Hibernation evolved in some small mammals that live in cold environments, presumably to conserve energy when food supplies are low. Throughout the winter, hibernators cycle spontaneously between torpor, with low metabolism and near-freezing body temperatures, and euthermia, with high metabolism and body temperatures near 37°C. Understanding the mechanisms underlying this natural model of extreme metabolic plasticity is important for fundamental and applied science. During entrance into torpor, reductions in metabolic rate begin before body temperatures fall, even when thermogenesis is not active, suggesting active mechanisms of metabolic suppression, rather than passive thermal effects. Mitochondrial respiration is suppressed during torpor, especially when measured in liver mitochondria fuelled with succinate at 37°C in vitro. This suppression of mitochondrial metabolism appears to be invoked quickly during entrance into torpor when body temperature is high, but is reversed slowly during arousal when body temperature is low. This pattern may reflect body temperaturesensitive, enzyme-mediated post-translational modifications of oxidative phosphorylation complexes, for instance by phosphorylation or acetylation.KEY WORDS: Heat, Body temperature, Thermoregulation, Thermogenesis, Oxidative phosphorylation, Post-translational modification, Acetylation, Phosphorylation
IntroductionMost mammals are strict endotherms, i.e. they maintain fairly constant body temperatures (T b ) near 37°C using heat derived primarily from endogenous metabolism. In cold environments, mammals retain some of this heat by regulating insulation (using underfur and/or subcutaneous fat), peripheral blood circulation (using vasoconstriction and/or countercurrent heat exchangers) and ventilatory evaporation. At very cold temperatures, the high gradient between T b and ambient temperature (T a ) causes large heat loss by radiation and conduction, which is also affected by convection of water or air. For small mammals, the high body surface area, relative to volume, results in greater mass-specific rates of heat loss. While large land mammals can increase insulation by changing the quality (microstructure) and quantity (length, density) of underfur, this option is limited for small mammals. Imagine a 70 mm long lemming growing fur 100 mm long; it might stay warm, but when it tripped over that fur it would be easy prey.Despite the challenges, many small mammals thrive in cold environments. To compensate for high heat loss, these mammals upregulate their capacities to produce metabolic heat. Though COMMENTARY Department of Biology, University of Western Ontario, London, ON, Canada, N6A 5B8.*Author for correspondence (jfstaple@uwo.ca) effective, this strategy requires large quantities of fuel during winter when food availability is typically low. Many small mammals use stored food to power thermogenic metabolism. For example, American red squirrels (Tamiascurius hudsonicus) cache spruce cones and feed on the seeds throughout winter, allowing th...