Territoriality and testosterone in an equatorial population of rufous-collared sparrows, Zonotrichia capensis

Territorial aggression, displayed by male vertebrates in a reproductive context, is generally thought to be mediated by testosterone. The challenge hypothesis predicts that in socially monogamous species, territorial challenges should induce an increase in plasma testosterone concentrations, which will enhance aggressive behaviour and territory defence. This hypothesis is based on northern latitude birds and needs to be tested in tropical birds before it can be universally accepted. We tested the challenge hypothesis in an equatorial population of rufous-collared sparrows in Papallacta, Ecuador. This population shows an extended breeding period during which males aggressively guard territories. During the early breeding season, males were challenged with conspecific or heterospecific simulated territorial intrusions (STIs) lasting 10 min. Conspecific-challenged males responded more aggressively than heterospecific-challenged males. However, there was no increase in plasma testosterone in response to the conspecific STI. During the breeding season, males were challenged with conspecific STIs lasting 0, 10 or 30 min. Males behaved aggressively regardless of STI duration, and did not differ in plasma testosterone or luteinizing hormone concentrations. During the breeding season, males were implanted with testosterone-filled or empty silastic tubes and subsequently challenged with a conspecific STI. Testosterone implants significantly raised plasma testosterone concentrations, but testosterone-implanted males were not more aggressive than blank-implanted controls. Combined, these findings suggest that testosterone concentrations above breeding baseline are not related to territorial aggression in this population and therefore do not support the challenge hypothesis.     

European robins (Erithacus rubecula) lack an increase in testosterone during simulated territorial intrusions

The challenge hypothesis (Wingfield et al. in Am Nat 136:829–846, 1990) predicts that circulating testosterone increases when socially monogamous male birds are challenged during breeding. Although the challenge hypothesis has been confirmed in large-scale interspecific comparisons of seasonal hormone profiles, experimental tests of the challenge hypothesis are still uncommon and the results equivocal. We tested one of the predictions of the challenge hypothesis by investigating the behavioural and hormonal response of free-living European robins during simulated territorial intrusions (STIs) in the breeding season. We conducted STIs by placing a stuffed decoy in a territory and playing robin song. After the behaviour of the focal male had been recorded for at least 10 min, it was captured and a blood sample was taken immediately. Controls were caught within 10 min of the first response of the territory owner. Hormone concentrations were measured by radio-immunoassay. Although previous studies have shown that testosterone has an impact on aggression, European robins do not respond to STIs by increasing circulating levels of testosterone.     

Mediation of a corticosterone-induced reproductive conflict

Current research in birds suggests that a conflict should exist during reproduction for the role of the glucocorticoid corticosterone (CORT). While elevated levels have been correlated with the increased energetic demand of raising offspring, elevated CORT levels have traditionally been implicated in reproductive abandonment. We examined the relationship between CORT and nest desertion in breeding wild female European starlings (Sturnus vulgaris) incorporating analyses of both total circulating levels and 'free', unbound CORT through analysis of corticosteroid-binding globulin (CBG). Free baseline CORT levels of nest-abandoning birds were significantly higher than nonabandoning birds within each stage, with chick-rearing birds exhibiting the highest free baseline CORT levels, while concurrently remaining the most resistant stage to nest desertion. Elevated free baseline CORT levels in chick-rearing birds were not due to increased total CORT secretion, but rather to a decrease in CBG levels. Overall, our results suggest that CORT and CBG interact to play a role in mediating the increased energetic demand of offspring, while minimizing the chances of nest desertion, thereby alleviating any potential behavioral conflict for CORT during reproduction. Furthermore, these results demonstrate that the traditional view of the role of CORT during reproduction is much more complex than previously appreciated. Together with mounting evidence, we suggest that elevated corticosteroid levels are an inherent and necessary part of reproduction in nonmammalian tetrapods.     

Short-term fasting affects locomotor activity,corticosterone, and corticosterone binding globulin in a migratory songbird

Unpredictable events such as severe storms lead to an increase in circulating corticosterone (CORT) in breeding birds. This increase is often accompanied by elevations in foraging and irruptive behavior. We were interested in determining if acute food restriction (such as might occur during inclement weather) is a sufficient cue to elicit an increase in locomotor activity, increase CORT secretion, and/or decrease circulating levels of corticosterone binding globulin (CBG) in white-crowned sparrows (Zonotrichia leucophrys gambelii). Male Z.l. gambelii were housed individually in environmental chambers on long days (LD 20:4) to simulate breeding season daylength. Birds were fed ad libitum, and on select days, food was removed 2 h after lights on (fasted treatment), or was removed and replaced (control). We analyzed CORT and CBG levels after 1, 2, 6, 22 (lights on), and 23 h under fasted and control conditions. We also measured activity during the 23-h experiment. Activity levels were increased under fasted conditions during the daytime relative to control conditions, but activity levels did not differ between treatments during the night. Fasting as little as 1, 2, and 6 h significantly increased total CORT levels above baseline (control), although after 22 h, total CORT levels under fasted conditions matched those under control conditions. Plasma CBG decreased after the 22-h fast, and remained low after the 23-h fast. This change was sufficient to significantly elevate free CORT levels in fasted birds relative to ad libitum food conditions, despite the lack of difference in total CORT levels.     

The natural plasma testosterone profile of male blue tits during the breeding season and its relation to song output

In male birds, the gonadal hormone testosterone (T) is known to influence territorial and mating behaviour. Plasma levels of T show seasonal fluctuations which vary in relation to mating system and social instability. First, we determined the natural T profile of male blue tits Parus caeruleus during the breeding season. We found that plasma levels of T increased at the onset of nest building. Thus, the increase in circulating T was not associated with territory establishment, nor with the fertile period of the males' mates. In most individuals, T levels dropped to values close to zero during the period of chick feeding. Second, we investigated the relationship between plasma levels of T and male age, size, and singing behaviour. During the mating period, T levels did not differ between 1 yr old and older males and did not correlate with body size or condition. However, song output during the dawn chorus tended to be positively correlated with T levels. Therefore, if high T levels are costly, song output might be an honest indicator of male quality in blue tits. Finally, we show that plasma levels of T are significantly higher during the night than during the day. This pattern has also been observed in captive non-passerine birds, but its functional significance remains unknown.     

Stress responses of testosterone and corticosterone-binding globulin in a multi-brooded species,Eurasian Tree Sparrows (Passer montanus): Does CBG function as a mediator?

In avian plasma, testosterone (T) and corticosterone (CORT) compete to bind with corticosterone-binding globulin (CBG). Elevation of CBG may function to "buffer" the tissues against high circulating levels of T and stress-induced levels of CORT. To demonstrate the effects of acute stress on CBG and T levels and their biological functions, we investigated seasonal changes of baseline and stress-induced T and CBG levels in Eurasian Tree Sparrows (Passer montanus) during different life stages using the capture-handling-restraint stress method. Our results show that (1) male sparrows had significantly higher baseline T levels and CBG capacities during the nest building, the first egg-laying, and the first nestling stages, and significantly decreased stress-induced T levels only during the nest building and the first egg-laying stages. They also expressed significantly increased stress-induced CBG capacities during the second nestling stage. (2) Females showed significantly higher baseline CBG capacities but significantly decreased stress-induced CBG capacities during the nest building stage, and females also showed significantly increased stress-induced CBG capacities during the second egg-laying and the second nestling stages. Therefore, the seasonal fluctuations of baseline CBG in both sexes and baseline T in males reflect their adaptive strategies for optimizing their physiological and behavioral states to the life history cycle. The different patterns of stress-induced CBG in females suggest CBG functions as an essential mediator in regulating stress response to unpredictable perturbations. Our results highlight the need for future studies of stress-induced CBG and T levels on a wide range of vertebrate species that vary in different life history stages to gain a full understanding of the mechanisms that underlie biological functions of CBG and T for unpredictable stressors.     

Effects of aggressive encounters on plasma corticosteroid-binding globulin and its ligands in white-crowned sparrows

In birds, corticosteroid-binding globulin (CBG) binds corticosterone, progesterone and testosterone. The concentration of each ligand can alter the binding of the other ligands through competitive interactions. Thus, an increase in corticosterone or progesterone may displace testosterone bound to CBG, leading to an increase in bioactive free testosterone levels without affecting total testosterone levels in the circulation. Aggressive interactions increase plasma total testosterone levels in some birds but not in others. Here, we tested the hypothesis that aggressive encounters in the late breeding season would not increase total testosterone levels in plasma, but would alter CBG, total corticosterone or total progesterone levels in such a way as to modify the number of available binding sites and therefore occupancy by testosterone. A marked decrease in CBG occupancy by testosterone would indirectly suggest an increase in free testosterone levels in plasma. Wild male white-crowned sparrows were exposed to a simulated territorial intrusion (STI) or control for 30 min. Subjects were then caught and bled. We measured CBG using a ligand-binding assay and corticosterone, progesterone and testosterone using highly sensitive radioimmunoassays. STI significantly increased aggressive behaviors but did not affect plasma total testosterone levels. STI significantly increased plasma CBG and total corticosterone levels and decreased plasma total progesterone levels. We predict that CBG occupancy by corticosterone will increase slightly following an aggressive encounter. However, this small change is unlikely to increase free testosterone levels, because of the large number of seemingly unoccupied CBG binding sites in these subjects.     

Nutritional stress affects corticosterone deposition in feathers of Caspian tern chicks

Stressful environmental conditions affect the adrenocortical function of developing animals, which can have consequences for their fitness. Discovery of the avian stress hormone corticosterone (CORT) in feathers has the potential to broaden the application of endocrine research in ecological and evolutionary studies of wild birds by providing a long‐term measure of CORT secretion. Mechanisms of CORT deposition in feathers are not well known and few studies have related feather CORT to circulating plasma CORT during feather growth. Our objective was to experimentally test the validity of using feather CORT as a measure of CORT secretion in developing birds experiencing nutritional stress. Caspian tern Hydroprogne caspia chicks were fed ad libitum or restricted (35% less than ad libitum) diets for four weeks. We measured CORT in feathers from these chicks to examine the relationship between feather CORT concentrations and nutritional limitation, circulating plasma CORT, and feather development. We found that feather CORT was higher in controls fed ad libitum than in restricted individuals, despite higher levels of plasma CORT in restricted chicks compared to controls. Feather mass and growth rates were strongly and positively related to feather CORT concentrations in both treatments. This is the first experimental study to show that feather CORT concentrations can be lower in response to nutritional stress, even when plasma CORT concentrations are elevated. Our results indicate that CORT deposition in feathers may be confounded when feather mass and growth rates are compromised by nutritional stress. We conclude that feather CORT can be used for assessing nutritional stress in growing birds, but the direction of response depends on how strongly stress affects feather development.     

Changes in plasma levels of testosterone during male-male interactions in the song sparrow,Melospiza melodia: time course and specificity of response

Summary Previous investigations have shown that malemale interactions over territory boundaries result in an increase in plasma levels of testosterone (T). In order to determine the time course of this increase following an agonistic challenge, male song sparrows, Zonotrichia (= Melospiza) melodia, were exposed to simulated territorial intrusions. Responding males were captured at intervals after onset of the intrusion for up to 60 min. Plasma levels of T were elevated by 10 min after onset of the challenge, and remained high in birds sampled 10–60 min after onset of the intrusion. In addition, plasma levels of luteinizing hormone (LH) increased and were significantly elevated after 10 min of stimulation. The results suggest that at least the prolonged elevation of T secretion precipitated by a male-male interaction was mediated via an increase in LH release. However, the initial surge of T levels in plasma may be LH independent. p Investigations of captive birds confirmed that circulating LH and T levels increased when male song sparrows were challenged by a conspecific male. Challenges involving a male house sparrow, Passer domesticus, had no effect suggesting that responses to male intrusions were species characteristic and not a result of general arousal. Further, auditory stimuli (vocalizations) alone, and visual stimuli (a devocalized male) alone were less effective in stimulating an increase in secretion of T than a combination of auditory and visual stimuli.Abbreviations DHT dihydrotestosterone - LH luteinizing hormon; - T testosterone     

Testosterone, reproductive stage, and territorial behavior of male and female European stonechats Saxicola torquata

We investigated territorial behavior and circulating testosterone (T) levels in a multiple-brooded population of the European stonechat, a socially monogamous passerine bird with biparental care. Between arrival at and departure from the breeding territories, we (1) quantified behavior of both sexes in response to a simulated territorial intrusion (STI) of a male conspecific and (2) measured plasma T concentrations in males and females. Male response scores to a STI and male T concentrations varied with phase, but there was no temporal association between plasma T levels and the intensity of territorial behavior. During both two sexual and two parental phases, at least half of the tested males showed aggressive responses. About 20% of the tested males responded with courtship prior to laying of the first clutch, but none of the males courted during later phases. Age had a positive overall effect on male plasma T. Females also reacted to the STI of a male, but their responses did not vary with breeding phase. Female plasma T varied with phase, being elevated during production of the first but not of the second clutch. As with males, female responses to the STI were not correlated with T levels. Responses of pair partners were positively correlated with each other. We conclude that modulation of male territorial aggression with breeding phase is not regulated by changes of plasma T concentrations. In light of other studies showing reduced male aggression by pharmacological inhibition of cellular actions of T, we propose that T is permissive for male territorial aggression, but does not mediate short-term changes associated with breeding phase. The function of the high female plasma T concentrations during formation of first clutches could be related to the production of eggs with high concentrations of androgens.     

Context-specific territorial behavior in urban birds: no evidence for involvement of testosterone or corticosterone

Testosterone produced by the gonads is a primary mediator of seasonal patterns of territoriality and may directly facilitate territorial behavior during an encounter with a potential intruder. Costs and benefits associated with territoriality can vary as a function of habitat, for example through differences in resource distribution between areas occupied by different individuals. We investigated behaviors in response to simulated territorial intrusions (hereafter territorial behaviors) in urban (Phoenix, Arizona) and nearby desert populations of two Sonoran Desert birds (Curve-billed Thrasher and Abert's Towhee). We also examined the degree to which these behaviors are mediated by testosterone (T) and the adrenal steroid, corticosterone (CORT), which can interact with T in territorial contexts. In both species, urban birds displayed more territorial behaviors than their desert conspecifics, but this difference was not associated with variation in either plasma total or in plasma free (i.e., unbound to binding globulins) T or CORT. In addition, neither plasma T nor plasma CORT changed as a function of duration of the simulated territorial intrusion. Urban Abert's Towhees displayed more territorial behaviors in areas where their population densities were high than in areas of low population densities. Urban Curve-billed Thrashers displayed more territorial behaviors in areas with a high proportion of desert-type vegetation, particularly in areas that differed in vegetation composition from nearby randomly sampled areas, than in areas with a high proportion of exotic or non-desert type vegetation. Associations between territorial behavior and habitat characteristics were not related to plasma T or CORT. Understanding the hormonal processes underlying these associations between behavior and habitat may provide insight into how free-ranging animals assess territorial quality and alter their defensive behavior accordingly.     

Acute embryonic exposure to corticosterone alters physiology,behaviour and growth in nestlings of a wild passerine

Maternally-derived glucocorticoids can modify the normal development of young animals. To date, little is known about maternal effects that are mediated by acute embryonic exposure to glucocorticoids. In birds, elevated maternal transmission of corticosterone (CORT) to egg albumen is mainly dependent on acute stress. In this study, we increased CORT levels in the egg albumen of a wild passerine, the great tit (Parus major), breeding in favourable deciduous and less suitable coniferous habitat. Subsequently we measured the somatic growth, baseline and acute glucocorticoid responses, immunity and behaviour of prenatally manipulated offspring with respect to control siblings. We found that prenatally CORT-exposed nestlings had lower baseline CORT levels, a more rapid decline in CORT during recovery from a standardized stressor, and a reduced heterophil/lymphocyte ratio compared with controls. Although stress-induced total CORT levels remained unchanged, free CORT levels were significantly lower and the levels of corticosteroid binding globulins (CBG) significantly higher in experimental offspring. Prenatally CORT-exposed offspring begged longer after hatching than controls. Stress-induced behavioural activity of fledglings did not differ between treatments, while its association with baseline CORT levels was significant in the control group only. The body mass and tarsus length of fledglings was positively affected by manipulation in unfavourable coniferous habitat only. We conclude that maternal effects related to elevated levels of albumen CORT modify diverse aspects of offspring phenotype and potentially increase offspring performance in resource poor environments. Moreover, our results indicate that maternal glucocorticoids may suppress the effect of hormones on behavioural responses.     

Year-to-year patterns of circulating levels of testosterone and corticosterone in relation to breeding density, experience, and reproductive success of the polygynous red-winged blackbird.

To determine across-year patterns in plasma testosterone (T) and corticosterone (B) levels in free-living birds, we took blood samples in the same 2-week breeding period during 4 consecutive years from territorial male red-winged blackbirds. We used our data to test predictions of the "challenge hypothesis" of T secretion and also to examine hormonal correlations with age, breeding experience, breeding density, and reproductive success. Average T and B levels across years were not significantly different. Within individuals, T levels between years were, in general, highly variable. T levels of males with territories in high-density breeding areas were significantly higher than those of males with territories in low-density areas. T levels were positively correlated with harem size and there was a trend for males with high T levels to fledge more offspring. We found no relationships between T and B levels and male age or breeding experience. Some results of the study were consistent with the predictions of the challenge hypothesis. This study constitutes one of the few examinations of across-year patterns in circulating hormone levels in wild populations.     

Extra-pair copulations can insure female blue tits against male infertility

Mating outside the pair-bond occurs frequently in socially monogamous birds, but the benefits that females gain from this behaviour remain debated. One hypothesis is that females engage in extra-pair copulations (EPCs) to ensure that their clutch is fertilised in case their own mate is infertile, but evidence for this idea is scarce. We report on a case of an infertile male blue tit that bred in three successive years with three different females. In the first year, all eggs were sired by an extra-pair male whereas in the second year all eggs were unfertilised and contained no sperm. In the third year, the female produced two clutches that were fertilised by an extra-pair male, but – unlike ‘normal’ clutches – sperm numbers on the perivitelline membrane decreased rapidly over the laying sequence. Our findings show that blue tit females mated to an infertile male can escape reproductive failure by engaging in EPCs and support previous suggestions that EPCs in blue tits cease after the onset of egg laying.     

Testosterone elevation and response to gonadotropin-releasing hormone challenge by male northern cardinals (Cardinalis cardinalis) following aggressive behavior

There is much discrepancy about the relationship between testosterone (T) and male aggressive behavior. For example, in birds, males of many species significantly elevate T levels during inter-male conflict. However, this is not universal, and in species where males typically do not elevate T during aggressive interactions, concentrations of the hormone are often assumed to be circulating at maximum levels. We examined if male northern cardinals (Cardinalis cardinalis) significantly elevated T during simulated territorial intrusions (STIs). We also examined if individuals had the capacity to further elevate T levels in response to gonadotropin-releasing hormone (GnRH) injections immediately after an aggressive encounter. Our results indicate that male cardinals do not significantly elevate T levels in response to STIs, but have the physiological capacity to significantly elevate T in response to GnRH injections following aggressive interactions. This implies that T levels of individuals captured during STIs were not at maximum concentrations. However, additional findings in this study also suggest the possibility that prolonged social instability could elicit significant elevations in T in males of this species, warranting further investigation.     

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.     

Disparity in the response of sex steroid-binding protein and corticosteroid-binding globulin to thyroxine in the primate

There is uncertainty regarding the differential effects of thyroid hormone on the circulating levels of sex steroid-binding protein (SBP) and corticosteroid-binding globulin (CBG). Therefore, we studied the effects of elevated thyroxine (T4) on SBP and CBG concentrations in serum in 4 baboon (Papio anubis) infants between 6 and 7 mo of age. The infants were given levothyroxine in increasing doses (25 to 600 micrograms) over a 1-mo period until the peripheral T4 levels were 3-4 times higher than baseline values. Each animal served as its own control. Blood samples were obtained at 2- to 3-day intervals prior to and during treatment. Serum T4 was measured by RIA. SBP and CBG were measured by diethylaminoethyl cellulose filter paper assays. Elevations in serum T4 resulted in a dose-response increase in circulating SBP concentrations with a maximal increase at the 600-micrograms dose of T4 (p less than 0.001). In contrast, significant elevations in peripheral CBG levels occurred with 50 and 100 micrograms of T4 (p less than 0.025 and p less than 0.01, respectively), but decreased thereafter with higher doses of T4. At the 600-micrograms dose of T4, CBG concentrations returned to baseline values. We conclude that in the baboon, T4 increases the circulating levels of SBP in a graded fashion, while the effect on CBG is biphasic.     

Timing of fledging is influenced by glucocorticoid physiology in Laysan Albatross chicks

Fledging is a major life transition for birds, when juveniles move from the safety of a nest into an environment where they must find food and avoid predators. The timing of fledging within a season can have significant effects on future survival and breeding success. Proximate triggers of fledging are unknown: though wing development is likely a primary factor, other physiological changes, such as elevated plasma corticosterone (CORT), may affect fledging behavior. Laysan Albatross (Phoebastria immutabilis) chicks have an extended post−hatching period during which they reach 150% of adult mass. However, approaching fledging, chicks fast for days to weeks and lose mass while still putting energy into feather growth. We evaluated chick morphology and physiology to elucidate proximate triggers of fledging. As in some other species, CORT increased as chicks fasted and lost body mass. At the same time, corticosteroid binding globulin (CBG) declined, thus amplifying free CORT prior to fledging. Once chicks reached a morphological threshold, free CORT levels predicted how long they stayed at the colony: chicks with higher free CORT fledged sooner. To perturb the relationship between body condition, endocrine physiology, and fledging behavior, we supplementally fed chicks for the month before fledging. Fed birds had a slower decrease in body mass, slower decrease in CBG, slower increase in free CORT, and stayed at the colony longer after reaching a morphological threshold. Our study suggests that as chicks lose mass, free CORT acts as a signal of energetic or nutritional state to adjust the timing of fledging.     

Corticosterone metabolites in blue tit and pied flycatcher droppings: effects of brood size, ectoparasites and temperature

The hypothalamic-pituitary-adrenal (HPA) axis of birds induces the secretion of corticosterone (CORT) as a response to different ecological variables. In this study we tested experimentally if manipulations of brood size or ectoparasitism led to subsequent differences in the concentration of excreted CORT metabolites of adult and nestling blue tits (Cyanistes caeruleus). No significant effect of the manipulation of brood size was detected in adults or nestlings. No significant effect of ectoparasitism was detected in males or nestlings, although females from uninfested nests showed lower concentrations of excreted CORT metabolites. In addition, we analysed if weather conditions had an influence on the concentration of excreted CORT metabolites of blue tits and pied flycatchers (Ficedula hypoleuca) breeding in the same forest. We detected no effect of weather conditions on adults, but nestlings of both species showed a negative correlation between their excreted CORT metabolites and the average mean temperatures they were subjected to during their growth. This effect was not found in blue tits in a colder year, suggesting that the sensitivity of the HPA axis to ambient temperature may be subjected to interannual variation. Moreover, we found a positive effect of the maximum temperature on the day of sampling on the concentration of CORT metabolites of blue tit nestlings in one of the years. These results suggest that weather conditions may act as environmental stressors to which the HPA axis of blue tit and pied flycatcher nestlings may be sensitive.     

Interaction of testosterone, corticosterone and corticosterone binding globulin in the white-throated sparrow (Zonotrichia albicollis)

Plasma binding globulins bind steroid hormones and are thought to regulate hormone access to tissues. Mammals have both sex steroid binding globulin (SSBG) and corticosteroid binding globulin (CBG). Birds, however, have no detectable SSBG, leading to the early conclusion that birds have no plasma regulation of sex steroids. CBG, however, can bind androgens with relatively high affinity. In birds, therefore, the control of androgenic effects may be tightly regulated by glucocorticoid physiology because glucocorticoids compete with androgens for CBG binding sites. We report levels of total testosterone (T), total corticosterone, CBG, and estimated free T in the males, the more aggressive morph had higher levels of total T; female morphs did not differ. Approximately 96% of T was bound to CBG, but a lack of morph or sex-specific differences in corticosterone titers or CBG capacity caused patterns of free T to mirror those of total T. While CBG has the potential to greatly influence T availability to tissues, in this species interactions between T, CBG and corticosterone do not appear to alter general patterns of T availability to tissues.