The recently introduced technique of measuring corticosterone in feathers currently provides the longest-term measure of corticosterone in birds. This review examines the strengths, weaknesses, and unresolved technical issues of the feather corticosterone technique. Feather corticosterone's major strengths are that it provides: a retrospective assessment of corticosterone physiology, including information from absent (unseen) or dead (e.g. museum specimens) individuals; a long-term measure of corticosterone exposure over the period of feather growth (days-weeks), integrating both baseline and responses to stressors; and flexible, minimally-invasive, sampling. However, researchers considering this technique should be aware of its limitations. Feather corticosterone only reflects hormone exposure during feather growth and, when sampling during molt, corticosterone titers and ecological conditions may not be representative of the majority of the annual cycle. Synchronization of molt is often unknown for a population, requiring assumptions when making inter-individual comparisons. Additionally, unresolved technical issues include: assessing whether corticosterone is the only hormone measured by assays; determining deposition dynamics to fully understand connections between feather and plasma corticosterone titers; studying the longevity and stability of corticosterone in the feather; establishing the impact of feather size and color on corticosterone deposition; and understanding the causes and implications of corticosterone variation along the length of the feather. Notwithstanding the above limitations and technical challenges, determining corticosterone titers in feathers is proving to be a useful technique for exploring some ecological and physiological correlates in individual birds. Given the unique perspective that feather corticosterone offers, we suggest that this measure complement, not replace, plasma measurements.

SUMMARYIntegrated measures of corticosterone (CORT), such as from feathers (CORT f ), have intuitive appeal because they incorporate both the duration and amplitude of glucocorticoid secretion. An association between CORT f and plasma CORT has never been shown in wild birds, and it is unclear as to when and whether these measures should be correlated, given that they are fundamentally different yet related measures of physiology. We hypothesized that CORT f should correlate with instantaneous measurements of plasma CORT when the latter reflect sustained changes in the activity of the hypothalamic-pituitary-adrenal (HPA) axis. To test this, we experimentally manipulated levels of plasma CORT in wild nestling tree swallows (Tachycineta bicolor) using 5day time-release CORT pellets, and measured plasma CORT and growth parameters before, during and at the end of hormone manipulation (days 7, 9 and 11 post-hatch, respectively). CORT f and plasma CORT were significantly positively related only when the latter was at its highest and most variable among individuals (day 9). A similar relationship was expected at day 11, but plasma CORT had returned to near-original levels. Nestlings with higher CORT f were smaller, lighter and less likely to fledge, but we did not detect seasonal effects on CORT f . Our results clearly demonstrate that CORT f from free-living birds can reflect plasma CORT, but correlations may not always be expected, especially if elevations in plasma CORT are relatively modest and of short duration. Our work suggests that CORT f is best used to study the activity of the HPA axis over relatively long time frames and can be used effectively to advance avian ecology. Supplementary material available online at

Enrichment is widely used as tool for managing fearfulness, undesirable behaviors, and stress in captive animals, and for studying exploration and personality. Inconsistencies in previous studies of physiological and behavioral responses to enrichment led us to hypothesize that enrichment and its removal are stressful environmental changes to which the hormone corticosterone and fearfulness, activity, and exploration behaviors ought to be sensitive. We conducted two experiments with a captive population of wild-caught Clark's nutcrackers (Nucifraga columbiana) to assess responses to short- (10-d) and long-term (3-mo) enrichment, their removal, and the influence of novelty, within the same animal. Variation in an integrated measure of corticosterone from feathers, combined with video recordings of behaviors, suggests that how individuals perceive enrichment and its removal depends on the duration of exposure. Short- and long-term enrichment elicited different physiological responses, with the former acting as a stressor and birds exhibiting acclimation to the latter. Non-novel enrichment evoked the strongest corticosterone responses of all the treatments, suggesting that the second exposure to the same objects acted as a physiological cue, and that acclimation was overridden by negative past experience. Birds showed weak behavioral responses that were not related to corticosterone. By demonstrating that an integrated measure of glucocorticoid physiology varies significantly with changes to enrichment in the absence of agonistic interactions, our study sheds light on potential mechanisms driving physiological and behavioral responses to environmental change.

Although altricial young are dependent on their parents during early life, they must respond to environmental variation to maintain homeostasis. The hormone corticosterone (CORT) may be an important link between environment and phenotype during early life; however, no previous study has experimentally assessed the sensitivity of CORT to nest microclimate in altricial birds beginning to thermoregulate. We tested the hypothesis that microclimate influences CORT by cross-fostering nestling Tree Swallows (Tachycineta bicolor (Vieillot, 1808)) between thicker-walled “aspen” nest boxes and thinner-walled “plywood” nest boxes. Quantification of CORT in nestling feathers allowed us to consider hormone secreted over days, rather than instantaneously from blood samples. In agreement with our hypothesis, we detected significant positive relationships between feather CORT and increased temperature variability and maximum, but not minimum, nest box temperatures. This could reflect the energetic challenge of warmer temperatures or positive developmental effects on the nestling hypothalamic–pituitary–adrenal axis. Feather CORT was significantly lower in chicks hatched in aspen nest boxes compared with plywood ones, but cross-fostering did not influence nestling CORT. This suggests that the influence of natal nest box environment on feather CORT was likely greater than the influence of the foster nest box environment. The relationships we detected highlight the sensitivity of feather CORT to environmental variation and contribute insight into nestling responses to environmental change.

Carotenoids produce many of the red, orange and yellow signal traits of birds, and individuals must trade off utilizing carotenoids for physiological processes versus ornamentation. Proximate mechanisms regulating this trade-off are poorly understood, despite their importance for expression of color signals. Corticosterone (CORT) may play a significant mechanistic role in signal expression because it mobilizes energy substrates and influences foraging behavior. We used a unique feather-based approach to test whether CORT mediates expression of carotenoid-based coloration. First, we investigated relationships between levels of CORT from feathers (CORT(f)) and carotenoid-based plumage signals in common redpolls (Acanthis flammea). Then, we determined how the width of growth bars and probability of having fault bars on feathers varied with CORT(f), specifically whether these metrics reflected developmental costs of elevated CORT ("stress" hypothesis) or represented an individual's quality ("quality" hypothesis). CORT(f) correlated positively with the strength of carotenoid signals, but only in adult males. However, also in adult males, CORT(f) was positively related to width of feather growth bars and negatively with probability of having fault bars, providing support for the quality hypothesis. Overall, CORT(f) was lower in adult males than in females or young males, possibly due to dominance patterns. Our results indicate that CORT may indirectly benefit feather quality, potentially by mediating the expression of carotenoid signals. We place our sex-specific findings into a novel framework that proposes that the influences of CORT in mediating carotenoid-based plumage traits will depend on the extent to which carotenoids are traded off between competing functions.

Offspring of long-lived species should face costs of parental trade-offs that vary with overall energetic demands encountered by parents during breeding. If sex differences exist in how parents make the trade-off, sex-specific differences may exist in the contribution of each parent to those costs. Adaptations of offspring facing such costs are not well understood, but the hormone corticosterone probably plays a role. We manipulated breeding effort in Cory's shearwaters (Calonectris diomedea) to increase costs to offspring and used an integrated measure of corticosterone from chick feathers to investigate how experimental variation in parental investment influences offspring physiology. Average foraging trip duration and foraging efficiency (FE) of breeding pairs were not related to chick corticosterone, but sex biases in FE were. Adult male investment was more strongly related to chick corticosterone than was female investment. Importantly, we show for the first time suppression of adrenocortical activity in nestling Procellariiform seabirds, and explain how our results indicate an adaptive mechanism invoked by chicks facing increased costs of parental trade-offs.

Diet during the non‐breeding period influences condition and subsequent reproduction. Physiological mechanisms underlying such carry‐over effects are poorly understood but could be clarified by studying physiological responses to variation in diet during non‐breeding. The hormone corticosterone provides a functional link between diet and survival and reproduction, but methodological limitations have prevented previous studies from testing the hypothesis that, on an individual level, avian corticosterone levels during the non‐breeding period reflect broader patterns in feeding ecology during that time. Using museum specimens (1859–2002) and live birds (2012), we found that corticosterone from feathers (CORTf) is negatively related to trophic position (TP) inferred from feather stable‐nitrogen isotope values (δ15N) in Leach's Storm‐petrels Oceanodroma leucorhoa. CORTf was not related to stable‐carbon isotope values (δ13C). We detected no temporal trends in CORTf or δ15N, and neither was related to a large‐scale index of winter climate, suggesting a general ecological phenomenon rather than a reflection of historical environmental changes. However, we detected a temporal trend in feather δ13C, and δ13C was related to δ15N. Our findings suggest a physiological benefit of feeding at higher TPs, either through increased nutritional value or reduced foraging costs associated with higher TP prey, and future research should aim to distinguish between these two explanations. Nevertheless, ours is the first evidence of a correlation between individual endocrine levels and foraging ecology, and demonstrates non‐lethal variation in a physiological mediator in turn related to variation in resource use.

Reproductive success often declines with breeding date in diverse taxa, including temperate-nesting bird species. The date hypothesis predicts that seasonally deteriorating environmental quality drives this pattern. While mechanisms are not fully understood, a seasonal increase in parasitism may contribute to the declining quality of nestlings hatched later in the season. We examined the effect of ectoparasites on seasonal variation in indices of nestling quality and survival in Tree Swallows (Tachycineta bicolor (Vieillot, 1808)) at breeding sites in British Columbia (PG) and Saskatchewan (SDNRA). In a parasite removal experiment, we detected no seasonal trend in flea abundance and, contrary to expectation, there were fewer blow flies in nests of late breeders. Negative effects of parasites on nestlings were documented at PG, where lengths of primary feather and head–bill were affected. Parental, rather than environmental, quality had the greatest effect on reproductive success at PG, as nestling survival declined seasonally regardless of treatment and despite seasonally increasing food biomass. Nestlings in control (i.e., parasitized) nests at SDNRA had elevated feather corticosterone levels, but no other effects of parasites were detected possibly because parent birds were of higher quality or parents and offspring had greater access to food such that nestlings were capable of coping with parasite-related challenges.