Acclimation to different thermal conditions in a northerly wintering shorebird is driven by body mass-related changes in organ size

Seasonal acclimatization and experimental acclimation to cold in birds typically results from increased shivering endurance and elevated thermogenic capacity leading to improved resistance to cold. A wide array of physiological adjustments, ranging from biochemical transformations to organ mass variations, are involved in this process. Several studies have shown that improved cold endurance is accompanied by increases in summit metabolic rate (Msum), a measure of maximal heat production and an indicator of the level of sustainable thermogenic capacity. However, improved endurance to cold can also be achieved without significant changes in Msum. The same is true for basal metabolic rate (BMR), which is known to increase in association with cold acclimatization or acclimation in some species but not in others. We investigated cold acclimation in a migrant shorebird known for extreme physiological flexibility, the red knot (Calidris canutus, the northerly wintering subspecies islandica). We measured BMR and Msum over two months in birds caught in the wild and transferred to experimentally controlled conditions representative of aspects of their seasonal thermal environment ( two groups at constant 25 degrees C, one group at constant 4 degrees C and two groups experiencing variable outdoor temperatures). Birds maintained in both cold and variable ambient temperatures showed a 14-15% higher body mass, 33-45% higher food intake, and 26% and 13% elevations in BMR and Msum, respectively, compared with birds kept at thermoneutrality. These results, together with data on alimentary tract size and pectoral muscle thickness measured by ultrasonography, suggest that red knots acclimate to cold primarily through modulation of ( lean) body mass components. Heavier individuals have larger muscles, which allow higher maximal heat production and better thermal compensation. Cold acclimation effects on BMR are most probably due to changes in the size of visceral organs, although not the alimentary tract in this specific case. The liver, known for its thermogenic capacity, is a probable candidate. Overall, our results indicate that relatively small changes in body mass and muscle size allow enough reserve capacity in terms of heat production to cope with typical wintering ambient temperature variations as measured on the red knot's wintering grounds.