Data logging of body temperatures provides precise information on phenology of reproductive events in a free-living arctic hibernator

Precise measures of phenology are critical to understanding how animals organize their annual cycles and how individuals and populations respond to climate-induced changes in physical and ecological stressors. We show that patterns of core body temperature (T _b) can be used to precisely determine the timing of key seasonal events including hibernation, mating and parturition, and immergence and emergence from the hibernacula in free-living arctic ground squirrels (Urocitellus parryii). Using temperature loggers that recorded T _b every 20 min for up to 18 months, we monitored core T _b from three females that subsequently gave birth in captivity and from 66 female and 57 male ground squirrels free-living in the northern foothills of the Brooks Range Alaska. In addition, dates of emergence from hibernation were visually confirmed for four free-living male squirrels. Average T _b in captive females decreased by 0.5–1.0°C during gestation and abruptly increased by 1–1.5°C on the day of parturition. In free-living females, similar shifts in T _b were observed in 78% (n = 9) of yearlings and 94% (n = 31) of adults; females without the shift are assumed not to have given birth. Three of four ground squirrels for which dates of emergence from hibernation were visually confirmed did not exhibit obvious diurnal rhythms in T _b until they first emerged onto the surface when T _b patterns became diurnal. In free-living males undergoing reproductive maturation, this pre-emergence euthermic interval averaged 20.4 days (n = 56). T _b-loggers represent a cost-effective and logistically feasible method to precisely investigate the phenology of reproduction and hibernation in ground squirrels.     

Energetics of arousal episodes in hibernating arctic ground squirrels

Arctic ground squirrels overwintering in northern Alaska experience average soil temperature of −10°C. To examine energetic costs of arousing from hibernation under arctic compared to temperate conditions, captive ground squirrels were maintained in ambient temperatures (T _a) of 2, −5 and −12°C. Rates of oxygen consumption and carbon dioxide production were used to estimate metabolic rate and fuel use during the three phases of arousal episodes: rewarming, euthermia, and recooling. Respiratory quotient comparisons suggest exclusive use of lipid during rewarming and mixed fuel use during euthermia. Animals rewarming from torpor at T _a −12°C took longer, consumed more oxygen, and attained higher peak rates of oxygen consumption when compared to 2°C. T _a had no significant effect on cost or duration of the euthermic phase. Animals recooled faster at −12°C than at 2°C, but total oxygen consumption was not different. T _a had no significant effect on the total cost of arousal episodes when all three phases are included. Arousal episodes account for 86% of estimated costs of a complete hibernation cycle including torpor when at 2°C and only 23% at −12°C. Thus, due to the higher costs of steady-state metabolism during torpor, proportional metabolic costs of arousal episodes at T _a characteristic of the Arctic are diminished compared to relative costs of arousals in more temperate conditions.     

Winter body temperature patterns in free-ranging Cape ground squirrel, Xerus inauris: no evidence for torpor

The body temperature (T _b) of Cape ground squirrels (Xerus inauris, Sciuridae) living in their natural environment during winter has not yet been investigated. In this study we measured abdominal T _b of eight free-ranging Cape ground squirrels over 27 consecutive days during the austral winter. Mean daily T _b was relatively stable at 37.0 ± 0.2°C (range 33.4 to 40.2°C) despite a marked variation in globe temperature (T _g) (range −7 to 37°C). Lactating females (n = 2) consistently had a significantly higher mean T _b (0.7°C) than non-lactating females (n = 3) and males. There was a pronounced nychthemeral rhythm with a mean active phase T _b of 38.1 ± 0.1°C and a mean inactive phase T _b of 36.3 ± 0.3°C for non-lactating individuals. Mean daily amplitude of T _b rhythm was 3.8 ± 0.2°C. T _b during the active phase closely followed T _g and mean active phase T _b was significantly correlated with mean active phase T _g (r ² = 0.3–0.9; P < 0.01). There was no evidence for daily torpor or pronounced hypothermia during the inactive phase, and mean minimum inactive phase T _b was 35.7 ± 0.3°C for non-lactating individuals. Several alternatives (including nocturnal huddling, an aseasonal breeding pattern and abundant winter food resources) as to why Cape ground squirrels do not employ nocturnal hypothermia are discussed.     

Overwinter body temperature patterns in captive jerboas (<Emphasis Type="Italic">Jaculus orientali</Emphasis>s): influence of sex and group

The jerboa (Jaculus orientalis) has been described in the past as a hibernator, but no reliable data exist on the daily and seasonal rhythmicity of body temperature (T _b). In this study, T _b patterns were determined in different groups of jerboas (isolated males and females, castrated males and grouped animals) maintained in captivity during autumn and winter, and submitted to natural variations of light and ambient temperature (T _a). T _b and T _a variations were recorded with surgically implanted iButton temperature loggers at 30-min intervals for two consecutive years. About half (6/13) of isolated female jerboas hibernated with a T _b < 33°C, with hibernation bouts interspersed with short periods of normothermy from November to February. Hibernation bout durations were longer (4–5 days) than those of normothermia phases (1–4 days). During hibernation, the minimum T _b was low (T _bmin ~10.7°C). In contrast, one of the 12 isolated males showed short hibernation bouts of ca. 2 days late in the hibernation season, February–March. The males had T _bmin values of 15.1°C. In contrast to predictions, no castrated males hibernated. When jerboas were grouped, females and males exhibited concomitant torpor bouts. In males, the longest bouts were observed during the late hibernation season. These data suggest complex regulation of hibernation in jerboas.     

Comparison of hibernation, estivation and daily torpor in the edible dormouse, Glis glis

Three major forms of dormancy in mammals have been classified: hibernation in endotherms is characterised by reduced metabolic rate (MR) and body temperature (T _b) near ambient temperature (T _a) over prolonged times in the winter. Estivation is a similar form of dormancy in a dry and hot environment during summertime. Daily torpor is defined as reduced MR and T _b lower than 32 °C, limited to a duration of less than 24 h. The edible dormouse (Glis glis) is capable for all three distinct forms of dormancy. During periods of food restriction and/or low T _a, daily torpor is displayed throughout the year, alternating with hibernation and estivation in winter and summer respectively. We recorded T _b, O₂-consumption and CO₂-production in unrestrained dormice at different T _a's for periods of up to several months. Cooling rate and rate of metabolic depression during entrance into the torpid state was identical in all three forms of dormancy. The same was true for thermal conductance, maximum heat production, duration of arousal and cost of an arousal. The only difference between hibernation and daily torpor was found in the bout duration. A daily torpor bout lasted 3–21 h, a hibernation bout 39–768 h. As a consequence of prolonged duration, MR, T _b and also the T _b − T _a gradient decreased to lower values during hibernation bouts when compared to daily torpor bouts. Our findings suggest that all three forms of dormancy are based on the same physiological mechanism of thermal and metabolic regulation. Accepted: 27 June 2000     

Global analysis of circulating metabolites in hibernating ground squirrels

Hibernation in mammals involves major alterations in nutrition and metabolism that would be expected to affect levels of circulating molecules. To gain insight into these changes we conducted a non-targeted LC–MS based metabolomic analysis of plasma using hibernating ground squirrels in late torpor (LT, T_b ~ 5 °C) or during an interbout arousal period (IBA, T_b ~ 5 °C) and non-hibernating squirrels in spring (T_b ~ 37 °C). Several metabolites varied and allowed differentiation between hibernators and spring squirrels, and between torpid and euthermic squirrels. Methionine and the short-chain carnitine esters of propionate and butyryate/isobutyrate were reduced in LT compared with the euthermic groups. Pantothenic acid and several lysophosphatidylcholines were elevated in LT relative to the euthermic groups, whereas lysophosphatidylethanolamines were elevated during IBA compared to LT and spring animals. Two regulatory lipids varied among the groups: sphingosine 1-phosphate was lower in LT vs. euthermic groups, whereas cholesterol sulfate was elevated in IBA compared to spring squirrels. Levels of long-chain fatty acids (LCFA) and total NEFA tended to be elevated in hibernators relative to spring squirrels. Three long-chain acylcarnitines were reduced in LT relative to IBA; free carnitine was also lower in LT vs. IBA. Our results identified several biochemical changes not previously observed in the seasonal hibernation cycle, including some that may provide insight into the metabolic limitations of mammalian torpor.     

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.     

Thermoregulation in free‐ranging ground woodpeckers Geocolaptes olivaceus: no evidence of torpor

Heterothermic responses characterised by pronounced hypometabolism and reductions in body temperature (T_b) are one of the most effective ways in which small endotherms can offset the energetic cost of endothermic homeothermy. It remains unclear, therefore, why daily torpor and hibernation are restricted to only a subset of avian lineages. To further our understanding of the phylogenetic distribution of avian torpor, we investigated winter thermoregulation in the southern African ground woodpecker Geocolaptes olivaceus. We considered this species a good candidate for heterothermy, because it is resident year‐round in mountainous regions with cold winters and reliant on small ectothermic prey. We recorded T_b patterns in free‐ranging individuals and measured T_b and metabolic rates in captive individuals. Neither free‐ranging nor captive woodpeckers showed any indication of daily torpor or even shallow rest‐phase hypothermia. All birds maintained bimodally distributed T_b characteristic of classic endothemic homeothermy, with a mean rest‐phase T_b of 37.9 ± 0.2°C and no data below 37.0°C. The mean circadian amplitude of T_b was 4.2°C, equivalent to approximately twice the expected value. There was some evidence of seasonal acclimatisation in T_b, with a small decrease in rest‐phase T_b with the onset of the austral winter. Captive birds showed patterns of resting metabolic rate and T_b consistent with the classic model of endothermic homeothermy. The apparent absence of torpor in G. olivaceus supports the notion that, unlike the case in mammals, many avian taxa that may a priori be expected to benefit from deep heterothermy do not use it.     

Ecology of natural hibernation in the marsupial mountain pygmy-possum (Burramys parvus)

The hibernating marsupial mountain pygmy-possum (Burramys parvus, 40 g) has to raise its slow-growing offspring during a short alpine summer. Only females provide parental care, while after mating males emigrate to marginal habitats often at lower altitudes which can sustain only low possum densities. We predicted that the hibernation strategies in mountain pygmy-possums are distinct from those of similar-sized placental hibernators, because of the developmental constraints in marsupials and because hibernation differs between the sexes. Using temperature-sensitive radio transmitters, we studied the hibernation patterns of free-living male and female mountain pygmy-possums living in a north- and a south-facing boulder field (Kosciusko National Park) for two consecutive winters. Individual possums commenced hibernation several months before the snow season. As in other hibernators, torpor in the mountain pygmy-possum was interrupted by periodic arousals which occurred most often during the late afternoon. Torpor bouts initially lasted a few days when the hibernacula temperature (T _hib) ranged from 4 to 7°C. As the hibernation season progressed, torpor bouts became longer and possum body temperatures (T _b) approached 2°C. The T _bs of females were significantly lower and torpor bouts were longer in the second half of the hibernation season than in males. Between torpor bouts, both sexes were often active and left hibernacula for periods of up to 5 days. Especially during the first months of the hibernation season, possums also frequently changed hibernacula sites probably in an attempt to select a site with a more suitable microclimate. Emergence from hibernation was closely coupled with the disappearance of snow from the possum habitat (September 1995, October 1996) and the limited fat stores probably dictate an opportunistic spring emergence. However, in 1995, spring was early and males emerged significantly earlier than females. In 1996, when snow melt was delayed, this difference vanished. Testes are regressed in males during hibernation and the time needed for testes growth and spermatogenesis favours an earlier emergence for males which was probably achieved by their preference for the more sun exposed north-facing boulder field. A sexual dimorphism in hibernation strategies and spring emergence therefore enables mountain pygmy-possums to cope with their harsh alpine environment. Received: 22 May 1997 / Accepted: 21 August 1997     

Heterothermy in an Australian passerine, the Dusky Woodswallow (Artamus cyanopterus)

Information regarding passerine heterothermy and torpor is scant, although many species are small and must cope with a fluctuating food supply and presumably would benefit from energy savings afforded by torpor. We studied whether insectivorous Dusky Woodswallows (Artamus cyanopterus; ∼35 g) enter spontaneous torpor (food ad libitum) when held outdoors as a pair in autumn/winter. Woodswallows displayed pronounced and regular daily fluctuations in body temperature (T _b) over the entire study period. The mean T _b ranged from ∼39°C to 40°C (photophase, day time) and ∼33°C to 36°C (scotophase, night time). However, on 88% of bird nights, nocturnal T _b minima fell to < 35°C. The lowest T _b observed in air was 29.2°C. However, when a bird fell into water its T _b dropped further to ∼22°C; this T _b was regulated for several hours and the bird survived. Our observations suggest that heterothermy is a normal part of the daily thermal regime for woodswallows to minimise energy expenditure. Spontaneous nocturnal torpor in captive woodswallows suggests that torpor in the wild may be more pronounced than recorded here because free-living birds are likely challenged by both low food availability and adverse weather.     

Hibernation patterns of Turkish hamsters: influence of sex and ambient temperature

Turkish hamsters (Mesocricetus brandti) are a model organism for studies of hibernation, yet a detailed account of their torpor characteristics has not been undertaken. This study employed continuous telemetric monitoring of body temperature (T _b) in hibernating male and female Turkish hamsters at ambient temperatures (T _as) of 5 and 13 °C to precisely characterize torpor bout depth, duration, and frequency, as well as rates of entry into and arousal from torpor. Hamsters generated brief intervals of short (<12 h), shallow test bouts (T _b > 20 °C), followed by deep torpor bouts lasting 4–6 days at T _a = 5 °C and 2–3 days at T _a = 13 °C. Females at T _a = 5 °C had longer bouts than males, but maintained higher torpor T _b; there were no sex differences at T _a = 13 °C. Neither body mass loss nor food intake differed between the two T _as. Hamsters entered torpor primarily during the scotophase (subjective night), but timing of arousals was highly variable. Hamsters at both T _as generated short, shallow torpor bouts between deep bouts, suggesting that this species may be capable of both hibernation and daily torpor.     

Daily and seasonal cycles of body temperature and aspects of heterothermy in the hedgehog Erinaceus europaeus

Summary Intra-abdominal temperature-sensitive radio transmitters were used to collect more than 350 sets of body temperature (T _b ) data from 23 captive adult hedgehogs over a 3-year period. Each data set comprised measurements made every 1/2 h for 24-h periods. Between 20 and 60 such data sets were recorded every calendar month, and a total of 17400 measurements of T _b were collected. The hedgehogs were exposed to natural environmental conditions at 57°N in NE Scotland. Hedgehogs showed seasonal changes in mean daily euthermic T _b ,with a July maximum of 35.9±0.2°C, a September minimum of 34.7±0.9°C, and a marked circadian T _b cycle that correlates closely with photoperiod. Maximal T _b occurred within 2 h of midnight and this pattern of nocturnal maximum and diurnal minimum T _b was most marked between April and September. The circadian T _b cycle was least correlated with photoperiod during winter. Hibernal T _b during winter correlated with ambient temperature (T _a ),it was maximal in September (17.7±1.0°C) and minimal in December (5.2±0.9°C). Apart from the tracking of T _a and T _b during hibernal bouts, with a time-lag of 4–6 h, circadian rhythmicity of hibernal T _b was not evident. However, the T _b of hibernating hedgehogs rose significantly when T _a fell below — 5°C, although the animals did not neccessarily arouse. Although hibernal bouts occurred between September and April, 89.5% of such bouts were recorded between November and February. The mean time of entry into hibernation was 01:45±5.1 h GMT while the mean time of the start of spontaneous arousal from hibernation was 11:53±4.8 h GMT. Therefore, during hibernation hedgehogs were either fully aroused at night, when euthermic hedgehogs have maximalT _b ,or in deep hibernation around midday, when euthermic hedgehogs have minimal T _b .Since wild hedgehogs will feed during spontaneous arousal from hibernation, these timings are probably adaptive, and suggest that entry into, and arousal from, hibernation may be extensions of circadian cyclicity. Spontaneous bouts of transient shallow torpor (TST) were recorded throughout the year, with nearly 80% of observations occurring during August and September, at the start of the hibernal period. TST bouts lasted for 4.9±2.9 h, with T _b falling to 25.8±3.1 °C. Only 20% of TST bouts immediately preceded hibernation and their duration did not correlate with T _a or body mass. TST bouts started at 06:51±4.7 h GMT, significantly later than entry into hibernation, and ended at 13:04±5.4 h GMT. The function of TST bouts is unclear, but they may be preparation for the hibernation season or a further energy conservation strategy. When arousing from hibernation hedgehogs warmed at a rate of 1.9±0.4°C·h^-1, and when entering hibernation cooled at 7.9±1.9°C·h^-1. Warming rates were slightly higher during mid-winter when T _b and body mass were minimal, but cooling rates were 44% higher at the end of the hibernal period compared to the start. Cooling and warming rates were strikingly similar to those measured in hedgehogs at 31°N. These results demonstrate that thermoregulation in the hedgehog is closely regulated and changes on a seasonal basis, in meeting with requirements of surviving food shortages and low temperature during winter.Abbreviations T _a ambient temperature - T _b body temperature - CSD circular standard deviation - SWS slow wave sleep - TST transient shallow torpor     

Taking the heat: thermoregulation in Asian elephants under different climatic conditions

Some mammals indigenous to desert environments, such as camels, cope with high heat load by tolerating an increase in body temperature (T _b) during the hot day, and by dissipating excess heat during the cooler night hours, i.e., heterothermy. Because diurnal heat storage mechanisms should be favoured by large body size, we investigated whether this response also exists in Asian elephants when exposed to warm environmental conditions of their natural habitat. We compared daily cycles of intestinal T _b of 11 adult Asian elephants living under natural ambient temperatures (T _a) in Thailand (mean T _a ~ 30°C) and in 6 Asian elephants exposed to cooler conditions (mean T _a ~ 21°C) in Germany. Elephants in Thailand had mean daily ranges of T _b oscillations (1.15°C) that were significantly larger than in animals kept in Germany (0.51°C). This was due to both increased maximum T _b during the day and decreased minimum T _b at late night. Elephant’s minimum T _b lowered daily as T _a increased and hence entered the day with a thermal reserve for additional heat storage, very similar to arid-zone ungulates. We conclude that these responses show all characteristics of heterothermy, and that this thermoregulatory strategy is not restricted to desert mammals, but is also employed by Asian elephants.     

Timing of torpor bouts during hibernation in European hamsters (Cricetus cricetus L.)

Body temperature (T _b) of seven European hamsters maintained at constant ambient temperature (T _a = 8 °C) and constant photoperiod (LD 8:16) was recorded throughout the hibernating season using intraperitoneal temperature-sensitive HF transmitters. The animals spent about 30% of the hibernation season in hypothermia and 70% in inter-bout normothermy. Three types of hypothermia, namely deep hibernation bouts (DHBs), short hibernation bouts (SHBs), and short and shallow hibernation bouts (SSHBs), were distinguished by differences in bout duration and minimal body temperature (T _m). A gradual development of SSHBs from the diel minimum of T _b during normothermy could be seen in individual hamsters, suggesting a stepwise decrease of the homeostatic setpoint of T _b regulation during the early hibernation season. Entry into hibernation followed a 24-h rhythm occurring at preferred times of the day in all three types of hypothermia. DHBs and SHBs were initiated approximately 4 h before SSHBs, indicating a general difference in the physiological initiation of SSHBs on the one hand and DHBs and SHBs on the other. Arousals from SHBs and SSHBs also followed a 24-h rhythm, whereas spontaneous arousals from DHBs were widely scattered across day and night. Statistical analyses of bout length and the interval between arousals revealed evidence for a free-running circadian rhythm underlying the timing of arousals. The results clearly demonstrate that entries into hypothermia are linked to the light/dark-cycle. However, the role of the circadian system in the timing of arousals from DHBs remains unclear. Accepted: 11 December 1996     

Standard operative temperatures and cost of thermoregulation in the arctic ground squirrel,Spermophilus undulatus

Summary Body temperatures (T _b) and daily activity patterns of free-living arctic ground squirrells (Spermophilus undulatus) were determined via telemetry at a field site in northern Alaska. Simultaneous measurements were made of ambient temperature (T _a), wind speed (V), and incident solar radiation. The operative environmental temperature (T _e) for ground squirrels was obtained from fur-covered, thin metal taxidermic models of the animals. Standard operative temperature (T _es), a comparative index of heat flow, was calculated from T _e, V, and laboratory measurements of thermal conductivity.During the period of the study (August), S. undulatus were active for about 14 h per day (06.00 to 20.00 h). T _b was high throughout the daily cycle, averaging 38–39°C. Circadian variations in T _b were slight; average T _b values dropped <1°C at night. Daytime T _b fluctuations were not closely correlated to activity or to changes in environmental conditions. Air temperatures during the study were low, usually between 10 and 15°C during the day. However, T _es in exposed areas was normally higher, even though skies were generally overcast. During periods of sunshine, T _es may be as high as 34°C. The absence of nocturnal activity may result from increased costs of thermoregulation at night, which sharply reduces foraging efficiency. The high and stable body temperatures of S. undulatus probably result from thermoneutral daytime T _es, low activity levels, and the use of well-insulated nests.     

Hibernation enhancesd-glucose uptake by intestinal brush border membrane vesicles in ground squirrels

The ability to actively transport nutrients is maintained in intestinal tissues of hibernating ground squirrels compared with their active counterparts, and shows apparent upregulation in hibernators when transport rates are normalized to tissue mass. To identify the mechanisms responsible for the preservation of transport function during the extended fast of hibernation, we studiedd-glucose uptake into jejunal brush border membrane vesicles prepared from active and hibernating 13-lined ground squirrels. Hibernators were without food and showing regular bouts of torpor for at least 6 weeks before sacrifice. Electron micrographs indicated similar microvillus heights of jejunal enterocytes in the two activity states, whereas microvillus density was slightly greater in the hibernators. Glucose uptake into brush border membrane vesicles was inversely related to medium osmolarity, indicating negligible binding of substrate to brush border membrane vesicles surfaces, and intravesicular spaces were similar in hibernating and active squirrels. Glucose uptake showed strong Na⁺ dependency in both groups, with equivalent overshoot values in the presence of Na⁺. Kinetic analysis revealed a significant increase in the maximal velocity of transport (J _max) in hibernators (55.9±5.6 nmol·min^-1·mg^-1) compared with active squirrels (36.7±5.1 nmol·min^-1·mg^-1,P<0.05), with no change inK _m. Thus, the structure and absorptive capacity of the intestinal brush border persists in fasted hibernators, and the increase inJ _max for glucose uptake during hibernation likely contributes to the enhanced Na⁺-dependent glucose absorption previously observed at the tissue level.Abbreviations BBM brush border membrane(s) - BBMV brush border membranes vesicles - SGLT1 Na⁺-glucose transporter - 3-OMG 3-orthomethylglucose - J _max maximal velocity of transport - K _m transporter affinity for substrate - T _b body temperature     

Seasonal Rhythms of Body Temperature in the Free‐Ranging Raccoon Dog (Nyctereutes procyonoides) with Special Emphasis on Winter Sleep

The raccoon dog (Nyctereutes procyonoides) is the only canid with passive overwintering in areas with cold winters, but the depth and rhythmicity of wintertime hypothermia in the wild raccoon dog are unknown. To study the seasonal rhythms of body temperature (T_b), seven free‐ranging animals were captured and implanted with intra‐abdominal T_b loggers and radio‐tracked during years 2004–2006. The average size of the home ranges was 306±26 ha, and the average 24 h T_b was 38.0±<0.01°C during the snow‐free period (May–November). The highest and lowest T_b were usually recorded around midnight (21∶00–02∶00 h) and between 05∶00–11∶00 h, respectively, and the range of the 24 h oscillations was 1.2±0.01°C. The animals lost approximately 43±6% of body mass in winter (December–April), when the average size of the home ranges was 372±108 ha. During the 2–9‐wk periods of passivity in January–March, the average 24 h T_b decreased by 1.4–2.1°C compared to the snow‐free period. The raccoon dogs were hypothermic for 5 h in the morning (06∶00–11∶00 h), whereas the highest T_b values were recorded between 16∶00–23∶00 h. The range of the 24 h oscillations increased by approximately 0.6°C, and the rhythmicity was more pronounced than in the snow‐free period. The ambient temperature and depth of snow cover were important determinants of the seasonal T_b rhythms. The overwintering strategy of the raccoon dog resembled the patterns of winter sleep in bears and badgers, but the wintertime passivity of the species was more intermittent and the decrease in the T_b less pronounced.     

Regulation of glucokinase activity in liver of hibernating ground squirrel Spermophilus undulatus

The kinetic properties of glucokinase (GLK) from the liver of active and hibernating ground squirrels Spermophilus undulatus have been studied. Entrance of ground squirrels into hibernation from their active state is accompanied by a sharp decrease in blood glucose (Glc) level (from 14 to 2.9 mM) and with a significant (7-fold) decrease of GLK activity in the liver cytoplasm. Preparations of native GLK practically devoid of other molecular forms of hexokinase were obtained from the liver of active and hibernating ground squirrels. The dependence of GLK activity upon Glc concentration for the enzyme from active ground squirrel liver showed a pronounced sigmoid character (Hill coefficient, h = 1.70 and S _0.5 = 6.23 mM; the experiments were conducted at 25°C in the presence of enzyme stabilizers, K⁺ and DTT). The same dependence of enzyme activity on Glc concentration was found for GLK from rat liver. However, on decreasing the temperature to 2°C (simulation of hibernation conditions), this dependency became almost hyperbolic (h = 1.16) and GLK affinity for substrate was reduced (S _0.5 = 23 mM). These parameters for hibernating ground squirrels (body temperature 5°C) at 25°C were found to be practically equal to the corresponding values obtained for GLK from the liver of active animals (h = 1.60, S _0.5 = 9.0 mM, respectively); at 2°C sigmoid character was less expressed and affinity for Glc was drastically decreased (h = 1.20, S _0.5 = 45 mM). The calculations of GLK activity in the liver of hibernating ground squirrels based on enzyme kinetic characteristics and seasonal changes in blood Glc concentrations have shown that GLK activity in the liver of hibernating ground squirrels is decreased about 5500-fold.     

Daily torpor in the gray mouse lemur (Microcebus murinus) in Madagascar: energetic consequences and biological significance

Patterns and energetic consequences of spontaneous daily torpor were measured in the gray mouse lemur (Microcebus murinus) under natural conditions of ambient temperature and photoperiod in a dry deciduous forest in western Madagascar. Over a period of two consecutive dry seasons, oxygen consumption (VO₂) and body temperature (T _b) were measured on ten individuals kept in outdoor enclosures. In all animals, spontaneous daily torpor occurred on a daily basis with torpor bouts lasting from 3.6 to 17.6 h, with a mean torpor bout duration of 9.3 h. On average, body temperatures in torpor were 17.3±4.9°C with a recorded minimum value of 7.8°C. Torpor was not restricted to the mouse lemurs’ diurnal resting phase: entries occurred throughout the night and arousals mainly around midday, coinciding with the daily ambient temperature maximum. Arousal from torpor was a two-phase process with a first passive, exogenous heating where the T _b of animals increased from the torpor T _b minimum to a mean value of 27.1°C before the second, endogenous heat production commenced to further raise T _b to normothermic values. Metabolic rate during torpor (28.6±13.2 ml O₂ h^–1) was significantly reduced by about 76% compared to resting metabolic rate (132.6±50.5 ml O₂ h^–1). On average, for all M. murinus individuals measured, hypometabolism during daily torpor reduced daily energy expenditure by about 38%. In conclusion, all these energy-conserving mechanisms of the nocturnal mouse lemurs, with passive exogenous heating during arousal from torpor, low minimum torpor T _bs, and extended torpor bouts into the activity phase, comprise an important and highly adapted mechanism to minimize energetic costs in response to unfavorable environmental conditions and may play a crucial role for individual fitness. Received: 8 July 1999 / Accepted: 3 December 1999