Prolactin-induced parental hyperphagia in ring doves: are glucocorticoids involved?

Hyperphagia is a prominent component of the parental behavior repertoire in male and female ring doves and is necessary in order for parents to successfully provision their growing young. Although previous studies implicate both prolactin and the endogenous glucocorticoid, corticosterone, in parental hyperphagia, the functional interactions between these two hormones in regulating changes in feeding activity have not been characterized. These studies examined the possibility that prolactin's orexigenic effects are mediated through the increased secretion of corticosterone. Twice-daily intracerebroventricular (icv) injection of prolactin increased plasma corticosterone concentration in non-breeding doves of both sexes, with males exhibiting more pronounced effects than females. To further test the importance of glucocorticoid signaling in prolactin-induced feeding responses, changes in food intake were investigated in icv prolactin-treated, non-breeding doves following icv infusion of the glucocorticoid receptor antagonist RU38486 or propylene glycol vehicle. No attenuation of prolactin-induced hyperphagia was observed in either sex following co-administration of RU38486 at a dose shown previously to block dexamethasone-induced feeding in doves. These findings suggest that elevated corticosterone titers in blood may contribute to the hyperphagia observed in response to prolactin, but corticosterone signaling through a mammalian-type glucocorticoid receptor is not essential.     

Effect of parental feeding activity on squab-induced crop sac growth in ring doves (Streptopelia risoria)

Effects of parental regurgitation feeding activity on crop sac development were studied in mate-separated male and female ring doves given 2 hr of daily exposure to food-deprived or recently fed squabs, for 4 consecutive days during the early posthatching period of the breeding cycle. In both sexes, food-deprived squabs stimulated more squab-directed activity, more parental regurgitation feeding activity, and greater crop sac development than recently fed young. Crop sac weights of males in both groups tended to be positively correlated with one or more parental activities. Correlations obtained in males exposed to food-deprived young further suggested that tactile stimuli associated with regurgitation behavior may promote crop sac development. In contrast to males, crop sac weights of females in both groups were not highly correlated with any type of contact-related parental activity or group of activities. These results, together with previous findings, suggest that nontactile stimuli from young played some role in mediating female crop sac weight differences in the two exposure conditions.     

Facilitation of parental behavior in ring doves by systemic or intracranial injections of prolactin

Systemic administration of ovine prolactin (PRL) has been previously reported to stimulate parental feeding behavior toward 7-day-old foster squabs by nonbreeding ring doves with previous breeding experience. The first of the present experiments reexamined this claim in experienced, nonbreeding birds given test squabs of different ages. Each visually isolated male and female dove was given twice-daily subcutaneous injections of ovine PRL or vehicle for 7 days and then tested for parental responses toward a single 1-3, 6-8, or 11- to 13-day-old foster squab. Prolactin significantly increased the incidence or frequency of parental regurgitation-feeding episodes in tests with all three squab age groups and, in addition, increased the incidence of parental feeding invitations (squab-oriented bill openings) in tests with 6- to 8-day-old squabs. A second study explored the degree to which PRL can act directly on the central nervous system to facilitate parental activity in the absence of peripheral cues generated from PRL-induced changes in other target organs, such as those associated with crop sac growth and distension. In this experiment, 6- to 8-day-old test squabs were used to determine if parental behavior is enhanced by twice-daily intracerebroventricular (ICV) injections of PRL in doses below those required to stimulate peripheral target organs. Injection schedules and behavior testing procedures were the same as those used in Experiment 1. However, half of the ICV PRL-treated and ICV vehicle-treated birds were food deprived for 16 hr before and during the test in order to control for PRL-induced hyperphagia and resulting crop sac distension, which could confound the results by generating peripheral stimuli conductive to the display of regurgitation-feeding behavior. Intracranial injections of prolactin significantly increased the incidence of feeding behavior, parental feeding invitations, and crouching or sitting in the nest in food-deprived doves but not in freely fed animals. Empty crop sac weights of freely fed and food-deprived PRL-treated birds were not increased above control values, thus indicating that ICV PRL treatment did not result in significant stimulation of peripheral target organs. These results demonstrate a facilitative action of PRL on regurgitation-feeding responses and associated parental behaviors that is not restricted to squabs of one particular age range. They also indicate that PRL is capable of acting directly on the brain to promote these activities in the absence of PRL-induced changes in the crop sac and other peripheral target organs.     

Vasoactive intestinal polypeptide-like immunoreactivity during reproduction in doves: influence of experience and number of offspring

To assess the possibility that vasoactive intestinal polypeptide (VIP) plays a role in naturally occurring changes in prolactin secretion in ring doves, we used immunohistochemical techniques to measure VIP-like immunoreactivity in the brain as a function of stage of the reproductive cycle. Differences between parental and nonparental birds in VIP profiles were detected in the ventral portion of the infundibular region. More specifically, there is an increase in cell size and staining intensity in the ventral infundibulum of breeding birds compared to simultaneously processed tissue taken from control animals. In both sexes, an increase in size of VIP-like immunoreactive cells is detectable during courtship and early incubation, anticipating the increase in plasma prolactin levels. VIP cell size remains elevated from about Incubation Day 14 to Brooding Day 14, and a steady decrease is observed during the remaining posthatching period, as squab begin to feed independently. Compared to parents rearing one squab, those with two young have a prolonged interval of increased infundibular VIP immunoreactivity. Furthermore, doves with no previous experience of a breeding cycle exhibit prolonged VIP-like immunoreactivity compared to experienced parents, paralleling previously described differences between these groups in parental behavior.     

Effects of intracranial prolactin administration on maintenance of incubation readiness, ingestive behavior, and gonadal condition in ring doves

Systemic administration of prolactin (PRL) reportedly maintains readiness to sit on eggs, increases body weights and liver weights, and reduces gonadal and reproductive tract tissue weights in male and female ring doves. To determine if these effects are mediated by PRL-induced changes in the central nervous system, ring doves were isolated from their mates, nests, and eggs on Day 4 of incubation and given twice-daily intracerebroventricular injections of PRL (1 microgram, 0.031 IU) or saline vehicle (2 microliter) during the next 10 days. Food and water consumption were monitored daily during treatment and tests for incubation behavior and observation of organ weight changes were conducted at the end of the period. The incidence of incubation behavior in the PRL-treated group was not significantly different from that in the control group and was significantly lower than that reported in a previous study following subcutaneous PRL injections. However, gonadal and reproductive tract tissue weights were markedly reduced and food and water consumption were significantly elevated in PRL-treated birds. Body weights and liver weights were also increased following PRL administration. In contrast, crop sac weight did not increase in the PRL treated birds, thereby indicating that the effects of icv prolactin treatment were not mediated by increased plasma PRL levels and peripheral target organ stimulation. These results suggest that PRL can act directly on the brain to inhibit gonadotropin release and to stimulate ingestive behavior but central effects of PRL alone are not sufficient to maintain incubation readiness.     

Seasonal changes in testosterone and corticosterone levels in four social classes of a desert dwelling sociable rodent

Animals have to adjust their physiology to seasonal changes, in response to variation in food availability, social tactics and reproduction. I compared basal corticosterone and testosterone levels in free ranging striped mouse from a desert habitat, comparing between the sexes, breeding and philopatric non-breeding individuals, and between the breeding and the non-breeding season. I expected differences between breeders and non-breeders and between seasons with high and low food availability. Basal serum corticosterone was measured from 132 different individuals and serum testosterone from 176 different individuals of free living striped mice. Corticosterone and testosterone levels were independent of age, body weight and not influenced by carrying a transmitter. The levels of corticosterone and testosterone declined by approximately 50% from the breeding to the non-breeding season in breeding females as well as non-breeding males and females. In contrast, breeding males showed much lower corticosterone levels during the breeding season than all other classes, and were the only class that showed an increase of corticosterone from the breeding to the non-breeding season. As a result, breeding males had similar corticosterone levels as other social classes during the non-breeding season. During the breeding season, breeding males had much higher testosterone levels than other classes, which decreased significantly from the breeding to the non-breeding season. My results support the prediction that corticosterone decreases during periods of low food abundance. Variation in the pattern of hormonal secretion in striped mice might assist them to cope with seasonal changes in energy demand in a desert habitat.     

The breeding season duration hypothesis: acute handling stress and total plasma concentrations of corticosterone and androgens in male and female striped plateau lizards (Sceloporus virgatus)

Acute glucocorticoid elevations can be adaptations to short-term stressors. The breeding season hypothesis predicts reduced glucocorticoid responsiveness to acute stressors in populations or species with short breeding seasons. The striped plateau lizard (Sceloporus virgatus) has a short breeding season in Arizona. We measured plasma corticosterone and total androgen levels (dihydrotestosterone and testosterone) following one of the four stress-handling treatments (0, 10, 60, or 180 min). In both sexes, longer handling stress yielded higher corticosterone; females had higher corticosterone than males at all time points. Androgens did not vary with handling duration, in either sex. Combining treatments, plasma androgens correlated positively with corticosterone (CORT) in females but not in males; plasma CORT and body mass residuals were negatively correlated in both sexes, suggesting lizards in poor body condition and/or not investing heavily in reproduction (follicle mass) have higher acute corticosterone. Total plasma androgens and body mass residuals were positively associated in males, but showed no association in females. The maximal CORT elevation after handling stress in this single-clutching species was of comparable magnitude to responses in related multi-clutching lizard species with longer breeding seasons. Using data from studies of multiple populations of three Sceloporus species, we found no relationship between the relative magnitude of the CORT increase and either latitude or elevation, two variables in the literature correlated with duration of the breeding season, and only weak relationships with geographic elevation and actual (not relative) stress-elevated CORT values in this multi-population comparison.     

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

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

The Roles of Stimuli from Young, Previous Breeding Experience, and Prolactin in Regulating Parental Behavior in Ring Doves (Streptopelia risoria)

In addition to stimulating crop "milk" formation in ring doves, prolactin (PRL) may promote the parental regurgitation behavior that transfers this "milk" to the young at the time of hatching. Although earlier studies suggest that previous breeding experience is an important modulator of PRL-induced parental regurgitation behavior in ring doves, the ways in which experience, hormones, and stimuli from young interact to promote parental behavior have not been well characterized in this species. In the first study, untreated, nonbreeding female doves with and without previous breeding experience were given 10 daily parental behavior tests (2 h/day) with a hungry 5- to 10-day-old foster squab. Experienced females exhibited a higher incidence of regurgitation behavior, defensive behavior, and crouching or sitting in the nest than did inexperienced females. In a second study, nonbreeding females were given 10 daily tests for parental behavior while they received sc injections of ovine PRL or vehicle. Prolactin reduced squab-directed aggression and increased the incidence of regurgitation feeding behavior of foster squabs in both experienced and inexperienced females. However, the average number of regurgitation feeding acts displayed by those PRL-treated females that showed the behavior was over eight times higher in experienced females than in inexperienced females. Previous experience also enhanced the stimulatory effects of PRL on defensive behavior and crouching or sitting in the nest. The parental behavior exhibited by nonbreeding, PRL-treated experienced females was qualitatively and quantitatively similar to that observed in normally breeding females during a single test with their own hungry 5- to 10-day-old squabs. These findings indicate that PRL and previous breeding experience both enhance the parental responsiveness of nonbreeding female doves and that under optimal hormonal, experiential, and squab exposure conditions, nonbreeding doves exhibit levels of parental activity that rival those of normally breeding parents.     

Gonadal and hormonal characteristics of captive adult mourning doves during the nesting cycle

Paired mourning doves ($\text{Zenaida}$$\text{macroura}$) of a captive breeding colony were selected and sacrificed on specific day intervals postlaying and posthatching during 1976–1977. Blood plasma samples were analyzed for androgens, estradiol, and progestins using radioimmunoassays. Testes, ovary and oviduct weights, spermatozoan numbers, diameter of the three largest follicles in the ovary, the presence or absence of eggs in the reproductive tract, and crop gland characteristics also were recorded. Differences (P<0.05) existed between nonbreeding and breeding pairs in testes weights, spermatozoan numbers, and ovary and oviduct weights. No distinct cyclic pattern was observed in gonadal variables among breeding males and females during the nesting cycle, although trends were observed. No differences (P > 0.05) in androgen and progestin concentrations in males, or estradiol and progestin concentrations in females, between nonbreeding and breeding pairs, or among breeding pairs during different stages of the nesting cycle, were observed. Analysis of reproductive variables by stage of crop gland activity tended to indicate an inverse relationship between gonadal and crop gland activity in both male and female doves during the nesting cycle.     

Size of ornament is negatively correlated with baseline corticosterone in males of a socially monogamous colonial seabird

The Goymann–Wingfield model predicts that glucocorticoid levels in social animals reflect the costs of acquiring and maintaining social status. The crested auklet is one of the few avian colonial species where a mutual ornament in males and females is used in both sexual and aggressive displays. Previous studies of the crested auklet support the notion that the crest ornament is a badge of status in this species. Here, we examined the relationship between the crest ornament size and the adrenocortical function in breeding crested auklets. Crest length was negatively correlated with corticosterone at baseline in males, but not in females. Baseline corticosterone in females (but not in males) was negatively correlated with body condition index. Although male and female crested auklets are monomorphic in their ornamental traits, our results suggest that the socially mediated physiological costs associated with status signaling may differ between the sexes.     

Corticosterone manipulations alter morph-specific nestling provisioning behavior in male white-throated sparrows, Zonotrichia albicollis

In the polymorphic white-throated sparrow (Zonotrichia albicollis), tan-striped males provision nestlings at higher rates than do white-striped males. In a previous study, we found that tan-striped males had lower baseline corticosterone levels than white-striped males during the nestling stage. To determine if this variation in corticosterone influences morph-specific differences in nestling provisioning behavior, we used intraperitoneal osmotic pumps to increase baseline corticosterone levels in tan-striped males (TS CORT) and administer RU486, a glucocorticoid receptor antagonist, in white-striped males (WS RU486). These manipulations essentially reversed morph-specific nestling provisioning behavior in males. TS CORT males fed nestlings at lower rates than TS controls (vehicle-only implant), and at similar rates to WS controls (vehicle-only implant), while WS RU486 males fed nestlings at higher rates than WS controls, and at similar rates to TS controls. These results demonstrate that (1) increases in baseline corticosterone (i.e., below concentrations associated with the adrenocortical response to stress) can directly or indirectly inhibit nestling provisioning behavior, and (2) corticosterone influences morph-specific variation in parental behavior in male white-throated sparrows. This study contributes to the growing evidence that modulating baseline CORT mediates parental care and self-maintenance activities in birds, and thus may serve as a mechanism for balancing current reproductive success with survival.     

Corticosterone manipulations alter morph-specific nestling provisioning behavior in male white-throated sparrows,Zonotrichia albicollis

In the polymorphic white-throated sparrow (Zonotrichia albicollis), tan-striped males provision nestlings at higher rates than do white-striped males. In a previous study, we found that tan-striped males had lower baseline corticosterone levels than white-striped males during the nestling stage. To determine if this variation in corticosterone influences morph-specific differences in nestling provisioning behavior, we used intraperitoneal osmotic pumps to increase baseline corticosterone levels in tan-striped males (TS CORT) and administer RU486, a glucocorticoid receptor antagonist, in white-striped males (WS RU486). These manipulations essentially reversed morph-specific nestling provisioning behavior in males. TS CORT males fed nestlings at lower rates than TS controls (vehicle-only implant), and at similar rates to WS controls (vehicle-only implant), while WS RU486 males fed nestlings at higher rates than WS controls, and at similar rates to TS controls. These results demonstrate that (1) increases in baseline corticosterone (i.e., below concentrations associated with the adrenocortical response to stress) can directly or indirectly inhibit nestling provisioning behavior, and (2) corticosterone influences morph-specific variation in parental behavior in male white-throated sparrows. This study contributes to the growing evidence that modulating baseline CORT mediates parental care and self-maintenance activities in birds, and thus may serve as a mechanism for balancing current reproductive success with survival.     

The transition from sitting on eggs to sitting on young in ring doves, Streptopelia risoria: squab-egg preferences during the normal cycle.

Ring doves of both sexes sit on young squabs after hatching in much the same manner as they sit on eggs before hatching, but this study demonstrates that the preferred stimulus varies with the state of the animal. A simultaneous, squab-egg choice test was given on days 1, 4, 10, and 13 of incubation and on the day following hatching in normal reproductive cycles of experienced and naïve male and female ring doves. Naïve doves were more likely than experienced doves to choose eggs throughout the cycle (P<0·005) and, overall, eggs were more likely to be chosen during early incubation and squabs posthatching (P<0·005).     

Failure of glucocorticoid feedback during breeding in the male red-tailed phascogale Phascogale calura (Marsupialia: Dasyuridae)

An investigation was made into the factors which lead to an elevation in plasma free cortisol concentration during the last weeks of life of males in natural populations of the red-tailed phascogale Phascogale calura. The dexamethasone suppression-test was employed to examine the glucocorticoid feedback control of plasma cortisol both before and during the breeding season. In both sexes ACTH alone or in combination with dexamethasone caused an elevation in the plasma concentration of cortisol, corticosterone and free cortisol. Dexamethasone administration in both males and females resulted in significant decreases in the plasma concentration of each of the glucocorticoid groups both before and during the first week of the breeding season (June and early July), however during the last week of breeding (late July) dexamethasone decreased the plasma glucocorticoid concentration of females but not of males. Administration of ACTH caused a significant elevation in the plasma cortisol concentration in all groups. However, the magnitude of this response diminished with time in both sexes. Dexamethasone treatment resulted in a decrease in the plasma testosterone concentration in males before and early in the breeding season however toward the end of breeding this effect was abolished. It is apparent that towards the end of the breeding season and during the last week of life of the males, glucocorticoid feedback control of ACTH is almost abolished. These changes, which occur only in the males late in the breeding season and near the time of their disappearance from the population, are consistent with a condition known as end organ resistance to steroid hormones.     

The discovery of hatching and transition to feeding young by male Mountain Bluebirds

In many bird species, only females incubate the eggs, but both sexes feed nestlings. The means by which males of such species discover hatching and transition to feeding their offspring remains almost completely unexplored. Of particular interest are species with nests whose contents are concealed from view. During June and early July 2015 in the Bighorn Mountains of Wyoming, we used continuous video‐recording of nests of cavity‐nesting Mountain Bluebirds (Sialia currucoides) to document the transition to feeding young by males. We saw no evidence that females used distinct vocal or visual displays to signal hatching to males. Observing mates carrying eggshells away from, or food into, nest boxes did not appear to trigger provisioning by males. Rather, at all 24 nests observed, males did not begin feeding until they had come to nest boxes and presumably sensed the presence of hatchings directly. Individual males varied, however, in both the manner in which they inspected nest contents and the number of times they did so before starting to feed young. Although most males fully entered nest boxes where they could see, touch, hear, and possibly smell hatchlings (or eggshell parts), other males may have detected hatchlings only by sound or possibly smell while perched at a nest‐box entrance. Based on past studies of mice and doves, we suggest that, for provisioning behavior to begin, some kind of direct sensation of offspring may be necessary to activate relevant neurons in the medial preoptic area of the hypothalamus of males, an area of the brain important in parental care. Additional research is necessary to test this, and to examine the effects of factors such as hormone levels and breeding experience on the means and rapidity by which males discover hatching and transition to nestling provisioning.     

Stress hormones and sociality: integrating social and environmental stressors

In cooperatively breeding species, reproductive decisions and breeding roles may be influenced by environmental (food resources) or social factors (reproductive suppression of subordinates by dominants). Studies of glucocorticoid stress hormones in cooperatively breeding species suggest that breeding roles and hormone levels are related to the relative costs of dominance and subordination, which are driven primarily by social interactions. Few studies, however, have considered how environmental factors affect glucocorticoid levels and breeding roles in cooperative breeders, even though environmental stressors modulate seasonal glucocorticoid release and often influence breeding roles. I examined baseline and stress-induced levels of the glucocorticoid corticosterone (CORT) across 4 years in the plural breeding superb starling, Lamprotornis superbus, to determine whether (i) environmental factors (namely rainfall) directly influence breeding roles or (ii) environmental factors influence social interactions, which in turn drive breeding roles. Chronic baseline and maximal stress-induced CORT changed significantly across years as a function of pre-breeding rainfall, but dominant and subordinate individuals responded differently. Pre-breeding rainfall was also correlated directly with breeding roles. The results are most consistent with the hypothesis that environmental conditions influenced the relative costs of dominance and subordination, which in turn affected the degree and intensity of social interactions and ultimately reproductive decisions and breeding roles.     

Behavioural and hormonal stress responses during chick rearing do not predict brood desertion by female in a small Arctic seabird

We examined behavioural and hormonal stress responses in a small seabird (little auk, Alle alle), which exhibits a transition from biparental to male-only care towards the end of the nesting period, in order to understand the mechanisms underlying this parental strategy. We hypothesized that the male staying with the chick should be less sensitive to stressors. As such the male might offer the offspring more efficient protection during the fledging period than the female. We tested this hypothesis by observing male and female behaviour in a neophobia test. We also measured the birds' baseline and stress-induced levels of corticosterone and prolactin using the standardized capture-and-restraint protocol. Both sexes respond rapidly to foreign objects, delaying the entry time to the nest with food, consuming the food load, and/or temporarily abandoning feeding. However, we did not find any differences between the sexes in the frequency of each behaviour or in the time of the first reaction to the experimental treatment. Level of both corticosterone and prolactin increased after the experimental treatment. However, we did not find sex differences in baseline and stress-induced hormone levels. The results indicate that the males are as much sensitive to the stress situation as the females. Thus, the pattern of male and female behavioural and hormonal responses to stress does not predict their behaviour at the final breeding stage.     

Blood metabolite levels in normal and handicapped pied flycatchers rearing broods of different sizes

We measured levels of select metabolites (glucose, triglycerides, free fatty acids, glycerol, uric acid) and corticosterone in the blood plasma of adult pied flycatchers Ficedula hypoleuca while they were rearing broods whose sizes were modified experimentally. We also made it more difficult than normal for some pairs of birds to forage by removing certain wing and tail feathers (handicapping them). Both procedures have been shown previously to change parental workload. We did this in order to determine if the birds alter their use of nutrients in response to differences in their workload. Metabolite levels were not influenced by handicapping or brood size. However, the concentration of free fatty acids in the plasma of females and of triglycerides in the plasma of males was directly related to the frequency with which the adults fed their nestlings. These findings suggest that the two sexes have different ways of coping with the work associated with rearing the brood: females apparently undergo brief daily fasts while feeding their chicks, whereas males take more time to feed themselves while providing food for their young, and spend more time doing so as their workload increases. The flycatchers exhibited high concentrations of uric acid and corticosterone in the blood plasma; corticosterone and glycerol were positively correlated in females; and corticosterone and triglyceride levels were negatively correlated in males; all of which suggest that gluconeogenesis provides some of the energy required for their parental activities.     

Structural coloration signals condition, parental investment, and circulating hormone levels in Eastern bluebirds (Sialia sialis)

Many of the brilliant plumage coloration displays of birds function as signals to conspecifics. One species in which the function of plumage ornaments has been assessed is the Eastern bluebird (Sialia sialis). Studies of a population breeding in Alabama (USA) have established that plumage ornaments signal quality, parental investment, and competitive ability in both sexes. Here we tested the additional hypotheses that (1) Eastern bluebird plumage ornamentation signals nest defense behavior in heterospecific competitive interactions and (2) individual variation in plumage ornamentation reflects underlying differences in circulating hormone levels. We also tested the potential for plumage ornaments to signal individual quality and parental investment in a population breeding in Oklahoma (USA). We found that Eastern bluebirds with more ornamented plumage are in better condition, initiate breeding earlier in the season, produce larger clutches, have higher circulating levels of the stress hormone corticosterone, and more ornamented males have lower circulating androgen levels. Plumage coloration was not related to nest defense behavior. Thus, plumage ornamentation may be used by both sexes to assess the physiological condition and parental investment of prospective mates. Experimental manipulations of circulating hormone levels during molt are needed to define the role of hormones in plumage ornamentation.