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We tested the adaptive stress hypothesis that male arctic ground squirrels (Urocitellus parryii) exhibit a stress response over the course of the breeding season that is characterized by increasing free cortisol concentrations, increasing mobilization of stored energy, and decreasing physical condition. We assessed the functioning of the hypothalamic-pituitary-adrenal axis by measuring cortisol levels in response to the stress of capture and in response to a hormone challenge protocol (dexamethasone suppression and adrenocorticotropic hormone stimulation). We measured blood glucose levels, free fatty acids, white blood cells, and hematocrit to assess the downstream physiological responses to cortisol. Immediately after spring emergence, male arctic ground squirrels had ample free abdominal fat and few signs of wounding. By the end of the breeding season 3 wk later, visible fat reserves were almost entirely gone, and most males had extensive wounds. Total plasma cortisol concentrations increased over this period, but so did corticosteroid-binding capacity, resulting in no change in the free cortisol response to capture. We found no significant changes in how the animals responded to our hormone challenges, contrary to our prediction that the stress axis should increase free cortisol production. Even though we found no change in the functioning of the stress axis, all of the downstream measures suggested that male arctic ground squirrels are chronically exposed to high cortisol concentrations. Over the breeding season, blood glucose increased, fat stores and circulating free fatty acids were depleted, and both hematocrit levels and white blood cell counts decreased significantly. Our data suggest that a more complex relationship between the stress axis and downstream measures of stress exists than that proposed by the adaptive stress hypothesis. We propose several nonexclusive, testable mechanisms that could explain our observations. [ABSTRACT FROM AUTHOR]

Stress responses play a critical role in the ecology and demography of wild animals, and the analysis of fecal hormone metabolites is a powerful noninvasive method to assess the role of stress. We characterized the metabolites of injected radio-labeled cortisol in the urine and feces of Columbian ground squirrels and validated an enzyme immunoassay for measuring fecal cortisol metabolites (FCM) with a 5α-3β, 11β-diol structure by stimulation and suppression of adrenocortical activity and by evaluation of the circadian pattern of FCM excretion. In addition, we also evaluated the impact of capture, handling, and acclimation to the laboratory on FCM. Cortisol is highly metabolized, with virtually none being excreted, and of the radiolabeled cortisol injected, 31% was recovered in urine and 6.5% in feces. The lag time between cortisol injection and its appearance in urine and feces was 4.5 ± 0.82 (SE) h and 7.0 ± 0.53 (SE) h, respectively. FCM levels varied over the day, reflecting circadian variation in endogenous cortisol. Dexamethasone decreased FCM levels by 33%, and ACTH increased them by 255%. Trapping and housing initially increased FCM levels and decreased body mass, but these reversed within 3-7 d, indicating acclimation. Finally, FCM levels were modestly repeatable over time (r = 0.57) in wild, livetrapped, nonbreeding animals, indicating that FCMs provide a measure of the squirrel's stress-axis state. This assay provides a robust non-invasive assessment of the stress response of the Columbian ground squirrel and will facilitate an integration of its life history and physiology. [ABSTRACT FROM AUTHOR]

Hormones such as glucocorticoids (colloquially referred to as "stress hormones") have important effects on animal behavior and life-history traits, yet most of this understanding has come through correlative studies. While experimental studies offer the ability to assign causality, there are important methodological concerns that are often not considered when manipulating hormones, including glucocorticoids, in wild animals. In this study, we examined how experimental elevations of cortisol concentrations in wild North American red squirrels (Tamiasciurus hudsonicus) affected their hypothalamic-pituitary-adrenal (HPA) axis reactivity and life-history traits, including body mass, litter survival, and adult survival. The effects of exogenous cortisol on plasma cortisol concentrations depended on the time between treatment consumption and blood sampling. In the first 9 h after consumption of exogenous cortisol, individuals had significantly higher true baseline plasma cortisol concentrations, but adrenal gland function was impaired as indicated by their dampened response to capture and handling and to injections of adrenocorticotropic hormone compared to controls. Approximately 24 h after consumption of exogenous cortisol, individuals had much lower plasma cortisol concentrations than controls, but adrenal function was restored. Corticosteroid-binding globulin (CBG) concentrations were also significantly reduced in squirrels treated with cortisol. Despite these profound shifts in the functionality of the HPA axis, squirrel body mass, offspring survival, and adult survival were unaffected by experimental increases in cortisol concentrations. Our results highlight that even short-term experimental increases in glucocorticoids can affect adrenal gland functioning and CBG concentrations but without other side effects. [ABSTRACT FROM AUTHOR]

Interest in the ecology of stress in wild populations has triggered the development of noninvasive methods for quantifying stress hormones. Measurement of fecal corticosteroid metabolites (FCMs) is one such method, but it is still unclear whether FCMs can be a reliable proxy of free plasma glucocorticoids. To assess the validity of this assumption, we carried out a robust assessment on brown lemmings (Lemmus trimucronatus) from Bylot Island, Nunavut, Canada, that were hand captured and anesthetized and related plasma glucocorticoid levels to fecal metabolite glucocorticoid levels. We examined endogenous factors that could explain interindividual variability. Blood corticosterone was measured from samples obtained on capture and 30 min later, and FCM levels were measured from animals kept in captivity for 72 h. Plasma free corticosterone increased 135-fold over baseline values 30 min after capture, which confirmed that initial handling was perceived as a stressor. We found that FCM levels were highly related with free (marginal Rm2 = 0.53) but not with total (Rm2 = 0.02) corticosterone levels, regardless of age, sex, and reproductive condition. FCM levels started increasing 2 h after capture and reached maximum levels 4 h after capture. No circadian rhythm in FCMs was found. Plasma total corticosterone levels were much higher in adult females compared with adult males, but this difference was much smaller when measuring free corticosterone levels and FCM levels. Our results suggest that FCM levels are good measures of stress by being closely related to plasma free corticosterone levels in brown lemmings. [ABSTRACT FROM AUTHOR]

Monitoring fecal glucocorticoid metabolites in wild animals, using enzyme immunoassays, enables the study of endocrinological patterns relevant to ecology and evolution. While some researchers use antibodies against the parent hormone (which is typically absent from fecal samples), others advocate the use of antibodies designed to detect glucocorticoid metabolites. We validated two assays to monitor fecal cortisol metabolites in the eastern chipmunk (Tamias striatus). We compared an antibody produced against cortisol and one produced against 5a-pregnane-3b, 11b, 21-triol-20-one using a radiometabolism study and an injection with adrenocorticotropic hormone (ACTH). Most cortisol metabolites were excreted in the urine (~83%). Peak excretion in the feces occurred 8 h after injection. Both assays detected an increase in fecal cortisol metabolite levels after injection of ACTH. Males, but not females, exhibited a circadian variation in metabolite levels. The sexes did not exhibit any difference over the time course and route of excretion or the relative increase in fecal cortisol metabolite levels after ACTH injection. The cortisol assay displayed higher reactivity to ACTH injection relative to baseline than did the metabolite assay. While both antibodies gave comparable results, the cortisol antibody was more sensitive to changes in plasma cortisol levels in eastern chipmunks. [ABSTRACT FROM AUTHOR]