Lower Baseline Corticosterone Research Articles

Light received by embryos promotes postnatal junior phenotypes in a seabird

Abstract Light is a salient and variable ecological factor that can impact developmental trajectories of vertebrate embryos, yet whether prenatal light environment can act as an anticipatory cue preparing organisms to cope with postnatal conditions is still unclear. In asynchronous birds, last-laid eggs are particularly exposed to sunlight as parental incubation behavior becomes intermittent after the hatching of senior chicks. Here, we explore whether natural variations in prenatal light exposure shape the distinctive phenotype showed by last-hatched chicks of a semi-precocial seabird, the yellow-legged gull (Larus michahellis), potentially preparing them to cope with the postnatal competitive context. To do this, we manipulated the amount of light received by last-laid eggs (within a natural range) during last stages of embryonic development. Prenatal exposure to light cues promoted the development of the resilient “junior phenotype” exhibited by last-hatched gull chicks, characterized by accelerated hatching, increased begging behavior and a slower growth rate. These developmental and behavioral adjustments were accompanied by down-regulation of genes involved in metabolism and development regulation (SOD2 and TRalpha), as well as changes in the HPA-axis functioning (lower baseline corticosterone and robust adrenocortical response). Junior chicks exposed to light cues during the embryonic development showed longer telomeres during the early postnatal period, suggesting that light-induced adjustments could allow them to buffer the competitive disadvantages associated with hatching asynchrony. Our study provides evidence that postnatal junior phenotypes are, at least in part, prenatally shaped by light cues that act during a critical temporal window of developmental sensitivity.

The macroecology and evolution of avian competence forBorrelia burgdorferi

AbstractAimPrediction of novel reservoirs of zoonotic pathogens would be improved by the identification of interspecific drivers of host competence (i.e., the ability to transmit pathogens to new hosts or vectors). Tick‐borne pathogens can provide a useful model system, because larvae become infected only when feeding on a competent host during their first blood meal. For tick‐borne diseases, competence has been studied best forBorrelia burgdorferisensu lato (Bbsl), which causes Lyme borreliosis. Major reservoirs include several small mammal species, but birds might play an under‐recognized role in human risk given their ability to disperse infected ticks across large spatial scales. Here, we provide a global synthesis of the ecological and evolutionary factors that determine the ability of bird species to infect larval ticks withBbsl.LocationGlobal.Time period1983–2019.Major taxa studiedBirds.MethodsWe compiled a dataset ofBbsl competence across 183 bird species and applied meta‐analysis, phylogenetic factorization and boosted regression trees to describe spatial and temporal patterns in competence, characterize its phylogenetic distribution across birds, reconstruct its evolution and evaluate the trait profiles associated with competent avian species.ResultsHalf of the sampled bird species show evidence of competence forBbsl. Competence displays moderate phylogenetic signal, has evolved multiple times across bird species and is pronounced in the genusTurdus. Trait‐based analyses distinguished competent birds with 80% accuracy and showed that such species have low baseline corticosterone, exist on both ends of the pace‐of‐life continuum, breed and winter at high latitudes and have broad migratory movements into their breeding range. We used these trait profiles to predict various likely but unsampled competent species, including novel concentrations of avian reservoirs within the Neotropics.Main conclusionOur results can generate new hypotheses for how birds contribute to the dynamics of tick‐borne pathogens and help to prioritize surveillance of likely but unsampled competent birds. Our findings also emphasize that birds display under‐recognized variation in their contributions to enzootic cycles ofBbsl and the broader need to consider competence in ecological and predictive studies of multi‐host pathogens.

Cortisol is the predominant glucocorticoid in the giant paedomorphic hellbender salamander (Cryptobranchus alleganiensis)

Corticosterone is widely regarded to be the predominant glucocorticoid produced in amphibians. However, we recently described unusually low baseline and stress-induced corticosterone profiles in eastern hellbenders (Cryptobranchus alleganiensis alleganiensis), a giant, fully aquatic salamander. Here, we hypothesized that hellbenders might also produce cortisol, the predominant glucocorticoid used by fishes and non-rodent mammals. To test our hypothesis, we collected plasma samples in two field experiments and analyzed them using multiple analytical techniques to determine how plasma concentrations of cortisol and corticosterone co-varied after 1) physical restraint and 2) injection with adrenocorticotropic hormone (ACTH), the pituitary hormone responsible for triggering the release of glucocorticoids from amphibian interrenal glands. Using liquid chromatography-mass spectrometry, we found that baseline and restraint-induced plasma concentrations of cortisol were more than five times those of corticosterone. We then demonstrated that plasma concentrations of both glucocorticoids increased in response to ACTH in a dose-dependent manner, but cortisol concentrations were consistently higher (up to 10-fold) than corticosterone. Cortisol and corticosterone concentrations were not correlated with one another at basal or induced conditions. The extremely low plasma concentrations of corticosterone in hellbenders suggests that corticosterone could simply be a byproduct of cortisol production, and raises questions as to whether corticosterone has any distinct physiological function in hellbenders. Our results indicate that hellbenders produce cortisol as their predominant glucocorticoid, supporting a small and inconclusive body of literature indicating that some other amphibians may produce appreciable quantities of cortisol. We hypothesize that the use of cortisol by hellbenders could be an adaptation to their fully aquatic life history due to cortisol’s ability to fulfill both mineralocorticoid and glucocorticoid functions, similar to its functions in fishes. Given the large number of amphibian species that are fully aquatic or have aquatic life stages, we suggest that the broadly held assumption that corticosterone is the predominant glucocorticoid in all amphibians requires further scrutiny. Ultimately, multi-species tests of this assumption will reveal the ecological factors that influenced the evolution of endocrine adaptations among amphibian lineages, and may provide insight into convergent evolution of endocrine traits in paedomorphic species.

Posthatching Parental Care and Offspring Growth Vary with Maternal Corticosterone Level in a Wild Bird Population

Corticosterone is the primary metabolic steroid in birds and is vital for maintaining homeostasis. However, the relationship between baseline corticosterone and reproduction is unclear, and we lack an understanding of how differences in baseline corticosterone at one stage of the breeding cycle influence reproductive effort at later stages. In a wild population of house wrens, we quantified the concentration of corticosterone in yolks of freshly laid eggs as an integrated measure of maternal physiology and related this to a behavioral measure of stress reactivity made during the nestling period, namely, the latency with which females resumed parental activities following a standardized disturbance at their nest (setting up a camera to record provisioning). Females that recently produced eggs containing higher corticosterone concentrations, which were significantly repeatable within females, took longer to resume activity related to parental care (i.e., feeding and brooding young) following the disturbance. Moreover, a female's latency to resume parental activities negatively predicted her provisioning of nestlings with food and the condition of these young at fledging but did not predict the number fledged. We cross-fostered offspring prior to hatching so these effects on maternal behavior are independent of any prenatal maternal effects on nestlings via the egg. These results are consistent with earlier findings, suggesting that females with higher baseline corticosterone during egg laying or early incubation tend to prioritize self-maintenance over reproduction compared with females with lower baseline corticosterone and suggest that a female's latency to return to her nest and resume parental care following a disturbance might represent a simple, functional measure of maternal stress reactivity.

Migratory Species Show Distinct Patterns in Corticosterone Levels during Spring and Autumn Migrations

Abstract Twice a year billions of birds migrate between breeding and wintering grounds. To facilitate migrations, birds develop migratory disposition, a complex suite of behavioral and physiological adjustments. Glucocorticoid hormone corticosterone is involved in the regulation of migratory behavior and physiology, however no consensus on its exact role in controlling avian migration exists. Using a large dataset on seven songbird species (long- and short-distance migrants) obtained during eleven consecutive migratory seasons on the Courish Spit of the Baltic Sea, we showed the general tendency of similar baseline corticosterone concentrations during both migrations, although stress-induced levels were generally much higher during spring. No difference between long- and short-distance migrants was found in either baseline or stress-induced levels, while there was substantial between-species variation, especially in baseline concentrations. The distinct patterns of corticosterone secretion during seasonal migrations even in ecologically similar species indicate that it is likely to be a species-specific trait. Thus, our study corroborates the inconsistency found in earlier studies and demonstrates how scientific understanding of the role of corticosterone during migration is still evolving. Rather low baseline corticosterone concentrations observed in this study emphasize that birds in both migratory seasons were not in a “stressed” state before capture.

Hormones and Fitness: Evidence for Trade-Offs in Glucocorticoid Regulation Across Contexts

Glucocorticoid hormones are important regulators of metabolic processes, and of the behavioral and physiological responses to stressors. Within-population variation in circulating glucocorticoids has been linked with both reproductive success and survival, but the presence and direction of relationships vary. Although conceptual models suggest the potential for interactions between glucocorticoid secretion under acute stress and non-acute stress contexts to influence phenotype and fitness, very little is known about the presence or implications of such interactions. Here we use a large data set from breeding tree swallows (Tachycineta bicolor; n=215) to test the predictors of reproductive success and annual survival probability in females and males over a four-year period. Across years and life history stages, glucocorticoids predicted fitness in female tree swallows. Under challenging conditions, females that maintained low baseline corticosterone levels during incubation had higher reproductive success. During the nestling provisioning period, interactions between baseline glucocorticoids and the stress response suggest that females may face trade-offs in the regulation of glucocorticoids across contexts. Reproductive success was highest among females that maintained low baseline glucocorticoids coupled with a strong acute stress response, and among females with high baseline glucocorticoids and a weak acute stress response. Females with low or high glucocorticoid levels across contexts (baseline and acute stress) fledged fewer young. Glucocorticoid levels did not predict fledging success in males. None of the models of annual survival probability in females received strong support; model comparisons suggested weak negative effects of stress responsiveness during provisioning, and positive effects of body condition and age, on survival. Male survival probability was not predicted by breeding phenotype. Within and across years the glucocorticoid stress response was individually repeatable; across-year repeatability was particularly high during the nestling provisioning period. Both measures of the acute stress response – circulating stress-induced corticosterone levels and their stress-induced increase – showed similar repeatability, but baseline corticosterone was not repeatable within or across years. Overall, our results suggest that taking into account the potential for individual differences in glucocorticoid trait expression in one context to influence optimal endocrine expression in other contexts could be important for understanding the evolution of endocrine systems.

Elevated temperatures are associated with stress in rooftop-nesting Common Nighthawk (Chordeiles minor) chicks.

Grasslands and riparian forests in southeastern South Dakota have been greatly reduced since historical times, primarily due to conversion to row-crop agriculture. Common Nighthawk (Chordeiles minor) nesting habitat includes grasslands, open woodlands and urban rooftops, but nesting sites in southeastern South Dakota are confined to rooftops, as natural nesting habitat is limited. Nighthawks nesting on exposed rooftop habitats may encounter thermal conditions that increase operative temperatures relative to vegetated land cover types. Mean humidity has increased and mean wind speed and cloud cover have decreased during the nighthawk breeding season from 1948 to 2016 in southeastern South Dakota. These changes might contribute to increasing operative temperatures at exposed rooftop nest sites and this could influence chick condition. We studied nest micro-climate and the plasma stress response for 24 rooftop-nesting nighthawk chicks from 17 nests during 2015 and 2016. High humidity prior to blood collection reduced both baseline and stress-induced plasma corticosterone (CORT). In contrast, high maximum temperatures during the day before sampling increased stress-induced CORT. The magnitude of the chick stress response was significantly negatively related to maximum wind speed for the week prior to CORT measurement. Other weather and micro-climate variables were not significant effectors of CORT metrics. Most chicks had low baseline CORT and were able to mount a stress response, but a subset of chicks (n = 4) showed elevated baseline CORT and a negative association between the magnitude of stress response and ambient temperature. For this subset, mean ambient temperature for the day before sampling was significantly higher (2.3°C) than for chicks with typical baseline CORT levels. These data suggest that regional climate change trends could affect the ability of nighthawk chicks to mount a stress response, which, in turn, might influence the susceptibility of nighthawk chicks to climate change in the Northern Prairie region.

Wound-healing ability is conserved during periods of chronic stress and costly life history events in a wild-caught bird

Chronic stress, potentially through the actions of corticosterone, is thought to directly impair the function of immune cells. However, chronic stress may also have an indirect effect by influencing allocation of energy, ultimately shifting resources away from the immune system. If so, the effects of chronic stress on immune responses may be greater during energetically-costly life history events. To test whether the effects of chronic stress on immune responses differ during expensive life history events we measured wound healing rate in molting and non-molting European starlings (Sturnus vulgaris) exposed to control or chronic stress conditions. To determine whether corticosterone correlated with wound healing rates before starting chronic stress, we measured baseline and stress-induced corticosterone and two estimates of corticosterone release and regulation, negative feedback (using dexamethasone injection), and maximal capacity of the adrenals to secrete corticosterone (using adrenocorticotropin hormone [ACTH] injection). After 8days of exposure to chronic stress, we wounded both control and chronically stressed birds and monitored healing daily. We monitored nighttime heart rate, which strongly correlates with energy expenditure, and body mass throughout the study. Measures of corticosterone did not differ with molt status. Contrary to work on lizards and small mammals, all birds, regardless of stress or molt status, fully-healed wounds at similar rates. Although chronic stress did not influence healing rates, individuals with low baseline corticosterone or strong negative feedback had faster healing rates than individuals with high baseline corticosterone or weak negative feedback. In addition, wound healing does appear to be linked to energy expenditure and body mass. Non-molting, chronically stressed birds decreased nighttime heart rate during healing, but this pattern did not exist in molting birds. Additionally, birds of heavier body mass at the start of the experiment healed wounds more rapidly than lighter birds. Finally, chronically stressed birds lost body mass at the start of chronic stress, but after wounding all birds regardless of stress or molt status started gaining weight, which continued for the remainder of the study. Increased body mass could suggest compensatory feeding to offset energetic or resource demands (e.g., proteins) of wound healing. Although chronic stress did not inhibit healing, our data suggest that corticosterone may play an important role in mediating healing processes and that molt could influence energy saving tactics during periods of chronic stress. Although the experiment was designed to test allostasis, interpretation of data through reactive scope appears to be a better fit.

Hypothalamic–pituitary–adrenal axis activity is not elevated in a songbird (Junco hyemalis) preparing for migration

During spring, increasing daylengths stimulate gonadal development in migratory birds. However, late-stage reproductive development is typically postponed until migration has been completed. The hypothalamic–pituitary–adrenal (HPA) axis regulates the secretion of glucocorticoids, which have been associated with pre-migratory hyperphagia and fattening. The HPA-axis is also known to suppress the hypothalamic–pituitary–gonadal (HPG) axis, suggesting the possibility that final transition into the breeding life history stage may be slowed by glucocorticoids. We hypothesized that greater HPA-axis activity in individuals preparing for migration may foster preparation for migration while simultaneously acting as a “brake” on the development of the HPG-axis. To test this hypothesis, we sampled baseline corticosterone (CORT), stress-induced CORT, and negative feedback efficacy of Dark-eyed Juncos (Junco hyemalis) in an overwintering population that included both migratory (J.h. hyemalis) and resident (J.h. carolinensis) individuals. We predicted that compared to residents, migrants would have higher baseline CORT, higher stress-induced CORT, and weaker negative feedback. Juncos were sampled in western Virginia in early March, which was about 2–4wk before migratory departure for migrants and 4–5wk before first clutch initiation for residents. Contrary to our predictions, we found that migrants had lower baseline and stress-induced CORT and similar negative feedback efficacy compared with residents, which suggests that delayed breeding in migrants is influenced by other physiological mechanisms. Our findings also suggest that baseline CORT is not elevated during pre-migratory fattening, as migrants had lower baseline CORT and were fatter than residents.

Variation in plasma corticosterone in migratory songbirds: a test of the migration-modulation hypothesis.

Physiological mechanisms underlying migration remain poorly understood, but recent attention has focused on the role of the glucocorticoid hormone corticosterone (CORT) as a key endocrine regulator of migration. The migration-modulation hypothesis (MMH) proposes that baseline plasma CORT levels are elevated in migratory birds to facilitate hyperphagia and lipogenesis and that further elevation of CORT in response to acute stress is suppressed. Consequently, CORT may be a poor indicator of individual condition or environmental variation in migratory birds. We tested the MMH by measuring baseline and stress-induced CORT in common yellowthroats (Geothlypis trichas) during fall migration over 2 consecutive years in the Revelstoke Reach drawdown zone, a migratory stopover site affected by local hydroelectric operations. Birds had low baseline CORT at initial capture (<5 ng/mL) and then showed a robust stress response, with CORT increasing to ca. 50 ng/mL within 10-20 min. Our data therefore do not support the MMH. Baseline CORT did not vary with body mass, time of capture, Julian day, or year, suggesting that variable flooding regimes did not affect baseline CORT. Individual variation in the rate of increase in CORT was correlated with Julian day, being higher later in the migration period. Our data suggest that plasma CORT can be a useful metric in migration studies.

Endocrine phenotype, reproductive success and survival in the great tit, Parus major

A central goal in evolutionary ecology is to characterize and identify selection patterns on the optimal phenotype in different environments. Physiological traits, such as hormonal responses, provide important mechanisms by which individuals can adapt to fluctuating environmental conditions. It is therefore expected that selection shapes hormonal traits, but the strength and the direction of selection on plastic hormonal signals are still under investigation. Here, we determined whether, and in which way, selection is acting on the hormones corticosterone and prolactin by characterizing endocrine phenotypes and their relationship with fitness in free-living great tits, Parus major. We quantified variation in circulating concentrations of baseline and stress-induced corticosterone and in prolactin during the prebreeding (March) and the breeding season (May) for two consecutive years, and correlated these with reproductive success (yearly fledgling number) and overwinter survival in female and male individuals. In both years, individuals with high baseline corticosterone concentrations in March had the highest yearly fledgling numbers; while in May, individuals with low baseline corticosterone had the highest yearly reproductive success. Likewise, individuals that displayed strong seasonal plasticity in baseline corticosterone concentrations (high in March and low in May) had the highest reproductive success in each year. Prolactin concentrations were not related to reproductive success, but were positively correlated to the proximity to lay. Between-year plasticity in stress-induced corticosterone concentrations of males was related to yearly variation in food abundance, but not to overall reproductive success. These findings suggest that seasonally alternating directional selection is operating on baseline corticosterone concentrations in both sexes. The observed between-year consistency in selection patterns indicates that a one-time hormone sample in a given season can allow the prediction of individual fitness.

Corticosterone secretion patterns prior to spring and autumn migration differ in free-living barn swallows (Hirundo rustica L.)

Recent studies of long-distance migratory birds show that behavioural and physiological changes associated with predictable or unpredictable challenges during the annual cycle are distinctively regulated by hormones. Corticosterone is the primary energy regulating hormone in birds. Corticosterone levels are elevated during stresses but they are also modulated seasonally according to environmental conditions and life-history demands. We measured the baseline and stress-induced levels of corticosterone in the barn swallow (Hirundo rustica L.) just before spring and autumn migrations in South Africa and Finland, respectively. Barn swallows completing their pre-breeding moult had low body condition (residual body mass) and high baseline corticosterone levels in the wintering grounds. In contrast, baseline corticosterone levels in Finland were low and not related to residual mass. These data contradict the first prediction of the migration modulation hypothesis (MMH) by showing no association with baseline corticosterone levels and pre-migratory fuelling. Yet, the adrenocortical response to the capture and handling stress was notably blunted in South Africa compared to a strong response in Finland. Further, individuals that had started fuelling in Finland showed a reduced response to the handling stress. Taken together, elevated baseline corticosterone levels and high residual mass may blunt the adrenocortical response in long-distance migrants and aerial feeders such as the barn swallow. This observation lends support to the second prediction of the MMH.

Energetic constraints and parental care: Is corticosterone indicative of energetic costs of incubation in a precocial bird?

Suppression of the adrenocortical response (e.g., corticosterone release) to an acute stressor is a physiological adjustment thought to decrease the likelihood of avian parents abandoning their nests. However, some periods of parental care, like incubation, are energetically costly, thus corticosterone could increase during these stages to allow incubating parents to utilize energy reserves. Wood ducks (Aix sponsa) have ~30day incubation periods and only the female incubates the eggs. We hypothesized that corticosterone would be important in regulating energy availability during incubation in this species. Because resources invested in reproduction increase with clutch size, we also hypothesized that clutch size would influence plasma corticosterone during incubation. We measured baseline and stress-induced corticosterone in incubating females during early and late stages of incubation. At both stages of incubation all hens had low baseline corticosterone levels. However, we found that stress-induced corticosterone was 105% greater late in incubation than early in incubation. We also detected a significant negative correlation between female body mass and stress-induced corticosterone late in incubation, but not during the early stages of incubation. Furthermore, we found a significant positive relationship between stress-induced corticosterone and clutch size. These lines of evidence support the hypothesis that incubation in wood ducks is energetically costly and corticosterone is important in supporting the energetic demands of incubating hens. Our findings suggest that corticosterone's role in supporting parental care behaviors are dynamic and are influenced by several factors and that there is a greater physiological cost associated with incubating larger clutches.

Urinary corticosterone metabolites and chytridiomycosis disease prevalence in a free-living population of male Stony Creek frogs (Litoria wilcoxii)

The emerging amphibian disease chytridiomycosis, which is caused by the fungal pathogen (Batrachochytrium dendrobatidis, Bd), has caused mass mortalities of native amphibian populations globally. There have been no previous studies on the relationships between stress hormones in free-living amphibians and Bd infections. In this study, we measured urinary corticosterone metabolite concentrations and Bd infections within free-living populations of male Stony Creek frog (Litoria wilcoxii) in Queensland, Australia. Prevalence of Bd zoospores from frog skin swabs was quantified using a real-time quantitative PCR technique. A urinary corticosterone enzyme-immunoassay (EIA) was validated using adrenocorticotropic hormone (ACTH) challenge. Urinary corticosterone concentrations of male frogs increased within 1–2days after ACTH challenge and returned to baseline levels within 3days post-ACTH injection. None of the frogs showed any rise in urinary corticosterone after saline injections. Individual male frogs showed either low or high baseline corticosterone concentrations. Male frogs identified as positive for Bd infection had significantly higher baseline urinary corticosterone concentrations in comparison to Bd negative male frogs. Urinary corticosterone EIA provides a reliable indication of stress in this frog species and this non-invasive physiological tool can be used to further assess the dynamics of Bd infections and physiological stress responses in other native amphibians.

Hormone levels predict individual differences in reproductive success in a passerine bird

Hormones mediate major physiological and behavioural components of the reproductive phenotype of individuals. To understand basic evolutionary processes in the hormonal regulation of reproductive traits, we need to know whether, and during which reproductive phases, individual variation in hormone concentrations relates to fitness in natural populations. We related circulating concentrations of prolactin and corticosterone to parental behaviour and reproductive success during both the pre-breeding and the chick-rearing stages in both individuals of pairs of free-living house sparrows, Passer domesticus. Prolactin and baseline corticosterone concentrations in pre-breeding females, and prolactin concentrations in pre-breeding males, predicted total number of fledglings. When the strong effect of lay date on total fledgling number was corrected for, only pre-breeding baseline corticosterone, but not prolactin, was negatively correlated with the reproductive success of females. During the breeding season, nestling provisioning rates of both sexes were negatively correlated with stress-induced corticosterone levels. Lastly, individuals of both sexes with low baseline corticosterone before and high baseline corticosterone during breeding raised the most offspring, suggesting that either the plasticity of this trait contributes to reproductive success or that high parental effort leads to increased hormone concentrations. Thus hormone concentrations both before and during breeding, as well as their seasonal dynamics, predict reproductive success, suggesting that individual variation in absolute concentrations and in plasticity is functionally significant, and, if heritable, may be a target of selection.

Stress and the timing of breeding: Glucocorticoid-luteinizing hormones relationships in an arctic seabird

In birds, stressful environmental conditions delay the timing of breeding but the underlying mechanisms are poorly understood. The stress hormone corticosterone appears to be a good candidate for mediating the decision to breed and when to start egg-laying, via a possible inhibition of luteinizing hormone (LH) and sex-steroids production. We used luteinizing hormone releasing hormone (LHRH) challenge in pre-laying male and female Black-legged kittiwakes ( Rissa tridactyla) to test whether LH and testosterone secretion were depressed by elevated corticosterone levels. Females bearing high baseline corticosterone levels showed reduced baseline LH levels and a low ability to release LH, following LHRH challenge. Further, females bearing low baseline LH levels and elevated baseline corticosterone levels were more likely to skip breeding. However, non-breeding females were physiologically primed for breeding, since they mounted high LHRH-induced LH release. Egg-laying date was advanced in good body condition females but was unaffected by hormones secretion. In males, corticosterone levels had no effect on LH and/or testosterone secretion and did not affect their decision to breed. Interestingly, males with high LHRH-induced testosterone release bred early. Our study highlights clear sex-differences in the HPG sensitivity to stress hormones in pre-laying kittiwakes. Because females have to store body reserves and to build up the clutch, they would be more sensitive to stress than males. Moreover, intrasexual competition could force male kittiwakes to acquire reproductive readiness earlier in the season than females and to better resist environmental perturbations. We suggest that high testosterone releasing ability would mediate behavioural adjustments such as courtship feeding, which would stimulate early egg-laying in females.

Arrival date and territorial behavior are associated with corticosterone metabolite levels in a migratory bird

Glucocorticoids promote the mobilization of energy stores and they may facilitate the expression of energetically expensive functions. Early arrival on the breeding grounds in migratory species and territorial competition are energetically demanding activities that may be supported by elevated baseline glucocorticoid levels. Here, we evaluated the associations between the baseline levels of excreted corticosterone (CORT) metabolites of male Pied Flycatchers (Ficedula hypoleuca) just after arriving on their breeding area and timing of arrival, considering ornamental traits indicative of social status, like forehead patch size and black plumage coloration, as well as heat shock protein levels (HSP60). We observed a positive association of CORT metabolites with HSP60 levels, which are synthesized under several environmental challenges affecting cell homeostasis. Our data showed a negative association between arrival date and CORT metabolite levels, possibly as a result of the higher energetic demands imposed by the hard environmental conditions experienced at the time of an early arrival after migration. We observed a negative relationship of forehead patch dimensions and CORT metabolite levels, suggesting that dominance is associated with low baseline CORT metabolites. Also, males that expressed a higher degree of territorial behaviour when exposed to a playback song of a conspecific at their nest-box showed higher CORT metabolites upon arrival than males that expressed a lower degree of territorial behavior. This may indicate that elevated baseline CORT metabolite levels may facilitate an intense territorial competition in males. Thus, male–male competition may be a factor affecting observed baseline glucocorticoid levels in migratory birds.

Baseline and acute levels of corticosterone in Florida Scrub-Jays ( Aphelocoma coerulescens): Effects of food supplementation, suburban habitat, and year

Recent work has shown that Florida Scrub-Jays ( Aphelocoma coerulescens) living in suburban habitats, as well as birds that reside in a natural habitat (hereafter referred to as wildland birds) provided supplemental food during the pre-breeding period, have lower baseline corticosterone (CORT) levels than nonsupplemented wildland birds, suggesting that predictable and abundant foods can lower stress levels. Here, we investigate whether the acute CORT response to capture stress reflects our earlier findings. During the pre-breeding seasons in 2004 and 2005 we captured over 200 Florida Scrub-Jays and collected four blood samples from each bird over a 30 min time period. We compared hormonal and morphometric data from birds in four treatment groups: wildland birds that were provided with supplemental food during the pre-breeding period, wildland birds that were provided with supplemental food during the winter and pre-breeding period, nonsupplemented wildland birds, and suburban birds that effectively have access to ad libitum human-source food year-round. Suburban birds had lower baseline levels, and a more acute initial CORT response (i.e., rate of increase) than wildland birds, but we did not observe any differences among the treatment groups of wildland birds. Baseline CORT concentrations were unusually low across treatments in 2005, a year that was characterized by increased adult body mass, superior body condition, and a high fledgling success rate, all of which suggest favorable environmental conditions. Thus, abundant or more reliable food sources in 2005 may have overwhelmed any potential benefits of supplemental feeding and explain the absence of any strong experimental effects on CORT for that year.

Chronic stress in free-living European starlings reduces corticosterone concentrations and reproductive success

Chronic increases in stress hormones such as glucocorticoids are maladaptive, yet studies demonstrating a causal relationship among chronic stress, increases in glucocorticoid concentrations, and subsequent fitness costs in free-living animals are lacking. We experimentally induced chronic psychological stress in female European starlings ( Sturnus vulgaris) by subjecting half of the females at our study site to a chronic stress protocol consisting of 4, 30 min stressors (loud radio, predator calls, a novel object, or predator decoys including a snake, rat, and owl) administered in random order daily for 8 days after clutch completion. Experimental females were captured at the end of the chronic stress protocol (9 days after the onset of the chronic stress protocol), and unstressed control females were captured at the same stage of the nesting cycle. Chronically stressed females had lower baseline corticosterone (CORT, the avian glucocorticoid) concentrations and lower reproductive success than unstressed females. Furthermore, surviving nestlings in experimentally stressed broods showed sensitization of the CORT response to acute stress, which is a physiological change that could persist to adulthood. Attenuation of baseline CORT concentrations in adult females is contrary to the general assumption that elevated CORT concentrations indicate stress, suggesting that more research is necessary before CORT concentrations can be used to accurately assess chronic stress in field studies.