Cold-hearted bats: uncoupling of heart rate and metabolism during torpor at sub-zero temperatures.

Many hibernating animals thermoregulate during torpor and defend their body temperature ( T b ) near 0°C by an increase in metabolic rate. Above a critical temperature ( T crit ), animals usually thermoconform. We investigated the physiological responses above and below T crit for a small tree-dwelling bat ( Chalinolobus gouldii , ∼14 g) that is often exposed to sub-zero temperatures during winter. Through simultaneous measurement of heart rate ( f H ) and oxygen consumption ( V̇ O2 ), we show that the relationship between oxygen transport and cardiac function is substantially altered in thermoregulating torpid bats between 1 and -2°C, compared with thermoconforming torpid bats at mild ambient temperatures ( T a 5-20°C). T crit for this species was at a T a of 0.7±0.4°C, with a corresponding T b of 1.8±1.2°C. Below T crit , animals began to thermoregulate, as indicated by a considerable but disproportionate increase in both f H and V̇ O2 The maximum increase in f H was only 4-fold greater than the average thermoconforming minimum, compared with a 46-fold increase in V̇ O2 The differential response of f H and V̇ O2  to low T a was reflected in a 15-fold increase in oxygen delivery per heart beat (cardiac oxygen pulse). During torpor at low T a , thermoregulating bats maintained a relatively slow f H and compensated for increased metabolic demands by significantly increasing stroke volume and tissue oxygen extraction. Our study provides new information on the relationship between metabolism and f H in an unstudied physiological state that may occur frequently in the wild and can be extremely costly for heterothermic animals.