Thermal biology,torpor and behaviour in sugar gliders: a laboratory-field comparison

Most studies on animal physiology and behaviour are conducted in captivity without verification that data are representative of free-ranging animals. We provide the first quantitative comparison of daily torpor, thermal biology and activity patterns, conducted on two groups of sugar gliders (Petaurus breviceps, Marsupialia) exposed to similar thermal conditions, one in captivity and the other in the field. Our study shows that activity in captive gliders in an outdoor aviary is restricted to the night and largely unaffected by weather, whereas free-ranging gliders omit foraging on cold/wet nights and may also forage in the afternoon. Torpor occurrence in gliders was significantly lower in captivity (8.4% after food deprivation; 1.1% for all observations) than in the field (25.9%), mean torpor bout duration was shorter in captivity (6.9 h) than in the field (13.1 h), and mean body temperatures during torpor were higher in captivity (25.3°C) than in the field (19.6°C). Moreover, normothermic body temperature as a function of air temperature differed between captive and free-ranging gliders, with a >3°C difference at low air temperatures. Our comparison shows that activity patterns, thermal physiology, use of torpor and patterns of torpor may differ substantially between the laboratory and field, and provides further evidence that functional and behavioural data on captive individuals may not necessarily be representative of those living in the wild.     

Daily torpor and hibernation in birds and mammals

Many birds and mammals drastically reduce their energy expenditure during times of cold exposure, food shortage, or drought, by temporarily abandoning euthermia, i.e. the maintenance of high body temperatures. Traditionally, two different types of heterothermy, i.e. hypometabolic states associated with low body temperature (torpor), have been distinguished: daily torpor, which lasts less than 24 h and is accompanied by continued foraging, versus hibernation, with torpor bouts lasting consecutive days to several weeks in animals that usually do not forage but rely on energy stores, either food caches or body energy reserves. This classification of torpor types has been challenged, suggesting that these phenotypes may merely represent extremes in a continuum of traits. Here, we investigate whether variables of torpor in 214 species (43 birds and 171 mammals) form a continuum or a bimodal distribution. We use Gaussian‐mixture cluster analysis as well as phylogenetically informed regressions to quantitatively assess the distinction between hibernation and daily torpor and to evaluate the impact of body mass and geographical distribution of species on torpor traits. Cluster analysis clearly confirmed the classical distinction between daily torpor and hibernation. Overall, heterothermic endotherms tend to be small; hibernators are significantly heavier than daily heterotherms and also are distributed at higher average latitudes (～35°) than daily heterotherms (～25°). Variables of torpor for an average 30 g heterotherm differed significantly between daily heterotherms and hibernators. Average maximum torpor bout duration was >30‐fold longer, and mean torpor bout duration >25‐fold longer in hibernators. Mean minimum body temperature differed by ～13°C, and the mean minimum torpor metabolic rate was ～35% of the basal metabolic rate (BMR) in daily heterotherms but only 6% of BMR in hibernators. Consequently, our analysis strongly supports the view that hibernators and daily heterotherms are functionally distinct groups that probably have been subject to disruptive selection. Arguably, the primary physiological difference between daily torpor and hibernation, which leads to a variety of derived further distinct characteristics, is the temporal control of entry into and arousal from torpor, which is governed by the circadian clock in daily heterotherms, but apparently not in hibernators.     

Confirmation of pleisiomorphic daily torpor in mammals: the round-eared elephant shrew Macroscelides proboscideus (Macroscelidea)

The characteristics of daily torpor were measured in the round-eared elephant shrew Macroscelides proboscideus (Macroscelidea) in response to ambient temperature and food deprivation. Elephant shrews are an ancient mammal order within a superordinal African clade including hyraxes, elephants, dugongs and the aardvark. M. proboscideus only employed torpor when deprived of food; torpor did not occur under an ad libitum diet at ambient temperatures of 10, 15 and 25?°C. Torpor bout duration ranged from <1?h to ca. 18?h. The times of entry into torpor were restricted to the scotophase, despite normothermic body temperature patterns indicating a rest phase coincident with the photophase. Full arousal was always achieved within the first 3?h of the photophase. When food deprived, the onset of the rest phase, and hence torpor, advanced with respect to the experimental photoperiod. The lowest torpor body temperature measured was 9.41?°C. Daily torpor in M. proboscideus confirms a pleisiomorphic origin of daily heterothermy. Torpor facilitates risk-averse foraging behaviour in these small omnivores by overcoming long-term energy shortfalls generated by the inherent variability of food availability in their semi-arid, El Niño-afflicted habitats.     

Influence of Host Deprivation and Egg Expenditure on the Patch and Host-Finding Behavior of the Parasitoid Wasp <Emphasis Type="Italic">Mastrus ridibundus</Emphasis>

The influence of host deprivation and egg expenditure on the host location behavior ofMastrus ridibundus was evaluated under controlled conditions in a walk-in field cage in a greenhouse. The activity of females peaked at a temperature of 25–27°C generating two peaks of activity through the day. Increased host deprivation did not influence patch and host-finding success: females responded to host stimuli even after 9 days of host deprivation. In contrast, the longer M. ridibundus was exposed to excess hosts, the greater was the reduction in patch and host-finding success. Egg depletion expressed as the percentage of realized lifetime fecundity expended, rather than egg load, offered a good match for this decline. These results are discussed in the context of biological control introductions and the choice of pre-release treatment.     

Torpor characteristics and energy requirements of furless Siberian hamsters.

After approximately 10 wk of exposure to decreasing day lengths, Siberian hamsters (Phodopus sungorus) begin to display spontaneous torpor bouts several times each week. Torpor is associated with reduced daily energy expenditure and lower food consumption and ameliorates the thermoregulatory challenges of winter. We tested the extent to which the energy savings conferred by daily torpor depend on the presence of an insulative pelage. Female hamsters were housed in a winter day length (8L:16D) at 5 degrees C; daily food intake and torpor characteristics were recorded for 5 wk in shaved (furless) or normal hamsters. Torpor-bout incidence decreased by 62% in furless hamsters, but the duration of individual bouts and the minimum body temperature attained during torpor were unaffected by loss of pelage. Body temperature declined more rapidly during entry into torpor and increased more slowly during arousal from torpor in furless than in control hamsters. Energy savings per torpor bout, assessed by the amount of food consumed on days that included a torpor bout, was substantially greater in normal than in furless hamsters (16.0% vs. 3.3%); this difference likely reflects the increased cost of thermoregulation during torpor, as well as the increased caloric expenditure incurred by furless hamsters during arousal from torpor. An insulative pelage may be a prerequisite for the energetic benefits derived from heterothermy in this species.     

Energy and water balance during torpor and hydropenia in the pigmy gerbil,Gerbillus pusillus

Summary Under ad lib. food conditions,G. pusillus used 107% of the predicted daily energy expenditure. This increased significantly with water deprivation to 116% of the allometric expected value, thereby reflecting an increase in activity in search of food with high moisture content, a shift to metabolic water for meeting the animal's water requirements and a change in the state of hydration of the animal. Economic water expenditure through efficient kidney concentrating ability, reduced pulmocutaneous evaporation, reduced faecal water loss, and tolerance to haemoconcentration, was sufficient to impart virtual independence of exogenous water. Gerbillus pusillus responded to food shortage by abandoning homeothermy. In doing so, it was able to maintain a fairly stable body mass by monitoring energy intake and ensuring that energy expenditure did not exceed intake. There was no significant difference in energy expenditure between gerbils on a restricted energy intake and ad lib. water and those gerbils on a restricted energy intake and no water. Both these experimental groups were able to maintain a positive water balance, for whilst less metabolic water was available, water loss primarily through evaporation decreased concomittantly with reduced body temperature. In addition, the volume of urine excreted when torpid accounted for approximately 5% of the volume excreted by coenothermic rodents deprived of free water. Urine, although less concentrated than that of non-torpid gerbils deprived of water, was ten times more concentrated than the plasma. Daily torpor, although uniquely suited to energy conservation, is not without cost. A reduced body temperature resulted in a decline in ‘apparent assimilation efficiency’ (AE). It seems therefore that the cost of maintaining an elevated body temperature for the full duration of digestion is impractical. The cost of employing torpor as calculated from the reduced AE of food (0.5 kJ·day⁻¹), was however insignificant in comparison to the savings accured (32.25 kJ·day⁻¹) by the use of torpor.     

That’s hot: golden spiny mice display torpor even at high ambient temperatures

Golden spiny mice (Acomys russatus) living in the Judean desert are exposed to extended periods of food and water shortage. We investigated their thermal and metabolic response to three weeks of 50 % food reduction at ambient temperatures of 23, 27, 32 and 35 °C by long term records of metabolic rate and body temperature in the laboratory. At all ambient temperatures, A. russatus responded to starvation by a reduction of daily energy expenditure. At 32 and 35 °C, this metabolic adjustment fully compensated the reduced food availability and they maintained their energy balance at a slightly reduced body mass. At lower ambient temperatures, they could not fully compensate for the reduced food availability and kept a negative energy balance. The reduction of daily energy expenditure was largely achieved by the occurrence of daily torpor. Torpor even occurred at high ambient temperatures of 32 and 35 °C during which metabolic depression was not associated with a marked decrease of body temperature. The results show that the occurrence of daily torpor is not necessarily linked to cold exposure and the development of a pronounced hypothermia, but may even occur as depression of metabolic rate in a hot environment.     

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.     

Seasonal changes in daily torpor patterns of free-ranging female and male Daubenton’s bats (Myotis daubentonii)

Daily torpor can provide significant energy and water savings in bats during cold ambient temperatures and food scarcity. However, it may reduce rates of foetal and juvenile development. Therefore, reproductive females should optimize development by minimizing times in torpor. To test this hypothesis, the use of torpor by female and male free-ranging Daubenton’s bats (Myotis daubentonii) during reproduction (gestation, lactation, and post-lactation period) was investigated in 1998 and 1999. Temperature-sensitive radio transmitters were attached to the bats to measure skin temperature. Simultaneously, ambient temperature was recorded. While both sexes became torpid during daytime, male bats used daily torpor (>6°C below individual active temperature) significantly more often during reproductive period (mean: 78.4 % of day time in May and 43 % in June) than females. Female bats went into daily torpor, particularly in late summer when juveniles were weaned (mean: 66.6 % of daytime). Lowest skin temperatures occurred in a female bat with 21.0°C during post-lactation. Skin temperatures of male bats fluctuated from 16.8°C in torpor to 37.2°C during times of activity. There was a significant effect of reproductive period on skin temperature in females whereas mean ambient temperature had no significant effect. However, mean ambient temperature affected mean skin temperatures in males. Our findings indicate that female Daubenton’s bats adopt their thermoregulatory behaviour in particular to optimize the juvenile development.     

How to budget metabolic energy: torpor in a small Neotropical mammal

Neotropical nectar-feeding bats (Glossophaginae) are highly specialized in the exploitation of floral nectar and have one of the highest mass-specific metabolic rates among mammals. Nevertheless, they are distributed throughout the tropics and subtropics over a wide elevational range, and thus encounter many extreme and energetically challenging environmental conditions. Depressing their otherwise high metabolic rate, e.g., in situations of food restriction, might be an important adaptive physiological strategy in these dietary specialists. We investigated the thermoregulatory behavior of captive 10-g nectar feeding bats (Glossophaga soricina; Chiroptera, Phyllostomidae) under variable ambient temperatures (T _a) and feeding regimes and predicted that bats would use torpor as an energy-conserving behavior under energetic constraints. All tested animals entered torpor in response to energetic restrictions and the depth of torpor was dependent on the body condition of the animals and hence on their degree of physiological constraints. Periods of torpor with body temperatures (T _b) below 34°C were precisely adjusted to the photoperiod. The median length of diurnal torpor was 11.43 h. The lowest T _b measured was 21°C at a T _a of 19°C. Estimated energy savings due to torpor were considerable, with reductions in metabolic rate to as low as 5% of the metabolic rate of normothermic bats at the same T _a. However, contrary to temperate zone bats that also employ diurnal torpor, G. soricina regulated their T _b to the highest possible levels given the present energetic supplies. To summarize, G. soricina is a precise thermoregulator, which strategically employs thermoregulatory behavior in order to decrease its energy expenditure when under energetic restrictions. This adaptation may play a crucial role in the distribution and the assembly of communities of nectar-feeding bats and may point to a general capacity for torpor in tropical bats.     

Development of thermoregulation and torpor in a marsupial: energetic and evolutionary implications

Altricial mammals and birds become endothermic at about half the size of adults and presumably would benefit energetically from entering torpor at that time. Because little is known about torpor during development in endotherms, we investigated whether after the establishment of endothermic thermoregulation (i.e. the ability to maintain a high body temperature during cold exposure), Sminthopsis macroura, a small (∼25 g) insectivorous marsupial, is capable of entering torpor and whether torpor patterns change with growth. Endothermic thermoregulation was established when the nest young reached a body mass of ∼10 g, and they were capable of entering torpor early during development at ∼10–12 g, lending some support to the view that torpor is a phylogenetically old mammalian trait. Torpor bout length shortened significantly and the minimum metabolic rate during torpor increased as juveniles approached adult size, and consequently total daily energy expenditure increased steeply with age. Relationships between total daily energy expenditure and body mass during development of S. macroura (slope ∼1.3) differed substantially from the relationship between basal metabolism and body mass in adult endotherms (slope ∼0.75) suggesting that the energy expenditure–size relationship during the development differs substantially from that in adults under thermo-neutral conditions. Our study shows that while torpor can substantially reduce energy expenditure during development of endotherms and hence is likely important for survival during energy bottlenecks, it also may enhance somatic growth when food is limited. We therefore hypothesize that torpor during the development in endotherms is far more widespread than is currently appreciated.     

Interactive effects of temperature and photoperiod on the daily activity and energy metabolism of pouched mice (Saccostomus campestris: Cricetidae) from southern Africa

The daily activity and energy metabolism of pouched mice (Saccostomus campestris) from two localities in southern Africa was examined following warm (25 °C) and cold (10 °C) acclimation under long (LD 14:10) and short (LD 10:14) photoperiol. There was no differential effect of photoperiod on the daily activity or metabolism of pouched mice from the two localities examined, which suggests that reported differences in photoresponsivity between these two populations were not the result of differences in daily organisation. Neverthe-less, there was a significant increase in metabolism at 10 °C, irrespective of photoperiod, even though seven cold-acclimated animals displayed bouts of spontaneous torpor and saved 16.4–36.2% of their daily energy expenditure. All but one of these bouts occurred under short photoperiod, which suggests that short photoperiod facilitated the expression of torpor and influenced the daily energy metabolism of these individuals. As expected for a noctureal species, the amount of time spent active increased following acclimation to short photoperiod at 25 °C. However, there was a reduction in mean activity levels under short photoperiod at 10 °C, possibly because the stimulation of activity by short photoperiod was masked by a reduction in activity during bouts of spontaneous torpor. Cold temperature clearly had an overriding effect on the daily activity and metabolism of this species by necessitating an increase in metabolic heat production and eliciting spontaneous torpor which overrode the effect of short photoperiod on activity at an ambient temperature of 10 °C.Abbreviations 3-ANOVA three-way analysis of variance - %ACT percentage of time spent active - ADMR average daily metabolic rate - M _b body mass - MR metabolic rate - MR_dark metabolic rate recorded during the dark phase - MR_light metabolic rate recorded during the light phase - NST non-shivering thermogenesis - RQ respiratory quotient - STPD standard temperature and pressure, dry - T _a ambient temperature - T _b body temperature - VO₂ oxygen consumption     

Seasonal changes in energetics and torpor patterns in the subtropical blossom-bat Syconycteris australis (Megachiroptera)

Little is known about how animals from tropical and subtropical climates adjust their energy expenditure to cope with seasonal changes of climate and food availability. To provide such information, we studied the thermal physiology, torpor patterns and energetics of the nocturnal blossom-bat (Syconycteris australis 18 g) from a subtropical habitat in both summer and winter. In both seasons, S. australis frequently entered daily torpor at ambient temperatures between 12 and 25°C when food and water were withheld. Unlike patterns observed in temperate animals, mean minimum metabolic rates during torpor were lower in summer (0.47 ± 0.07 ml O₂ g⁻¹ h⁻¹) than in winter (0.75 ± 0.11 ml O₂ g⁻¹ h⁻¹). Body temperatures during torpor were regulated at 19.3 ± 1.0°C in summer and at 23.4 ± 2.0°C in winter. Torpor bout duration was significantly longer in summer (7.3 ± 0.6 h) than in winter (5.5 ± 0.3 h), but in both seasons, bout duration was not affected by ambient temperature. Consequently, average daily metabolic rates were also significantly lower in summer than in winter. Body temperatures and metabolic rates in normothermic bats did not change with season. Our findings on seasonal changes of torpor in this bat from the subtropics are opposite to those made for many species from cold climates which generally show deeper and longer torpor in winter and are often entirely homeothermic in summer. More pronounced torpor in subtropical S. australis in summer may be due to low or unpredictable nectar availability, short nights which limit the time available for foraging, and long days without access to food. Thus, the reversed seasonal response of this subtropical bat in comparison to temperate species may be an appropriate response to ecological constraints. Received: 6 May 1997 / Accepted: 19 October 1997     

Torpor patterns,arousal rates,and temporal organization of torpor entry in wildtype and UCP1-ablated mice

In eutherian mammals, uncoupling protein 1 (UCP1) mediated non-shivering thermogenesis from brown adipose tissue (BAT) provides a mechanism through which arousal from torpor and hibernation is facilitated. In order to directly assess the magnitude by which the presence or absence of UCP1 affects torpor patterns, rewarming and arousal rates within one species we compared fasting induced torpor in wildtype (UCP1^+/+) and UCP1-ablated mice (UCP^−/−). Torpor was induced by depriving mice of food for up to 48 h and by a reduction of ambient temperature (T _a) from 30 to 18°C at four different time points after 18, 24, 30 and 36 h of food deprivation. In most cases, torpor bouts occurred within 20 min after the switch in ambient temperature (30–18°C). Torpor bouts expressed during the light phase lasted 3–6 h while significantly longer bouts (up to 16 h) were observed when mice entered torpor during the dark phase. The degree of hypometabolism (5–22 ml h⁻¹) and hypothermia (19.5–26.7°C) was comparable in wildtype and UCP1-ablated mice, and both genotypes were able to regain normothermia. In contrast to wildtype mice, UCP1-ablated mice did not display multiple torpor bouts per day and their peak rewarming rates from torpor were reduced by 50% (UCP1^+/+: 0.24 ± 0.08°C min⁻¹; UCP1^−/−: 0.12 ± 0.04°C min⁻¹). UCP1-ablated mice therefore took significantly longer to rewarm from 25 to 32°C (39 vs. 70 min) and required 60% more energy for this process. Our results demonstrate the energetic benefit of functional BAT for rapid arousal from torpor. They also suggest that torpor entry and maintenance may be dependent on endogenous rhythms.     

Torpor patterns in the pouched mouse (Saccostomus campestris ; Rodentia): a model animal for unpredictable environments

Patterns of spontaneous and induced daily torpor were measured in the Afrotropical pouched mouse (77–115?g), Saccostomus campestris, in response to photoperiod, temperature, and food deprivation, using temperature telemetry. Photoperiod had no influence on the incidence, depth, or duration of daily torpor in either males and females. Although the testis size index decreased in response to food deprivation and photoperiod by a maximum of 24%, full testis regression did not occur. Torpor bout duration was, on average, 5.3?h, independent of photoperiod and ambient temperature. Males did not enter torpor in response to food deprivation but did in response to low ambient temperature, though significantly less frequently than females. At normothermia, the body temperatures (daily minimum, mean, maximum) of males were significantly lower than those of females. Minimum body temperatures of both males and females during torpor did not fall below 20?°C at an ambient temperature of 15?°C. The patterns of torpor measured here differ from those observed in species from strongly seasonal environments. They suggest adaptation to an environment rendered unpredictable by the El Niño Southern Oscillations. As an aseasonal, opportunistic breeder capable of year-round adaptive hypothermia, the pouched mouse represents an excellent model animal for research on physiological and behavioral adaptations to unpredictable environments.     

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.     

Basking and torpor in a rock-dwelling desert marsupial: survival strategies in a resource-poor environment

Australian deserts are characterized by unpredictability, low primary productivity, and high temperature fluctuations. Despite these adverse conditions the diversity of small insectivorous marsupials of the family Dasyuridae is surprisingly high. We quantified the thermal biology of the dasyurid Pseudantechinus madonnellensis (body mass ∼30 g) in the wild to gain some understanding of whether the success of dasyurids in the arid zone may be related to some extent to their use of energy conservation strategies. In winter, most free-ranging Pseudantechinus frequently (58.3% of 131 animal days) entered daily torpor after midnight (mean 0157 hours) in rock crevices when outside ambient temperatures (T _a) were low. Most animals remained torpid until the next morning when they moved while still torpid from rock crevices to sun-exposed basking sites. We visually observed basking during rewarming from torpor (mean commencement at 0943 hours) at body temperatures (T _b) as low as 19.3°C when radiant heat was high and T _a was rising. Basking continued for the rest of the day. Torpor use was not strongly correlated with T _a, but the temporal organization of daily torpor and activity were apparently linked to the thermal characteristics of basking sites. Our study suggests that by frequently employing daily torpor and basking and by appropriately coordinating their thermal biology with that of specific locations in their environment, Pseudantechinus can reduce daily energy expenditure and thus can live and reproduce in a challenging environment. It is likely that the success of other small dasyurids and perhaps many other small mammals living in deserts is linked to employment of torpor and basking for energy conservation.     

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     

Naloxone inhibition of stress-induced daily torpor

$\text{Peromyscus maniculatus}$, deermice, were induced into daily torpor by restricting food to one-half daily ration. Intraperitoneal injection of naloxone (20 mg/kg) into mice habituated to daily IP injections of saline inhibited or modified the expression of daily torpor. In those individuals demonstrating long duration/ deep bouts (greater than 300 min/body temperature 20°C or below) naloxone administration resulted in 1) a significant decrease in the duration of torpor, 2) a significant elevation in minimum body temperatures attained during torpor and 3) a significant delay in the initiation time of torpor. In those individuals demonstrating short duration/shallow bouts (less than 300 min/ body temperatures above 20°C), naloxone administration resulted only in a significant delay of initiation time. Upon subsequent return to saline administration, however, these mice displayed a significant increase in the duration and depth of torpor. The results suggest that the endogenous opiates modulate the state of daily torpor.