Stress response patterns of plasma corticosterone, prolactin, and growth hormone in the rat, following handling or exposure to novel environment.

Corticosterone, prolactin, and growth hormone responses to 5 s of handling or 3 min of novel environment were compared in rats at crest and trough of the diurnal adrenal rhythm 0, 5, 15, 30, and 60 min after stimulation. All hormones responded to stimulation, corticosterone and prolactin with a dramatic rise, and growth hormone with a precipitous fall. Resting corticosterone levels evidenced the expected diurnal variation, and prolactin but not growth hormone also showed a baseline diurnal variation of small magnitude at the times studied. Growth hormone response characteristics were unaffected by time of day or type of stimulation. Both corticosterone and prolactin response profiles differed at both times of day and following both types of stimulation. Corticosterone and prolactin levels were highly correlated and each was negatively correlated with growth hormone levels. This study confirms that hormone responses to stress are complex and depend not only on the stimulus but the context of stimulation.     

Regulation of protein breakdown and adrenocortical response to stress in birds during migratory flight

During long-term fasting at rest, protein utilization is maintained at low levels until it increases at a threshold adiposity. This study examines 1) whether such a shift in energy substrate use also occurs during endurance exercise while fasting, 2) the role of corticosterone, and 3) the adrenocortical response to an acute stressor. Ten species of migrating birds caught after an endurance flight over at least 500 km were examined. Plasma uric acid and corticosterone levels were low in birds with fat stores >5% of body mass and high in birds with smaller fat stores. Corticosterone levels were very high in birds with no visible fat stores and emaciated breast muscles. Corticosterone levels increased with handling time only in birds with large fat stores. These findings suggest that 1) migrating birds with appreciable fat stores are not stressed by endurance flight, 2) a metabolic shift (increased protein breakdown), regulated by an endocrine shift (medium corticosterone levels), occurs at a threshold adiposity, as observed in birds at rest, 3) adrenocortical response to an acute stressor is inhibited after this shift, and 4) an adrenocortical response typical for an emergency situation (high corticosterone levels) is only reached when muscle protein is dangerously low.     

Removal of a nest-mate elicits an age-dependent increase in plasma corticosterone of nestling Black-legged Kittiwakes

ABSTRACT.   Plasma corticosterone concentrations in birds often increase about 3 min after exposure to a stressor such as capture and handling. When measuring adrenal responsiveness of nestlings in broods with more than one nestling, standardizing capture protocols to equalize the stressor among nestlings if they simultaneously perceive the presence and activity of a researcher as a stressor is logistically difficult. The objective of our study was to determine if nestling Black-legged Kittiwakes ( Rissa tridactyla ) in broods of two mount a corticosterone response when the first nestling is removed from the nest or, alternatively, if each initiates a corticosterone response only at the time it is handled. We obtained blood samples from one nestling within 3 min of initial disturbance of the nest, and then removed and sampled its sibling 10 min later. For younger nestlings, we found no difference in corticosterone levels between those sampled at 3 min and their sibling sampled at 10 min. In contrast, older nestlings sampled at 10 min after initial nest disturbance had elevated corticosterone levels compared to those sampled within 3 min. In addition, nestlings sampled within 3 min of capture had elevated corticosterone when exposed to protracted periods of investigator disturbance at nearby nests. Our results suggest that it is necessary to treat initial disturbance of the nest as the onset of the stress response for all nestlings in multi-nestling broods when handling older nestlings or nestlings of unknown age. In addition, for species that nest in dense colonies, the presence of an investigator at one nest may be a stressor for nestlings in adjacent nests.     

Modulation of prolactin but not corticosterone responses to stress in relation to parental effort in a long-lived bird

We tested the hypothesis that parental effort modulates the magnitude of corticosterone and prolactin responses to stress in a long-lived bird, the Black-legged kittiwake (Rissa tridactyla). To do so, we compared corticosterone and prolactin responses to capture/restraint stress between chick-rearing birds and failed breeders (no parental effort). We predicted that (1) the increase in plasma corticosterone levels in response to stress should be lower in chick-rearing birds, (2) the decrease in plasma prolactin levels in response to stress should be lower in chick-rearing birds, and (3) as both sexes care for the chick, there should be no sex difference in the hormonal response to stress. Baseline plasma corticosterone and prolactin levels were higher in chick-rearing birds and were not influenced by body condition. Failed breeders were in better condition than chick-rearing individuals. Corticosterone response to stress was unaffected by parental effort as both chick-rearing and failed birds exhibited a robust corticosterone increase. Prolactin response to stress was however clearly influenced by parental effort: chick-rearing birds showed a modest 9% prolactin decrease whereas in failed birds prolactin concentrations fell by 41%. Body condition did not influence hormonal responses to stress. When facing stressful condition, breeding kittiwakes attenuate their prolactin response to stress while enhancing their secretion of corticosterone. Increasing corticosterone secretion triggers foraging efforts and diminishes nest attendance whereas an attenuation of prolactin response to stress maintains parental behavior. We suggest that this hormonal mechanism facilitates a flexible time-budget that has been interpreted as a buffer against environmental variability.     

Repeated thermal stressor causes chronic elevation of baseline corticosterone and suppresses the physiological endocrine sensitivity to acute stressor in the cane toad (Rhinella marina)

Extreme environmental temperature could impact the physiology and ecology of animals. The stress endocrine axis provides necessary physiological stress response to acute (day–day) stressors. Presently, there are no empirical evidences showing that exposure to extreme thermal stressor could cause chronic stress in amphibians. This could also modulate the physiological endocrine sensitivity to acute stressors and have serious implications for stress coping in amphibians, particularly those living in fragmented and disease prone environments. We addressed this important question using the cane toad (Rhinella marina) model from its introduced range in Queensland, Australia. We quantified their physiological endocrine sensitivity to a standard acute (capture and handling) stressor after exposing the cane toads to thermal shock at 35 °C for 30 min daily for 34 days. Corticosterone (CORT) responses to the capture and handling protocol were measured on three sampling intervals (days 14, 24, and 34) to determine whether the physiological endocrine sensitivity was maintained or modulated over-time. Two control groups (C1 for baseline CORT measurement only and C2 acute handled only) and two temperature treatment groups (T1 received daily thermal shock up to day 14 only and a recovery phase of 20 days and T2 received thermal shock daily for 34 days). Results showed that baseline CORT levels remained high on day 14 (combined effect of capture, captivity and thermal stress) for both T1 and T2. Furthermore, baseline CORT levels decreased for T1 once the thermal shock was removed after day 14 and returned to baseline by day 29. On the contrary, baseline CORT levels kept on increasing for T2 over the 34 days of daily thermal shocks. Furthermore, the magnitudes of the acute CORT responses or physiological endocrine sensitivity were consistently high for both C1 and T1. However, acute CORT responses for T2 toads were dramatically reduced between days 24 and 34. These novel findings suggest that repeated exposure to extreme thermal stressor could cause chronic stress and consequently suppress the physiological endocrine sensitivity to acute stressors (e.g. pathogenic diseases) in amphibians.     

Daily and photoperiod variations of basal and stress-induced corticosterone concentrations in house sparrows (Passer domesticus)

Corticosterone concentrations were measured in captive house sparrows (Passer domesticus) and found to vary both daily and with different photoperiods. Basal corticosterone was highest during the dark hours of the daily cycle and lowest during the light hours. This trend remained constant when the birds were held on short-day and long-day light cycles, and while the birds were undergoing a prebasic molt. At all times, corticosterone concentrations significantly increased in response to the stress of handling and restraint. Stress-induced corticosterone concentrations, however, only reflected a daily rhythm when the birds were held on short-days. Furthermore, even though mean basal corticosterone concentrations were equivalent over the short-day, long-day, and molt, total corticosterone output in response to stress was lower in molting birds, especially at night. Therefore, these data indicate that captive house sparrows modulate corticosterone in daily cycles that change in response to photoperiod.     

Corticosterone responses to capture and restraint in emperor and Adelie penguins in Antarctica

Birds respond to capture, handling and restraint with increased secretion of corticosterone, a glucocorticoid hormone that helps birds adjust to stressful situations. Hoods are reported to calm birds, but possible effects of hoods on corticosterone responses have not been reported for any bird. Corticosterone responses to restraint in Adelie penguins held by their legs with their head covered by a hood were markedly lower than responses of penguins restrained in a mesh bag inside a cardboard box (corticosterone at 30 min 15.69+/-1.72 cf. 28.32+/-2.75 ng/ml). The birds restrained by the two methods were sampled at the same location but in different years, so the differences in corticosterone responses cannot unequivocally be ascribed to an effect of hoods to reduce corticosterone responses. Corticosterone responses have been measured in some penguins, but not in the largest, the emperor penguin (Aptenodytes forsteri). The relationship between body mass and corticosterone responses to capture and restraint in penguins was examined in emperor penguins captured on sea ice in McMurdo Sound and Adelie penguins (Pygoscelis adeliae) captured at Cape Bird, Ross Island, Antarctica. Total integrated corticosterone responses were higher in the emperor than the Adelie penguins, but corrected integrated corticosterone responses, which represent the increase in corticosterone from initial concentrations and hence the corticosterone response to restraint, were the same. The results for the emperor and Adelie penguins, together with data from other penguin species, suggest that there is no relationship between the size of corticosterone responses and body mass in penguins.     

Comparison of plasma ACTH and corticosterone levels after embryo bursectomy

Plasma levels of both adrenocorticotropic hormone (ACTH) and corticosterone (B) were determined in embryos (day 15 of incubation), chicks (day 3 after hatch) and young chickens (8 weeks). Experimental animals were bursectomized at 80 hr of incubation, i.e., before any anlage of the bursa of Fabricius could develop. Bursectomized (BFX) animals were compared to sham-operated controls (T), in basal, resting condition and 7 (ACTH) or 14 min (B) after ether stress was delivered for 30 sec. Basal B and ACTH levels seemed not to be significantly modified in BFX embryos, chicks and chickens. Hypophysial and adrenocortical response to stress appeared more precociously in BFX embryos (day 15 of incubation) than in intact ones (day 19). The non stress-responsive period that was observed for one week after hatch of T birds did not appear in 3-day-old BFX chicks whose both B and ACTH stress-induced levels were as high as in intact adults. In contrast, adrenocortical and pituitary corticotropic responses to stress were markedly impaired (by 50%) in adult BFX chickens as compared to intact controls.     

Experimental enhancement of corticosterone levels positively affects subsequent male survival

Corticosterone is an important hormone of the stress response that regulates physiological processes and modifies animal behavior. While it positively acts on locomotor activity, it may negatively affect reproduction and social activity. This suggests that corticosterone may promote behaviors that increase survival at the cost of reproduction. In this study, we experimentally investigate the link between corticosterone levels and survival in adult common lizards (Lacerta vivipara) by comparing corticosterone-treated with placebo-treated lizards. We experimentally show that corticosterone enhances energy expenditure, daily activity, food intake, and it modifies the behavioral time budget. Enhanced appetite of corticosterone-treated individuals compensated for increased energy expenditure and corticosterone-treated males showed increased survival. This suggests that corticosterone may promote behaviors that reduce stress and it shows that corticosterone per se does not reduce but directly or indirectly increases longer-term survival. This suggests that the production of corticosterone as a response to a stressor may be an adaptive mechanism that even controls survival.     

Speed of exploration and risk-taking behavior are linked to corticosterone titres in zebra finches

The existence of consistent individual differences in behavioral strategies ("personalities" or coping styles) has been reported in several animal species. Recent work in great tits has shown that such traits are heritable and exhibit significant genetic variation. Free-living birds respond to environmental stresses by up-regulating corticosterone production. Behavior during mild stress can occur in accordance to two types of coping styles, i.e. active and passive. Using artificially selected lines of zebra finches that vary in the amount of corticosterone produced in response to a manual restraint stressor we ran three "personality" experiments. We show that birds in the different corticosterone lines differ in their exploratory and risk-taking behaviors. There was an increase in exploratory behavior as corticosterone titre increased but only in the low corticosterone line. Birds in high corticosterone line showed greater risk-taking behavior than birds in the other lines. Thus, in general, higher levels of circulating corticosterone following a mild stress result in greater exploratory behavior and greater risk taking. This study shows that lines of animals selected for endocrine hormonal responses differ in their "coping" styles or "personalities".     

Changes in masculine sexual behavior, corticosterone and testosterone in response to acute and chronic stress in male rats

Chronic exposure to stressors increases HPA axis activity and concomitantly reduces HPG axis activity. This antagonistic relationship between both these axes has been proposed to underlie the inhibition of reproductive function due to stress. Sexual behavior in males may be the most vulnerable aspect of male reproduction to acute and chronic stress and it has been suggested that alterations in sexual behavior during stress are due to the antagonistic relationship between testosterone and corticosteroids. However, only in a few studies has a correlation between the levels of testosterone and corticosterone, and sexual behavior been made. In this study, we evaluated the effects of different stressors, applied both acute and chronically, on masculine sexual behavior and whether or not these effects on sexual behavior are accompanied by changes in plasma levels of corticosterone and testosterone. Additionally, we evaluated the effect of testosterone treatment on the effects of stress on sexual behavior. Sexually experienced male rats were exposed to one of the following stressors: immobilization (IMB), electric foot shocks (EFS) or immersion in cold water (ICW). Sexual behavior and plasma levels of testosterone and corticosterone were assessed on days 1, 5, 10, 15, and 20 of stress. In a second experiment, males were castrated, treated with 3 different doses of testosterone propionate (TP) and exposed to ICW for 20 consecutive days. Sexual behavior was assessed on days 1, 5, 10, 15, and 20 and steroids were evaluated on day 20. Parameters of masculine sexual behavior were modified depending on the characteristics of each stressor. Mount, intromission and ejaculation latencies increased significantly, the number of mounts increased, and ejaculations decreased significantly in males exposed to EFS and to ICW but not in males exposed to IMB. Associated with these effects, testosterone decreased in the EFS and ICW groups on days 1, 15, and 20. However, corticosterone increased only in males exposed to ICW. In castrated males, TP treatment failed to block the effects of stress by ICW on sexual behavior and corticosterone. These results indicate that the effects of stress on sexual behavior depend on the characteristics of each stressor, and these effects, as well as the decrease in testosterone are not necessarily associated with the increase in corticosterone. The fact that testosterone treatment did not prevent the effects of stress on sexual behavior suggests that other mediators could be involved in the alterations of sexual behavior caused by stress.     

Trapping initiates stress response in breeding and non-breeding house sparrows Passer domesticus: implications for using unmonitored traps in field studies

In response to a variety of unpredictable conditions, birds secrete the steroid hormone corticosterone, which has numerous effects on physiology and behavior. A standardized protocol of handling and restraint has been demonstrated to elicit a robust corticosterone response in many species of birds. In contrast, comparatively little is known about the effects of capture technique on corticosterone secretion in wild birds. Setting up multiple live traps checked at regular intervals allows field researchers to capture many birds in a short period of time. However, one potential drawback of this technique is that birds may spend unknown lengths of time in traps prior to sampling. Many birds appear to remain calm and/or feed during this period, potentially leading researchers to assume that corticosterone secretion is unaffected by trapping. We assessed the impact of being left in traps for up to 30 minutes on baseline corticosterone and subsequent corticosterone responses to restraint in non‐breeding and breeding house sparrows Passer domesticus. Traps were baited with seed, and birds were either removed immediately after entry (controls), or left in the trap undisturbed for 15 or 30 min. Upon removal, birds were subjected to a standardized handling/restraint protocol in which blood samples were collected within 3 min, and again at regular intervals for 60 min. Analysis of blood samples revealed that both non‐breeding and breeding sparrows that were held in the traps had significantly higher baseline corticosterone than controls, and showed no further increase in corticosterone secretion in response to handling. However, corticosterone responses to trapping differed seasonally. Our study indicates that although birds did not exhibit prolonged escape behavior while trapped, entry into a walk‐in trap initiated a robust stress response. Taken together with data from a previous study, our data suggest that ornithologists should consider species‐specific and stage‐specific effects of trapping on physiology when designing field studies.     

Heritability of corticosterone response and changes in life history traits during selection in the zebra finch

Vertebrates respond to environmental stressors through the neuro-endocrine stress response, which involves the production of glucocorticoids. We have selected independent, duplicate divergent lines of zebra finches for high, low and control corticosterone responses to a mild stressor. This experiment has shown that over the first four generations, the high lines have demonstrated a significant realized heritability of about 20%. However, the low lines have apparently not changed significantly from controls. This asymmetry in response is potentially because of the fact that all birds appear to be showing increased adaptation to the environment in which they are housed, with significant declines in corticosterone response in control lines as well as low lines. Despite the existence of two- to threefold difference in mean corticosterone titre between high and low lines, there were no observed differences in testosterone titre in adult male birds from the different groups. In addition, there were no consistent, significant differences between the lines in any of the life history variables measured--number of eggs laid per clutch, number of clutches or broods produced per pair, number of fledglings produced per breeding attempt, nor in any of egg, nestling and fledgling mortality. These results highlight the fact that the mechanisms that underlie variation in the avian physiological system can be modified to respond to differences between environments through selection. This adds an additional level of flexibility to the avian physiological system, which will allow it to respond to environmental circumstances.     

The effect of althesin on plasma corticosterone levels

Althesin in doses which produced anesthesia (4 and 6 ml kg-1, i.p.) produced biphasic changes in plasma corticosterone levels. Plasma corticosterone showed an increase (p less than 0.05) due to the stress of injection but returned to basal levels by 30 min. Subsequent to the anesthetic effect (approximately 30 min) corticosterone levels increased markedly (p less than 0.01). Althesin's effectiveness showed time of day effects, i.e., Althesin was more effective in the A.M. Rats given 6 ml kg-1 Althesin showed graded plasma corticosterone responses to stresses of varying intensity. Blood withdrawal and surgical stress evoked significant increases in plasma corticosterone but a 2-min holding stress had no effect on plasma corticosterone levels. Instrumented rats receiving supplemental injections (i.p.) presented patterns of plasma corticosterone which were different from those receiving supplemental infusions (i.a.). Whereas plasma corticosterone levels of rats receiving the continuous infusion of Althesin remained relatively constant, corticosterone levels of those which received supplemental injections tended to increase. Collectively, these data suggest that Althesins usefulness as an experimental anesthetic is limited to those studies which are not compromised by stress-induced pituitary-adrenal activity.     

Exposure to chronic stress downregulates corticosterone responses to acute stressors

We used captive European starlings (Sturnus vulgaris) to test whether corticosterone responses differed in birds held under normal laboratory conditions or conditions of chronic stress. Surprisingly, both basal corticosterone concentrations and corticosterone responses to acute stress were significantly reduced when birds were chronically stressed. To determine the mechanism underlying this reduced response, animals under both conditions were injected with lactated Ringer's solution (control), adrenocorticotropin (ACTH), arginine vasotocin (AVT), or dexamethasone (DEX). ACTH increased corticosterone concentrations above stress-induced levels in both cases, although maximum responses were lower in chronically stressed birds. AVT did not augment the corticosterone response under nonchronically stressed conditions, but it did under chronically stressed conditions. DEX reduced maximal corticosterone concentrations in both cases. Neither ovine nor rat corticotropin-releasing factor (CRF) altered normal stress responses. These data indicate that changes in responsiveness of the hypothalamic-pituitary-adrenal axis to ACTH and AVT serve to downregulate corticosterone responses during chronic stress. Furthermore, these data lead to the following hypothesis: ACTH output from the pituitary limits maximum corticosterone concentrations under normal conditions, but reduced AVT release from the hypothalamus regulates lower corticosterone concentrations under chronic stress conditions.     

Sex and migratory strategy influence corticosterone levels in winter-grown feathers,with positive breeding effects in a migratory pelagic seabird

To overcome unpredictable stressful transitory events, animals trigger an allostatic response involving the hypothalamic–pituitary–adrenal cortex. This hormonal response, which involves the release of glucocorticoids which in turn mediate between the main physiological mechanisms that regulate the energetic demands and resource allocation trade-off with behavioural responses to environmental perturbations and may ultimately lead to variation in fitness. We have used the Cory’s shearwater Calonectris borealis, a sexually dimorphic pelagic seabird with a partial migratory strategy, as a model bird species to analyse a number of traits related to the stress response. We investigated whether the activation of a stressful response, mediated by corticosterone, during the wintering period (1) correlated with the previous breeding success, (2) was affected by the migratory behaviour of male birds and (3) had consequences in the fitness of the birds. Corticosterone levels in feathers grown overwinter were analysed in 61 adult birds during three consecutive migratory periods (2009–2012) and in 14 immature birds in the wintering period 2010–2011. Moreover, the levels of corticosterone were analysed in experimental birds which were freed from their reproductive duties and compared with control birds which raised fledglings to the end of the breeding period. The results show that the levels of corticosterone were sex dependent, differed between years and were affected by the migratory strategy performed by the birds. The activation of the stressful response over the wintering period generated residual carry-over effects that positively affected the reproductive output in the subsequent breeding stage, a phenomenon previously undescribed in a long-lived pelagic seabird. Our study provides evidence that the analysis of corticosterone from feathers is a useful tool to evaluate carry-over effects in birds far away from breeding sites, opening new possibilities for future studies in this field.     

Psychological stressors as a model of maternal adversity: diurnal modulation of corticosterone responses and changes in maternal behavior

Maternal adversity is associated with long-lasting consequences on cognitive development, behavior and physiological responses in rat offspring. Few studies have examined whether repeated maternal stress produces repeated activation of the hypothalamus-pituitary-adrenal (HPA) axis in mothers and whether it modifies maternal behavior. Here, we tested a novel model of perinatal stress using repeated exposure to "purely" psychological stressors throughout the gestation and lactation periods in rats. We first tested the diurnal influences of repeated 1-h strobe light exposure on maternal corticosterone secretion. Despite the hyporesponsiveness to stress documented in late pregnant and lactating mothers, we observed an enhanced response to strobe light in the afternoon compared to the morning in stressed mothers during lactation. Next, dams were exposed to 24-h forced foraging followed by 10-h wet bedding during the diurnal peak of corticosterone secretion. Although no corticosterone responses to forced foraging and wet bedding were observed, the combination of both stressors had a significant effect on maternal behavior. Mother-pup interactions were significantly altered during the first 8 days of lactation. Taken together, these findings suggest that lactating mothers maintain responsiveness to specific and repeated psychological stressors, in particular at the time of the diurnal peak in corticosterone secretion. Depending on the stressor applied, either neuroendocrine activation or changes in maternal behavior might be important determinants of the long-term consequences in the offspring. The combination of forced foraging, wet bedding and strobe light might represent a novel model of mild maternal adversity using "purely" psychological stressors.     

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.     

Development of stress response in nestling pied flycatchers

Birds respond to unpredictable events by secreting corticosterone, which induces various responses to cope with stressful situations. However, the evidence is still elusive whether altricial nestlings perceive and respond to external stressors. We investigated the development of adrenocortical stress response to handling-related stressor in nestlings of a small passerine bird, the pied flycatcher (Ficedula hypoleuca). Nestlings were held in isolation from their parents during the experiment to ensure that they indeed respond to handling, not to parental alarm calls. We found that both 9- and 13-day-old nestlings were able to elicit hormonal stress response. Although baseline as well as stress-induced corticosterone levels rose slightly with age, the magnitude of difference between the control and stress-induced levels remained similar in both age groups. However, comparison with adults showed that the stress response of nestlings prior to fledging was still incomplete and significantly lower than in adults. Overall, our results indicate that altricial nestlings do respond to acute stressors, but on the contrary to previous predictions the development of corticosterone stress response during growth period is not gradual and varies remarkably between different passerine species.     

Acute corticosterone elevation enhances antipredator behaviors in male tree lizard morphs

In response to stressful events, most vertebrates rapidly elevate plasma glucocorticoid levels. Corticosterone release stimulates physiological and behavioral responses that can promote survival while suppressing behaviors that are not crucial to immediate survival. Corticosterone also has preparatory effects for subsequent stressors. Using male tree lizards (Urosaurus ornatus), we tested our prediction that elevated corticosterone is important for mediating and enhancing antipredator behaviors. Male tree lizards express developmentally fixed polymorphisms that are mediated by early organizational actions of steroid hormones, and thus we also tested the hypothesis that morph-specific differences in antipredator behaviors of adults are independent of circulating corticosterone levels. Plasma corticosterone levels were elevated exogenously for 12-16 h using non-invasive dermal patches, and we then compared the behavioral responses of these corticosterone-patched males to control-patched males during a simulated encounter with a caged predator (collared lizard, Crotaphytus nebrius) in outdoor enclosures. Elevating corticosterone did not alter the antipredator behavioral repertoire of each male morph, but did enhance their responses during the predator encounter: all corticosterone-patched males responded more quickly, hid longer, and displayed more toward the predator than control-patched males. With the corticosterone patch, the non-territorial and wary orange morph was still behaviorally the most wary morph, responding more quickly and hiding longer than either the bolder orange-blue or mottled morphs. Smaller males were generally warier than larger males, regardless of the endocrine treatment or color morph type. In sum, elevated circulating corticosterone enhances antipredator responses for all male tree lizard morphs, without altering morph-specific or size-specific differences in their behavioral responses.