Corticosterone Stress Response Research Articles

Does short-term fasting lead to stressed-out parents? A study of incubation commitment and the hormonal stress responses and recoveries in snow petrels

The hormonal stress response is flexible and can be modulated by individuals according to its costs and benefits. Therefore, it is predicted that parents in poor body condition should modify their hormonal stress response, and thus, redirect energy allocation processes from parental care to self-maintenance when stressors occur. To test this prediction, most studies on free-living vertebrates have only focused on the stress response while the stress recovery – how quickly hormonal levels return to baseline values – has been neglected. Moreover, most studies have only focused on corticosterone – the primary mediator of allostasis – without paying attention to prolactin despite its major role in mediating parental behaviors. Here, we examined the effect of a short-term fasting event on the corticosterone and prolactin stress responses and recoveries, and we subsequently explored their relationships with parental decision in the snow petrel (Pagodroma nivea). By comparing the hormonal profiles of fasting and non-fasting snow petrels, we showed that parents modulate their corticosterone (but not prolactin) stress response according to their energetic status. We also described for the first time the hormonal stress recoveries in wild birds and found that they did not differ between fasting and non-fasting birds. Importantly, egg neglect was negatively correlated with circulating prolactin but not corticosterone levels in this species, demonstrating therefore a complex link between body condition, parental behavior and circulating corticosterone and prolactin levels. We suggest that both corticosterone and prolactin play a major role in the way parents adjust to stressors. This multiple signaling may allow parents to fine-tune their response to stressors, and especially, to activate specific allostasis-related mechanisms in a timely manner.

Stress and success: Individual differences in the glucocorticoid stress response predict behavior and reproductive success under high predation risk

A fundamental element of how vertebrates respond to stressors is by rapidly elevating circulating glucocorticoid hormones. Individual variation in the magnitude of the glucocorticoid stress response has been linked with reproductive success and survival. But while the adaptive value of this response is believed to stem in part from changes in the expression of hormone-mediated behaviors, it is not clear how the behavior of stronger and weaker glucocorticoid responders differs during reproduction, or during exposure to ecologically relevant stressors. Here we report that in a population of barn swallows (Hirundo rustica erythrogaster) experiencing high rates of nest predation, circulating levels of corticosterone (the primary avian glucocorticoid) during exposure to a standardized stressor predict aspects of subsequent behavior and fitness. Individuals that mounted a stronger corticosterone stress response during the early reproductive period did not differ in clutch size, but fledged fewer offspring. Parents with higher stress-induced corticosterone during the early reproductive period later provisioned their nestlings at lower rates. Additionally, in the presence of a model predator stress-induced corticosterone was positively associated with the latency to return to the nest, but only among birds that were observed to return. Model comparisons revealed that stress-induced hormones were better predictors of the behavioral and fitness effects of exposure to transient, ecologically relevant stressors than baseline corticosterone. These findings are consistent with functional links between individual variation in the hormonal and behavioral response to stressors. If such links occur, then selection on the heritable components of the corticosterone stress response could promote adaptation to novel environments or predation regimes.

Distinct responses of baseline and stress-induced corticosterone levels to genetic and environmental factors

Glucocorticoid (GC) hormones, i.e. corticosterone (CORT) in birds, support physiological homeostasis and facilitate adaptations to stressful situations. However, maintaining high GC levels are energetically costly and interfere with other physiological processes. To keep the balance of costs and benefits of GC hormones, various mechanisms act to adapt GC levels to environmental conditions on different timescales, i.e. over generations, between parents and their offspring and within the life-time of a single individual. We elucidated whether two strains (domesticated and wild) of grey partridges (Perdix perdix) differed in the developmental trajectories of baseline and stress response CORT throughout the first 80days of life. We also explored the potential of prenatal and postnatal factors, e.g. parental origin, predictable vs. unpredictable food treatments, individual and social factors to modify these trajectories. Baseline CORT was similar between strains and unaffected by perinatal food treatments. It was negatively related to body size and body condition. Conversely, the CORT stress response was not markedly affected by physiological condition. It was stronger in wild than in domesticated birds and it increased with age. Birds subjected to prenatal unpredictable food supply exhibited an accelerated development of the CORT stress response which could reflect an adaptive maternal effect. We conclude that the vital role of baseline CORT may allow little adaptive scope since changes can quickly become detrimental. In contrast, the CORT stress response may show considerable adaptive potential which might ultimately support homeostasis in a changing environment.

Corticosterone stress response shows long-term repeatability and links to personality in free-living Nazca boobies

The concept of “coping styles”, or consistently different responses to stressors, is of broad interest in behavioral ecology and biomedicine. Two critical predictions of this concept are individual consistency of neurophysiological and behavioral responses (relative to population variability) and a negative relationship between aggression/proactivity and hypothalamic–pituitary–adrenal axis reactivity. Recent studies failed to provide strong support for these predictions, especially outside of strictly controlled conditions, and long-term measures to test the first prediction are rare. Here, we demonstrate individual repeatability across 2–3years of maximum circulating corticosterone concentration [CORT] and area under the [CORT] response curve (AUCI) during a standard capture-restraint test in wild, free-living adult Nazca boobies (Sula granti). We also show that the stress response predicts the personality traits aggression and anxiety in these birds (measured in the wild); however, the strength of these results was weak. Maximum [CORT] and AUCI showed higher repeatability between years than baseline [CORT]. After controlling breeding status, sex, mass, date sampled, and their interactions, baseline [CORT] was most closely related to personality traits, followed by AUCI, and then maximum [CORT]. The direction of these relationships depended on whether the testing context was social or non-social. [CORT] parameters had little to no relationship with cross-context plasticity in personality traits. Our results generally affirm two critical predictions of coping styles, but match the emerging trend that these relationships are weak in the wild, and may depend on testing context.

Role of corticosteroid binding globulin in emotional reactivity sex differences in mice.

Sex differences exist for stress reactivity as well as for the prevalence of depression, which is more frequent in women of reproductive age and often precipitated by stressful events. In animals, the differential effect of stress on male's and female's emotional behavior has been well documented. Crosstalk between the gonadal and stress hormones, in particular between estrogens and glucocorticoids, underlie these sex differences on stress vulnerability. We have previously shown that corticosteroid binding globulin (CBG) deficiency in a mouse model (Cbg k.o.) leads, in males, to an increased despair-like behavior caused by suboptimal corticosterone stress response. Because CBG displays a sexual dimorphism and is regulated by estrogens, we have now investigated whether it plays a role in the sex differences observed for emotional reactivity in mice. By analyzing Cbg k.o. and wild-type (WT) animals of both sexes, we detected sex differences in despair-like behavior in WT mice but not in Cbg k.o. animals. We showed through ovariectomy and estradiol (E2) replacement that E2 levels explain the sex differences found in WT animals. However, the manipulation of E2 levels did not affect the emotional behavior of Cbg k.o. females. As Cbg k.o. males, Cbg k.o. females have markedly reduced corticosterone levels across the circadian cycle and also after stress. Plasma free corticosterone levels in Cbg k.o. mice measured immediately after stress were blunted in both sexes compared to WT mice. A trend for higher mean levels of ACTH in Cbg k.o. mice was found for both sexes. The turnover of a corticosterone bolus was increased in Cbg k.o. Finally, the glucocorticoid-regulated immediate early gene early growth response 1 (Egr1) showed a blunted mRNA expression in the hippocampus of Cbg k.o. mutants while mineralocorticoid and glucocorticoid receptors presented sex differences but equivalent mRNA expression between genotypes. Thus, in our experimental conditions, sex differences for despair-like behavior in WT mice are explained by estrogens levels. Also, in both sexes, the presence of CBG is required to attain optimal glucocorticoid concentrations and normal emotional reactivity, although in females this is apparent only under low E2 concentrations. These findings suggest a complex interaction of CBG and E2 on emotional reactivity in females.

Physiological and Behavioral Effects of Repeated Handling and Short-Distance Translocations on Free-Ranging Northern Pacific Rattlesnakes (Crotalus oreganus oreganus)

Abstract Translocation, a management and conservation strategy used commonly in which animals are moved from their sites of origin to other localities, has proven controversial. We examined the physiological and behavioral impacts of repeated handling and short-distance translocation on rattlesnakes, which are often translocated from areas of human use because of a perceived threat to people. Northern Pacific Rattlesnakes (Crotalus oreganus oreganus) were radiotracked for 2 months, during which time one of three treatments was imposed weekly: translocation, walk and release at that day's capture site (handling control), and undisturbed control. At both the beginning (spring) and the end (summer) of the study, blood samples were obtained before and after an acute handling stressor, and plasma concentrations of corticosterone (CORT) and testosterone (T) were determined. All rattlesnakes showed a CORT stress response, but baseline and stressed concentrations of neither hormone were affected by either translo...

Roads are associated with a blunted stress response in a North American pit viper

Whereas numerous studies have examined roads as anthropogenic stressors in birds and mammals, comparatively few studies have been undertaken on reptiles. We investigated plasma corticosterone (CORT) levels at baseline and following 30min of restraint stress in free-ranging copperhead snakes (Agkistrodon contortrix) captured within the forest interior or while in contact with public roads. There was no difference in baseline CORT levels between snakes in the forest and on roads. Copperheads responded to restraint stress by increasing plasma levels of CORT; however snakes on roads exhibited a lower CORT stress response compared to forest snakes. Additionally, among snakes captured on roads there was a negative association between road traffic and baseline CORT, stressed CORT, and the magnitude of the CORT response. Our results suggest that roads are associated with a blunted stress response in copperheads. Reduced stress responses may be indicative of acclimation, the inhibited ability to mount a stress response in the face of prolonged chronic stress, or that road environments select for individuals with lower CORT responsiveness. Either scenario could result in increased road mortality if snakes do not perceive roads as a potential threat.

Acute Thermal Stressor Increases Glucocorticoid Response but Minimizes Testosterone and Locomotor Performance in the Cane Toad (Rhinella marina)

Climatic warming is a global problem and acute thermal stressor in particular could be considered as a major stressor for wildlife. Cane toads (Rhinella marina) have expanded their range into warmer regions of Australia and they provide a suitable model species to study the sub-lethal impacts of thermal stressor on the endocrine physiology of amphibians. Presently, there is no information to show that exposure to an acute thermal stressor could initiate a physiological stress (glucocorticoid) response and secondly, the possible effects on reproductive hormones and performance. Answering these questions is important for understanding the impacts of extreme temperature on amphibians. In this study, we experimented on cane toads from Queensland, Australia by acclimating them to mildly warm temperature (25°C) and then exposing to acute temperature treatments of 30°, 35° or 40°C (hypothetical acute thermal stressors). We measured acute changes in the stress hormone corticosterone and the reproductive hormone testosterone using standard capture and handling protocol and quantified the metabolites of both hormones non-invasively using urinary enzyme-immunoassays. Furthermore, we measured performance trait (i.e. righting response score) in the control acclimated and the three treatment groups. Corticosterone stress responses increased in all toads during exposure to an acute thermal stressor. Furthermore, exposure to a thermal stressor also decreased testosterone levels in all toads. The duration of the righting response (seconds) was longer for toads that were exposed to 40°C than to 30°, 35° or 25°C. The increased corticosterone stress response with increased intensity of the acute thermal stressor suggests that the toads perceived this treatment as a stressor. Furthermore, the results also highlight a potential trade-off with performance and reproductive hormones. Ultimately, exposure acute thermal stressors due to climatic variability could impact amphibians at multiple eco-physiological levels through impacts on endocrine physiology, performance and potentially fitness traits (e.g. reproductive output).

Perinatal exposure to low-dose bisphenol A affects the neuroendocrine stress response in rats

Bisphenol A (BPA) is an estrogen-mimicking endocrine disruptor. Early-life exposures to low doses of BPA exert long-lasting effects on animals' reproductive and brain physiology. However, little is known about the effects of BPA on the stress-response system. Given the interaction of sex and stress hormones, we examined the effect of a low perinatal BPA exposure on the function of the hypothalamic-pituitary-adrenal (HPA) axis at rest and upon application of acute stress. Throughout pregnancy and lactation rats received daily 40 μg BPA/kg body weight orally via cornflakes. We studied the effect of this low but chronic exposure to BPA in the male and female offspring at puberty. BPA exposure led to abnormal adrenal histology including reduced zona reticularis especially in male offspring, hyperplasia of zona fasciculata in both sexes, and increased adrenal weight in female offspring. BPA-treated females had increased basal corticosterone and reduced hypothalamic glucocorticoid receptors (GR) levels. Stressed BPA-exposed females exhibited anxiety-like behavioral coping, a less rigorous corticosterone response, and did not downregulate GR in the hypothalamus, compared with control females. BPA-exposed males exhibited a heightened corticosterone stress response compared with females; they also displayed increased pro-opiomelanocortin mRNA levels and retained the prestress levels of pituitary corticotropin-releasing hormone-receptor 1, compared with control males. We found that perinatal chronic exposure to a low dose of BPA perturbs the basal and stress-induced activity of the HPA axis in a sexually dimorphic manner at adolescence. Exposure to BPA might contribute to increased susceptibility to stress-related disorders in later life.

Sight of a Predator Induces a Corticosterone Stress Response and Generates Fear in an Amphibian

Amphibians, like other animals, generate corticosterone or cortisol glucocorticoid responses to stimuli perceived to be threatening. It is generally assumed that the corticosterone response of animals to capture and handling reflects the corticosterone response to stimuli such as the sight of a predator that are thought to be natural stressors. Fijian ground frogs ( Platymantisvitiana ) are preyed upon by the introduced cane toads ( Rhinellamarina ), and we used ground frogs to test the hypothesis that the sight of a predator will induce a corticosterone stress response in an amphibian. Urinary corticosterone metabolite concentrations increased in male ground frogs exposed to the sight of a toad for 1, 3 or 6 h, whereas corticosterone did not change in frogs exposed to another male ground frog, a ball, or when no stimulus was present in the test compartment. The frogs exposed to a toad initially moved towards the stimulus then moved away, whereas frogs exposed to another frog moved towards the test frog and remained closer to the frog than at the start of the test. Tonic immobility (TI) was measured as an index of fearfulness immediately after the test exposure of the frogs to a stimulus. The duration of TI was longer in frogs exposed to a toad than to another frog or to a ball. The results provide novel evidence that the sight of a predator can induce a corticosterone response and lead to increased fearfulness in amphibians. In addition, they show that endemic frogs can recognise an introduced predator as a threat.

Influence of temperature on the corticosterone stress–response: An experiment in the Children’s python (Antaresia childreni)

To cope with environmental challenges, organisms have to adjust their behaviours and their physiology to the environmental conditions they face (i.e. allostasis). In vertebrates, such adjustments are often mediated through the secretion of glucocorticoids (GCs) that are well-known to activate and/or inhibit specific physiological and behavioural traits. In ectothermic species, most processes are temperature-dependent and according to previous studies, low external temperatures should be associated with low GC concentrations (both baseline and stress-induced concentrations). In this study, we experimentally tested this hypothesis by investigating the short term influence of temperature on the GC stress response in a squamate reptile, the Children’s python (Antaresia childreni). Snakes were maintained in contrasting conditions (warm and cold groups), and their corticosterone (CORT) stress response was measured (baseline and stress-induced CORT concentrations), within 48h of treatment. Contrary to our prediction, baseline and stress-induced CORT concentrations were higher in the cold versus the warm treatment. In addition, we found a strong negative relationship between CORT concentrations (baseline and stress-induced) and temperature within the cold treatment. Although it remains unclear how cold temperatures can mechanistically result in increased CORT concentrations, we suggest that, at suboptimal temperature, high CORT concentrations may help the organism to maintain an alert state.

Repeatability of Baseline Corticosterone and Acute Stress Responses to Capture, and Patterns of Reproductive Hormones in Vitellogenic and Non‐Vitellogenic Female Fijian Ground Frog (Platymantis vitiana)

The survival of animal species and individuals is largely determined by their ability to express physiological stress responses to predictable and unpredictable environmental challenges. Currently, there is no empirical evidence presenting the stress endocrine responses of female frogs during breeding between different reproductive groups. In this study, non-invasive urine sampling and standard capture and handling protocol were used to quantify baseline and short-term corticosterone stress responses in vitellogenic and non-vitellogenic female Fijian ground frog (Platymantis vitiana) during the annual breeding period. Urinary estrogen and progesterone metabolites were also quantified in the same frog urine samples. Repeated sampling of the female frogs (n = 20) on three occasions enabled repeatability (r) of reproductive and stress hormones to be quantified. All female frogs generated urinary corticosterone responses to the standard capture and handling stressor. Both baseline and short-term corticosterone responses were significantly higher in magnitude in the vitellogenic females in comparison to the non-vitellogenic female frogs. Vitellogenic females also showed significantly higher levels of urinary estrogen and progesterone metabolites in comparison to the non-vitellogenic females. Baseline urinary corticosterone, short-term corticosterone responses, urinary estrogen, and progesterone metabolites were highly repeatable for both female groups. The results highlight the importance of reproductive and stress hormones during the breeding period in female ground frogs. Future studies should determine the role of potential biological stressors (such as interactions with invasive species) that could be mediating the observed differences in stress endocrine responses of the vitellogenic and non-vitellogenic female frogs.

Repeatability of baseline corticosterone and short-term corticosterone stress responses, and their correlation with testosterone and body condition in a terrestrial breeding anuran (Platymantis vitiana)

Repeatability of physiological response variables, such as the stress hormone corticosterone, across numerous sampling occasions is an important assumption for their use as predictors of behaviour, reproduction and fitness in animals. Very few studies have actually tested this assumption in free-living animals under uncontrolled natural conditions. Non-invasive urine sampling and standard capture handling protocol have enabled the rapid quantification of baseline corticosterone and short-term corticosterone stress responses in anuran amphibians. In this study, established non-invasive methods were used to monitor physiological stress and urinary testosterone levels in male individuals of the terrestrial breeding Fijian ground frog (Platymantis vitiana). Adult male frogs (n=20) were sampled at nighttime on three repeated occasions at intervals of 14days during their annual breeding season on Viwa Island, Fiji. All frogs expressed urinary corticosterone metabolite responses to the capture and handling stressor, with some frogs showing consistently higher urinary corticosterone responses than others. Ranks of corticosterone values at 0, 4 and 8h, and the corrected rank were highly significant (r=0.75–0.99) between the three repeated sampling occasions. Statistical repeatabilities were high for baseline corticosterone (r=0.973) and for corticosterone values at 2h (r=0.862), 4h (r=0.861), 6h (r=0.820) and 8h (r=0.926), and also for the total (inclusive of baseline corticosterone values) and the corrected integrated responses (index of the acute response) [r=0.867 and r=0.870]. Urinary testosterone levels also showed high statistical repeatability (r=0.78). Furthermore, variation in baseline and short-term corticosterone stress responses was greater between individuals than within individuals. Baseline urinary corticosterone was significantly negatively correlated with the corrected integrated corticosterone response (r=−0.3, p<0.001) but non-significantly with body-condition (r=−0.04) and baseline urinary testosterone (r=−0.07). In contrast, the corrected integrated corticosterone response was positively correlated (non-significantly) with baseline urinary testosterone (r=+0.04) and body-condition (r=+0.08). Urinary testosterone levels and body-condition were significantly negatively correlated (r=−0.23, p<0.001). The results suggest that male frogs with higher levels of testosterone could have depleted energy reserve during the breeding period. The acute corticosterone responses help in replenishing energy that is needed for breeding and survival. The results also provide some support to the ‘cort-fitness’ hypothesis as highlighted by the negative correlation between baseline corticosterone and body-condition. It is most likely that the acute corticosterone response is adaptive and linked positively with reproductive fitness and survival in male anurans.

Stress responsiveness predicts individual variation in mate selectivity

Steroid hormones, including glucocorticoids, mediate a variety of behavioral and physiological processes. Circulating hormone concentrations vary substantially within populations, and although hormone titers predict reproductive success in several species, little is known about how individual variation in circulating hormone concentrations is linked with most reproductive behaviors in free-living organisms. Mate choice is an important and often costly component of reproduction that also varies substantially within populations. We examined whether energetically costly mate selection behavior in female Galápagos marine iguanas (Amblyrhynchus cristatus) was associated with individual variation in the concentrations of hormones previously shown to differ between reproductive and non-reproductive females during the breeding season (corticosterone and testosterone). Stress-induced corticosterone levels – which are suppressed in female marine iguanas during reproduction – were individually repeatable throughout the seven-week breeding period. Mate selectivity was strongly predicted by individual variation in stress-induced corticosterone: reproductive females that secreted less corticosterone in response to a standardized stressor assessed more displaying males. Neither baseline corticosterone nor testosterone predicted variation in mate selectivity. Scaled body mass was not significantly associated with mate selectivity, but females that began the breeding period in lower body condition showed a trend towards being less selective about potential mates. These results provide the first evidence that individual variation in the corticosterone stress response is associated with how selective females are in their choice of a mate, an important contributor to fitness in many species. Future research is needed to determine the functional basis of this association, and whether transient acute increases in circulating corticosterone directly mediate mate choice behaviors.

Inter-island differences in the corticosterone responses of North Island Saddlebacks (Philesturnus rufusater) in New Zealand do not suggest selective effects of translocation

North Island Saddlebacks (Philesturnus rufusater) have been the subject of a well-documented sequence of translocations to isolated islands around New Zealand. We measured corticosterone stress responses in North Island Saddlebacks derived from the ancestral population and from a series of three sequential translocations. Each translocation might impose a selective pressure on the founder populations of North Island Saddleback. This selective filter might favour a particular corticosterone response linked through an associated coping ability. Evidence of selection would be reflected in a progressive shift in the mean corticosterone response related to the number of sequential translocations. Mean corticosterone responses to a standardised stressor were highest in birds descended from a population subject to two sequential translocations. However, responses measured at localities representing the original source population and populations derived from one and three sequential translocations were not significantly different from each other. Accordingly, our data suggest that translocations do not necessarily cause directional selection on the stress responses of these vulnerable populations of birds.

Effects of repeated adolescent stress and serotonin transporter gene partial knockout in mice on behaviors and brain structures relevant to major depression

In humans, exposure to stress during development is associated with structural and functional alterations of the prefrontal cortex (PFC), amygdala (AMY), and hippocampus (HC) and their circuits of connectivity, and with an increased risk for developing major depressive disorder particularly in carriers of the short (s) variant of the serotonin transporter (5-HTT) gene-linked polymorphic region (5-HTTLPR). Although changes in these regions are found in carriers of the s allele and/or in depressed patients, evidence for a specific genotype × developmental stress effect on brain structure and function is limited. Here, we investigated the effect of repeated stress exposure during adolescence in mice with partial knockout of the 5-HTT gene (HET) vs. wildtype (WT) on early-adulthood behavioral measures and brain structure [using magnetic resonance imaging (MRI)] relevant to human major depression. Behaviorally, adolescent stress (AS) increased anxiety and decreased activity and did so to a similar degree in HET and WT. In a probabilistic reversal learning task, HET-AS mice achieved fewer reversals than did HET-No-AS mice. 5-HTT genotype and AS were without effect on corticosterone stress response. In terms of structural brain differences, AS reduced the volume of two long-range white matter tracts, the optic tract (OT) and the cerebral peduncle (CP), in WT mice specifically. In a region-of-interest analysis, AS was associated with increased HC volume and HET genotype with a decreased frontal lobe volume. In conclusion, we found that 5-HTT and AS genotype exerted long-term effects on behavior and development of brain regions relevant to human depression.

Mineralocorticoid receptor blockade during a rat’s first violent encounter inhibits its subsequent propensity for violence.

In individuals naïve to serious conflict in an unfamiliar environment, violence has long-lasting effects on subsequent aggressive behavior. This effect of the stressful experience of a first violent conflict occurs in victims as well as offenders. The authors study in the male rat as offender the role of a rapid corticosterone signal mediated by brain mineralocorticoid receptors (MR) in adjusting the threshold of aggressive responses. For this purpose, the authors have applied electrical stimulation of the brain's aggression circuit via the hypothalamic attack area or HAA. Using this paradigm, they found that in inexperienced rats, retesting of the animals on subsequent days facilitated aggression. Hypothalamic attack thresholds decreased to about 50% of their initial level. However, blocking the MR once with the mineralocorticoid antagonist spironolactone, during the very first evoked attacks, permanently prevented attack facilitation in subsequent conflicts in that same environment. The MR-mediated effect blocked by the antagonist occurred within an hour following the start of the first aggression tests only. A later MR blockade was not effective. These findings suggest that the corticosterone stress response during a very first serious conflict initializes an enhanced propensity for violent aggression through the brain MR.

Corticosterone stress response of Greater Rhea (Rhea americana) during short-term road transport

The effect of transport stress on blood corticosterone levels in captive Greater Rheas was investigated. Twelve adult individuals (7 males; 5 females) were loaded in pairs inside wooden crates and transported along a paved road for 30 min. Blood samples were taken before the individuals were introduced into the crate (baseline value) and immediately after they were unloaded (30 min after capture). To assess whether corticosterone levels were affected by the blood sampling procedure per se, another 6 (nontransport) control birds (3 males; 3 females) were also captured and sampled at the same times as their transported counterparts. Plasma corticosterone concentrations were measured using a commercially available corticosterone (125)I radio-immunoassay kit. Baseline corticosterone levels were similar in the control and transported birds (9.0 ± 1.6 and 10.4 ± 0.8 ng/mL, respectively). Transportation induced a highly significant (P < 0.001), more than 40-fold increase in the corticosterone levels (433.6 ± 35.4 ng/mL) that was about 5 times higher (P < 0.001) than in their nontransported counterparts (88.4 ± 14.8 ng/mL). The present findings suggest that Greater Rhea is a species highly sensitive to stressful manipulations. Both blood sampling and transportation induced highly significant adrenocortical responses. Considering that transportation is one of the unavoidable common practices in the management of Greater Rheas and, as shown in the present study, that it induces a significant 40-fold corticosterone stress response, efforts should focus on helping to generate management transport standards for optimization of the welfare of this ratite.

Modulation of the prolactin and the corticosterone stress responses: Do they tell the same story in a long-lived bird, the Cape petrel?

Over the last decades, the corticosterone stress response has been suggested as a major physiological tool to understand what strategy an individual might adopt in response to environmental perturbations. More recently, another hormone related to parental care – prolactin – has been suggested as a complementary tool to investigate this question. Indeed, both of these hormones are affected by stressors and are involved in parental decisions, such as deserting the nest. Because of these similarities, it remains unclear what the functional distinction between the prolactin and corticosterone stress responses is. Here, we investigated whether natural variations of the corticosterone and prolactin stress responses are functionally linked in free-living Cape petrel (Daption capense) parents. If prolactin and corticosterone mediate the same functional response to a stressor and are the proxies of the same response, we predict that corticosterone and prolactin stress responses (1) will be modulated according to the same factors; (2) will affect reproductive performances in the same way; and, (3) of course, will be correlated. Contrary to these predictions, we found that the corticosterone and prolactin stress responses were respectively modulated according to body condition and breeding status. Moreover, prolactin levels, but not corticosterone levels, were related to hatching success in this species. Finally, we did not find any significant correlation between these two stress responses under any circumstances (failed breeders, incubating or chick rearing birds) and this result was overall supported by a review of the existing literature. Therefore, these two stress responses do not seem to be tightly linked and we believe that they may provide complementary pieces of information on parental investment in birds.

Macrostress: do large‐scale ecological patterns exist in the glucocorticoid stress response of vertebrates?

SummaryDisturbances, or stressors, are pervasive in nature and profoundly influence fitness of individuals and, ultimately, species extinction risk. Glucocorticoid (GC) hormones affect multiple physiological and behavioural traits regulating the responses of vertebrates to stressors. Current understanding of what influencesGCstress response variation among species is poor. However, identifying environmental causes of species‐level variation in theGCstress response is important to understanding macroevolutionary patterns in this key physiological system and potentially offers a metric for evaluating species risk to global change.We tested whetherGCstress response of 22 reptile and 66 bird species exposed to a uniform capture stress protocol could be predicted from aGCstressor space model. This model predicts that species‐level variation inGCs results from multivariate selection from environmental gradients and species‐specific metabolic sensitivities. Thus, these multivariate selection interactions delimit the cumulative selection intensity causingGCstress response variation among vertebrates.Using phylogenetic generalized least square regression analysis, we ranked models incorporating putative variables pertaining to the corticosterone stress response (CSR) for reptiles and birds separately. The top‐ranked models explained between 14% and 33% of species variation in theCSR. Models indicated that combinations of variables including increasing body mass, net primary productivity and latitude influenced GC stress responsiveness in reptiles and birds.We conclude thatGCstress response variation among species represents, in part, evolution to species‐specific traits and large‐scale environmental variation. Further improvement to predicting species variation inGCstress response could be achieved by expanding our stressor space model to integrate microevolutionary and ecological processes.There is a pressing need to identify physiological traits that predict species responses to environmental change. Conceptually, the use of plasmaGClevels as a species‐specific marker for environmental tolerance could be plausible. However, we identify three major knowledge gaps and ensuing research areas to aid understanding of the utility of variation inGCstress response as a metric for inferring how species might respond to global change.