Baseline CORT Research Articles

Relationships between Spatial Biology and Physiological Ecology in the Gopher Tortoise, Gopherus polyphemus.

AbstractThe overlap between spatial and physiological ecology is generally understudied, yet both fields are fundamentally related in assessing how individuals balance limited resources. Herein, we quantified the relationships between spatial ecology using two parameters of home range (annual home range area and number of burrows used in 1 yr) and four measures of physiology that integrate stress and immunity (baseline plasma corticosterone [CORT] concentration, plasma lactate concentration, heterophil-to-lymphocyte [H∶L] ratio, and bactericidal ability [BA]) in a wild free-ranging population of the gopher tortoise (Gopherus polyphemus) to test the hypothesis that space usage is correlated with physiological state. We also used structural equation models (SEMs) to test for causative relationships between the spatial and physiological parameters. We predicted that larger home ranges would be negatively correlated with traditional biomarkers of stress and positively correlated with immunity, consistent with our hypothesis that home ranges are determined based on individual condition. Males had larger home ranges, used more burrows, and had higher baseline CORT than females. We found significant negative correlations between lactate and home range (, , ). CORT was negatively correlated with the number of burrows used in both sexes (, , , adjusted ). No correlations were observed between space use and BA or, notably, H∶L ratio. SEMs suggested that variation in the number of burrows used was a result of variation in baseline CORT. The lack of a relationship between H∶L ratio and home range suggests that home range differences are not associated with differences in chronic stress, despite the pattern between baseline CORT and number of burrows used. Instead, this study indicates that animals balance trade-offs in energetics, likely by way of baseline corticosteroid, in such a way as to maintain function across continuously variable home range strategies.

Male plumage signal expression is related to feather corticosterone concentration in the Pied Flycatcher Ficedula hypoleuca

Male signals may express the capacity to sustain environmental challenges. In some migratory birds like the Pied Flycatcher Ficedula hypoleuca, plumage ornaments are molted in the winter quarters shortly before spring migration while most feathers are replaced shortly after the breeding season in the breeding areas. The concentration of corticosterone in feathers (CORTf) may relate to baseline CORT levels at the time of molt which could be expressed through plumage signals. Male Pied Flycatchers present white patches on forehead feathers and tertials which are molted before spring migration and on secondaries and primaries replaced after breeding. They also express a variable degree of melanisation of head and back feathers molted in the wintering areas. All these plumage traits have been previously shown to function in social contests and/or mate attraction. Here we have collected tertials on the two wings and two tail feathers, molted in wintering and breeding areas respectively, of males in a Spanish montane population and analysed CORTf in the laboratory with standard enzyme immunoassays. There is no correlation within individuals between CORTf in the two types of feathers, although levels are similar. The size of the forehead patch is negatively associated with CORTf in tail feathers, mainly in small males, while the blackness of head and back is negatively related to CORTf in tertials, mainly in large males. The size of the wing patch composed of patches on feathers molted both in wintering (tertials) and breeding areas (primaries and secondaries) is not related to CORTf in any type of feather. Different male plumage traits thus may reflect circulating CORT levels during molt processes occurring in the wintering respectively breeding range as expressed by CORTf in different types of feathers.

Are glucocorticoids good indicators of disturbance across populations that exhibit cryptic variation in contaminant tolerance?

AbstractGlucocorticoids (corticosterone/cortisol; cort) are frequently used in conservation as biomarkers of disturbance in wild populations. However, the context‐dependent nature of cort means that it may not always accurately reflect disturbance. For example, there is growing evidence that wildlife populations can evolve or acclimate to human‐induced environmental change (i.e. contaminants) by expressing higher levels of tolerance. Mechanisms that allow for populations to achieve higher contaminant tolerance can be related to cort and thereby impact the reliability of cort as an indicator of disturbance. This study asks: (1) do wildlife populations that differ in tolerance to contaminants differentially express baseline and stress‐induced cort and (2) is cort a viable indicator of disturbance across populations that differ in tolerance to contaminants? Toward this goal, we identified three wood frog Rana sylvatica populations with relatively high NaCl tolerance and three populations with relatively low NaCl tolerance. Tadpoles from these populations were reared to metamorphosis in either an environmentally relevant concentration of NaCl (0.5 g L−1 NaCl) or a control (0 g L−1 NaCl). At metamorphosis we used a non‐invasive waterborne assay to measure baseline and stress‐induced cort release rates and measured fitness‐related metrics. We found that contaminant tolerance influences cort levels. More tolerant populations had lower baseline cort and higher fitness compared to less tolerant populations. However, despite variation in cort across populations with different levels of tolerance, cort still represents a viable indicator of condition as our results show a consistent negative relationship between cort and fitness. Lastly, we found that levels of cort were consistent regardless of whether amphibians were reared in NaCl contaminated or non‐contaminated environments. Overall, we emphasize the importance of recognizing population‐level variation in cort due to contaminant tolerance when using cort as a biomarker for conservation purposes.

Feather, But Not Plasma, Glucocorticoid Response to Artificial Light at Night Differs between Urban and Forest Blue Tit Nestlings.

Urbanization drives phenotypic variation in many animal species. This includes behavioral and physiological traits such as activity patterns, aggression, and hormone levels. A current challenge of urban evolutionary ecology is to understand the environmental drivers of phenotypic variation in cities. Moreover, do individuals develop tolerance to urban environmental factors, which underlie adaptative responses and contribute to the evolution of urban populations? Most available evidence comes from correlative studies and rare experiments where a single urban-related environmental factor has been manipulated in the field. Here we present the results of an experiment in which we tested for differences in the glucocorticoid (CORT) response of urban and rural blue tits nestlings (Cyanistes caeruleus) to artificial light at night (ALAN). ALAN has been suggested to alter CORT response in several animal species, but to date no study has investigated whether this effect of ALAN differs between urban and rural populations. Immediately after hatching, urban and forest broods were either exposed to 2 lux of ALAN (using an LED source mounted inside the nestbox) or received no treatment (dark control). The experiment lasted until the chicks fledged. When the chicks were 13 days old plasma samples were collected to measure baseline CORT concentrations, and feather samples to provide an integrative measure of CORT during growth. Forest birds had higher plasma CORT (pCORT) concentrations than their urban counterparts, irrespective of whether they were exposed to ALAN or not. Conversely, we found population-specific responses of feather CORT to ALAN. Specifically, urban birds that received ALAN had increased feather CORT compared with the urban dark controls, while the opposite was true for the forest birds. pCORT concentrations were negatively associated to fledging success, irrespective of population and treatment, while feather CORT was positively associated to fledging success in broods exposed to ALAN, but negatively in the dark control ones. Our results demonstrate that ALAN can play a role in determination of the glucocorticoid phenotype of wild animals, and may thus contribute to phenotypic differences between urban and rural animals.

Social isolation enhances cued-reinstatement of sucrose and nicotine seeking, but this is reversed by a return to social housing

Physical or perceived (i.e. loneliness) social isolation is increasing in Western cultures. Unfortunately, social isolation is associated with a range of negative physical and mental health outcomes, including increased incidence of obesity and smoking. Here we monitored the impact of social isolation on a range of physical measures, and then tested whether social isolation in adult rats changes how reward-related stimuli motivate sucrose- or nicotine-seeking. Socially isolated rats showed elevated baseline CORT, gained significantly less weight across the study, were more active in response to a novel or familiar environment. Isolated rats also acquired nose-poking for a food pellet more rapidly, and showed increased susceptibility to cue-, but not reward-induced reinstatement. Notably, these effects are partially mitigated by a return to group housing, suggesting that they are not necessarily permanent, and that a return to a social setting can quickly reverse any deficits or changes associated with social isolation. This study advances our understanding of altered reward-processing in socially isolated individuals and reiterates the importance of socialisation in the treatment of disorders such as overeating and addiction.

Evaluating corticosterone as a biomarker for amphibians exposed to increased salinity and ambient corticosterone.

Physiological biomarkers are commonly used to assess the health of taxa exposed to natural and anthropogenic stressors. Glucocorticoid (GC) hormones are often used as indicators of physiological stress in wildlife because they affect growth, reproduction and survival. Increased salinity from human activities negatively influences amphibians and their corticosterone (CORT; the main amphibian GC) physiology; therefore, CORT could be a useful biomarker. We evaluated whether waterborne CORT could serve as a biomarker of salt stress for three free-living amphibian species that vary in their sensitivity to salinity: boreal chorus frogs (Pseudacris maculata), northern leopard frogs (Rana pipiens) and barred tiger salamanders (Ambystoma mavortium). Across a gradient of contamination from energy-related saline wastewaters, we tested the effects of salinity on baseline and stress-induced waterborne CORT of larvae. Stress-induced, but not baseline, CORT of leopard frogs increased with increasing salinity. Salinity was not associated with baseline or stress-induced CORT of chorus frogs or tiger salamanders. Associations between CORT and salinity were also not related to species-specific sensitivities to salinity. However, we detected background environmental CORT (ambient CORT) in all wetlands and spatial variation was high within and among wetlands. Higher ambient CORT was associated with lower waterborne CORT of larvae in wetlands. Therefore, ambient CORT likely confounded associations between waterborne CORT and salinity in our analysis and possibly influenced physiology of larvae. We hypothesize that larvae may passively take up CORT from their environment and downregulate endogenous CORT. Although effects of some hormones (e.g. oestrogen) and endocrine disruptors on aquatic organisms are well described, studies investigating the occurrence and effects of ambient CORT are limited. We provide suggestions to improve collection methods, reduce variability and avoid confounding effects of ambient CORT. By making changes to methodology, waterborne CORT could still be a promising, non-invasive conservation tool to evaluate effects of salinity on amphibians.

Survival is negatively associated with glucocorticoids in a wild ungulate neonate.

It is unknown how ungulate physiological responses to environmental perturbation influence overall population demographics. Moreover, neonatal physiological responses remain poorly studied despite the importance of neonatal survival to population growth. Glucocorticoid (GC) hormones potentially facilitate critical physiological and behavioral responses to environmental perturbations. However, elevated GC concentrations over time may compromise body condition and indirectly reduce survival. We evaluated baseline salivary cortisol (CORT; a primary GC in mammals) concentrations in 19 wild neonatal white-tailed deer (Odocoileus virginianus) in a northern (NS) and southern (SS) area in Pennsylvania. After ranking survival models consisting of variables hypothesized to influence neonate survival (i.e. weight, sex), the probability of neonate survival was best explained by CORT concentrations, where elevated CORT concentrations were associated with reduced survival probability to 12 weeks of age. Cortisol concentrations were greater in the SS where predation rates and predator densities were lower. As the first evaluation of baseline CORT concentrations in an ungulate neonate to our knowledge, this is also the first study to demonstrate CORT concentrations are negatively associated with ungulate survival at any life stage. Glucocorticoid hormones could provide a framework in which to better understand susceptibility to mortality in neonatal white-tailed deer.

Developing a Stopover-CORT hypothesis: Corticosterone predicts body composition and refueling rate in Gray Catbirds during migratory stopover

Migratory flight is energetically challenging, requiring alternating phases of fuel catabolism and fuel accumulation, accompanied by dramatic changes in body composition and behavior. Baseline corticosterone (CORT; the primary glucocorticoid in birds) is thought to underlie transitions between fuel catabolism during flight, fuel deposition during stopover, and the initiation of migratory flight. However, studies of CORT on stopover physiology and behavior remain disparate efforts, lacking the cohesion of a general hypothesis. Here we develop a Stopover-CORT hypothesis formalizing the relationships among CORT, body condition, and refueling rate in migratory birds. First we expect body mass to increase with triglycerides (TRIG) as birds refuel. Second, based on a synthesis of previous literature, we predict a U-shaped CORT curve over the course of stopover, postulating that elevated CORT at arrival is reactive, responding to poor body condition, while CORT elevation before departure is preparative, driving changes in behavior and body condition. We tested these predictions in Gray Catbirds (Dumetella carolinensis) following a trans-Gulf flight during spring migration. We found baseline CORT was negatively correlated with body condition and TRIG, corresponding with our predictions for arriving and refueling—but not departing—birds. It is possible catbirds undergo regional habitat translocations rather than complete the entire stopover phase at our study site. We propose the Stopover-CORT hypothesis as a useful predictive framework for future studies of the mechanistic basis of stopover physiology. By studying the regulation of stopover refueling and departure, we may better understand physiological limitations to overall migration rate and improve assessments of habitat quality for refueling birds.

Across time and space: Hormonal variation across temporal and spatial scales in relation to nesting success.

There is a renewed interest in investigating individual variation in hormone levels in relation to fitness metrics, as hormones act as mediators of life-history trade-offs. Hormone concentrations, however, are labile, responding to both internal and external stimuli, so the relationship between hormones and fitness can be non-consistent. One explanation of this inconsistent relationship is that a single hormone sample may not be representative of individual phenotypes in a free-living species. We addressed this issue by repeatedly sampling a free-living population of mountain white-crowned sparrows, Zonotrichia leucophrys oriantha, for baseline and stress-induced corticosterone (cort) and testosterone (T) across different stages of the breeding season. We measured (co)variation using three different methods, taking into account inter- and intra-individual variances, to determine whether hormone levels and the stress response are repeatable. We documented the temporal (over 3months) and spatial (home-range) variation of individual hormone phenotypes and investigated how these components related to nesting success. At the population level, we found significant repeatability in male stress-induced cort concentrations but no repeatability in male or female baseline cort or male T concentrations. Using a new metric of intra-individual variance focusing on the stress response (profile repeatability), we found a wide range of variance scores, with most individuals showing high variation in their stress response. Similarly, we found a low level of repeatability of the reaction norm intercept and slope for the stress response across different life-history stages. Males with higher concentrations of stress-induced cort had more central home-ranges. Males with higher body condition had larger home-ranges; however, home-range size did not relate to male hormone concentrations or nesting success. We also did not find any significant relationship between variation in hormone levels and nesting success. We recommend that future studies combine both physiological and environmental components to better understand the relationship between hormones and fitness.

Living on the edge: Glucocorticoid physiology in desert iguanas (Dipsosaurus dorsalis) is predicted by distance from an anthropogenic disturbance, body condition, and population density

Ecological factors, such as habitat quality, influence the survival and reproductive success of free-living organisms. Urbanization, including roads, alters native habitat and likely influences physiology, behavior, and ultimately Darwinian fitness. Some effects of roads are clearly negative, such as increased habitat fragmentation and mortality from vehicle collision. However, roads can also have positive effects, such as decreasing predator density and increased vegetation cover, particularly in xeric habitats due to increased water run-off. Glucocorticoids are metabolic hormones that reflect baseline metabolic needs, increase in response to acute challenges, and may mediate endogenous resource trade-offs between survival and reproduction. Here we examined circulating concentrations of corticosterone (baseline and stress-induced) in desert iguanas (Dipsosaurus dorsalis) in relation to the distance from a major anthropogenic disturbance, a high-traffic road in Palm Springs, CA. Additionally, we analyzed body condition and population density as additional predictors of glucocorticoid physiology. Surprisingly, we found lower baseline CORT levels closer to the road, but no effect of distance from road on stress-induced CORT or stress responsiveness (difference between baseline and stress-induced concentrations). Both population density and body condition were negative predictors of baseline CORT, stress-induced CORT, and stress responsiveness. Given the known effect of roads to increase run-off and vegetation density, increased water availability may improve available forage and shade, which may then increase the carrying capacity of the habitat and minimize metabolic challenges for this herbivorous lizard. However, it is important to recognize that surfaces covered by asphalt are not usable habitat for iguanas, likely resulting in a net habitat loss.

Metabolic Rates Predict Baseline Corticosterone and Reproductive Output in a Free-Living Passerine.

SynopsisOrganisms continuously face environmental fluctuations, and allocation of metabolic investment to meet changing energetic demands is fundamental to survival and reproductive success. Glucocorticoid (GC) hormones (e.g., corticosterone [CORT]) play an important role in energy acquisition and allocation in the face of environmental challenges, partly through mediation of energy metabolism. Although GCs and metabolic rate are expected to covary, surprisingly few empirical studies have demonstrated such relationships, especially in wild animals. Moreover, studies testing for associations between GCs and fitness generally do not account for among-individual differences in energy expenditure or energy allocation. We measured CORT (baseline and stress-induced) and metabolic traits (resting metabolic rate [RMR], cold-induced VO2max [Msum], and aerobic scope [the difference between Msum and RMR]) in female tree swallows (Tachycineta bicolor) during chick-rearing, and tested for their associations with several variables of reproductive performance. We found a positive relationship between RMR and baseline CORT, but no consistent associations between stress-induced CORT (SI-CORT) and Msum. This suggests that while baseline CORT may be a good indicator of an individual’s baseline metabolic investment, SI-CORT responses are not associated with aerobic scope or the upper limits of aerobic performance. Furthermore, we found that metabolic traits were associated with reproductive performance: females with higher reproductive output showed higher Msum, and also tended to show higher RMR. Overall, these results suggest that metabolic traits are better predictors of reproductive output in tree swallows than CORT concentrations. They further point to the maximal aerobic capacity being higher in females investing more heavily in a current reproductive event, but whether this association reflects trade-offs between current and future reproductive efforts remains to be tested.

Influence of temperature, size and confinement on testosterone and corticosterone levels in breeding male spined toads (Bufo spinosus)

Winter breeding amphibians opportunistically engage in reproductive behaviour when environmental conditions become favourable. In such explosive breeding systems, males strongly compete for reproductive females. Although most research has been oriented on species in which males use mating calls to attract females, many high-density explosive breeding amphibians do not rely on mating calls. In such systems, larger and stronger males are thought to have significant advantages to access reproductive females. Testosterone (T) is expected to increase with the physical attributes that facilitate access to females, while increased corticosterone levels (CORT) are needed to sustain the energetic requirements associated with mating behaviour. In this study, we investigated how environmental temperature, and body size influence testosterone and corticosterone in males of an explosive winter-breeding species with low investment in mating call, the spined toad (Bufo spinosus). We found that both baseline CORT and T were positively correlated with environmental temperature. Interestingly, despite a remarkable range of variation in CORT and T, there was no evidence that either hormone was correlated with body size. Finally, we found no effect of confinement (13 h of captivity) on circulating CORT and T levels. This suggests that breeding male toads may be relatively insensitive to stress in order to maintain reproductive effort during their short mating period. Future studies should investigate both the influence of the phenology of breeding and the social interactions on these hormonal levels.

Corticosterone profiles in northern cardinals (Cardinalis cardinalis): Do levels vary through life history stages?

As animals move through life history stages, energy requirements for each stage will vary. Both daily and annual variation in the glucocoriticoid hormones (specifically corticosterone, or CORT, in birds) helps provide the variable energy needed through life history stages. In many bird species, CORT is higher in the breeding season when energy demands can be quite high and is often lower in the non-breeding season. Additionally, CORT has a role to play in the response to stressful stimuli and the level to which CORT is elevated following stressful events can vary through the annual cycle as well. Here we report on baseline and stress-induced CORT levels in both sexes of northern cardinals, Cardinalis cardinalis, a non-migrating, year-round territorial species across life history stages. Corticosterone is overall higher in the non-breeding season than the breeding season in both sexes. Males tend to have higher levels of stress-induced CORT than females, although the observed patterns are complex. Our findings differ from one of the more common profile reported in songbirds where breeding season CORT tends to be higher than non-breeding levels. A strong influence may be the prolonged breeding season seen in cardinals; lower levels of CORT during breeding may guard against adverse maternal effects, interruptions in breeding behaviors, or egg production. Additional investigation of species with similar ecologies to northern cardinals, and more populations of cardinals, may show that annual glucocorticoid profiles are more labile than previously appreciated.

Circulating corticosterone levels vary during exposure to anthropogenic stimuli and show weak correlation with behavior across an urban gradient in house finches (Haemorhous mexicanus)

Urban environments are rapidly expanding and presenting animal populations with novel challenges, many of which are thought to be stressors that contribute to low biodiversity. However, studies on stress responses in urban vs rural populations have produced mixed results, and many of these studies use a standard stressor that cannot be replicated in the wild (e.g. restraining an animal in a bag). Pairing physiological and behavioral measurements in response to urban-related stressors improves our understanding of the mechanism underlying animal success in human-dominated landscapes. Here, we examined the physiological stress (plasma corticosterone, CORT) responses of a songbird species (the house finch, Haemorhous mexicanus) to two different anthropogenic stimuli – (1) the presence of a human and (2) a captive environment containing man-made objects. During three field seasons (summer 2012, winter 2014, and winter 2015), we captured birds at six sites along an urban gradient in Phoenix, Arizona, USA and measured plasma CORT levels both before and after each trial. Though CORT levels did increase post-human exposure, though not during exposure to novel environment, indicating only one of the treatments caused a physiological response, baseline or post-trial plasma CORT levels did not differ between finches between urban and rural birds in 2012 or 2014. However, rural birds demonstrated relatively low pre- and post-trial plasma CORT levels during the human-exposure trials in 2015. Furthermore, we found few correlations between behavioral and physiological responses. A significant positive correlation was only detected between activity behavior after human approach and post-trial plasma CORT levels in 2012. Taken together, our results reveal a weak, conditional relationship between stress physiology, behavioral responses, and urbanization in house finches.

Predicting Cortisol Exposure from Paediatric Hydrocortisone Formulation Using a Semi-Mechanistic Pharmacokinetic Model Established in Healthy Adults

Optimisation of hydrocortisone replacement therapy in children is challenging as there is currently no licensed formulation and dose in Europe for children under 6 years of age. In addition, hydrocortisone has non-linear pharmacokinetics caused by saturable plasma protein binding. A paediatric hydrocortisone formulation, Infacort® oral hydrocortisone granules with taste masking, has therefore been developed. The objective of this study was to establish a population pharmacokinetic model based on studies in healthy adult volunteers to predict hydrocortisone exposure in paediatric patients with adrenal insufficiency. Cortisol and binding protein concentrations were evaluated in the absence and presence of dexamethasone in healthy volunteers (n=30). Dexamethasone was used to suppress endogenous cortisol concentrations prior to and after single doses of 0.5, 2, 5 and 10mg of Infacort® or 20 mg of Infacort®/hydrocortisone tablet/hydrocortisone intravenously. A plasma protein binding model was established using unbound and total cortisol concentrations, and sequentially integrated into the pharmacokinetic model. Both specific (non-linear) and non-specific (linear) protein binding were included in the cortisol binding model. A two-compartment disposition model with saturable absorption and constant endogenous cortisol baseline (Baseline cort,15.5nmol/L) described the data accurately. The predicted cortisol exposure for a given dose varied considerably within a small body weight range in individuals weighing <20kg. Our semi-mechanistic population pharmacokinetic model for hydrocortisone captures the complex pharmacokinetics of hydrocortisone in a simplified but comprehensive framework. The predicted cortisol exposure indicated the importance of defining an accurate hydrocortisone dose to mimic physiological concentrations for neonates and infants weighing <20 kg. EudraCT number: 2013-000260-28, 2013-000259-42.

Stress-responsiveness influences baseline glucocorticoid levels: Revisiting the under 3 min sampling rule

Plasma glucocorticoid (CORT) levels collected within 3min of capture are commonly believed to reflect pre-stressor, baseline CORT levels. Differences in these “baseline” values are often interpreted as reflecting differences in health, or the amount of social and environmental stress recently experienced by an individual. When interpreting “baseline” values it is generally assumed that any effect of capture-and-handling during the initial sampling period is small enough and consistent enough among individuals to not obscure pre-capture differences in CORT levels. However, plasma CORT increases in less than 3min post-capture in many free-living, endothermic species in which timing has been assessed. In addition, the rate of CORT secretion and the maximum level attained (i.e., the degree of stress-responsiveness) during a severe stressor often differs among individuals of the same species. In Florida scrub-jays (Aphelocoma coerulescens), an individual’s stress-responsiveness during a 30min post-capture stressor is correlated with CORT levels in samples collected within 1.5min of capture, suggesting there is an intrinsic connection between stress-responsiveness and pre-capture CORT levels. Although differences in stress-responsiveness accounted for just 11% of the variance in these samples, on average, higher stress-responsive jays (top third of individuals) had baseline values twice that of lower stress-responsive jays (bottom third). Further, plasma CORT levels begin to increase around 2min post-capture in this species, but the rate of increase between 2 and 3min differs markedly with CORT increasing more rapidly in jays with higher stress-responsiveness. Together, these data indicate that baseline CORT values can be influenced by an individual’s stress response phenotype and the differences due to stress-responsiveness can be exaggerated during sample collection. In some cases, the effects of differences in stress-responsiveness and the increase in CORT during sample collection could obscure, or supersede, differences in pre-capture plasma CORT levels that are caused by extrinsic factors.

Stressed snakes strike first: Hormone levels and defensive behavior in free ranging cottonmouths (Agkistrodon piscivorus)

Stress is believed to be an important factor mediating animal behavior. Here we explore the relationship between concentrations of a stress hormone and defensive behavior of a snake. The cottonmouth (Agkistrodon piscivorus) is an abundant, large-bodied pitviper that is well known for its intense defensive behaviors. The defensive behaviors and hormonal ecology of cottonmouths have been studied extensively, but the interaction between these is not well understood. We conducted field trials, recording the snake’s behavior and obtaining blood samples to quantify plasma CORT concentrations, both upon first encountering a snake and after a 30min standardized confinement stressor. We found that snakes with elevated levels of baseline CORT at first encounter were more likely to strike than exhibit a threat display when approached in the field. However, this behavior was not related to the magnitude of the snake’s CORT increase following confinement, suggesting that more stress-prone snakes are not more defensive. Post-stressor antipredator behavior was also not related to any of our CORT measures. This study suggests that baseline CORT levels can be important correlates of defensive behavior. If this is a causative relationship, environmental challenges that increase baseline stress levels of populations may elevate cottonmouth defensive behavior. This would increase costs associated with defensive behavior (energetic, lost opportunity, etc.) and have important consequences for animal-human interactions.

Success despite the stress: violet‐green swallows increase glucocorticoids and maintain reproductive output despite experimental increases in flight costs

Summary Glucocorticoid steroid hormones play a central role in regulating the metabolic state of animals, especially when they cope with unanticipated stressors in their environment. The cort‐adaptation hypothesis predicts that baseline concentrations of glucocorticoids are adjusted upward to match energetic needs and promote fitness when individuals are faced with physiological challenges, including those associated with reproduction. We tested the cort‐adaptation hypothesis in the violet‐green swallow (Tachycineta thalassina) by experimentally increasing flight costs during the offspring rearing period. Individual females were assigned to one of three treatments: light feather clipping (removal of four wing feathers), heavy feather clipping (removal of eight wing feathers) or a sham‐clip control. We measured baseline corticosterone and body mass immediately prior to implementing treatments during the offspring rearing period and then 10 days after initial manipulations took place. We also quantified risk‐taking behaviour, offspring feeding rate and the number of offspring fledged. Finally, we examined how treatments influenced offspring phenotype via measurements of nestling body mass and baseline corticosterone, as both measures have been associated with post‐fledging survival. We found that handicapped females significantly increased baseline corticosterone between the two sampling periods, with the magnitude of change in the light clipping and heavy clipping treatments 2·5× and 6·1× greater than controls, respectively. All individuals lost mass between the two sampling periods, but the degree of loss was greater for females in both clipping treatments relative to unmanipulated controls. In contrast, we found no evidence of treatment differences in female risk‐taking, offspring provisioning or in the number of offspring fledged. Offspring raised by females in both handicapped treatments did have significantly elevated baseline corticosterone relative to those in control broods, but we detected no treatment differences in offspring body mass. Our study found that handicapped females increased circulating glucocorticoids and were able to maintain critical parental care behaviours and raise a similar number of offspring as unmanipulated controls. Thus, increases in baseline cort of handicapped females appeared to have allowed them to maintain fitness despite increased physiological challenges, providing support for the cort‐adaptation hypothesis. A Lay Summary is available for this article.

The role of medial prefrontal corticosterone and dopamine in the antidepressant-like effect of exercise

Despite the well-documented beneficial effect of exercise on stress coping and depression treatment, its underlying neurobiological mechanism remains unclear. This is further complicated by a ‘side effect’ of exercise: it increases basal glucocorticoid (CORT), the stress hormone, which has been shown to be a mediator linking stress to depressive disorders. Here we show that three weeks of voluntary wheel running reduced rats’ immobility in the forced swim test (FST), an antidepressant-like effect. Monitoring extracellular fluids in the medial prefrontal cortex PFC (mPFC) using microdialysis we found that, wheel running was associated with higher baseline CORT, but lower FST-responsive CORT. Further, wheel running resulted in a higher dopamine (DA) both at baseline and following FST. Interestingly, the antidepressant-like effect of wheel running was completely abolished by intra-mPFC pre-microinjection of a D2R (haloperidol) but not D1R (SCH23390) antagonist, at a dose that does not affect normal rats’ performance in the FST. It suggests that exercise exerts antidepressant-like effect through upregulated DA and in a D2R dependent way in the mPFC. Importantly, the antidepressant-like effect of wheel running was also abolished by intra-mPFC pre-microinjection of a GR antagonist (RU486). Finally, intra-mPFC pre-microinjection of RU486 also downregulated the originally elevated basal and FST-responsive DA in the mPFC of exercise rats. These results suggest a causal pathway linking CORT, GR, DA, and D2R, to the antidepressant-like effect of exercise. In conclusion, exercise achieves antidepressant-like effect through the CORT-GR-DA-D2R pathway and that the increased basal CORT by exercise itself may be beneficial rather than detrimental.

Baseline glucocorticoids are drivers of body mass gain in a diving seabird.

Life‐history trade‐offs are influenced by variation in individual state, with individuals in better condition often completing life‐history stages with greater success. Although resource accrual significantly impacts key life‐history decisions such as the timing of reproduction, little is known about the underlying mechanisms driving resource accumulation. Baseline corticosterone (CORT, the primary avian glucocorticoid) mediates daily and seasonal energetics, responds to changes in food availability, and has been linked to foraging behavior, making it a strong potential driver of individual variation in resource accrual and deposition. Working with a captive colony of white‐winged scoters (Melanitta fusca deglandi), we aimed to causally determine whether variation in baseline CORT drives individual body mass gains mediated through fattening rate (plasma triglycerides corrected for body mass). We implanted individuals with each of three treatment pellets to elevate CORT within a baseline range in a randomized order: control, low dose of CORT, high dose of CORT, then blood sampled and recorded body mass over a two‐week period to track changes in baseline CORT, body mass, and fattening rates. The high CORT treatment significantly elevated levels of plasma hormone for a short period of time within the biologically relevant, baseline range for this species, but importantly did not inhibit the function of the HPA (hypothalamic–pituitary–adrenal) axis. Furthermore, an elevation in baseline CORT resulted in a consistent increase in body mass throughout the trial period compared to controls. This is some of the first empirical evidence demonstrating that elevations of baseline CORT within a biologically relevant range have a causal, direct, and positive influence on changes in body mass.