We have analyzed seasonal shifts of energy and time allocation in a population of golden-mantled ground squirrels (Spermophilus saturatus) by directly measuring total daily energy expenditure (DEE) with an isotopic technique ("doubly labeled water"=dlw), and by estimating components of total DEE through an integration of field behavioral observations with laboratory-measured rates of energy expenditure (oxygen consumption) associated with major behavioral and physiological states. Hibernation laster about 7 1/2 months, and the 4 1/2-month activity season consisted of mating, a 28-d gestation of 3-5 young, 5 1/2 weeks of postnatal growth building to a peak in lactation just before the young emerged above ground, an additional 2-3-week period of maternal care before dispersal, and finally restoration of body mass preceding hibernation. Although the hibernation season comprised nearly two-thirds of the year, it involved only 13-17% of annual energy expenditure, leaving about 85% of energy expenditure for the active season. Ground squirrels were actually present on the surface for only about 11% of the year's time, and the foraging time required to obtain the total annual energy supply amounted to only about 2% of the year's time. The squirrels fed mainly on herbs in the early season and hypogeous fungi later; both were used extensively during peak lactation when female energy expenditure and demand were maximal. Average daily foraging time increased steadily throughout the season to a maximum of 28% of aboveground time as availability of greens diminished and fungus predominated in the diet; time availability did not limit foraging since the animals sat on average for 65% of the daily surface time of about 7 h. Timing of reproduction is apparently optimized such that peak reproductive energy demands are matched with maximal food availability and moderate thermal conditions that minimize energy demand. Despite the greater body mass of males, the greatest total DEE (measured by dlw) of any squirrels at any time of year was that of females during peak lactation. For production of young and lactation through above-ground emergence of an average litter of 2.7, females required a total energy increase of 24% above annual nonreproductive metabolism. Yearling females all bred and performed similarly to older females, yet some costs were greater because the yearlings began and ended hibernation at smaller mass, compensated by giving birth later, and finally showed a greater absolute increase in body mass over the active season than older females. Annual metabolic energy expenditure of breeding males was about 18% greater than that of females, due to greater male body mass. Yet the annual energy intake requirement for both sexes was essentially identical (about 42MJ) due to the greater reproductive export by females in the form of newborn and milk. During the mating season males showed wide-ranging exploratory behavior and social interactions, including aggression, that involved considerable locomotory energy expenditures. Al...

Male and female redpolls (Acanthis flammea) showed marked increases in circulating corticosterone up to 1 hour after exposure to a common stress-apture, handling and restraint-indicating that their hypothalamo-pituitary-adrenal axis responded to acute stress in a manner similar to that of other vertebrates. We used this protocol as a measure of responsiveness of the adrenocortical cells to acute stress in general and for comparison with gender and across seasons. In both sexes the adrenocortical response to stress was reduced in January (at Fairbanks, 64"N) and maximal when birds were breeding in June at Tbolik Lake (69"N). The elevation of circulating corticosterone following capture and handling in breeding males at Barrow (71'") was significantly less than in breeding males at Toolik Lake. There were also considerable variations among individuals in the intensity of the adrenocortical responses, particularly in the maximum levels of corticosterone attained. This individual variation correlated significantly with fat score andor body mass in both sexes only in breeding birds at Barrow. This difference may be explained by generally lower, and thus reduced variability in body fat and mass in birds sampled in the warmer climate of Toolik Lake. A similar trend was seen in non-breeding birds, but this was not significant. Additionally, in January, baseline cortisterone levels in males were correlated with body mass, although this relationship did not hold when both sexes were considered. Body mass and fat score in winter were similar to those of redpolls sampled at Barrow in June. These data suggest that redpolls may be able to adjust their responsiveness to acute stresses in relation t o fat stores. Those with greater fat depots had reduced responsiveness to stress.

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