainment of a functional biological clock. Although 58M previously appeared photoperiod insensitive in 12-h photoperiods, extending the photoperiod up to 24 h delayed floral initiation for up to 2 weeks but did not much affect shoot elongation. Thus, although 58M flowers early in intermediate photoperiods, a residual photoperiod sensitivity remains that presumably is not due to the missing 123-kD phytochrome. Since rapid shoot elongation persists in 58M under extended photoperiods despite delayed floral initiation, long photoperiods uncouple those processes. The observed absence of a red light-high irradiance response in 58M, in contrast to the presence of the response in 100M, strengthens the suggestion that the 123-kD phytochrome missing from 58M is a phyB.

Journal Article↗

Short photoperiod and cold activate brown fat lipoprotein lipase in the Djungarian hamster.

Activation of lipoprotein lipase (LPL) in brown fat of the Djungarian hamster ensures substrate supply for nonshivering thermogenesis. Cold acclimation [5 degrees C ambient temperature (Ta)] in long photoperiod [light-dark (LD) 16:8] increased brown fat LPL activity from 4.7 to 22.7 nmol oleate.mg-1.min-1 within 1 day. Slight reduction of this high LPL activity was observed during prolonged cold exposure for up to 4 wk. Deacclimation (transfer from 5 to 23 degrees C Ta) caused total inactivation of brown fat LPL within 4 days. Short photoperiod (LD 8:16) also stimulates brown fat LPL activity. Its effect can be summarized by three different observations. At thermoneutrality short photoperiod elevated LPL activity to 7.1 instead of 4.7 nmol.mg-1.min-1 observed in long photoperiod. Second, at low ambient temperature short photoperiod reinforces the effect of cold acclimation. Maximum LPL activity of cold-exposed Djungarian hamsters averaged 23.7 nmol.mg-1.min-1 in long photoperiod and was elevated to 35.5 nmol.mg-1.min-1 in short photoperiod. Third, inactivation of LPL during deacclimation was delayed in short photoperiod. These results demonstrate that photoperiod as well as ambient temperature may be cooperatively used as environmental cues for seasonal acclimation of brown fat substrate supply for nonshivering thermogenesis.

Acclimatization↗

Pre- and postnatal effects of photoperiod on collared lemmings (Dicrostonyx groenlandicus).

We investigated the hypothesis that photoperiod information received during the prenatal and/or early postnatal periods influences subsequent development in collared lemmings. Pregnant lemmings were exposed to either a short [8:16-h light-dark cycle (8:16)] or long (22:2) photoperiod throughout gestation. On the day of birth, pups were cross-fostered to dams housed in either short or long photoperiod. After an 11-day experimental lactational period (LACT), all animals were transferred to an intermediate photoperiod (16:8), the response to which depends on prior photoperiod exposure. Pups remained on this photoperiod until death at postnatal day 90. Information received during gestation (GEST) influenced offspring growth, pelage color, guard hair length, presence of the bifid claw at weaning, testes mass at 10 wk postweaning, and serum prolactin (PRL). Parameters measured at weaning reflected directly the photoperiod experienced during GEST (e.g., short-day GEST pups had white pelage). Conversely, parameters measured at 10 wk postweaning reflected the change in photoperiod experienced between GEST and 16:8 (e.g., short-day GEST young had gray pelage, indicative of an increase in photoperiod). Information received during LACT influenced growth and the presence of the bifid claw at weaning.

Aging↗

Neural pathways involved in the photoperiodic control of reproductive physiology and behavior in female hamsters (Mesocricetus auratus).

Female hamsters received horizontal knife cuts to investigate the role of hypothalamic connections in the photoperiodic control of female reproductive functions. Knife cuts placed ventral to or through the paraventricular nucleus (PVN), but dorsal to the suprachiasmatic nucleus (SCN), prevented photoperiod-induced acyclicity and uterine regression in animals maintained under a nonstimulatory photoperiod for 10 weeks. The animals were then ovariectomized and tested for lordosis behavior following subcutaneous injections of ovarian hormones to investigate the photoperiodic modulation of female sexual behavior. Animals exposed to a nonstimulatory photoperiod were less behaviorally sensitive to treatment with estradiol benzoate (EB) alone, but did not differ from animals maintained under a stimulatory photoperiod when EB was combined with progesterone. The effect of photoperiod and behavioral sensitivity to hormone replacement was independent of the surgical condition. The results are consistent with the hypothesis that dorsal projections from the SCN to the PVN mediate gonadal responses to short photoperiods. They also indicate that photoperiod-induced changes in behavioral sensitivity to gonadal steroids may be mediated by neural pathways distinct from those that mediate the gonadal changes.

Animals↗

Photoperiodism in Neurospora crassa.

Plants and animals use day or night length for seasonal control of reproduction and other biological functions. Overwhelming evidence suggests that this photoperiodic mechanism relies on a functional circadian system. Recent progress has defined how flowering time in plants is regulated by photoperiodic control of output pathways, but the underlying mechanisms of photoperiodism remain to be described. The authors investigate photoperiodism in a genetic model system for circadian rhythms research, Neurospora crassa. They find that both propagation and reproduction respond systematically to photoperiod. Furthermore, a nonreproductive light-regulated function is also enhanced under certain photoperiodic conditions. All of these photoperiodic responses require a functional circadian clock, in that they are absent in a clock mutant. Night break experiments show that measuring night length is one of the mechanisms used for photoperiod assessment. This represents the first formal report of photoperiodism in the fungi.

Biological Clocks↗

Photoperiodic control of body weight and energy metabolism in Syrian hamsters (Mesocricetus auratus): role of pineal gland, melatonin, gonads, and diet.

The effects of photoperiod and the pineal hormone melatonin on the regulation of body weight and energy metabolism were examined in Syrian hamsters, Mesocricetus auratus. Short photoperiod-housed female and male hamsters showed increases in body weight gain, feed efficiency, carcass lipid, brown adipose tissue mass, and thermogenic capacity. These effects of short photoperiods were mimicked by afternoon melatonin injections to hamsters in long-day photoperiods and were exaggerated in hamsters fed high fat diets. To determine the role of the gonads in these effects, ovariectomized hamsters were treated similarly and found to exhibit changes in body weight and energy metabolism that were 80-90% of those in gonadally intact hamsters. The role of the pineal gland in short photoperiod-induced body weight gain was examined in sham-pinealectomized and pinealectomized hamsters. Short photoperiod-induced increases in body weight were seen in both pinealectomized and sham-pinealectomized hamsters. Thus, pinealectomy blocks the effects of short photoperiods on reproductive function and thyroid activity, but not on body weight regulation. These results suggest that the effects of short photoperiods on body weight and energy metabolism are mediated by multiple, redundant mechanisms involving decreases in gonadal hormone secretion, changes in melatonin secretion, and gonad- and pineal-independent changes. All of the effects of short photoperiods are exaggerated in hamsters fed a high fat diet. These changes may represent adaptive preparatory responses that enhance winter survival in Syrian hamsters.

Adipose Tissue, Brown↗

Prenatal photoperiod influences neonatal prolactin secretion in the sheep.

This study tested the hypothesis that the fetal sheep can respond to photoperiod cues. Pregnant Suffolk ewes were maintained in artificial photoperiod of either long days [16 h of light, 8 h of dark (16L:8D)] or short days (8L:16D) from approximately 100 days of gestation until term at approximately 147 days. On the day of birth, all lambs and their mothers were transferred to an intermediate photoperiod of 12L:12D; both groups were housed together. To provide an index of response to photoperiod, serum PRL concentrations were measured in blood samples collected daily 3-4 h after lights on. In lambs (n = 8 male; n = 7 female) born to mothers on long days, serum PRL concentrations were high (greater than 200 ng/ml) for the first few days after birth, but fell rapidly to low levels (less than 50 ng/ml) within 14 days postnatally in 12L:12D. Conversely, lambs (n = 8 male; n = 7 female) born to mothers on short days initially had low PRL concentrations, but these gradually increased in the postnatal 12L:12D photoperiod to 150 ng/ml by 32 days of age. Thus, serum PRL concentrations in lambs at birth reflect the photoperiodic treatment of their mother, and the subsequent PRL response to an intermediate photoperiod of 12L:12D depends on the photoperiodic history received in utero. We infer from these findings that the fetal lamb receives and responds to information about day length in utero and begins developing a seasonal photoperiod history before birth.

Animals↗

Influence of photoperiod on the seasonal pattern of secretion of luteinizing hormone and testosterone and on the antler cycle in roe deer (Capreolus capreolus).

Annual variations in concentrations of luteinizing hormone (LH) and testosterone in plasma were analysed in relation to the antler cycle in six adult male roe deer exposed to a natural photoperiod (latitude 46 degrees 10'N) and in four adult males maintained in a constant short-day photoperiod (8 h light: 16 h dark) for a year, from the winter solstice at which time both groups of animals had antlers in velvet. The animals were sampled, every 15 min for 2 or 4 h at intervals of one month for a year. Under both natural and experimental conditions, LH concentrations were high from January to March, but in the experimental conditions they decreased between April and May-June, whereas in the natural conditions they increased. Plasma LH concentration was lowest between July and November in animals under natural photoperiod, whereas under 8 h light:16 h dark photoperiod a second increase in plasma LH occurred between August and September. Between March and August, concentrations of plasma testosterone increased under natural photoperiod, whereas under experimental photoperiod there was a biphasic pattern of plasma testosterone with peaks between February and May and between September and November. Under natural photoperiod, antlers were cast in November, 369 +/- 6 days after the previous antlers were cast. Under experimental photoperiod, antlers were cast after 193 +/- 10 days, and a new set developed. The sexual cycle of the male appears to be initiated by an endogenous rhythm in winter and is then maintained by hormonal changes resulting from increasing photoperiod in spring.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Growth and performance of growing-finishing lambs exposed to long or short photoperiods.

The effects of photoperiod (8 hr light: 16 hr dark [8L:16D] or 16L:8D) on performance of growing-finishing lambs were evaluated in an experiment with 24 rams and 24 wethers. All lambs were caged in groups of two in a controlled environment and fed a pelleted diet ad libitum for the duration of the study (10 to 22 weeks of age). Performance and carcass data showed that both photoperiod and sex of lamb affected growth rate (P less than .01) and feed efficiency (P less than .05) but that only sex of lamb affected carcass quality (P less than .05) and yield (P less than .10). A sex by photoperiod interaction not observed for any of the traits measured. Average daily gain (grams/per day) and feed efficiency (feed/gain) for lambs in the four treatment groups tested were: rams (16L:8D), 410 and 4.3; rams (8L:16D), 340 and 4.5; wethers (16L:8D), 345 and 4.6, and wethers (8L:16D(, 300 and 4.8 respectively. Carcass weight, like slaughter weight, was affected by photoperiod (P less than .01) and sex of the lamb (P less than .05); both rams and wethers exposed to long photoperiods had the heaviest carcasses. Ram carcasses were leaner, had better yield grades and were heavier than wether carcasses, yet both ran and wether carcasses graded Choice. Testosterone concentrations in serum were undetectable (less than .2 ng/ml) at 22 weeks of age in wethers but were elevated (P less than .01) in ram lambs exposed to short (3.4 +/- 1.4 ng/ml) or long (.8 +/- .2 ng/ml) photoperiods. Prolactin concentrations, on the other hand, were not influenced by sex of lamb but were influenced (P less than .01) by photoperiod; i.e., prolactin concentrations were high in rams (190 +/- 34 ng/ml) and wethers (178 +/- 28 ng/ml) exposed to long photoperiods and low in rams (43 +/- 9 ng/ml) and wethers (27 +/- 11 ng/ml) exposed to short photoperiods. These results suggest that testosterone and prolactin may independently affect growth and performance of growing-finishing lambs.

Animals↗

Effect of photoperiod on milk production and prolactin of Holstein dairy cows.

Milk yield and composition were measured in 74 Holstein cows and first-calf heifers in two experiments comparing varying lengths of photoperiod. High pressure sodium vapor lamps provided light intensity at cow eye level of 254 +/- 26 lx during the day and 132 +/- 9 lx at night in a free stall barn open on two sides. The first experiment compared continuous light with 18 h light for 16 wk during the winter in animals previously exposed to continuous light. Photoperiod had no effect on milk yield or basal concentration of serum prolactin in either age group. Peak concentration of prolactin after injection of thyrotropin releasing hormone was not affected by photoperiod in heifers but was higher in cows exposed to 18 h light. The second experiment compared natural photoperiod (9 to 12 h light) with 18 h light for 9 wk during the winter in animals previously exposed to a natural photoperiod. Photoperiod had no effect on 4% fat-corrected milk in heifers whereas there was a significant interaction of photoperiod by week for 4% fat-corrected milk in cows. Basal concentration of serum prolactin was lower in cows than in heifers although there were no differences due to age or photoperiod after injection of thyrotropin releasing hormone. We conclude that multipara Holstein cows produce about 7% more 4% fat-corrected milk during winter months when exposed to 18 h light providing they were previously in a natural photoperiod environment. For reasons not understood, 18 h light failed to enhance milk yield of cows previously in a continuous light environment. Milk yield and composition in heifers were not affected by 18 h or continuous light.

Animals↗

[Effect of the photoperiod of the site of oviposition of Aedes mariae (Diptera,Culicidae)].

The overwintering biology of Aedes (Ochlerotatus) mariae (one of the sibling species of the mariae complex) was studied in populations from the Tyrrhenian coast of Central Italy (Sperlonga and Formia). Ae. mariae has in this zone several generations per year and its larvae are commonly found from March to October exclusively in rock pools along the coast. Field and laboratory observations show in these populations an embryonic winter diapause induced by short-day photoperiods (9-12 hours). The photoperiod acts primarily on the parental females during their preimaginal life. The expression of diapause is influenced by temperatures and photoperiods experienced by the embryos. Full embryonic diapause is observed when the eggs are incubated at relatively low temperatures (less than 16 degrees C) and at short photoperiods. Short-day photoperiods also induce a remarkable change in the oviposition behavior of Ae. mariae. Laboratory observations show that the adult females readily oviposit on water surfaces when originating from larvae reared at long-day photoperiod while they are very reclutant to oviposit in the same situation when reared at short day photoperiod. Choice experiments involving four alternative oviposition sites (see Plate 1) demonstrate a preference for outside free water in long-day mosquitoes and for inside moist surfaces in short-day mosquitoes. The behavioral difference persists in successive gonotrophic cycles and it is not apparently affected by the photoperiod acting on the adult females. The above evidence together with recovery of overwintering eggs in holes and crevices of rock pools suggest that the shift in oviposition site shown in the laboratory reflects a similar shift occurring in nature. Such photoperiodically induced change in oviposition behavior seems to have an important adaptive significance in providing more constant microclimates to the diapausing eggs and in protecting them from the mechanical action of winter storms.

Aedes↗

Carry-over effects of long and short photoperiods on body fattening and gonadal weight of blackheaded bunting (Emberiza melanocephala).

A study was made of the carry-over effect of long photoperiods followed by short photoperiods ou the fattening and gonadal response in a photoperiodic migratory species, the Blackheaded Bunting (Emberiza melanocephala). The effect was studied in photosensitive and then in photostimulated birds. Two experiments were performed: experiment I with photosensitive males and experiment II with photosensitive males exposed to 30 long photoperiods. In both of them, five groups were submitted to the following treatments for 30 days: Group S, a short daily photoperiod; Group L, a long daily photoperiod; Group LS, alternating long and short daily photoperiods; Group L 2S, one long with two short daily photoperiods; Group L 3S, one long with three short daily photoperiods. The results showed that the inductive effect of long days or the inhibitory effect of short days was affected by intervening reversed daylengths.

Adipose Tissue↗