Winter Phenotype Research Articles

Metabolic Processes and Physiological Parameters Involved with the Ischemia/Reperfusion Protective Phenotype in Hibernators

The physiology of a hibernating mammal is capable of profound plasticity. Each year hibernators cycle though summer homeothermic and winter heterothermic seasons bracketed by fall and spring transition periods in which physiological and environmental parameters are mixed. Within winter heterothermy, the animals repeatedly cycle through bouts of cold torpor that are punctuated by rapid rewarming during arousal to a brief interbout period of warm body temperature (Tb), During entrance into torpor, the animals dramatically depress their metabolic, heart, and respiratory rates, allowing Tb approach freezing. Organ blood perfusion is also decreased by ~90% during torpor (ischemia) and then rapidly re‐established during arousal (reperfusion). Thus, unlike most mammals, hibernators must be resistant to ischemia/reperfusion (I/R) injury during their winter heterothermic season. The medical applications of this resistance are numerous, including protection against tissue damage that occurs during stroke, cardiac arrest, and trauma/hemorrhagic shock. However, to be able to confer benefit from the use of a torpor‐like physiology, one must first understand the molecular mechanisms underpinning such extreme physiology. Neither the molecular drivers underlying I/R resistance nor those responsible for the physiological changes that occur during the fall transition that enable these animals to enter torpor have been elucidated. In the current study, we collected UHPLC‐MS plasma metabolomics at eleven physiologically distinct points across the hibernator's year to address these deficits in our knowledge. These metabolite data were then compared to metabolites linked to I/R damage to identify potentially protective responses in the hibernators. We then used the diverse fall transition animals to: 1) test which changing physiological and environmental factors in the fall transition period correlate to the metabolome of the protective winter phenotype; and 2) elucidate if experiencing I/R alters their metabolome. We found that during the arousal process, there were changes in succinate concentrations, consistent with its utilization during reperfusion. There were no increases in lactate, a non‐specific indicator of I/R injury, or known biomarkers of I/R organ‐specific tissue damage. However, there were increases in metabolic measures to assuage oxidative stress and decrease utilization of glucose. In the fall transition animals, Tb was the only measured physiological or environmental factor that correlated with the percent winter/protective metabolome, and there was no difference in metabolome between animals that had experienced I/R from the previous use of torpor followed by arousal and animals that had not. Results suggest that the I/R protective phenotype may be due in part to a combination of reduced glycolytic energy production and increases in oxidative stress mitigation that are independent of experiencing an initial torpor bout in the fall but are instead related to changes in Tb.Support or Funding InformationThis project was funded by University of Colorado Fund AEF CDB SIRC BRIDGE‐Martin.

When to initiate torpor use? Food availability times the transition to winter phenotype in a tropical heterotherm

Timing of winter phenotype expression determines individual chances of survival until the next reproductive season. Environmental cues triggering this seasonal phenotypic transition have rarely been investigated, although they play a central role in the compensation of climatic fluctuations via plastic phenotypic adjustments. Initiation of winter daily torpor use-a widespread energy-saving phenotype-could be primarily timed according to anticipatory seasonal cues (anticipatory cues hypothesis), or flexibly fine-tuned according to actual energy availability (food shortage hypothesis). We conducted a food supplementation experiment on wild heterothermic primates (grey mouse lemurs, Microcebus murinus) at the transition to the food-limited dry season, i.e. the austral winter. As expected under the food shortage hypothesis, food-supplemented individuals postponed the seasonal transition to normal torpor use by 1-2 month(s), spent four times less torpid, and exhibited minimal skin temperature 6 °C higher than control animals. This study provides the first in situ experimental evidence that food availability, rather than abiotic cues, times the launching of torpor use. Fine-tuning of the timing of seasonal phenotypic transitions according to actual food shortage should provide heterotherms with a flexible adaptive mechanism to survive unexpected environmental fluctuations.

Environmental, ecological and mechanistic drivers of avian seasonal metabolic flexibility in response to cold winters

Small birds inhabiting regions with cold winter climates show seasonally flexible metabolic phenotypes, with the winter phenotype characterized by increments of summit metabolic rate (Msum) and cold tolerance. In the study reported here, we focused on variations in Msum as a metric of metabolic performance because it is positively correlated with cold tolerance in birds and positively related to overwinter survival in small mammals, although the latter has yet to be demonstrated in birds. Temperature appears to be a prominent driver of seasonal metabolic phenotypes in birds, as evidenced by the correlation between inter- and intra-seasonal variation in Msum and temperature variation, by recent temperature variables serving as better predictors of Msum variation than long-term climate variables, and by the induction of Msum variation by experimental cold exposure. In contrast, photoperiod and social status do not appear to be prominent drivers of metabolic flexibility in birds studied to date. Because skeletal muscle is the primary thermogenic tissue in birds, studies of the mechanistic underpinnings of metabolic flexibility have focused on skeletal muscles, particularly flight muscles. At the level of the skeletal muscle, two potential mechanisms exist for increasing thermogenic capacity, namely, muscle hypertrophy and elevated cellular metabolic intensity. Correlative studies suggest consistent winter increments in flight muscle size, with a potential regulatory role for the muscle growth inhibitor myostatin. Recent experimental studies in small birds, including modification of flight costs, cold acclimation, and exercise training, also suggest that muscle size is an important driver of metabolic flexibility in birds. Therefore, the focus of our study was on the seasonal regulation of muscle size and its contribution to metabolic flexibility. Future studies should address fitness consequences of Msum variation, the relative roles of muscle hypertrophy, and other factors (e.g., oxygen and substrate transport, cellular metabolic intensity) promoting Msum variation, as well as the molecular mechanisms underlying seasonal phenotypes.

Phenotypic flexibility in passerine birds: Seasonal variation in fuel storage, mobilization and transport

Winter acclimatization in small birds living in cold climates produces a winter phenotype characterized by upregulation of metabolic rates to meet enhanced thermoregulatory demands. We measured several key aspects of fuel storage, mobilization and transport in summer and winter to determine whether black-capped chickadees (Poecile atricapillus), white-breasted nuthatches (Sitta carolinensis), and house sparrows (Passer domesticus) seasonally modulate these attributes to meet enhanced winter thermoregulatory demands. In addition, we exposed birds to thermoneutral (control) and severe cold exposure treatments to determine whether acute cold exposure influenced fuel storage, mobilization or transport. Carcass lipid mass and pectoralis intramuscular lipid did not vary significantly between seasons or temperature treatments for any of the study species. Muscle glycogen varied significantly seasonally only for chickadee supracoracoideus and leg muscles, and did not vary among warm or cold treatments for any species. Pectoralis fatty acid binding protein (FABPc) was significantly elevated in winter for chickadees and nuthatches, but not for sparrows. Plasma metabolites showed little consistent variation in response to season or acute cold exposure. Thus, fuel storage and mobilization do not appear to be major targets of adjustment associated with seasonal metabolic flexibility in these species, but modulation of intracellular lipid transport by FABPc may be an important contributor to seasonal phenotypes in some species of small birds.

Relative roles of temperature and photoperiod as drivers of metabolic flexibility in dark-eyed juncos

Seasonal phenotypic flexibility in small birds produces a winter phenotype with elevated maximum cold-induced metabolic rates (=summit metabolism, Msum). Temperature and photoperiod are candidates for drivers of seasonal phenotypes, but their relative impacts on metabolic variation are unknown. We examined photoperiod and temperature effects on Msum, muscle masses and activities of key catabolic enzymes in winter dark-eyed juncos (Junco hyemalis). We randomly assigned birds to four treatment groups varying in temperature (cold=3°C; warm=24°C) and photoperiod [short day (SD)=8 h:16 h light:dark; long day (LD)=16 h:8 h light:dark] in a two-by-two design. We measured body mass (Mb), flight muscle width and Msum before and after 3 and 6 weeks of acclimation, and flight muscle and heart masses after 6 weeks. Msum increased for cold-exposed, but not for warm-exposed, birds. LD birds gained more Mb than SD birds, irrespective of temperature. Flight muscle size and mass did not differ significantly among groups, but heart mass was larger in cold-exposed birds. Citrate synthase, carnitine palmitoyl transferase and β-hydroxyacyl Co-A dehydrogenase activities in the pectoralis were generally higher for LD and cold groups. The cold-induced changes in Msum and heart mass parallel winter changes for small birds, but the larger Mb and higher catabolic enzyme activities in LD birds suggest photoperiod-induced changes associated with migratory disposition. Temperature appears to be a primary driver of flexibility in Msum in juncos, but photoperiod-induced changes in Mb and catabolic enzyme activities, likely associated with migratory disposition, interact with temperature to contribute to seasonal phenotypes.

Seasonal phenotypic flexibility of flight muscle size in small birds: a comparison of ultrasonography and tissue mass measurements

Changes in flight muscle size are important mediators of phenotypic flexibility in birds, so the ability to track such changes over time in individual birds is a valuable tool for investigating phenotypic flexibility. Ultrasonography has been used to track changes in flight muscle size in shorebirds, but has not been previously used to track such changes in small birds, despite variation in flight muscle size being an important contributor to phenotypic flexibility in these birds. One prominent avian example of phenotypic flexibility is the seasonal phenotypes of small birds in response to climatic variation. The winter phenotype in these birds is characterized by increases in organismal metabolic rates and pectoralis muscle mass. We measured seasonal flight muscle size in House Sparrows (Passer domesticus, 25–30 g) using both ultrasonography and wet muscle mass and tested the correlation between ultrasonographic measures of breast muscle thickness and muscle mass. We further tested whether ultrasonographic measures of muscle thickness were sufficiently precise to detect seasonal variation in flight muscle mass. Muscle mass was significantly and positively associated with ultrasonographic measurements of breast muscle thickness for short-axis (SA), long-axis (LA), and combined SA and LA measurements. Breast muscle mass was significantly greater in winter than in summer (17.5 %) and muscle thickness also increased significantly in winter for both SA (9.1 %) and LA (7.5 %) measures. Thus, these data confirm that winter elevations of flight muscle mass consistently contribute to the winter phenotype in House Sparrows and that ultrasonography is effective in detecting seasonal changes in muscle mass in small birds.

Sequence Polymorphism in Candidate Genes for Differences in Winter Plumage between Scottish and Scandinavian Willow Grouse (Lagopus lagopus)

BackgroundPopulation variation in the degree of seasonal polymorphism is rare in birds, and the genetic basis of this phenomenon remains largely undescribed. Both sexes of Scandinavian and Scottish Willow grouse (Lagopus lagopus) display marked differences in their winter phenotypes, with Scottish grouse retaining a pigmented plumage year-round and Scandinavian Willow grouse molting to a white morph during winter. A widely studied pathway implicated in vertebrate pigmentation is the melanin system, for which functional variation has been characterised in many taxa.Methodology/Principal FindingsWe sequenced coding regions from four genes involved in melanin pigmentation (DCT, MC1R, TYR and TYRP1), and an additional control involved in the melanocortin pathway (AGRP), to investigate the genetic basis of winter plumage in Lagopus. Despite the well documented role of the melanin system in animal coloration, we found no plumage-associated polymorphism or evidence for selection in a total of ∼2.6 kb analysed sequence.Conclusions/SignificanceOur results indicate that the genetic basis of alternating between pigmented and unpigmented seasonal phenotypes is more likely explained by regulatory changes controlling the expression of these or other loci in the physiological pathway leading to pigmentation.

Role of melatonin on the regulation of ovarian function: new findings in Danio rerio

Event Abstract Back to Event Role of melatonin on the regulation of ovarian function: new findings in Danio rerio Oliana Carnevali1*, Giorgia Gioacchini1, Francesca Maradonna1, Ike Olivotto1 and Beatrice Migliarini1 1 Polytechnic University of Marche, Department of Marine Sciences, Italy Melatonin is involved in a number of activities including reproduction acting on hypothalamus , pituitary and gonads . The mechanism by which melatonin acts on the reproductive axis is still unknown although this hormone does not appear to have a direct influence on GnRH neurons. The nature of relationship between pineal melatonin and reproductive physiology remains controversial, in mammals the administration of melatonin may induce or inhibit reproduction depending on the species reproductive strategy: melatonin inhibits long day summer phenotype and induces a short day winter phenotype reproduction. In these latter species, melatonin is referred to as being progonadotrophic .Conflicting conclusions on the role of melatonin in the neuroendocrine system has been also found in teleost; administration of melatonin decreases LH β and FSHβ secretion in the ell and inhibits reproduction in catfish, trout, etc…while LH β is induced in kroacker, sea bass and carp. Recently, the antioxidant activity of melatonin have been found in the mammalian oocyte evidencing the beneficial effects of this hormone on reproductive physiology by improving oocyte quality. In zebrafish, we have demonstrated a clear positive role of melatonin in follicle growth and maturation. The large variability of the impact of melatonin on brain, pituitary and on gonadal factors observed among vertebrates will be discussed. Keywords: comparative endocrinology Conference: 25th Conference of the European Comparative Endocrinologists, Pécs, Hungary, 31 Aug - 4 Sep, 2010. Presentation Type: Conference Presentation Topic: Comparative endocrinology Citation: Carnevali O, Gioacchini G, Maradonna F, Olivotto I and Migliarini B (2010). Role of melatonin on the regulation of ovarian function: new findings in Danio rerio. Front. Endocrinol. Conference Abstract: 25th Conference of the European Comparative Endocrinologists. doi: 10.3389/conf.fendo.2010.01.00011 Copyright: The abstracts in this collection have not been subject to any Frontiers peer review or checks, and are not endorsed by Frontiers. They are made available through the Frontiers publishing platform as a service to conference organizers and presenters. The copyright in the individual abstracts is owned by the author of each abstract or his/her employer unless otherwise stated. Each abstract, as well as the collection of abstracts, are published under a Creative Commons CC-BY 4.0 (attribution) licence (https://creativecommons.org/licenses/by/4.0/) and may thus be reproduced, translated, adapted and be the subject of derivative works provided the authors and Frontiers are attributed. For Frontiers’ terms and conditions please see https://www.frontiersin.org/legal/terms-and-conditions. Received: 26 Aug 2010; Published Online: 29 Aug 2010. * Correspondence: Dr. Oliana Carnevali, Polytechnic University of Marche, Department of Marine Sciences, Ancona, Italy, o.carnevali@univpm.it Login Required This action requires you to be registered with Frontiers and logged in. To register or login click here. Abstract Info Abstract The Authors in Frontiers Oliana Carnevali Giorgia Gioacchini Francesca Maradonna Ike Olivotto Beatrice Migliarini Google Oliana Carnevali Giorgia Gioacchini Francesca Maradonna Ike Olivotto Beatrice Migliarini Google Scholar Oliana Carnevali Giorgia Gioacchini Francesca Maradonna Ike Olivotto Beatrice Migliarini PubMed Oliana Carnevali Giorgia Gioacchini Francesca Maradonna Ike Olivotto Beatrice Migliarini Related Article in Frontiers Google Scholar PubMed Abstract Close Back to top Javascript is disabled. Please enable Javascript in your browser settings in order to see all the content on this page.

Dietary palmitate and linoleate oxidations, oxidative stress, and DNA damage differ according to season in mouse lemurs exposed to a chronic food deprivation

This study investigated the extent to which the increase in torpor expression in the grey mouse lemur, due to graded food restriction, is modulated by a trade-off between a whole body sparing of polyunsaturated dietary fatty acids and the related oxidative stress generated during daily torpor. We measured changes in torpor frequency, total energy expenditure (TEE), linoleate (polyunsaturated fatty acid) and palmitate (saturated fatty acid) oxidation, hexanoyl-lysine (HEL; the product of linoleate peroxidation), and 8-hydroxydeoxyguanosine (8OHdG; a marker of DNA damage). Animals under summer-acclimated long days (LD) or winter-acclimated short days (SD) were exposed to a 40% (LD40 and SD40) and 80% (LD80 and SD80) 35-day calorie restriction (CR). During CR, all groups reduced their body mass, but LD80 animals reached survival-threatened levels at day 22 and were then excluded from the CR trial. Only SD mouse lemurs increased their torpor frequency with CR and displayed a decrease in their TEE adjusted for fat-free mass. After CR, SD40 mouse lemurs shifted the dietary fatty acid oxidation toward palmitate and spared linoleate. Such a shift was not observed in LD animals and during severe CR, during which oxidation of both dietary fatty acids was increased. Concomitantly, HEL increased in both LD40 and SD80 groups, whereas DNA damage was only seen in SD80 food-restricted animals. HEL correlated positively with linoleate oxidation confirming in vivo the substrate/product relationship demonstrated in vitro, and negatively with TEE adjusted for fat-free mass, suggesting higher oxidative stress associated with increased torpor expression. This suggests a seasonal-dependant, cost-benefit trade-off between maximizing torpor propensity and minimizing oxidative stress that is associated with a shift toward sparing of dietary polyunsaturated fatty acids that is dependent upon the expression of a winter phenotype.

Circadian Rhythms of Photorefractory Siberian Hamsters Remain Responsive to Melatonin

Short day lengths increase the duration of nocturnal melatonin (Mel) secretion, which induces the winter phenotype in Siberian hamsters. After several months of continued exposure to short days, hamsters spontaneously revert to the spring-summer phenotype. This transition has been attributed to the development of refractoriness of Mel-binding tissues, including the suprachiasmatic nucleus (SCN), to long-duration Mel signals. The SCN of Siberian hamsters is required for the seasonal response to winter-like Mel signals, and becomes refractory to previously effective long-duration Mel signals restricted to this area. Acute Mel treatment phase shifts circadian locomotor rhythms of photosensitive Siberian hamsters, presumably by affecting circadian oscillators in the SCN. We tested whether seasonal refractoriness of the SCN to long-duration Mel signals also renders the circadian system of Siberian hamsters unresponsive to Mel. Males manifesting free-running circadian rhythms in constant dim red light were injected with Mel or vehicle for 5 days on a 23.5-h T-cycle beginning at circadian time 10. Mel injections caused significantly larger phase advances in activity onset than did the saline vehicle, but the magnitude of phase shifts to Mel did not differ between photorefractory and photosensitive hamsters. Similarly, when entrained to a 16-h light/8-h dark photocycle, photorefractory and photosensitive hamsters did not differ in their response to Mel injected 4 h before the onset of the dark phase. Activity onset in Mel-injected hamsters was masked by light but was revealed to be significantly earlier than in vehicle-injected hamsters upon transfer to constant dim red light. The acute effects of melatonin on circadian behavioral rhythms are preserved in photorefractory hamsters.

Developmental plasticity of mating calls enables acoustic communication in diverse environments

Male calls of the katydid Neoconocephalus triops exhibit substantial developmental plasticity in two parameters: (i) calls of winter males are continuous and lack the verse structure of summer calls and (ii) at equal temperatures, summer males produce calls with a substantially higher pulse rate than winter males. We raised female N. triops under conditions that reliably induced either summer or winter phenotype and tested their preferences for the call parameters that differ between summer and winter males. Neither generation was selective for the presence of verses, but females had strong preferences for pulse rates: only a narrow range of pulse rates was attractive. The attractive ranges did not differ between summer and winter females. Both male pulse rate and female preference for pulse rate changed with ambient temperature, but female preference changed more than the male calls. As a result, the summer call was attractive only at 25 degrees C, whereas the slower winter call was attractive only at 20 degrees C. Thus, developmental plasticity of male calls compensates for differences in temperature dependency between calls and preferences and enables the communication system to function in heterogeneous environments. The potential role of call plasticity during the invasion of new habitats is discussed.

Melatonin acts at the suprachiasmatic nucleus to attenuate behavioral symptoms of infection.

In common with reproduction, immune function exhibits strong seasonal patterns, which are driven by annual changes in day length (photoperiod) and melatonin secretion. Whereas changes in melatonin communicate seasonal time into the reproductive axis via subcortical receptors, the relevant melatonin targets for communicating seasonal time into the immune system remain unspecified. The authors report that melatonin implants targeting the hypothalamic suprachiasmatic nuclei (SCN) induced a winter phenotype in the immune system. SCN melatonin implants attenuated infection-induced anorexia and cachexia, indicating that the SCN mediate the effects of melatonin on these behavioral and metabolic symptoms of infection. However, SCN melatonin implants failed to induce winter-like peripheral leukocyte concentrations or behavioral thermoregulatory responses to infection. In contrast, subcutaneous melatonin implants induced winter-like changes in all behavioral and immunological parameters. Melatonin acts directly at the SCN to induce seasonal changes in neural-immune systems that regulate behavior. The data identify anatomical overlap between neural substrates mediating the effects of melatonin on the reproductive and immune systems but also suggest that the SCN are not the sole mediator of photoperiodic effects of melatonin on immunity.

Decoding the nightly melatonin signal through circadian clockwork

Photoperiod regulates the timing of seasonal cycles in reproduction, energy metabolism, moult and other seasonal characteristics, and the effects are transduced through changes in the duration of nocturnal melatonin secretion from the pineal gland. Short daily melatonin signals (4–8 h/day) activate a summer physiology, while long signals (>10 h/day) produce a winter phenotype. Decoding signal duration occurs in specific target cells in the brain and pituitary gland, each governing a different component of the seasonal adaptation. The pars tuberalis (PT) of the pituitary regulates prolactin release and provides a tractable model system to investigate the molecular decoding mechanism. In the PT, melatonin onset at dusk activates cryptochrone ( Cry1) gene expression and melatonin offset at dawn activates period ( Per1) gene expression, thus the Cry/ Per interval varies directly with nightlength, and inverse to daylength. It is proposed that photoperiod-induced changes in this phase-relationship dictates the level of CRY/PER protein heterodimer formation, and in turn, the level of transcriptional drive to the genes that control PT output – up-regulated under long days stimulating prolactin secretion and a summer physiology, and – down-regulated by short days in winter. The melatonin signal is thus decoded through circadian clock genes.

Dim nocturnal illumination alters coupling of circadian pacemakers in Siberian hamsters, Phodopus sungorus

The circadian pacemaker of mammals comprises multiple oscillators that may adopt different phase relationships to determine properties of the coupled system. The effect of nocturnal illumination comparable to dim moonlight was assessed in male Siberian hamsters exposed to two re-entrainment paradigms believed to require changes in the phase relationship of underlying component oscillators. In experiment 1, hamsters were exposed to a 24-h light-dark-light-dark cycle previously shown to split circadian rhythms into two components such that activity is divided between the two daily dark periods. Hamsters exposed to dim illumination (<0.020 lx) during each scotophase were more likely to exhibit split rhythms compared to hamsters exposed to completely dark scotophases. In experiment 2, hamsters were transferred to winter photoperiods (10 h light, 14 h dark) from two different longer daylengths (14 h or 18 h light daily) in the presence or absence of dim nighttime lighting. Dim nocturnal illumination markedly accelerated adoption of the winter phenotype as reflected in the expansion of activity duration, gonadal regression and weight loss. The two experiments demonstrate substantial efficacy of light intensities generally viewed as below the threshold of circadian systems. Light may act on oscillator coupling through rod-dependent mechanisms.

Molting in the Djungarian hamster (Phodopus sungorus Pallas): Seasonal or continuous process?

After transfer into a short daylight regimen, the brownish summer pelage of the Djungarian hamster (Phodopus sungorus) changes into the whitish winter phenotype. Although changes in serum prolactin levels are identified as the initiating hormonal signal, morphological data about molting in that species are sparse. The aim of this study was to characterize in detail the summer and winter pelage of the Djungarian hamster and to analyze the alterations in the skin and pelage induced by photoperiodic changes. The main difference between summer and winter hair types is the pattern of pigmentation. In contrast to other mammalian species showing seasonal changes, the winter coat of the Djungarian hamster is not characterized by an increase in hair density. Molting patches were observed at all times, even in the winter coat, showing that the light regimen does not control the process of molting itself but the pattern of pigmentation and eventually the loss of hair during the single molting wave.

Photoperiod and androgens act independently to induce spinal nucleus of the bulbocavernosus neuromuscular plasticity in the Siberian hamster, Phodopus sungorus.

In the Siberian hamster, Phodopus sungorus, short-day photoperiods induce the winter phenotype, which in males includes a decrease in the production of androgens and changes in physiology to inhibit reproduction. Motoneurones of the spinal nucleus of the bulbocavernosus (SNB) and their target muscles, the bulbocavernosus and the levator ani, a neuromuscular system involved in male copulation, also display seasonal plasticity in P. sungorus. It is not known whether the plasticity seen in the SNB system of gonadally intact hamsters is due to the effects of photoperiod per se, or to the photoperiod-induced changes in androgen production. To answer this question, we castrated adult male hamsters from long days and then implanted them with capsules containing either testosterone or blanks. Half of the hamsters from each hormone condition were moved into short photoperiod (8 : 16 h light/dark cycle) while the rest were maintained under long-day conditions (15 : 9 h light/dark cycle). After 15 weeks, many measures of the SNB system, such as somata size and weight of target muscles, responded only to androgen, not to photoperiod. However, there were effects of photoperiod on the neuromuscular junctions (NMJs) that were independent of androgen status. For example, the number of synaptic zones per NMJ and the area of the NMJs were significantly increased by short days and/or testosterone treatment. The two factors exerted an additive, rather than an interactive, effect on these measures. Another striated muscle, the extensor digitorum longus, which is present in both sexes and plays no specialized role in reproduction, displayed neither an effect of androgen nor of photoperiod on fibre size or NMJ structure. These results suggest that, in addition to androgenic effects on SNB plasticity, there is also an androgen-independent effect of photoperiod on the SNB neuromuscular system.

Low ambient temperature accelerates short-day responses in Siberian hamsters by altering responsiveness to melatonin.

Exposure to low ambient temperatures (Ta) accelerates appearance of the winter phenotype in Siberian hamsters transferred from long to short day lengths. Because melatonin transduces the effects of day length on the neuroendocrine axis, the authors assessed whether low Ta promotes the transition to winterlike traits by accelerating the onset of increased nocturnal melatonin secretion or by enhancing responsiveness to melatonin in short day lengths. Male hamsters were transferred from 16L (16 h light/day) to 8L (8 h light/day) photoperiods and held at 5 degrees C or 22 degrees C. Locomotor activity was recorded continuously, and body mass, testis size, and pelage color were determined biweekly for 8 weeks. The duration of nocturnal locomotion (alpha), a reliable indicator of the duration of nocturnal melatonin secretion, lengthened significantly earlier in hamsters exposed to a Ta of 5 degrees C than 22 degrees C. Cold exposure increased the proportion of hamsters that were photoresponsive: gonadal regression in short days increased from 44% at 22 degrees C to 81% at 5 degrees C (p < 0.05); low Ta did not, however, accelerate testicular regression in animals that were photoresponsive. Nonphotoresponsive animals at 5 degrees C temporarily had longer alphas during the first 4 weeks in short days and significant decreases in body mass and testicular size that were reversed during the ensuing weeks when alpha decreased. In a 2nd experiment, pinealectomized male hamsters infused for 10 h/day with melatonin for 2 weeks had significantly lower body and testes masses when maintained at 5 degrees C but not 22 degrees C. Low-ambient temperature appears to accelerate the appearance of the winter phenotype primarily by increasing target tissue responsiveness to melatonin and to a lesser extent by augmenting the rate at which the duration of nocturnal melatonin secretion increases in short day lengths.

Genetic and environmental influences on short-day responsiveness in Siberian hamsters (Phodopus sungorus).

Siberian hamsters are photoperiodic rodents that typically exhibit several physiological changes when exposed to a short-day photoperiod. However, development of the winter phenotype in short days is largely conditional on prior photoperiod history: Hamsters that have been reared in an exceptionally long day length (18 L) do not usually exhibit the winter phenotype after transfer to short days, whereas animals reared under "moderately" long days (16 L) are more variable in responsiveness to subsequent short-day exposure, with 20% to 30% generally failing to exhibit winter-type responses. Hamsters reared exclusively in an "intermediate" day length (14 L) are almost uniformly responsive to short photoperiod. In the present study, the authors examine the influence of photoperiod history on short-day responsiveness in a breeding line of hamsters that has been subjected to artificial selection for resistance to the effects of short days. The results demonstrate that photoperiod history is an important determinant of short-day responsiveness in both random-bred (UNS) hamsters and animals artificially selected and bred for nonresponsiveness to short photoperiod (PNR). The PNR hamsters have a reduced requirement for long-day exposure to evoke a state of unresponsiveness to short days. The results are discussed in relation to possible significance for the origin of population and species differences in photoperiod responsiveness.

Melatonin chimeras alter reproductive development and photorefractoriness in Siberian hamsters.

Nightly melatonin (MEL) durations > 8 h provoke gonadal regression and decreases in body mass, whereas signals < 7 h stimulate gonadal and somatic growth in male Siberian hamsters. The authors sought to determine the minimum frequency of short MEL signals sufficient to induce the long-day phenotype in several photoperiodic traits. D,L-propranolol (hereafter propranolol) injections shortened MEL signals on the night of treatment without altering MEL on the subsequent night; this permitted interpolation of short MEL signals at variable frequencies against a background of long MEL signals (chimeras). Hamsters kept in short days (10 h light/day, 10L) were injected with propranolol 6 h after dark onset for 28 consecutive weeks beginning at 30 days of age (Week 0) either every other day or once every 3, 6, or 9 days. Control animals were injected with saline or with propranolol during the light phase or were transferred to long days (16L) at Week 0. Hamsters in 16L underwent rapid gonadal development and increases in body mass and displayed summer pelage color, as did hamsters treated with propranolol every other day. Animals treated with propranolol less frequently than every other day uniformly maintained undeveloped gonads and winter-like body weights, but pelage color became proportionately darker with increased frequency of propranolol treatments. The onset of spontaneous testicular development in 10L was unaffected by propranolol injections. After termination of injections at Week 28, testicular regression was not observed in most 10L animals that previously had undergone spontaneous testicular development; however, 40% of hamsters that had been injected with propranolol every 3rd night did manifest the winter phenotype after Week 28. In an alternating sequence, short MEL signals completely override long signals and induce the summer phenotype. Threshold frequencies differ for MEL stimulation of long-day pelage and gonadal phenotypes. The timing and development of refractoriness to MEL does not depend in any simple manner on the number of long MEL signals or on the accumulation of a reaction product produced by long, and depleted by short, MEL signals.