Fat ablation and food restriction influence reproductive development and hibernation in ground squirrels

The role of white adipose tissue in development of the reproductive apparatus of male golden-mantled ground squirrels was assessed by surgical removal of fat (lipectomy) immediately prior to onset of hibernation or by manipulation of fat levels through food restriction for 5 wk preceding hibernation. Animals then were maintained without food at 6 degrees C, and they hibernated from November 1983 until April 1984. At that time, blood plasma was assayed for hormone levels, and body mass, body composition, and masses of the testes and seminal vesicle-prostate complex were determined. At autopsy, testes and sexual accessory organs were heavier in Control squirrels than in Food-Restricted or Lipectomized (LIPX) animals. Paired-testes mass was positively correlated with body mass. Testosterone, luteinizing hormone (LH), and follicle-stimulating hormone (FSH) levels were uniformly low in animals sampled during torpor, but were elevated in 2 animals that had regained the euthermic state. The Food-Restricted animals spent more time in torpor than did the other two groups, and the percentage of decrease in body mass over the hibernation season was less for Food-Restricted than for LIPX or Control animals. Adequate lipid stores may be essential for the normal development of the reproductive apparatus of male ground squirrels. In the field, animals with insufficient fat stores may forego reproduction and thereby conserve energy for survival.     

The influence of hibernation on testis growth and spermatogenesis in the golden-mantled ground squirrel, Spermophilus lateralis

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.     

Changes in body condition of hibernating bats support the thrifty female hypothesis and predict consequences for populations with white-nose syndrome

White-nose syndrome (WNS) is a new disease of bats that has devastated populations in eastern North America. Infection with the fungus, Geomyces destructans, is thought to increase the time bats spend out of torpor during hibernation, leading to starvation. Little is known about hibernation in healthy, free-ranging bats and more data are needed to help predict consequences of WNS. Trade-offs presumably exist between the energetic benefits and physiological/ecological costs of torpor, leading to the prediction that the relative importance of spring energy reserves should affect an individual''s use of torpor and depletion of energy reserves during winter. Myotis lucifugus mate during fall and winter but females do not become pregnant until after spring emergence. Thus, female reproductive success depends on spring fat reserves while male reproductive success does not. Consequently, females should be “thrifty” in their use of fat compared to males. We measured body condition index (BCI; mass/forearm length) of 432 M. lucifugus in Manitoba, Canada during the winter of 2009/2010. Bats were captured during the fall mating period (n = 200), early hibernation (n = 125), and late hibernation (n = 128). Adult females entered hibernation with greater fat reserves and consumed those reserves more slowly than adult males and young of the year. Consequently, adult females may be more likely than males or young of the year to survive the disruption of energy balance associated with WNS, although surviving females may not have sufficient reserves to support reproduction.     

Intraspecific differences in the pattern of hibernation in the ground squirrelSpermophilus beldingi

Summary Intraspecific differences in the patterns of heterothermy were found in captive Belding's ground squirrels that hibernated undisturbed at ambient temperatures of 5°C, 10°C, and 15°C. The timing of all entrances into and arousals from hibernation was determined from records of copper-constantan thermocouples that were mounted on the floor of each animal's nest box and connected to continuously recording potentiometers. In the absence of food, large adult males terminated hibernation spontaneously in the spring. In contrast, females and small non-breeding males (yearlings) did not stop hibernating but instead they shortened their bouts of torpor in the spring so that they aroused every three or four days. This interval of frequent arousals, termed the emergence period, lastel until the squirrels became emaciated, and it was only in the 2 or 3 weeks preceeding death (starvation period) that arousal frequency once again decreased towards midwinter values (Fig. 3). These animals terminated hibernation when fed during the emergence or starvation periods, but they were able to resume torpor if that feeding lasted less than a week.Termination of hibernation and reproductive development in males were related to the size of the animals, not their age. Males two years and older which did not deposit normal quantities of fat were like yearlings in that they had open-ended hibernation seasons and showed little testicular enlargement. Likewise, juvenile males that grew to near adult size in their first summer spontaneously terminated hibernation and had well developed testes like most older individuals. In addition, both the frequency and duration of arousals in the emergence period were related, in part, to the extent of the animals' fat reserves, such that large squirrels spent more time at high body temperatures than small individuals (Figs. 5, 6). This trend was most pronounced at high ambient temperatures.These intraspecific differences in hibernation physiology are consistent with the sex and agerelated differences in the timing of emergence above ground in nature. The increase in the time spent euthermic in the spring isinterpreted as an adaptation for increasing the opportunities for environmental assessment. The intraspecific differences in the extent of this euthermia appear to be associated with differences in the balance between the energy supplies available to an animal and its need to accelerate the use of that energy (i.e., arouse) in order to achieve an early and accurately-time emergence from the hibernaculum.     

The effects of caloric restriction on the body composition and hibernation of the golden-mantled ground squirrel (Spermophilus lateralis)

In preparation for hibernation, golden-mantled ground squirrels (Spermophilus lateralis) must deposit sufficient amounts of lipid during the summer to survive winter hibernation. We conducted an experiment from May 1998 to February 1999 to examine the effects of caloric restriction on the body composition (lipid and fat-free mass) and hibernation of golden-mantled ground squirrels. Ground squirrels were either provided with food ad lib. (controls) or with only enough food to maintain a constant body mass throughout the experiment (calorically restricted). Changes in body composition were followed using total body electrical conductivity (TOBEC). Implanted data loggers that recorded body temperature were used to determine when ground squirrels entered their first torpor bout and the lengths of torpor bouts. Body composition did not change in the calorically restricted ground squirrels between May and September, while both lipid and fat-free mass increased in the controls. However, from September to February, calorically restricted ground squirrels lost only fat-free mass, not lipid mass, but controls lost both lipid and fat-free mass. Calorically restricted ground squirrels entered their first torpor bout about 4 wk after controls, but the torpor bout duration (or length) during hibernation did not differ between the two groups. These results show that ground squirrels maintain body composition during caloric restriction, and the limited quantities of stored lipid have an effect on when hibernation begins but not on torpor bout length.     

Hibernating patterns of free-ranging Columbian ground squirrels

Summary The patterns of torpor and euthermy during hibernation was documented for 28 free-ranging Columbian ground squirrels (Spermophilus columbianus) fitted with temperature-sensitive radio transmitter collars. Adult males began hibernation earlier, were euthermic for a greater proportion of the hibernating season and emerged earlier than other age and sex classes. The patterns of hibernation of adult females did not differ significantly from those of juveniles. Emergence from the hibernaculum was preceded by a long (3–12 d) euthermic interval in adult males but not in adult females or juveniles. Changes in soil temperature did not appear to initiate emergence. The greater time spent euthermic by adult males is interpreted as a significantly greater energy cost of hibernation for adult males than for other age and sex classes. The benefits offsetting these costs may be increased reproductive potential in spring and avoidance of predation in late summer.     

Testis development and testosterone secretion in captive European ground squirrels before, during, and after hibernation

In some hibernating species, an extended euthermic period before spring emergence has been reported during which testicular maturation occurred. In this study, we investigated whether male European ground squirrels Spermophilus citellus (Linnaeus, 1766) develop scrotal testes before or after the termination of hibernation. The course of testis development and testosterone concentrations were examined in young-of-year male ground squirrels (n = 4) before, during, and after their first hibernation. In the environmental chamber, all animals showed regular hibernation patterns with decreasing torpor bout lengths prior to the end of hibernation. Four weeks before hibernation ended, testosterone levels were elevated during spontaneous arousals. Testosterone concentrations peaked during the first 2 months post heterothermy and decreased thereafter. In 2 experimental males, testes descended from the inguinal region into the scrotum shortly before the end of heterothermy. The state of testis development and testosterone secretion corresponded to that found in semi-free-living males at spring emergence. Testis size increased in active animals to maximum widths during weeks 3 and 4 post heterothermy. Scrotal skin pigmentation was absent until heterothermy ended and developed during the first month thereafter. The data suggest that male S. citellus do not need a pre-emergence euthermic period for reproductive development.     

Nutrient constraints in the feeding ecology of an omnivore in a seasonal environment

Summary Nutrient requirements of adult, nonreproductive, omnivorous antelope ground squirrels (Ammospermophilus leucurus) were compared with the nutritional value of their food resources. It was found that nutrient constraints would be important factors in ground squirrel feeding ecology primarily in winter. Potentially important constraints were the requirement for water and nitrogen, and a digestive requirement that average dry matter digestibility of the diet exceed ca. 50%. An unlikely constraint was the requirement for any specific mineral. A linear programming model was used to determine potential diets ground squirrels could consume which satisfied these nutritional requirements and also the ground squirrel's daily energy requirements. During spring ground squirrels could be strict herbivores, but during winter before winter rains ground squirrels had to eat some arthropods to satisfy water requirements.These ground squirrels are not energy maximizers because they spend only one third of their activity period feeding and do not accumulate excess energy as fat. Thus, optimum diets were predicted for winter and spring assuming the goal of feeding time minimization. The model correctly predicted that in wintertime ground squirrels would be primarily granivorous but would consume about 20% arthropods, and that they would switch to herbivory in springtime. Ground squirrels, however, selected a wider dietary range than predicted in both winter and spring. Possible reasons for this discrepancy include an inappropriate assumption that ground squirrels forage for food classes nonsimultaneously, and the possibility that ground squirrels employ sampling as part of their foraging behavior.     

Food constraints explain the restricted distribution of wintering Lesser White‐fronted Geese Anser erythropus in China

More than 90% of the Lesser White‐fronted Geese Anser erythropus in the Eastern Palearctic flyway population winter at East Dongting Lake, China. To explain this restricted distribution and to understand better the winter feeding ecology and habitat requirements of this poorly known species, we assessed their food availability, diet and energy budgets at this site through two winters. Lesser White‐fronted Geese maintained a positive energy budget when feeding on above‐ground green production of Eleocharis and Alopecurus in recessional grasslands in autumn and spring to accumulate fat stores. Such food was severely depleted by late November and showed no growth in mid‐winter. Geese fed on more extensive old‐growth Carex sedge meadows in mid‐winter where they were in energy deficit and depleted endogenous fat stores. Geese failed to accumulate autumn fat stores in one year when high water levels prevented the Geese from using recessional grassland feeding areas. Fat stores remained lower throughout that winter and Geese left for breeding areas later in spring than in the previous year, perhaps reflecting the need to gain threshold fat stores for migration. Sedge meadows are widespread at other Yangtze River floodplain wetlands, but recessional grasslands are rare and perhaps restricted to parts of East Dongting Lake, which would explain the highly localized distribution of Lesser White‐fronted Geese in China and their heavy use of these habitats at this site. Sympathetic management of water tables is essential to maintain the recessional grasslands in the best condition for Geese. Regular depletion of fat stores whilst grazing sedge meadows in mid‐winter also underlines the need to protect the species from unnecessary anthropogenic disturbances that enhance energy expenditure. The specialized diet of the Lesser White‐fronted Goose may explain its highly restricted winter distribution and global rarity.     

Extensive use of torpor in 13-lined ground squirrels in the fall prior to cold exposure

Mammalian hibernation is characterized by profound reductions in body temperature (T _b) and metabolic, heart and respiratory rates. These reductions are characteristic of torpor, which is temporally confined to winter. Hibernators including ground squirrels are heterothermic in winter, cycling between multiday periods of torpor with low T _b and brief periods of rewarming. In contrast, ground squirrels remain homeothermic during summer, like non-hibernating mammals. The transition between the homeothermic and heterothermic phases of the circannual rhythm of hibernation is often overlooked in hibernation studies. Here, we examined the use of torpor throughout the fall transition in laboratory-housed 13-lined ground squirrels by recording core body temperature with an implanted data logger. As is typical of laboratory-based hibernation studies, animals were kept in standard housing prior to being moved into a cold, dark room to simulate natural hibernation conditions. Significantly, the vast majority of both male and female ground squirrels expressed torpor in the fall while still housed conventionally and prior to cold exposure. The expression of torpor was not predicted by body weight or age, rather it appears to be preprogrammed in a time-dependent manner that is independent of, yet enhanced by, environmental cues. The timing and duration of these torpor bouts occurring prior to cold exposure were also remarkably sporadic. Thus, it is not possible to know with certainty which animals are torpor-naive before cold exposure in the absence of continuous measurement of body temperature. We conclude that fall animals encompass variable points in the transition between summer and winter phases of the circannual cycle of hibernation, thereby confounding studies in which they are used as non-hibernating controls. Conversely, these fall transition animals offer unique opportunities to define the molecular changes that accompany and enable hibernation.     

The influence of androgens on hibernation phenology of free-living male arctic ground squirrels

Free-living ground squirrel species are sexually dimorphic in hibernation phenology. The underlying causes of these differences are not yet known. Androgens, testosterone (T) in particular, inhibit hibernation. To determine the influence of endogenous androgens on annual timing of hibernation we first measured circulating levels of T and dehydroepiandrosterone (DHEA), an adrenal androgen implicated in non-mating season aggression in other species, in free-living male arctic ground squirrels (Urocitellus parryii, AGS). We also manipulated endogenous androgen levels by surgical castration, and consequently compared body temperature records from intact (n = 24) and castrated (n = 9) males to elucidate the influence of endogenous androgens on annual body temperature cycles. The highest T levels (0.53 ± 0.10 ng/mL) were in reproductively mature male AGS in spring; whereas, both immature males in spring and all males in late summer had T levels an order of magnitude lower (0.07 ± 0.01 and 0.06 ± 0.00 ng/mL, respectively). DHEA levels were higher in males during the late summer compared to reproductively mature males in spring (120.6 ± 18.9 and 35.9 ± 2.3 pg/mL, respectively). Eliminating gonadal androgens via castration resulted in males delaying euthermy by extending heterothermy significantly in spring (Apr 22 ± 2.9) than reproductive males (Mar 28 ± 3.9) but did not change the timing of hibernation onset (castrate: Oct 12 ± 1.0 vs. intact: Oct 3 ± 3.1). We conclude that while androgens play a significant role in spring hibernation phenology of males, their role in fall hibernation onset is unclear.     

Contrasting stress response of male arctic ground squirrels and red squirrels

A hormonal-challenge protocol was used to compare the stress response of males of Arctic ground squirrels and red squirrels during the breeding season (May). These squirrels live in the same boreal forest of the Yukon, but have very different life histories and utilize the forest in markedly different ways. Red squirrels had levels of total cortisol, maximum corticosteroid-binding capacity, and free cortisol that were 5, 7, and 2 times, respectively, those of Arctic ground squirrels. Red squirrels were resistant to suppression by an artificial glucocorticoid, dexamethasone (DEX); Arctic ground squirrels were not. Cortisol levels in red squirrels responded slowly but continuously to the ACTH injection; Arctic ground squirrels responded rapidly and then stabilized. Testosterone levels in red squirrels were extremely sensitive to the challenge, being suppressed by both DEX and ACTH; levels in Arctic ground squirrels were resistant to the challenge, being modestly suppressed by DEX and stimulated by ACTH. Energy mobilization, as measured by glucose and free fatty acid responses, was not affected. Red squirrels had four times the levels of white blood cells and higher proportions of lymphocytes and lower proportions of eosinophils than Arctic ground squirrels, indicating that the latter were in worse condition immunologically. Our evidence suggests that the functions associated with the hypothalamic-pituitary-adrenal axis are compromised in breeding male Arctic ground squirrels, but not in red squirrels. We propose that in male red squirrels this axis has evolved in the context of a stable social system based on long-lived animals with individual territories which are needed to deal with unpredictable winter food supplies. In contrast, Arctic ground squirrels escape the rigors of winter by hibernation and this hormonal axis has evolved in short-lived males in the context of intense intra-sexual competition in a social system based on female kin groups and regular male dispersal to avoid inbreeding. J. Exp. Zool. 286:390-404, 2000.     

Mating of captive thirteen-lined ground squirrels and the annual timing of estrus

Thirteen-lined ground squirrels (Spermophilus tridecemlineatus) have a single annual mating season in Michigan, beginning shortly after their spring emergence from hibernation. Captive females were studied during a 3-year period to determine relations among time of removal from the coldroom, vaginal estrus, and mating behavior. Following a 7-month period females spent in a coldroom, vaginal lavages were taken daily to monitor changes in estrous condition. Females removed from the coldroom about when free-living animals emerge from hibernation were in vaginal estrus within 24-48 hr and had an initial period of persistent estrus (ca. 3 weeks), followed by briefer (less than 1 week) and more sporadic estrous periods. Females left in the coldroom 3 weeks longer than normal had significantly briefer initial periods of vaginal estrus after being removed from the coldroom. Similarly, virgin Yearlings and virgin 2-Year-Olds had significantly briefer initial periods of estrus than nonvirgin Adults (greater than or equal to 2 years old). In 1985, eight of eight females paired with males mated within the first week after removal from the coldroom and subsequently produced litters. Mated females remained in vaginal diestrus from within a few days of mating until after parturition. In contrast, unmated females remained in prolonged vaginal estrus during this period. Females first paired with males 3 weeks after being taken from the coldroom failed to mate. In 1986, five of six females first paired with males 2 weeks after being removed from the coldroom similarly failed to mate. However, five of six females did mate that were removed from the coldroom 10 days after those in the previously described group and paired with a male 4 days after removal. This first report of reliable mating behavior in captive thirteen-lined ground squirrels should facilitate subsequent analysis of reproductive patterns in this species.     

Daily body temperature rhythms persist under the midnight sun but are absent during hibernation in free-living arctic ground squirrels

In indigenous arctic reindeer and ptarmigan, circadian rhythms are not expressed during the constant light of summer or constant dark of winter, and it has been hypothesized that a seasonal absence of circadian rhythms is common to all vertebrate residents of polar regions. Here, we show that, while free-living arctic ground squirrels do not express circadian rhythms during the heterothermic and pre-emergent euthermic intervals of hibernation, they display entrained daily rhythms of body temperature (T(b)) throughout their active season, which includes six weeks of constant sun. In winter, ground squirrels are arrhythmic and regulate core body temperatures to within ±0.2°C for up to 18 days during steady-state torpor. In spring, after the use of torpor ends, male but not female ground squirrels, resume euthermic levels of T(b) in their dark burrows but remain arrhythmic for up to 27 days. However, once activity on the surface begins, both sexes exhibit robust 24 h cycles of body temperature. We suggest that persistence of nycthemeral rhythms through the polar summer enables ground squirrels to minimize thermoregulatory costs. However, the environmental cues (zeitgebers) used to entrain rhythms during the constant light of the arctic summer in these semi-fossorial rodents are unknown.     

Long-term effects of pinealectomy on the annual cycle of golden-mantled ground squirrels,Spermophilus lateralis

Summary Pinealectomies and radical sham pinealectomies were performed on adult golden-mantled ground squirrels (Spermophilus lateralis) approximately one month prior to the beginning of the normal winter hibernation season. The cyclic components of the overwintering period and subsequent active season (the rhythms of homeothermy-heterothermy, body mass and testicular function) of the first year were not disrupted in either of the operated groups as compared with the intact animals. In the second year, the initiation of the hibernatory period occurred as usual in all intact, pinealectomized, and sham operated animals, but the pinealectomized group failed to show the progressive increase in the length of heterothermic bouts that was characteristic of the intact and sham-pinealectomized squirrels. The loss of body mass also was accelerated in the pinealectomized squirrels with respect to the intact and sham groups and appeared to be associated with terminal arousal which occurred approximately six weeks earlier in the second year after pinealectomy. Additionally, male squirrels showed a correspondingly early onset of testicular development as compared to the intact and sham operated males.     

Retention of social recognition after hibernation in Belding's ground squirrels

The retention of social memory during long periods of separation, such as hibernation or migration, has not been well documented, despite evidence for long-term social relationships in migrating species or in long-lived sedentary species. We investigated the ability of captive Belding's ground squirrels, Spermophilus beldingi, to remember previously familiar individuals as well as littermates after 9 months of isolation. Before hibernation, young ground squirrels discriminated between odours of familiar and unfamiliar individuals, as shown by greater investigation of a novel individual's odour. The following spring, these yearlings did not respond differentially to odours of previously familiar and unfamiliar individuals, suggesting that memory for familiar conspecifics was lost during hibernation. In contrast, both female and male yearlings continued to discriminate between odours of littermates and previously familiar nonlittermates. Thus, recognition of close kin was maintained during prolonged social isolation, but recognition of familiar, unrelated individuals was not. If re-establishment of familiarity is not costly or if adults rarely interact with the same individuals in successive years, then selection may not favour retention of individual memories of particular conspecifics over the winter. Even though males rarely encounter kin after dispersal, yearling males did recognize their siblings, suggesting that the relative costs of maintaining kin-recognition abilities year-round may be low. Possible mechanisms underlying the formation and maintenance of individual and kin recognition are discussed. Copyright 2000 The Association for the Study of Animal Behaviour.     

Annual cycle of energy and time expenditure in a golden-mantled ground squirrel population

Summary We have analyzed seasonal shifts of energy and time allocation in a population of golden-mantled ground squirrels (Spermophilus saturatus) by directly measuring total daily energy expenditure (DEE) with an isotopic technique (doubly labeled water=dlw), and by estimating components of total DEE through an integration of field behavioral observations with laboratory-measured rates of energy expenditure (oxygen consumption) associated with major behavioral and physiological states. Hibernation laster about 7 1/2 months, and the 4 1/2-month activity season consisted of mating, a 28-d gestation of 3–5 young, 5 1/2 weeks of postnatal growth building to a peak in lactation just before the young emerged above ground, an additional 2–3-week period of maternal care before dispersal, and finally restoration of body mass preceding hibernation. Although the hibernation season comprised nearly two-thirds of the year, it involved only 13–17% of annual energy expenditure, leaving about 85% of energy expenditure for the active season. Ground squirrels were actually present on the surface for only about 11% of the year's time, and the foraging time required to obtain the total annual energy supply amounted to only about 2% of the year's time. The squirrels fed mainly on herbs in the early season and hypogeous fungi later; both were used extensively during peak lactation when female energy expenditure and demand were maximal. Average daily foraging time increased steadily throughout the season to a maximum of 28% of aboveground time as availability of greens diminished and fungus predominated in the diet; time availability did not limit foraging since the animals sat on average for 65% of the daily surface time of about 7 h. Timing of reproduction is apparently optimized such that peak reproductive energy demands are matched with maximal food availability and moderate thermal conditions that minimize energy demand. Despite the greater body mass of males, the greatest total DEE (measured by dlw) of any squirrels at any time of year was that of females during peak lactation. For production of young and lactation through above-ground emergence of an average litter of 2.7, females required a total energy increase of 24% above annual nonreproductive metabolism. Yearling females all bred and performed similarly to older females, yet some costs were greater because the yearlings began and ended hibernation at smaller mass, compensated by giving birth later, and finally showed a greater absolute increase in body mass over the active season than older females. Annual metabolic energy expenditure of breeding males was about 18% greater than that of females, due to greater male body mass. Yet the annual energy intake requirement for both sexes was essentially identical (about 42MJ) due to the greater reproductive export by females in the form of newborn and milk. During the mating season males showed wide-ranging exploratory behavior and social interactions, including aggression, that involved considerable locomotory energy expenditures. Although we did not directly account for the energetics of these specific reproductive behaviors, they are critical to male reproductive success and on a daily basis they probably involved much greater energy expenditure than sperm production. Some yearling males avoided these costs by foregoing testicular development, yet they allocated four times as much energy to growth as older males, thereby increasing somatic condition for the future.     

Effect of winter high temperatures on reproduction and circannual rhythms in hibernating ground squirrels

We tested whether prevention of hibernation in ground squirrels by midwinter exposure to high ambient temperatures influenced timing of the spring phase of reproductive maturation and the phase and period of subsequent circannual rhythms of reproduction and body mass. Exposing hibernating adult male Spermophilus lateralis to 30 degrees C for 6 weeks beginning December 4 advanced the timing of testicular recrudescence by 4-5 weeks, compared to controls left at 4 degrees C. Males exposed to 30 degrees C for 6 weeks beginning at the average time of spontaneous end of hibernation (January 15) reached reproductive maturation at a time intermediate to those of controls and of the December 4 experimental group. However, neither the date of the subsequent fall's body mass peak, the date of the next year's reproductive maturation, nor the periods of circannual rhythms of body mass and reproduction differed among groups. Premature interruption of hibernation appears to allow early expression of reproduction, but does not affect the underlying timing mechanism.     

Seasonal variations in feeding range and flock structure of the Rook Corvus frugilegus in eastern Ireland

The feeding range and flock structure of Rooks showed temporal variations caused mainly by the dispersion of food and reproductive behaviour.
Feeding range was restricted in spring, autumn and early winter when food availability was high and Rooks were reproductively active. A large feeding range occurred in late summer and late winter, when food availability was either generally low or locally distributed and when Rooks were reproductively inactive. The occurrence of flocks common to several rookeries mirrored variations in feeding range; mixed rookery flocks were more common in late summer and late winter. Similarly, the size of the rookery (as measured by the number of nests) was related to feeding range only when Rooks were reproductively inactive and at such times larger rookeries had greater feeding ranges.
Flock structure showed similar seasonal variations; small widely spaced flocks predominated in summer, autumn, early winter and spring, whereas large dense flocks occurred in late winter. Diurnal variations in flock structure occurred within any one season. Both seasonal and diurnal variations in flock structure may be determined by the dispersion of the prey and the feeding strategy used to obtain it, reproductive behaviour and the risk of predation.     

Cholesterol and lipoprotein dynamics in a hibernating mammal

Hibernating mammals cease feeding during the winter and rely primarily on stored lipids to fuel alternating periods of torpor and arousal. How hibernators manage large fluxes of lipids and sterols over the annual hibernation cycle is poorly understood. The aim of this study was to investigate lipid and cholesterol transport and storage in ground squirrels studied in spring, summer, and several hibernation states. Cholesterol levels in total plasma, HDL and LDL particles were elevated in hibernators compared with spring or summer squirrels. Hibernation increased plasma apolipoprotein A-I expression and HDL particle size. Expression of cholesterol 7 alpha-hydroxylase was 13-fold lower in hibernators than in active season squirrels. Plasma triglycerides were reduced by fasting in spring but not summer squirrels. In hibernators plasma β-hydroxybutyrate was elevated during torpor whereas triglycerides were low relative to normothermic states. We conclude that the switch to a lipid-based metabolism during winter, coupled with reduced capacity to excrete cholesterol creates a closed system in which efficient use of lipoproteins is essential for survival.