Regulated torpor and unregulated hypothermia are both characterized by substantially reduced body temperature (Tb) and metabolic rate (MR), but they differ physiologically. Although the remarkable, medically interesting adaptations accompanying torpor (e.g., tolerance for cold and ischemia, absence of reperfusion injury, and disuse atrophy) often do not apply to hypothermia in homeothermic species such as humans, the terms "torpor" and "hypothermia" are often used interchangeably in the literature. To determine how these states differ functionally and to provide a reliable diagnostic tool for differentiating between these two physiologically distinct states, we examined the interrelations between Tb and MR in a mammal (Sminthopsis macroura) undergoing a bout of torpor with those of the hypothermic response of a similar-sized juvenile rat (Rattus norvegicus). Our data show that under similar thermal conditions, 1) cooling rates differ substantially (approximately fivefold) between the two states; 2) minimum MR is approximately sevenfold higher during hypothermia than during torpor despite a similar Tb; 3) rapid, endogenously fuelled rewarming occurs in torpor but not hypothermia; and 4) the hysteresis between Tb and MR during warming and cooling proceeds in opposite directions in torpor and hypothermia. We thus demonstrate clear diagnostic physiological differences between these two states that can be used experimentally to confirm whether torpor or hypothermia has occurred. Furthermore, the data can clarify the results of studies investigating the ability of physiological or pharmacological agents to induce torpor. Consequently, we recommend using the terms "torpor" and "hypothermia" in ways that are consistent with the underlying regulatory differences between these two physiological states.

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Endothermic arousal from torpor is an energetically costly process and imposes enormous demands on the cardiovascular system, particularly during early stage arousal from low body temperature (Tb). To minimize these costs many bats and other heterothermic endotherms rewarm passively from torpor using solar radiation or fluctuating ambient temperature (Ta). Because the heart plays a critical role in the arousal process in terms of blood distribution and as a source of heat production, it is desirable to understand how the function of this organ responds to passive rewarming and how this relates to changes in metabolism and Tb. We investigated heart rate (HR) in hibernating long-eared bats (Nyctophilus gouldi) and its relationship to oxygen consumption (V̇o₂) and subcutaneous temperature (Tsub) during exposure to increasing Ta compared with endogenous arousals at constant low Ta. During passive rewarming, HR and V̇o₂ remained low over a large Tsub range and increased concurrently with increasing Ta (Q₁₀ 2.4 and 2.5, respectively). Absolute values were higher than during steady-state torpor but below those measured during torpor entry. During active arousals, mean HR and V̇o₂ were substantially higher than during passive rewarming at corresponding Tsub. In addition, partial passive rewarming reduced the cost of arousal from torpor by 53% compared with entirely active arousal. Our data show that passive rewarming considerably reduces arousal costs and arousal time; we suggest this may also contribute to minimizing exposure to oxidative stresses as well as demands on the cardiovascular system.

While heart rate (f H ) has been used as an indicator of energy expenditure, quantitative data showing the relationship between these variables are only available for normothermic animals. To determine whether f H also predicts oxygen consumption (V · O2 ) during torpor, we simultaneously measured V · O2 , f H and subcutaneous body temperature (T sub ) of a hibernator, Gould's long-eared bats (Nyctophilus gouldi, 9 g, N=18), at ambient temperatures (T a ) between 0 and 25°C. At rest, f H of normothermic resting bats was negatively correlated with T a , with maximum f H of 803 beats min −1 (T a =5°C). During torpor, the relationship between f H and T a was curvilinear, and at low T sub (~6°C), f H fell to a minimum average of 8 beats min