Voluntary running in deer mice: speed, distance, energy costs and temperature effects

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Abstract
SUMMARY The energetics of terrestrial locomotion are of considerable interest to ecologists and physiologists, but nearly all of our current knowledge comes from animals undergoing forced exercise. To explore patterns of energy use and behavior during voluntary exercise, we developed methods allowing nearly continuous measurements of metabolic rates in freely behaving small mammals,with high temporal resolution over periods of several days. We used this approach to examine relationships between ambient temperature(Ta), locomotor behavior and energy costs in the deer mouse, a small mammal that routinely encounters a large range of temperatures in its natural habitat. We tested for individual consistency in running behavior and metabolic traits, and determined how locomotor costs vary with speed and Ta. Because of the importance of thermoregulatory costs in small mammals, we checked for substitution of exercise heat for thermostatic heat production at Ta below the thermal neutral zone and determined the fraction of the daily energy budget comprising exercise costs. Locomotor behavior was highly variable among individuals but had high repeatability, at least over short intervals. We found few temperature-related changes in speed or distance run, but Ta strongly affected energy costs. Partial substitution of exercise heat for thermogenic heat occurred at low Ta. This reduced energy expenditure during low-temperature running by 23–37%, but running costs comprised a fairly minor fraction of the energy budget, so the daily energy savings viasubstitution were much smaller. Deer mice did not adjust running speed to maximize metabolic economy, as they seldom used the high speeds that provide the lowest cost of transport. The highest voluntary speeds (4–5 km h-1) were almost always below the predicted maximal aerobic speed,and were much less than the species' maximal sprint speed. Maximum voluntarily attained rates of oxygen consumption(V̇O2) were highest at low Ta, but rarely approached maximal V̇O2 during forced treadmill exercise. Mean respiratory exchange ratios coincident with maximal voluntary V̇O2increased slightly as Ta declined, but were always below 1.0 (another indication that metabolic rate was less than the aerobic maximum). Individuals with high running performance (cumulative distance and running time) had high resting metabolism, which suggests a cost of having high capacity or propensity for activity.