Comparison of adrenocortical responses to acute stress in lowland and highland Eurasian tree sparrows (Passer montanus): similar patterns during the breeding, but different during the prebasic molt

Previous studies indicate most free-living avian species in both extreme and temperate environments seasonally modulate the adrenocortical responses to acute stress, and those breeding in harsh environments always express reduced adrenocortical responses, which may allow them to obtain maximal reproductive success. However, recent investigations showing a human commensal species, house sparrows (Passer domesticus), expressed similar corticosterone (CORT) responses in both benign and harsh environments. In this study, focusing on another human commensal species, Eurasian tree sparrows (P. montanus), we examined the adrenocortical response to acute stress in lowland populations, among the early and late breeding, the prebasic molt, and the wintering stages, and compared them with previously published data from populations on the Tibetan Plateau. Our results show: (1) similar to highland Eurasian tree sparrows, lowland populations show no differences in baseline CORT levels among life history stages, and the stress-induced CORT (maximal CORT, total and corrected integrated CORT) levels are lower during the early breeding and the prebasic molt stages than those in the late breeding and the wintering stages; (2) highland Eurasian tree sparrows show stronger adrenocortical responses during the prebasic molt stage than lowland populations, whereas there are no differences between the early and the breeding stages (except for maximal CORT). Our results suggest that Eurasian tree sparrows from both harsh and benign environments have similar patterns of adrenocortical responses in the breeding stage, whereas they are different in the prebasic molt stage. In highland birds, the increased maximal CORT levels during the late breeding and the small increases in adrenocortical responses during the prebasic molt are interesting but remain unexplained.     

Corticosterone treatment has no effect on reproductive hormones or aggressive behavior in free-living male tree sparrows, Spizella arborea

We examined the effect of corticosterone on plasma levels of reproductive hormones (testosterone, dihydrotestosterone, and luteinizing hormone) and territorial defense behavior in male tree sparrows, Spizella arborea. Birds receiving Silastic implants filled with corticosterone (B) had significantly higher plasma levels of B than control birds, which received empty implants, and exhibited pectoral muscle wastage and a decrease in body mass. We evaluated the hormonal and agonistic responses of the two implanted groups of birds using a simulated territorial intrusion (STI) 2 to 4 days after they were implanted. Corticosterone-treated and control birds did not differ in their circulating levels of reproductive hormones or in their behavioral responses to STI (latency to respond to intrusion, number of songs, and closest approach to a decoy and tape recording). Unlike previous studies of north temperate passerines, high physiological levels of exogenous B had no effect either on circulating levels of reproductive hormones or on territorial behaviors associated with breeding. Nonetheless, untreated tree sparrows do mount a robust adrenocortical response, having a two- to fourfold increase in plasma B levels during a 1-h period of capture. Thus, adrenocortical responsiveness is maintained in these birds, but elevated levels of glucocorticoids do not suppress reproductive hormones or associated behaviors. We believe that this hormonal and behavioral refractoriness to glucocorticoids-or uncoupling of the stress response from the reproductive axis-may be advantageous for species having extreme temporal constraints on their breeding schedules.     

Rapid behavioral response to corticosterone varies with photoperiod and dose

The magnitude of the adrenocortical response to stress can be modulated by a variety of environmental and physiological factors, such as daylength and body condition. To determine the consequences of this modulation for the organism, one also needs to investigate behavioral sensitivity to glucocorticoids. We examined the behavioral response of Gambel's white-crowned sparrows (Zonotrichia leucophrys gambelii) to elevated glucocorticoids. Using a behavioral assay in which a rapid and transient dose of corticosterone (CORT: the avian glucocorticoid) rapidly increases perch hopping, we first investigated the photoperiodic regulation of this behavioral response. Intermediate levels of CORT ( approximately 24 ng/ml), which increased activity in sparrows exposed to a long-day (breeding) photoperiod, had no behavioral effect in sparrows exposed to a short-day (winter) photoperiod. Hence, the neural mechanisms which regulate the behavioral response to CORT appear to be less sensitive under a winter photoperiod. Using the same behavioral assay, we also measured a dose-response curve for CORT's effects on activity in sparrows exposed to the long-day photoperiod. Intermediate levels (24 and 40 ng/ml) increased activity to threefold background levels, whereas high physiological levels (65 and 97 ng/ml) had no effect. Given that the behavioral response does not increase linearly with CORT, we can no longer assume that modulation of the adrenocortical response to stress will result in linear changes in behavior.     

Interspecific variation in avian blood parasites and haematology associated with urbanization in a desert habitat

Many avian species are negatively impacted by urbanization, but other species survive and prosper in urbanized areas. One factor potentially contributing to the success of some species in urban areas is the reduced presence of predators or parasite vectors in urban compared to rural areas. In addition, urban areas may provide increased food and water resources, which can enhance immune capacity to resist infection and the ability to eliminate parasites. We determined patterns of blood parasitism, body condition, and immune cell profiles in urban and rural populations of five adult male songbird species that vary in their relative abundance within urban areas. Urban birds generally exhibited less blood parasitism than rural birds. This difference was particularly evident for the urban-adaptable Abert's towhee Pipilo aberti . In contrast, no difference in haemoparasitism was seen between urban and rural populations of the curve-billed thrasher Toxostoma curvirostre , a less-urban adaptable species. In two closely related species, the curve-billed thrasher and the northern mockingbird Mimus polyglottos , urban birds had a higher leukocyte count and a higher heterophil to lymphocyte ratio, which is often associated with chronic stress or current infection, than rural birds. Urban northern mockingbirds were in better condition than rural counterparts, but no habitat-related differences in condition were detected for other species. Parasitic infection was correlated with body condition in only one species, the canyon towhee Pipilo fuscus . Parasitic infection in most species was correlated with changes in leukocyte abundance and profile. The findings suggest that interspecific differences in parasitic infection cannot be attributed entirely to differences in vector abundance or body condition. Interactions between immune function, parasite infection risk, and resource availability may contribute to determining the relative ability of certain species to adapt to cities.     

Variation in adrenocortical stress physiology and condition metrics within a heterogeneous urban environment in the song sparrow Melospiza melodia

In urban habitats, organisms face unique fitness challenges including disturbance from human activity and noise. One physiological mechanism that may be plastically or evolutionarily modified to ameliorate deleterious effects of anthropogenic disturbance is the adrenocortical stress response. Individuals in urban environments may display smaller stress responses, which may prevent pathologies associated with consistent elevation of stress hormones, and may also show differences in baseline corticosterone (CORT, the primary avian stress hormone), due to altered energetic demands or chronic stress. We examined whether stress physiology and condition metrics in male song sparrows Melospiza melodia vary as a function of discrete differences in anthropogenic disturbance level (activity centers and refuges) or with continuous variation in an urbanization score and noise environment. Males breeding in activity centers displayed lower maximal (acute) CORT levels than activity refuge males, and acute CORT also tended to negatively correlate with urbanization score. Baseline CORT did not differ between habitat types, and activity center males also showed no evidence of changes in body mass, hematocrit, or antioxidant capacity. Further, activity center males had higher quality feathers (indicative of higher condition at molt) than activity refuge males. We found no indication that the noise environment altered stress physiology or condition in song sparrows. Overall, results suggest that song sparrows are an urban adapter species, which are not detrimentally affected by unique selective pressures encountered in the urban environment.     

Environmental and seasonal adaptations of the adrenocortical and gonadal responses to capture stress in two populations of the male garter snake, Thamnophis sirtalis

Stress and reproduction are generally thought to work in opposition to one another. This is often manifested as reciprocal relationships between glucocorticoid stress hormones and sex steroid hormones. However, seasonal differences in how animals respond to stressors have been described in extreme environments. We tested the hypothesis that garter snakes, Thamnophis sirtalis, with limited reproductive opportunities will suppress their hormonal stress response during the breeding season relative to conspecifics with an extended breeding season. The red-sided garter snake, T.s. parietalis, of Manitoba, Canada, has a brief breeding season during which males displayed no change in either plasma levels of testosterone or corticosterone, which were both elevated above basal levels, in response to capture stress. During the summer, capture stress resulted in increased plasma corticosterone and decreased testosterone. During the fall, when mating can also occur, males exhibited a significant decrease in testosterone but no increase in corticosterone in response to capture stress. The red-spotted garter snake, T.s. concinnus, of western Oregon, has an extended breeding season during which males displayed a stress response of increased plasma corticosterone and decreased testosterone levels. The corticosterone response to capture stress was similar during the spring, summer, and fall. In contrast, the testosterone response was suppressed during the summer and fall when gametogenesis was occurring. These data suggest that male garter snakes, in both populations, seasonally adapt their stress response but for different reasons and by potentially different mechanisms. J. Exp. Zool. 289:99-108, 2001.     

Stress,reproduction, and adrenocortical modulation in amphibians and reptiles

While the hypothalamo-pituitary-adrenocortical (HPA) response to stress appears to be conserved in vertebrates, the manner in which it is activated and its actions vary. We examine two trends in the stress biology literature that have been addressed in amphibian and reptilian species: (1). variable interactions among stress, corticosterone, and reproduction and (2). adrenocortical modulation. In the first topic we examine context-dependent interactions among stress, corticosterone, and reproduction. An increasing number of studies report positive associations between reproduction and corticosterone that contradict the generalization that stress inhibits reproduction. Moderately elevated levels of stress hormones appear to facilitate reproduction by mobilizing energy stores. In contrast, pronounced activation of the HPA axis and extremely elevated levels of stress hormones appear to inhibit reproduction. Much of these contrasting effects of stress and reproduction can be explained by expanding the Energetics-Hormone Vocalization Model, proposed for anuran calling behavior, to other taxa. In the second topic, a number of amphibians and reptiles modulate their HPA stress response. Adrenocortical modulation can occur at multiple levels and due to a variety of factors. However, we have little information as to the physiological basis for the variability. We suggest that several ecologically based ideas, such as variability in the length of the breeding season and lifetime reproductive opportunities, can be used to explain the utility of adrenocortical modulation in these taxa.     

Coping with the extremes: stress physiology varies between winter and summer in breeding opportunists

Seasonal changes in stress steroid hormone secretions are thought to reflect investment in self-maintenance versus reproduction. The capricious conditions hypothesis (CCH) posits that reduced corticosterone (CORT) secretion during stress coincident with parental phases of breeding is necessary in harsh environments because a full response would otherwise trigger repeated nest abandonments. To test this hypothesis, we measured seasonal changes in stress physiology in free-living red crossbills (Loxia curvirostra), an opportunistically breeding songbird that regularly breeds in summer and winter. This species allows unique comparisons of breeding physiology under very different seasonal environmental conditions within locations. We found strong support for the CCH: red crossbills showed reduced CORT secretion only when in high reproductive condition in the winter, when compared with summer breeders and winter non-breeders. These data demonstrate that behavioural status and local environmental conditions interact to affect mechanisms underlying investment trade-offs, presumably in a way that maximizes lifetime reproductive success.     

Stress responses in tropical sparrows: comparing tropical and temperate Zonotrichia

Seasonal modulation of the adrenocortical response appears to be ubiquitous in mid- to high-latitude vertebrates but has not been investigated in tropical vertebrates. Previous studies demonstrate that temperate passerines show seasonality in corticosterone secretion and corticosteroid binding globulin capacities. We examined seasonal and sex differences in the stress response in an equatorial population of Zonotrichia capensis, the only Zonotrichia that breeds in the tropics, and compared the results with those of northern Zonotrichia. Seasonal differences in tropical Zonotrichia would presumably be independent of photoperiod and thus directly related to such activities as reproduction and feather molt. In addition, we investigated the possible role of binding globulin as a sex steroid binding globulin, as suggested for temperate passerines. Similar to northern congeners, Z. capensis show seasonal modulation in total corticosterone and binding globulin capacity with higher levels during breeding than molt. However, unlike many temperate passerines, there are no sex differences in corticosterone secretion or binding globulin capacity. Furthermore, the seasonal differences in total corticosterone diminish when the free levels are calculated. The contrast between equatorial and northern congeners indicates factors such as breeding environment and life-history strategy may play important roles in shaping stress response in these species.     

Arctic spring: hormone-behavior interactions in a severe environment

Arctic breeding birds arrive on their nesting grounds in spring when weather conditions may still be extreme (low temperature, snow). The brief Arctic summer requires that they begin breeding as early as possible to take advantage of the ephemeral abundance of food to feed young. Failure to adjust to the local phenology results in drastically reduced reproductive success. Hormone-behavior adaptations may have evolved that maximize survival and reproductive success in the Arctic. It has been shown that the interrelationship of testosterone and territorial aggression, as birds arrive on the Arctic breeding grounds, varies according to species and locality. In some, territoriality is extremely brief following which birds become apparently refractory to the effects of testosterone. Others are territorial throughout the breeding season, but the dependence of these behaviors upon activation by testosterone is lost. Extensive data also indicate that Arctic birds modulate the adrenocortical response to acute stress. Secretion of corticosterone in response to a standardized capture stress protocol, used to mimic acute stress as a function of local environmental conditions, varies with the stage in the breeding cycle. Arctic breeding birds modulate the sensitivity of the adrenocortical response to acute stress at both the population and individual levels. These modulations are thought to be adaptations to allow the onset of territorial behavior and breeding in the face of potentially stressful conditions. Behavioral and physiological responses to corticosterone treatment are also diminished. A combination of these two hormone-behavior interrelationships can form important components of the proximate mechanisms by which birds, and other vertebrates, breed successfully in a severe and often capricious environment.     

Baseline and stress-induced levels of corticosterone during different life cycle substages in a shorebird on the high arctic breeding grounds

After a migratory flight of several thousand kilometers to their high arctic breeding grounds, red knots (Calidris canutus islandica, Scolopacidae) showed high baseline concentrations of plasma corticosterone (58 ng/mL). Such high baseline corticosterone levels may be conditional for the right behavioral and metabolic adjustments to environmental and social stresses that shorebirds experience on arrival in an unpredictable tundra breeding environment. Despite the high baseline levels of corticosterone, red knots still showed a marked stress response during the postarrival period, with corticosterone concentrations increasing significantly during a 60-min period of confinement. Baseline levels of corticosterone declined as the breeding season progressed. Red knots with brood patches, that is, birds that had completed egg laying and commenced incubation, had a reduced adrenocortical response to the stress of confinement compared with red knots with no, or with half-developed, brood patches. This is consistent with the idea that birds breeding in extreme environments with short breeding seasons may exhibit a decreased adrenocortical response to stressful events to prevent high corticosterone concentrations from inducing interruptions of reproductive behavior.     

Integrating stress physiology,environmental change,and behavior in free-living sparrows

As weather deteriorates, breeding animals have a diverse array of options to ensure survival. Because of their mobility, birds can easily abandon territories to seek out benign conditions away from the breeding site. The timing of abandonment, however, may have repercussions for territory size, mate quality, reproductive success, and survival. There is a large body of evidence indicating that the adrenocortical response to stress plays a role in mediating the onset and maintenance of this behavioral switch. Here we develop a model describing the interactions of weather, food availability, body condition, and stress physiology in initiating departure from the breeding site. We tested the model using a population of white-crowned sparrows breeding at high elevation in the Sierra Nevada, where severe weather at the beginning of the breeding season often induces temporary abandonment of breeding territories and facultative altitudinal migration to lower elevation refugia. The data show that (1). during inclement weather, exogenous corticosterone delays return to the breeding site after territory abandonment; (2). during good weather, exogenous corticosterone alone does not induce territory abandonment, but does increase activity range around the breeding site; and (3). the magnitude of the corticosteroid response to stress is inversely related to body condition of the sparrow.     

Modulation of the adrenocortical response to acute stress with respect to brood value, reproductive success and survival in the Eurasian hoopoe

Reproducing parents face the difficult challenge of trading-off investment in current reproduction against presumed future survival and reproduction. Glucocorticoids are supposed to mediate this trade-off because the adrenocortical response to stress disrupts normal reproductive behaviour in favour of self-maintenance and own survival. According to the brood-value hypothesis, individuals with a low survival probability until the next reproductive season have to invest in current reproduction, a process driven by a down-regulation of their adrenocortical response. If the adrenocortical response to stress effectively mediates the trade-off between current reproduction versus future survival and reproduction, we expect a negative relationship with reproductive success and a positive correlation of the adrenocortical stress response with survival. We studied the relationship between corticosterone secretion in parents and their current brood value, reproductive success and survival in a short-lived multi-brooded bird, the Eurasian hoopoe Upupa epops. The adrenocortical response to acute handling stress was correlated with the brood value within the individual (first and second broods of the year) and between individuals. Birds breeding late in the season mounted a lower total corticosterone response to acute stress than birds breeding earlier, while females showed lower levels than males. We observed a negative relationship between the adrenocortical stress response and rearing success or fledging success in females, as predicted by the brood-value hypothesis. However, we could not evidence a clear link between the adrenocortical stress response and survival. Future research testing the brood-value hypothesis and trade-offs between current reproduction and future survival should also measure free corticosterone and carefully differentiate between cross-sectional (i.e. between-individual) and individual-based experimental studies.     

Changes in pituitary and adrenal sensitivities allow the snow bunting (Plectrophenax nivalis), an Arctic-breeding song bird, to modulate corticosterone release seasonally

Several free-living avian species have recently been shown to seasonally modulate corticosterone release in response to capture and restraint. We examined possible mechanisms underlying seasonal adrenocortical modulation in snow buntings (Plectrophenax nivalis), a species that breeds and molts (the energetically costly replacement of feathers) in the Alaskan Arctic. Snow buntings dramatically reduced baseline and maximal corticosterone titers during molt compared to the breeding season. This effect is not explained by changes in either corticosterone binding protein capacity or the overall condition of the bird (assessed by weight and fat storage). Although the adrenal's capacity to secrete corticosterone is reduced during molt, adrenal insensitivity does not fully explain reduced maximal output since exogenous adrenocorticotropic-hormone enhanced corticosterone release during both seasons. In contrast, no exogenous adrenocorticotropic hormone releasing factor (corticotrophin-releasing factor, arginine vasotocin or mesotocin) enhanced corticosterone secretion during molt. This suggests that the pituitary's endogenous adrenocorticotropic secretion was maximal in response to capture and handling, making the pituitary an important site regulating corticosterone levels. Taken together, these results indicate that seasonal modulation of corticosterone release in this species is controlled at both the adrenal and pituitary glands. Accepted: 16 February 1998     

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.     

Non-breeding feather concentrations of testosterone, corticosterone and cortisol are associated with subsequent survival in wild house sparrows

Potential mechanistic mediators of Darwinian fitness, such as stress hormones or sex hormones, have been the focus of many studies. An inverse relationship between fitness and stress or sex hormone concentrations has been widely assumed, although empirical evidence is scarce. Feathers gradually accumulate hormones during their growth and provide a novel way to measure hormone concentrations integrated over time. Using liquid chromatography-tandem mass spectrometry, we measured testosterone, corticosterone and cortisol in the feathers of house sparrows (Passer domesticus) in a wild population which is the subject of a long-term study. Although corticosterone is considered the dominant avian glucocorticoid, we unambiguously identified cortisol in feathers. In addition, we found that feathers grown during the post-nuptial moult in autumn contained testosterone, corticosterone and cortisol levels that were significantly higher in birds that subsequently died over the following winter than in birds that survived. Thus, feather steroids are candidate prospective biomarkers to predict the future survival of individuals in the wild.     

Adrenocortical stress responses influence an invasive vertebrate's fitness in an extreme environment

Continued range expansion into physiologically challenging environments requires invasive species to maintain adaptive phenotypic performance. The adrenocortical stress response, governed in part by glucocorticoid hormones, influences physiological and behavioural responses of vertebrates to environmental stressors. However, any adaptive role of this response in invasive populations that are expanding into extreme environments is currently unclear. We experimentally manipulated the adrenocortical stress response of invasive cane toads (Rhinella marina) to investigate its effect on phenotypic performance and fitness at the species'' range front in the Tanami Desert, Australia. Here, toads are vulnerable to overheating and dehydration during the annual hot–dry season and display elevated plasma corticosterone levels indicative of severe environmental stress. By comparing unmanipulated control toads with toads whose adrenocortical stress response was manipulated to increase acute physiological stress responsiveness, we found that control toads had significantly reduced daily evaporative water loss and higher survival relative to the experimental animals. The adrenocortical stress response hence appears essential in facilitating complex phenotypic performance and setting fitness trajectories of individuals from invasive species during range expansion.     

Effects of captivity and body condition on plasma corticosterone, locomotor behavior, and plasma metabolites in curve-billed thrashers

The acute stress response involves the secretion of catabolic glucocorticoids, such as corticosterone (CORT) in birds, that mobilize intrinsic energy stores primarily through a gluconeogenic pathway involving fat breakdown, thus linking body condition and stress. We measured changes in CORT and gluconeogenic metabolites (triglycerides, free glycerols, glucose) during handling stress in curve-billed thrashers Toxostoma curvirostre from two habitats (urban vs. desert) that may differ in food abundance in the wild, in captivity, and in response to both food restriction and subsequent recovery. Urban thrashers were heavier and secreted more CORT than desert birds in the field, but differences did not persist in captivity. Decreased access to food resulted in decreased body mass and a diminished ability to elevate plasma CORT in response to handling stress. However, the opposite effect was observed as these birds recovered from food restriction. Plasma levels of glucose and triglycerides did not change with stress. Food restriction also increased locomotor activity, which likely further exacerbated energy loss. These observations suggest that body condition and stress differences between urban and desert birds may be related to differences in their relative energetic states, possibly due to food availability. Body condition may affect the extent to which an individual can elevate CORT and use free glycerol as energy during acute stress.     

Female song sparrow, Melospiza melodia, response to simulated conspecific and heterospecific intrusion across three seasons

To investigate female responses to territorial intrusion I presented female song sparrows with either a simulated female song sparrow intrusion or a simulated spotted towhee, Pipilo maculatus, intrusion as a control during either the prebreeding, breeding or postmoult seasons. Aggressive and nonaggressive behaviours and vocalizations were compared between intrusion types and across seasons. Principle components analysis suggested that female responses fell into three categories: (1) responses directed towards the intruder, mostly aggressive; (2) responses directed towards the mate; and (3) lack of response to the intruder. In every season, females responded more aggressively to simulated female song sparrow intrusion than simulated towhee intrusion. Responses directed towards female song sparrow intruders dropped across the three seasons and were significantly higher in the prebreeding season than in the breeding or postmoult seasons. Copyright 2000 The Association for the Study of Animal Behaviour.     

Influences of weather on reproductive function in female Song sparrows, Melospiza melodia

Prolonged and severe periods of inclement weather can retard gonadal development in early spring or disrupt the breeding cycle in summer. Previous investigations on the endocrine responses of free-living avian species to storms have been restricted almost entirely to males. This communication presents the endocrine responses of free-living female Song sparrows, Melospiza melodia, during gonadal recrudescence after a severe winter, and during storms in early spring and summer. As in males, gonadal recrudescence was delayed by the severe winter weather, and this effect was exacerbated by an aseasonal snow storm in early spring. Unlike males, female Song sparrows did appear to be stressed by this snow storm, since body mass and fat deposit decreased, and plasma levels of corticosterone increased. A second storm in early summer did not, however, prove to be stressful to females, although males were apparently stressed by the same storm. At the time the early summer storm occurred, young had been fledged and were being fed mostly by the male. Females begin preparing for a second clutch of eggs and do not feed fledglings to the same extent as males. Thus it is possible that males are stressed by storms at this time, because they are feeding several young as well as themselves during a period of reduced availability of food, whereas females are feeding themselves only. These data suggest that, not only do the endocrine responses to inclement weather vary in relation to stage in the reproductive cycle when storms strike, but also that there are sex differences in those responses.