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.     

Absence of heterothermy in the European red squirrel (Sciurus vulgaris)

The European red squirrel Sciurus vulgaris inhabits areas which undergo profound seasonal declines in food availability and ambient temperature. We measured the body temperature (T_b) of free-ranging S. vulgaris over the course of one year to examine its thermoregulatory strategies and found no evidence of heterothermy, with T_b never dropping below 36.7 °C. A lower average T_b and a reduced active phase are likely to have resulted in some energetic savings, sufficient for survival during the particularly mild winter with unhindered access to food stores. We cannot exclude that a different T_b pattern may be seen in energetically more demanding years, but we can confirm that heterothermy is not an obligatory behaviour in this species to counter energetic bottlenecks. Either S. vulgaris is indeed a strict homeotherm, or the need for torpor is flexibly adjusted.     

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.     

Exogenous passive heating during torpor arousal in free-ranging rock elephant shrews, <Emphasis Type="Italic">Elephantulus myurus</Emphasis>

In the laboratory rock elephant shrews (Elephantulus myurus; mean body mass 56.6 g) displayed the lowest torpor T_{b min} yet recorded (ca. 5°C) in a placental daily heterotherm. It was unknown whether these low T_bs were characteristic of daily heterothermy in free-ranging animals. It was also unclear how cost effective these low T_bs were since considerable energy is required to arouse from low T_bs on a daily basis. We continuously measured body temperature once every hour for 85 days in 13 free-ranging E. myurus from May to August 2001 (winter) in Weenen Game Reserve, KwaZulu-Natal, South Africa. We recorded a total of 412 torpor bouts. Free-ranging E. myurus had a high propensity for torpor with females displaying higher torpor frequency than males. The lowest T_b recorded was 7.5°C at T_a=2.7°C and the minimum torpor T_b was strongly correlated with ambient temperature. Torpor arousal was tightly coupled with ambient temperature cycles. Low torpor T_{b min} at low T_as was therefore cost-effective because the animals offset the high cost of arousal through exogenous passive heating. Laboratory studies under constant ambient temperatures may therefore underestimate the energetic benefits of torpor in free-ranging small mammals that inhabit regions where seasonality is moderate.     

Body temperature variation in free-living and food-deprived yellow-necked mice sustains an adaptive framework for endothermic thermoregulation

In many mammalian species, variation in body temperature (T_b) exceeds the values suitable for defining homeothermy, making it justifiable and even necessary to resort to the term “heterothermic”. However, T_b data are only available for ca. 1% of extant mammalian species. We investigated variations in T_b in wild free-living and experimentally food-deprived yellow-necked mice Apodemus flavicollis, during the temperate-zone autumn-winter period. In line with the adaptive framework for endothermic thermoregulation, we hypothesised that T_b in the mice should be adjustable with the energetic cost-benefit trade-off associated with maintaining homeothermy. In laboratory conditions, mice clearly entered a state of daily torpor when food-deprived. Our study thus makes it clear that A. flavicollis is a heterothermic species in which food deprivation results in switching between endothermic and poikilothermic thermoregulation. We also assumed that, in free-living mice, heterothermy increases with elevated environmental challenges, e.g. when the ambient temperature (T_a) decreases. Consistent with this was the inverse correlation noted between variation in T_b in free-living mice and T_a, with most individuals clearly becoming torpid when T_a decreases below 0 °C. It is the increased cost of food hoarding under cold conditions that most likely triggers a state of torpor as a last result. Overall, our study indicates that yellow-necked mice can provide a further example of species sustaining an adaptive framework for endothermic thermoregulation.     

Thermal biology, torpor use and activity patterns of a small diurnal marsupial from a tropical desert: sexual differences

Many small desert dasyurids employ torpor almost daily during winter, because cold nights and low food availability impose high energetic costs. However, in Western Australia the arid zone extends into tropical, coastal regions, where winter temperature conditions are far less severe. We studied the thermal biology and activity patterns of free-ranging kaluta (~27 g), a dasyurid restricted to these tropical spinifex deserts, during the Austral winter (June–July) and in addition quantified activity patterns in captivity. Unlike most dasyurids, wild and captive kalutas were almost exclusively diurnal and retreated into underground burrows during the night. Despite being active during the warmer part of the day, kalutas entered torpor daily. However, torpor patterns differed remarkably between males and females. While females spent most of the night torpid at body temperatures (T _b) as low as 21°C, close to soil temperature, males entered multiple short and shallow bouts (T _b > 25°C) during the night. Males also maintained higher T _bs during the early morning when active, occupied larger home ranges and covered greater distances while foraging than females. Hence, males appear to expend more energy than the similar-sized females both while foraging and during the rest phase. We propose that physiological as well as behavioural preparations for the September mating season that culminate in a complete male die-off might already impose energetic costs on males during winter.     

Body temperature patterns in two syntopic elephant shrew species during winter

We measured body temperature (T_b) in free-ranging individuals of two species of elephant shrews, namely western rock elephant shrews (Elephantulus rupestris) and Cape rock elephant shrews (E. edwardii), during winter in a winter-rainfall region of western South Africa. These syntopic species have similar ecologies and morphologies and thus potential for large overlaps in diet and habitat use. Unexpectedly, they displayed different T_b patterns. Western rock elephant shrews were heterothermic, with all individuals decreasing T_b below 30 °C on at least 34% of nights. The level of heterothermy expressed was similar to other species traditionally defined as daily heterotherms and was inversely related to T_a, as is commonly seen in small heterothermic endotherms. In contrast, Cape rock elephant shrews rarely allowed their T_b to decrease below 30 °C. The level of heterothermy was similar to species traditionally defined as homeotherms and there was no relationship between the level of heterothermy expressed and T_a. In both species, the minimum daily T_b was recorded almost exclusively at night, often shortly before sunrise, although in some individuals minimum T_b occasionally occurred during the day. The interspecific variation in T_b patterns among Elephantulus species recorded to date reiterates the importance of ecological determinants of heterothermy that interact with factors such as body mass and phylogeny.     

Thermal physiology of pregnant and lactating female and male long-eared bats, Nyctophilus geoffroyi and N. gouldi

During roosting in summer, reproductive female bats appear to use torpor less frequently and at higher body temperatures (T _b) than male bats, ostensibly to maximise offspring growth. To test whether field observations result from differences in thermal physiology or behavioural thermoregulation during roosting, we measured the thermoregulatory response and energetics of captive pregnant and lactating female and male long-eared bats (Nyctophilus geoffroyi 8.9 g and N. gouldi 11.5 g) during overnight exposure to a constant ambient temperature (T _a) of 15°C. Bats were captured 1–1.5 h after sunset and measurements began at 21:22±0:36 h. All N. geoffroyi entered torpor commencing at 23:47±01:01 h. For N. gouldi, 10/10 males, 9/10 pregnant females and 7/8 lactating females entered torpor commencing at 01:10±01:40 h. The minimum T _b of torpid bats was 15.6±1.1°C and torpid metabolic rate (TMR) was reduced to 0.05±0.02 ml O₂ g⁻¹ h⁻¹. Sex or reproductive condition of either species did not affect the timing of entry into torpor (F=1.5, df=2, 19, P=0.24), minimum TMR (F=0.21, df=4, 40, P=0.93) or minimum T _b (F=0.76, df=5, 41, P=0.58). Moreover, sex or reproductive condition did not affect the allometric relationship between minimum resting metabolic rate and body mass (F=1.1, df=4, 37, P=0.37). Our study shows that under identical thermal conditions, thermal physiology of pregnant and lactating female and male bats are indistinguishable. This suggests that the observed reluctance by reproductive females to enter torpor in the field is predominantly because of ecological rather than physiological differences, which reflect the fact that females roost gregariously whereas male bats typically roost solitarily.     

Strategic (adaptive) hypothermia in bull dromedary camels during rut; could it increase reproductive success?

In this study of body temperatures (T_b) in free ranging dromedary camels, we found that bulls in rut start the days cooler. Daily minima during rut averaged 0.6°C lower than at other times (95% CI 0.27–0.94°C) and daily maxima averaged 0.45°C higher (95% CI −0.01 to –0.91°C), increasing the daily T_b cycle. Knut Schmidt-Nielsen described a similar pattern in captive dromedaries deprived of water in hot conditions, which he interpreted as a strategy to conserve water. Our observations were made in winter and with water freely available. Dromedaries can apparently employ heterothermy for more than just water conservation. In the strenuous daily contests between rival bulls in rut, a lower T_b early in the day should extend the time for which a contestant can challenge or defend before heat stress becomes a problem. Calculations show that lowering T_b by even 0.6°C extends that time by more than 30 min, and many daily minima during rut were lower than that. Because the eventual winner of contests gains or retains a herd of females, we speculate that cooler T_b at the start of daily contests confers an advantage which translates directly into increased reproductive success.     

Body temperature variation of free-ranging and captive southern brown bandicoots Isoodon obesulus (Marsupialia: Peramelidae)

To investigate patterns of thermoregulation in free-ranging and captive southern brown bandicoots Isoodon obesulus, we measured abdominal body temperature (T_b) of five free-ranging bandicoots over 42 days using implanted data loggers and T_b of three captive bandicoots over 3 months using implanted temperature-sensitive radio transmitters. Bandicoots in the wild had a mean T_b of 36.5±1.0 °C (range 33.4–39.8 °C) and showed a pronounced nychthemeral pattern with two distinct temperature phases. T_b increased at 13:30±2.6 h each day and remained high for 10.65±2.07 h, suggesting a crepuscular and early evening activity pattern. Daily T_b variation of I. obesulus would save considerable energy by reducing daytime thermoregulatory costs in the wild. Captive bandicoots had a similar mean body temperature (36.9±0.2°C) and range (33.0–39.9°C) as free-ranging bandicoots. However, the nychthemeral T_b pattern of captive bandicoots was different from free-ranging bandicoots, with a less pronounced daily cycle and the nocturnal rise in T_b occurring mainly at sunset and the daily decline occurring mainly at dawn.     

Thermoregulation in African Green Pigeons (Treron calvus) and a re-analysis of insular effects on basal metabolic rate and heterothermy in columbid birds

Columbid birds represent a useful model taxon for examining adaptation in metabolic and thermal traits, including the effects of insularity. To test predictions concerning the role of insularity and low predation risk as factors selecting for the use of torpor, and the evolution of low basal metabolic rate in island species, we examined thermoregulation under laboratory and semi-natural conditions in a mainland species, the African Green Pigeon (Treron calvus). Under laboratory conditions, rest-phase body temperature (T _b) was significantly and positively correlated with air temperature (T _a) between 0 and 35 °C, and the relationship between resting metabolic rate (RMR) and T _a differed from typical endothermic patterns. The minimum RMR, which we interpret as basal metabolic rate (BMR), was 0.825 ± 0.090 W. Green pigeons responded to food restriction by significantly decreasing rest-phase T _b, but the reductions were small (at most ~5 °C below normothermic values), with a minimum T _b of 33.1 °C recorded in a food-deprived bird. We found no evidence of the large reductions in T _b and metabolic rate and the lethargic state characteristic of torpor. The absence of torpor in T. calvus lends support to the idea that species restricted to islands that are free of predators are more likely to use torpor than mainland species that face the risk of predation during the rest-phase. We also analysed interspecific variation in columbid BMR in a phylogenetically informed framework and verified the conclusions of an earlier study which found that BMR is significantly lower in island species compared to those that occur on mainlands.     

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.     

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.     

Hibernation and non-shivering thermogenesis in the Hottentot golden mole (<Emphasis Type="Italic">Amblysomus hottentottus longiceps</Emphasis>)

Although heterothermy (hibernation and torpor) is a common feature among mammals, there is debate over whether it is a derived or ancestral trait relative to endothermic homeothermy. Determination of the physiological characteristics of primitive mammals is central to understanding the evolution of endothermy. Moreover, evaluation of physiological mechanisms responsible for endothermic heat production [e.g. non-shivering thermogenesis (NST)] is key to understanding how early mammals responded to historical climate changes and colonised different geographical regions. Here we investigated the capacity for NST and heterothermy in the Hottentot golden mole, a basal eutherian mammal. NST was measured as the metabolic response to injections of noradrenalin and heterothermy by recording body temperature in free-ranging animals. We found that hibernation and torpor occurred and that the seasonal phenotypic adjustment of NST capacity was similar to that found in other placental mammals. Using phylogenetically independent contrasts, we compared measured values of NST with those obtained from the literature. This showed that all variation in NST was accounted for by differences in phylogeny and not zoogeography. These findings lend support to the observation that NST and heterothermy occur in the Afrotheria, the basal placental mammalian clade. Furthermore, this work suggests that heterothermy, rather than homeothermy is a plesiomorphic trait in mammals and supports the notion that NST mechanisms are phylogenetically ancient.     

Continuous growth through winter correlates with increased resting metabolic rate but does not affect daily energy budgets due to torpor use

Small mammals that are specialists in homeothermic thermoregulation reduce their self-maintenance costs of normothermy to survive the winter. By contrast, heterothermic ones that are considered generalists in thermoregulation can lower energy expenditure by entering torpor. It is well known that different species vary the use of their strategies to cope with harsh winters in temperate zones; however, little is still known about the intraspecific variation within populations and the associated external and internal factors. We hypothesized that yellow-necked mice Apodemus flavicollis decrease their resting metabolic rate (RMR) from autumn to winter, and then increase it during spring. However, since the alternative for seasonal reduction of RMR could be the development of heterothermy, we also considered the use of this strategy. We measured body mass (m_b), RMR, and body temperature (T_b) of mice during 2 consecutive years. In the 1st year, mice decreased whole animal RMR in winter, but did not do so in the 2nd year. All mice entered torpor during the 2nd winter, whereas only a few did so during the first one. Mice showed a continuous increase of m_b, which was steepest during the 2nd year. The relationship between RMR and m_b varied among seasons and years most likely due to different mouse development stages. The m_b gain at the individual level was correlated positively with RMR and heterothermy. This indicates that high metabolism in winter supports the growth of smaller animals, which use torpor as a compensatory mechanism. Isotope composition of mice hair suggests that in the 1st year they fed mainly on seeds, while in the 2nd, they likely consumed significant amounts of less digestible herbs. The study suggests that the use of specialist or generalist thermoregulatory strategies can differ with environmental variation and associated differences in developmental processes.     

The tradeoff between torpor use and reproduction in little brown bats (Myotis lucifugus)

In mammals, reproduction, especially for females is energetically demanding. Therefore, during the reproductive period females could potentially adjust patterns of thermoregulation and foraging in concert to minimise the energetic constraints associated with pregnancy and lactation. We assessed the influence of pregnancy, lactation, and post-lactation on torpor use and foraging behaviour by female little brown bats, Myotis lucifugus. We measured thermoregulation by recording skin temperature and foraging by tracking bats which carried temperature-sensitive radio-tags. We found that individuals, regardless of reproductive condition, used torpor, but the patterns of torpor use varied significantly between reproductive (pregnant and lactating) females and post-lactating females. As we predicted, reproductive females entered torpor for shorter bouts than post-lactating females. Although all females used torpor frequently, pregnant females spent less time in torpor, and maintained higher skin temperatures than either lactating or post-lactating females. This result suggests that delayed offspring development which has been associated with torpor use during pregnancy, may pose a higher risk to an individual’s reproductive success than reduced milk production during lactation. Conversely, foraging behaviour of radio-tagged bats did not vary with reproductive condition, suggesting that even short, shallow bouts of torpor produce substantial energy savings, likely obviating the need to spend more time foraging. Our data clearly show that torpor use and reproduction are not mutually exclusive and that torpor use (no matter how short or shallow) is an important means of balancing the costs of reproduction for M. lucifugus.     

The energetic cost of arousal from torpor in the marsupial Sminthopsis macroura : benefits of summer ambient temperature cycles

The costs of arousal from induced torpor were measured in the striped-faced dunnart (Sminthopsis macroura; ca. 25 g) under two experimental ambient temperature cycles. The sinusoidal-type temperature cycles were designed to evaluate the effects of passive, ambient temperature heating during arousal from torpor in these insectivorous marsupials. It was hypothesised that diel ambient temperature cycles may offer significant energy savings during arousal in animals that employ daily torpor in summer as a response to unpredictable food availability. The cost of arousal in animals in which passive, exogenous heating occurred was significantly lower than that in animals not exposed to an ambient temperature cycle. The total cost of all three phases of torpor (entry, maintenance and arousal) was almost halved when animals were exposed to an ambient heating cycle from 15 °C to 25 °C over a 24-h period. In all animals, irrespective of the experimental ambient temperature cycle employed, the minimum torpor body temperature was 17–18 °C. The body temperature (T_b) of animals exposed to exogenous heating increased from the torpor T_b minimum to a mean value of 22.59 °C before endogenous heat production commenced. This relatively small increase in T_b of ca. 5 °C through `free' passive heating was sufficient to account for the significant ca. three-fold decrease in the cost of arousal and may represent an important energetic aid to free-ranging animals. Accepted: 4 October 1998     

Baby in the bathwater: Should we abandon the use of body temperature thresholds to quantify expression of torpor?

Boyles et al. (this issue) argue against the use of body temperature (T_b) thresholds to quantify the expression of torpor in endotherms and our purpose is to provide a counterpoint argument. We contend that T_b thresholds provide valuable information about ecological factors influencing the evolution of thermoregulation. We also point out shortcomings of the so-called heterothermy index proposed as an alternative. However, to be clear, we do agree with Boyles et al. (this issue) that the use of torpor thresholds can limit some aspects of the study of thermoregulation and applaud the more widespread incorporation of theoretical underpinnings proposed by Boyles et al. (this issue) and others.     

Cytoskeletal Regulation Dominates Temperature-Sensitive Proteomic Changes of Hibernation in Forebrain of 13-Lined Ground Squirrels

13-lined ground squirrels, Ictidomys tridecemlineatus, are obligate hibernators that transition annually between summer homeothermy and winter heterothermy – wherein they exploit episodic torpor bouts. Despite cerebral ischemia during torpor and rapid reperfusion during arousal, hibernator brains resist damage and the animals emerge neurologically intact each spring. We hypothesized that protein changes in the brain underlie winter neuroprotection. To identify candidate proteins, we applied a sensitive 2D gel electrophoresis method to quantify protein differences among forebrain extracts prepared from ground squirrels in two summer, four winter and fall transition states. Proteins that differed among groups were identified using LC-MS/MS. Only 84 protein spots varied significantly among the defined states of hibernation. Protein changes in the forebrain proteome fell largely into two reciprocal patterns with a strong body temperature dependence. The importance of body temperature was tested in animals from the fall; these fall animals use torpor sporadically with body temperatures mirroring ambient temperatures between 4 and 21°C as they navigate the transition between summer homeothermy and winter heterothermy. Unlike cold-torpid fall ground squirrels, warm-torpid individuals strongly resembled the homeotherms, indicating that the changes observed in torpid hibernators are defined by body temperature, not torpor per se. Metabolic enzymes were largely unchanged despite varied metabolic activity across annual and torpor-arousal cycles. Instead, the majority of the observed changes were cytoskeletal proteins and their regulators. While cytoskeletal structural proteins tended to differ seasonally, i.e., between summer homeothermy and winter heterothermy, their regulatory proteins were more strongly affected by body temperature. Changes in the abundance of various isoforms of the microtubule assembly and disassembly regulatory proteins dihydropyrimidinase-related protein and stathmin suggested mechanisms for rapid cytoskeletal reorganization on return to euthermy during torpor-arousal cycles.     

Mitochondrial metabolism during fasting-induced daily torpor in mice

During fasting, mice (Mus musculus) undergo daily bouts of torpor, considerably reducing body temperature (T_b) and metabolic rate (MR). We examined females of different laboratory strains (Balb/c, C57/6N, and CD1) to determine whether liver mitochondrial metabolism is actively reduced during torpor. In all strains, we found that state 3 (phosphorylating) respiration rate measured at 37 °C was reduced up to 35% during torpor for at least one of the substrates (glutamate and succinate) used to fuel respiration. The extent of this suppression varied and was correlated with T_b at sampling. This suggests that, at the biochemical level, the transition to and from a hypometabolic torpid state is gradual. In fasted non-torpid animals, T_b and MR still fluctuated greatly: T_b dropped by as much as 4 °C and MR was reduced up to 25% compared to fed controls. Changes in T_b and MR in fasted, non-torpid animals were correlated with changes in mitochondrial state 3 respiration rate measured at 37 °C. This suggests that fasting mice may conserve energy even when not torpid by occasionally reducing T_b and mitochondrial oxidative capacity to reduce MR. Furthermore, proton conductance was higher in torpid compared to non-torpid animals when measured at 15 °C (the lower limit of torpid T_b). This pattern is similar to that reported previously for daily torpor in Phodopus sungorus.