Metabolic adjustments to increasing foraging costs of starlings in a closed economy.

Knowledge of the physiological consequences of variation in food availability may be essential for understanding behavioural and life history responses to such variation. To study the physiological consequences of food availability animals are generally subjected to caloric restriction or starvation, thereby reducing the upper limit to the energy budget. The relevance of this approach to free-living animals is questionable, however, because under natural conditions low food availability often results in higher foraging costs, and everything else remaining equal this results in a higher energy budget. We manipulated food availability by varying the foraging costs and studied effects on daily energy expenditure (DEE) and energy allocation of captive starlings Sturnus vulgaris. Birds in a closed economy earned their food by flying between two perches 5 m apart. The probability of a reward was set at three different levels, thereby creating a 'poor', 'intermediate' and 'rich' environment. Compared with the rich environment, birds flew 4 times more (2.3 h per day) in the poor environment, and increased DEE by 43% to 220 kJ day-1 (3.7xBMR), within the range of free-living parents rearing young. To our knowledge this is the first study to show an increase in DEE with decreasing food availability. Body mass, basal metabolic rate (BMR) and pectoral muscle size were reduced in the poor environment. Nocturnal energy expenditure was further reduced by reaching BMR earlier in the night. Calculations show that the energy demands in the poor environment could not be met with the flight costs of 20.5 W that we measured previously in a rich environment. Flight costs derived indirectly from the energy budget were lower, at 17.5 W, probably due to lower body mass. By reducing body mass by 20%, and economising during sleep, the birds achieved savings of 37% in their DEE. Without these savings, a DEE substantially higher than measured in free-living parents rearing young would be required to remain in energy balance. Surprisingly little data exist to verify whether free-living animals use the same tactics to survive periods with low food availability.