Deep Torpor Research Articles

Extreme and variable torpor among high-elevation Andean hummingbird species.

Torpor is thought to be particularly important for small endotherms occupying cold environments and with limited fat reserves to fuel metabolism, yet among birds deep torpor is both rare and variable in extent. We investigated torpor in hummingbirds at approximately 3800 m.a.s.l. in the tropical Andes by monitoring body temperature (Tb) in 26 individuals of six species held captive overnight and experiencing natural air temperature (Ta) patterns. All species used pronounced torpor, with one Metallura phoebe reaching a minimum Tb of 3.26°C, the lowest yet reported for any bird or non-hibernating mammal. The extent and duration of torpor varied among species, with overnight body mass (Mb) loss negatively correlated with both minimum Tb and bout duration. We found a significant phylogenetic signal for minimum Tb and overnight Mb loss, consistent with evolutionarily conserved thermoregulatory traits. Our findings suggest deep torpor is routine for high Andean hummingbirds, but evolved species differences affect its depth.

Torpor expression is associated with differential spermatogenesis in hibernating eastern chipmunks.

Hibernators suppress physiological processes when expressing torpor, yet little is known about the effects of torpor on male reproductive physiology. Studies of hibernating mammals suggest that deep torpor negatively impacts spermatogenesis and that transitions between torpor and euthermic arousals increase cellular oxidative stress, with potentially damaging effects on sperm. Here, we hypothesize that variation in torpor expression affects the reproductive readiness of hibernators by impacting their sperm production. To test this, we examined the relationship between torpor expression and spermatogenesis in captive eastern chipmunks (Tamias striatus). We determined torpor depth with temperature data loggers and assessed its relationship with spermatogenesis by examining spermatogenic progression, cell division, sperm counts, sperm maturity, and DNA damage. We show that deep hibernators (high levels of torpor) largely halted spermatogenesis in late hibernation in comparison with shallow hibernators (low levels of torpor), where ongoing spermatogenesis was observed. Despite these differences in spermatogenic state during hibernation, spermatogenic progression, sperm numbers, and maturity did not differ in spring, potentially reflecting similar degrees of reproductive readiness. Interestingly, shallow hibernators exhibited higher rates of DNA damage in spermatogenic cells during hibernation, with this trend reversing in spring. Our results thus indicate that once heterothermy is terminated, deep hibernators resume spermatogenesis but are characterized by higher rates of DNA damage in spermatogenic cells at the seasonal stage when spring mating commences. Therefore, our study confirmed posthibernation recovery of sperm production but also a potential impact of deep torpor expression during winter on DNA damage in spring.

Always a price to pay: hibernation at low temperatures comes with a trade-off between energy savings and telomere damage.

We experimentally tested the costs of deep torpor at low temperatures by comparing telomere dynamics in two species of rodents hibernating at either 3 or 14°C. Our data show that hibernators kept at the warmer temperature had higher arousal frequencies, but maintained longer telomeres than individuals hibernating at the colder temperature. We suggest that the high-energy demand of frequent arousals is counteracted by a lower temperature differential between torpid and euthermic body temperature and that telomere length is restored during arousals when the body temperature is returned to normothermic values. Taken together, our study shows that hibernation at low body temperatures comes with costs on a cellular level and that hibernators need to actively counterbalance the shortening of telomeres.

Marmots

Walter Arnold introduces the biology of marmots.

Neuronal Activity in the Hibernating Brain.

Hibernation is a natural phenomenon in many species which helps them to survive under extreme ambient conditions, such as cold temperatures and reduced availability of food in the winter months. It is characterized by a dramatic and regulated drop of body temperature, which in some cases can be near 0°C. Additionally, neural control of hibernation is maintained over all phases of a hibernation bout, including entrance into, during and arousal from torpor, despite a marked decrease in overall neural activity in torpor. In the present review, we provide an overview on what we know about neuronal activity in the hibernating brain focusing on cold-induced adaptations. We discuss pioneer and more recent in vitro and in vivo electrophysiological data and molecular analyses of activity markers which strikingly contributed to our understanding of the brain’s sensitivity to dramatic changes in temperature across the hibernation cycle. Neuronal activity is markedly reduced with decreasing body temperature, and many neurons may fire infrequently in torpor at low brain temperatures. Still, there is convincing evidence that specific regions maintain their ability to generate action potentials in deep torpor, at least in response to adequate stimuli. Those regions include the peripheral system and primary central regions. However, further experiments on neuronal activity are needed to more precisely determine temperature effects on neuronal activity in specific cell types and specific brain nuclei.

Molecular control of protein synthesis, glucose metabolism, and apoptosis in the brain of hibernating thirteen-lined ground squirrels.

Thirteen-lined ground squirrels (Ictidomys tridecemlineatus) are excellent models for studying acute brain ischemia because they show high resistance to reductions in blood flow and oxygen delivery without evidence of neurological damage. In this study, we analyzed the insulin signaling pathway and regulation of mitochondrial substrate oxidation in three regions of ground squirrel brain (forebrain, cerebellum, and brainstem), comparing summer, late torpor, and interbout arousal conditions. We found select decreases in phospho-Akt in the cerebellum during torpor compared with summer animals, as well as select increases in the forebrain during interbout arousal, suggesting that Akt may influence either metabolism or cytoprotective pathways. The phosphoprotein abundance of glycogen synthase kinase 3 beta (GSK3β) showed the most consistent trend across all three brain regions, with peak increases observed during deep torpor, suggesting a crucial role for this protein during hibernation. Furthermore, all three regions of the brain showed increased phospho-protein abundance of pyruvate dehydrogenase at serine 232 during both deep torpor and interbout arousal, and serine 300 during interbout arousal only, whereas other phosphorylation sites showed a region-specific expression pattern. Information collected from these studies sheds light on the molecular controls governing insulin signaling and fuel utilization in the brain of hibernating ground squirrels.

Shallow Torpor Expression in Free-Ranging Common Hamsters With and Without Food Supplements

Energy expenditure during winter can be reduced by expressing torpor, which is characterized by decreased metabolic rate and body temperature. In addition to deep, multiday torpor bouts alternating with short arousal episodes during the hibernation period, common hamsters can also enter shallow torpor bouts (STBs), lasting for less than 24 h at a minimum body temperature between 30-20 °C. Food supplements provided shortly before winter have been shown to shorten hibernation in males, but did not affect hibernation duration in females. In the presented study, we analyzed the expression of STBs and compared supplemented to unsupplemented common hamsters. Body temperature during winter was recorded using subcutaneously implanted data loggers (iButtons). The results revealed that supplemented males showed more STBs and thus spent more time in shallow torpor than unsupplemented individuals. The duration of STBs, however, was shorter in supplemented males and both mean and minimum body temperature were significantly higher compared to unsupplemented males. In females, shallow torpor expression did not differ between individuals with and without food supplements. STBs were mainly expressed before the onset of the first deep, multiday torpor bout, but the number of STBs was not related to that of deep torpor bouts. These results indicate that in males, shallow torpor combined with feeding on food stores could be the more appropriate overwintering strategy when sufficient external energy reserves are available. Females generally cache more food than males and are therefore assumed to be less affected by the additional food stores. These results underline the flexibility of the species in the use of heterothermy and could enable adequate and rapid responses to changes in food availability.

Cell Death Patterns Due to Warm Ischemia or Reperfusion in Renal Tubular Epithelial Cells Originating from Human, Mouse, or the Native Hibernator Hamster.

Ischemia–reperfusion injury contributes to the pathogenesis of many diseases, with acute kidney injury included. Hibernating mammals survive prolonged bouts of deep torpor with a dramatic drop in blood pressure, heart, and breathing rates, interspersed with short periods of arousal and, consequently, ischemia–reperfusion injury. Clarifying the differences under warm anoxia or reoxygenation between human cells and cells from a native hibernator may reveal interventions for rendering human cells resistant to ischemia–reperfusion injury. Human and hamster renal proximal tubular epithelial cells (RPTECs) were cultured under warm anoxia or reoxygenation. Mouse RPTECs were used as a phylogenetic control for hamster cells. Cell death was assessed by both cell imaging and lactate dehydrogenase (LDH) release assay, apoptosis by cleaved caspase-3, autophagy by microtubule-associated protein 1-light chain 3 B II (LC3B-II) to LC3B-I ratio, necroptosis by phosphorylated mixed-lineage kinase domain-like pseudokinase, reactive oxygen species (ROS) fluorometrically, and lipid peroxidation, the end-point of ferroptosis, by malondialdehyde. Human cells died after short periods of warm anoxia or reoxygenation, whereas hamster cells were extremely resistant. In human cells, apoptosis contributed to cell death under both anoxia and reoxygenation. Although under reoxygenation, ROS increased in both human and hamster RPTECs, lipid peroxidation-induced cell death was detected only in human cells. Autophagy was observed only in human cells under both conditions. Necroptosis was not detected in any of the evaluated cells. Clarifying the ways that are responsible for hamster RPTECs escaping from apoptosis and lipid peroxidation-induced cell death may reveal interventions for preventing ischemia–reperfusion-induced acute kidney injury in humans.

Can bats sense smoke during deep torpor?

While torpor is a beneficial energy-saving strategy, it may incur costs if an animal is unable to respond appropriately to external stimuli, which is particularly true when it is necessary to escape from threats such as fire. We aimed to determine whether torpid bats, which are potentially threatened because they must fly to escape, can sense smoke and whether respiration rate (RR), heart rate (HR) and reaction time of torpid bats prior to and following smoke introduction is temperature-dependent. To test this we quantified RR and HR of captive Australian tree-roosting bats, Nyctophilus gouldi (n=5, ~10g), in steady-state torpor in response to short-term exposure to smoke from Eucalyptus spp. leaves between ambient temperatures (Ta) of 11 and 23°C. Bats at lower Ta took significantly longer (28-fold) to respond to smoke, indicated by a cessation of episodic breathing and a rapid increase in RR. Bats at lower Ta returned to torpor more swiftly following smoke exposure than bats at higher Ta. Interestingly, bats at Ta<15°C never returned to thermoconforming steady-state torpor prior to the end of the experimental day, whereas all bats at Ta≥15°C did, as indicated by apnoeic HR. This shows that although bats at lower Ta took longer to respond, they appear to maintain vigilance and prevent deep torpor after the first smoke exposure, likely to enable fast escape. Our study reveals that bats can respond to smoke stimuli while in deep torpor. These results are particularly vital within the framework of fire management conducted at Ta<15°C, as most management burns are undertaken during winter when bats will likely respond more slowly to fire cues such as smoke, delaying the time to escape from the fire.

Effects of food store quality on hibernation performance in common hamsters.

Hibernating animals can adjust torpor expression according to available energy reserves. Besides the quantity, the quality of energy reserves could play an important role for overwintering strategies. Common hamsters are food-storing hibernators and show high individual variation in hibernation performance, which might be related to the quality of food hoards in the hibernacula. In this study, we tested the effects of food stores high in fat content, particularly polyunsaturated fatty acids (PUFAs), on hibernation patterns under laboratory conditions. Control animals received standard rodent pellets only, while in the other group pellets were supplemented with sunflower seeds. We recorded body temperature during winter using subcutaneously implanted data loggers, documented total food consumption during winter, and analysed PUFA proportions in white adipose tissue (WAT) before and after the winter period. About half of the individuals in both groups hibernated and torpor expression did not differ between these animals. Among the high-fat group, however, individuals with high sunflower seeds intake strongly reduced the time spent in deep torpor. PUFA proportions in WAT decreased during winter in both groups and this decline was positively related to the time an individual spent in deep torpor. Sunflower seeds intake dampened the PUFA decline resulting in higher PUFA levels in animals of the high-fat group after winter. In conclusion, our results showed that common hamsters adjusted torpor expression and food intake in relation to the total energy of food reserves, underlining the importance of food hoard quality on hibernation performance.

Body temperatures of hibernating little brown bats reveal pronounced behavioural activity during deep torpor and suggest a fever response during white-nose syndrome.

Hibernating animals use torpor [reduced body temperature (T b) and metabolic rate] to reduce energy expenditure during winter. Periodic arousals to normal T b are energetically expensive, so hibernators trade off arousal benefits against energetic costs. This is especially important for bats with white-nose syndrome (WNS), a fungal disease causing increased arousal frequency. Little brown bats (Myotis lucifugus) with WNS show upregulation of endogenous pyrogens and sickness behaviour. Therefore, we hypothesized that WNS should cause a fever response characterized by elevated T b. Hibernators could also accrue some benefits of arousals with minimal T b increase, thus avoiding full arousal costs. We compared skin temperature (T sk) of captive Myotis lucifugus inoculated with the WNS-causing fungus to T sk of sham-inoculated controls. Infected bats re-warmed to higher T sk during arousals which is consistent with a fever response. Torpid T sk did not differ. During what we term "cold arousals", bats exhibited movement following T sk increases of only 2.2 ± 0.3°C, compared to >20°C increases during normal arousals. Cold arousals occurred in both infected and control bats, suggesting they are not a pathophysiological consequence of WNS. Fever responses are energetically costly and could exacerbate energy limitation and premature fat depletion for bats with WNS. Cold arousals could represent an energy-saving mechanism for both healthy and WNS-affected bats when complete arousals are unnecessary or too costly. A few cold arousals were observed mid-hibernation, typically in response to disturbances. Cold arousals may, therefore, represent a voluntary restriction of arousal temperature instead of loss of thermoregulatory control.

Changes in the phosphoproteome of brown adipose tissue during hibernation in the ground squirrel, Ictidomys tridecemlineatus.

Mammalian hibernation is characterized by metabolic rate depression and a strong decrease in core body temperature that together create energy savings such that most species do not have to eat over the winter months. Brown adipose tissue (BAT), a thermogenic tissue that uses uncoupled mitochondrial respiration to generate heat instead of ATP, plays a major role in rewarming from deep torpor. In the present study we developed a label-free liquid chromatography mass spectrometry (LC-MS) strategy to investigate both differential protein expression and protein phosphorylation in BAT extracts from euthermic vs. hibernating ground squirrels (Ictidomys tridecemlineatus). In particular, we incorporated the filter-assisted sample preparation protocol, which provides a more in-depth analysis compared with gel-based and other LC-MS proteomics approaches. Surprisingly, mitochondrial membrane and matrix protein expression in BAT was largely constant between active euthermic squirrels and their hibernating counterparts. Validation by immunoblotting confirmed that the protein levels of mitochondrial respiratory chain complexes were largely unchanged in hibernating vs. euthermic animals. On the other hand, phosphoproteomics revealed that pyruvate dehydrogenase (PDH) phosphorylation increased during squirrel hibernation, confirmed by immunoblotting with phospho-specific antibodies. PDH phosphorylation leads to its inactivation, which suggests that BAT carbohydrate oxidation is inhibited during hibernation. Phosphorylation of hormone-sensitive lipase (HSL) was also found to increase during hibernation, suggesting that HSL would be active in BAT to produce the fatty acids that are likely the primary fuel for thermogenesis upon arousal. Increased perilipin phosphorylation along with that of a number of other proteins was also revealed, emphasizing the importance of protein phosphorylation as a regulatory mechanism during mammalian hibernation.

Torpor patterns in common hamsters with and without access to food stores

Hibernating species significantly reduce energy expenditure during winter by entering torpor. Nevertheless, the various benefits of hibernation might be counteracted by negative effects of torpor such as immune depression, oxidative stress, or neuronal impairment. Considering these trade-offs, adequate energy reserves could allow animals to reduce the time spent in torpor or the extent of metabolic depression. Common hamsters use food stores during hibernation and previously documented high individual variations in body temperature patterns during winter could, therefore, be related to differences in external energy reserves. In this study, we manipulated the availability of food stores under laboratory conditions to investigate potential effects on hibernation patterns. Female hamsters were kept in artificial burrows in climate chambers and subcutaneous temperature was recorded using implanted data loggers. One group had access to large food stores, whereas another group received daily food portions which were removed on the next day if not consumed. Almost all hamsters without access to food stores hibernated, while less than half of the individuals with food stores entered deep torpor. Individuals without food hoards additionally expressed more short torpor bouts and exhibited lower minimum subcutaneous temperatures during torpor than those with food stores. Thus, individuals confronted with lacking food reserves were more likely to hibernate and additionally saved energy by entering short torpor bouts more frequently and remaining at lower subcutaneous temperature both during torpor and euthermic periods. In conclusion, our results demonstrate that food store availability affects torpor expression and also highlight variation in torpor patterns and energy-saving strategies in common hamsters.

Seasonal loss and resumption of circadian rhythms in hibernating arctic ground squirrels.

Circadian clocks are near universal among organisms and play a key role in coordinating physiological and metabolic functions to anticipate or coincide with predictable daily changes in the physical and social environment. However, whether circadian rhythms persist and are functionally important during hibernation in all mammals is currently unclear. We examined whether circadian rhythms of body temperature (T b) persist during multi-day, steady-state torpor and investigated the association between timing of animal emergence, exposure to light, and resumption of activity and T b rhythms in free-living and captive male arctic ground squirrels. High-resolution (0.02 °C) temperature loggers revealed that circadian rhythms of T b were not present during deep torpor in free-living arctic ground squirrels. Significant circadian rhythms of T b resumed, however, following the resumption of euthermia, but prior to emergence, though rhythms became much more robust coincident with aboveground emergence. Additionally, squirrels maintained in captivity under conditions of constant darkness spontaneously developed significant circadian rhythms of T b and activity soon after ending torpor. Exposing animals to a 5-s pulse of light within a week when they ended torpor increased the strength of rhythms. Our results are consistent with the hypothesis that circadian clock function is inhibited during hibernation in arctic ground squirrels, and we postulate that exposure to external stimuli, such as light in free-living animals, and meals or acute disturbance for captive squirrels, may enhance T b rhythmicity by synchronizing loosely coupled circadian oscillators within the suprachiasmatic nucleus.

Hibernation as a major determinant of life-history traits in marmots

During the downward trend of Quaternary global temperatures, crown-group marmots evolved to occupy harsh, seasonal environments at high latitudes and elevations. Marmots evolved hibernation and large body size, which allowed them to cope with a long, cold season with low, stressful ambient temperatures when food was unavailable. Harsh conditions during a short active season frequently cause reproductive failure and reproductive skipping in marmots. Seasonal phenology is controlled by a circannual rhythm that directs metabolic changes and fat accumulation for hibernation and reproduction. Metabolism and water balance in marmots are adapted to the low environmental temperatures of hibernation. Thus, marmots are subject to heat stress from solar radiation and high ambient temperatures during the active season. Energy conservation characterizes metabolism in marmots throughout the annual cycle: metabolic rates are lower than predicted from body size and decline during the active season, metabolism is actively suppressed to enter torpor, and most of the time during hibernation is spent at low metabolism in deep torpor. Young typically require 2 or more of the short active seasons to reach reproductive maturity and are retained at home, which leads to sociality based on kinship. Competition within the social group is expressed primarily as reproductive suppression, which delays the age of 1st reproduction that in turn strongly affects demography. Early snowmelt due to global warming may enable marmots to colonize new habitats or increase population growth. However, climate change most likely will diminish the ability of marmots to mobilize sufficient energy resources in a drier, warmer active season, which will reduce both survivorship and reproduction.

Heart rate dynamics in a marsupial hibernator.

The eastern pygmy possum (Cercartetus nanus) is a small marsupial that can express spontaneous short bouts of torpor, as well as multi-day bouts of deep hibernation. To examine heart rate (fH) control at various stages of torpor in a marsupial hibernator, and to see whether fH variability differs from that of deep placental hibernators, we used radiotelemetry to measure ECG and body temperature (Tb) while measuring the rate of O2 consumption and ventilation. fH and O2 consumption rate during euthermia were at a minimum (321±34 beats min-1, 0.705±0.048 ml O2 g-1 h-1) at an ambient temperature (Ta) of 31°C. fH had an inverse linear relationship with Ta to a maximum of 630±19 beats min-1 at a Ta of 20°C. During entry into torpor at a Ta of 20°C, fH slowed primarily as a result of episodic periods of cardiac activity where electrical activity of the heart occurred in groups of 3 or 4 heart beats. When Tb was stable at 24°C in these torpor bouts, the episodic nature of fH had disappeared (i.e. no asystoles) with a rate of 34±3 beats min-1 For multi-day bouts of deep torpor, Ta was lowered to 6.6±0.8°C. During these deep bouts of torpor, Tb reached a minimum of 8.0±1.0°C, with a minimum fH of 8 beats min-1 and a minimum O2 consumption rate of 0.029±0.07 ml O2g-1h-1 Shivering bouts occurred in deep torpor about every 8 min, during which ventilation occurred, and fH was elevated to 40 beats min-1 The duration of the QRS complex increased from 12 ms during euthermia to 69 ms at a Tb of 8°C. These findings demonstrate the dynamic functioning range of fH to be about 600 beats min-1 (∼80-fold), one of the largest known ranges in mammals. Our study shows that despite a separation of ∼160 million years, the control and function of the cardiac system seems indistinguishable in marsupial and placental hibernating mammals.

Deeply torpid bats can change position without elevation of body temperature

Because body temperature is tightly coupled to physiological function, hibernating animals entering deep torpor are typically immobile. We analysed thermal behaviour and locomotory activity of hibernating greater mouse-eared bats Myotis myotis and found two types of movement behaviour related to body temperature, i.e. movement at high fur temperature and at low fur temperatures (Tflow; <5°C). First Tflow movements appeared at the beginning of March and often occurred during long torpor bouts. In most cases, Tflow events represented slow displacements between clusters of bats. In several cases, however, departure or arrivals from and into clusters was also recorded without any elevation in body temperature. Distance travelled, flight duration and speed of locomotion during Tflow events was lower than in high fur temperature events. Such behaviour could allow bats to save energy long-term and prolong torpor bouts. Tflow movement in torpid bats significantly changes our understanding of basic hibernation principles and we strongly recommend further studies on the subject.

Hypometabolism during Daily Torpor in Mice is Dominated by Reduction in the Sensitivity of the Thermoregulatory System.

Some mammals enter a hypometabolic state either daily torpor (minutes to hours in length) or hibernation (days to weeks), when reducing metabolism would benefit survival. Hibernators demonstrate deep torpor by reducing both the sensitivity (H) and the theoretical set-point temperature (TR) of the thermogenesis system, resulting in extreme hypothermia close to ambient temperature. However, these properties during daily torpor remain poorly understood due to the very short steady state of the hypometabolism and the large variation among species and individuals. To overcome these difficulties in observing and evaluating daily torpor, we developed a novel torpor-detection algorithm based on Bayesian estimation of the basal metabolism of individual mice. Applying this robust method, we evaluated fasting induced torpor in various ambient temperatures (TAs) and found that H decreased 91.5% during daily torpor while TR only decreased 3.79 °C in mice. These results indicate that thermogenesis during daily torpor shares a common property of sensitivity reduction with hibernation while it is distinct from hibernation by not lowering TR. Moreover, our findings support that mice are suitable model animals to investigate the regulation of the heat production during active hypometabolism, thus suggesting further study of mice may provide clues to regulating hypometabolism in mammals.

Can hibernators sense and evade fires? Olfactory acuity and locomotor performance during deep torpor.

Increased habitat fragmentation, global warming and other human activities have caused a rise in the frequency of wildfires worldwide. To reduce the risks of uncontrollable fires, prescribed burns are generally conducted during the colder months of the year, a time when in many mammals torpor is expressed regularly. Torpor is crucial for energy conservation, but the low body temperatures (T b) are associated with a decreased responsiveness and torpid animals might therefore face an increased mortality risk during fires. We tested whether hibernators in deep torpor (a) can respond to the smell of smoke and (b) can climb to avoid fires at T bs below normothermic levels. Our data show that torpid eastern pygmy-possums (Cercartetus nanus) are able to detect smoke and also can climb. All males aroused from torpor when the smoke stimulus was presented at an ambient temperature (T a) of 15°C (T b ∼18°C), whereas females only raised their heads. The responses were less pronounced at T a 10°C. The first coordinated movement of possums along a branch was observed at a mean T b of 15.6°C, and animals were even able to climb their prehensile tail when they reached a mean T b of 24.4°C. Our study shows that hibernators can sense smoke and move at low T b. However, our data also illustrate that at T b ≤13°C, C. nanus show decreased responsiveness and locomotor performance and highlight that prescribed burns during winter should be avoided on very cold days to allow torpid animals enough time to respond.

Decreases in body temperature and body mass constitute pre-hibernation remodelling in the Syrian golden hamster, a facultative mammalian hibernator

Hibernation is an adaptive strategy for surviving during periods with little or no food availability, by profoundly reducing the metabolic rate and the core body temperature (Tb). Obligate hibernators (e.g. bears, ground squirrels, etc.) hibernate every winter under the strict regulation of endogenous circannual rhythms, and they are assumed to undergo adaptive remodelling in autumn, the pre-hibernation period, prior to hibernation. However, little is known about the nature of pre-hibernation remodelling. Syrian hamsters (Mesocricetus auratus) are facultative hibernators that can hibernate irrespective of seasons when exposed to prolonged short photoperiod and cold ambient temperature (SD-Cold) conditions. Their Tb set point reduced by the first deep torpor (DT) and then increased gradually after repeated cycles of DT and periodic arousal (PA), and finally recovered to the level observed before the prolonged SD-Cold in the post-hibernation period. We also found that, before the initiation of hibernation, the body mass of animals decreased below a threshold, indicating that hibernation in this species depends on body condition. These observations suggest that Syrian hamsters undergo pre-hibernation remodelling and that Tb and body mass can be useful physiological markers to monitor the remodelling process during the pre-hibernation period.