The effects of testosterone on a viral infection in greenfinches (Carduelis chloris): an experimental test of the immunocompetence-handicap hypothesis

The immunocompetence-handicap hypothesis suggests that the honesty of quality signals could be guaranteed if testosterone (T) suppresses immune function while enhancing male ornaments. In addition, it has been proposed that the cost of enhancing ornaments should be highest for males with small ornaments. Recently, the assertion that T causes obligate immunosuppression has been questioned. In this study, we tested whether elevated T levels would increase susceptibility to a viral infection, and whether this hypothesized effect would be most pronounced in males with small ornaments. We surgically inserted T implants into 15 male greenfinches (Carduelis chloris) and control implants into a further 15 males. All birds were then infected with a naturally occurring virus (Sindbis virus, Alphavirus genus), and each bird's daily viraemia (blood virus concentration) was measured for seven days. The specific antibody response was measured for eight weeks. T-implanted males did not exhibit increased viraemia or decreased antibody response, and males with small and large ornaments did not respond differently to T implantation. We did, however, find that T implantation decreased viraemia early in the course of the infection and increased viraemia late in the infection. Thus, our results demonstrate that T may act both to increase and to decrease viraemia.     

Shield characteristics are testosterone-dependent in both male and female moorhens

The immunocompetence handicap hypothesis proposes that the expression of secondary sexual characteristics is positively related to testosterone levels, but that elevated testosterone levels also impose costs from immune suppression. Hence, testosterone-dependent characteristics should accurately reflect male quality because only high-quality males are able to invest in large sexual characteristics without detrimental effects upon their own immune system. Most studies to date have focused on the role of testosterone in the expression of male ornaments and on the possible immunosuppressant effects of androgens in males. In the moorhen (Gallinula chloropus), a sexually monomorphic monogamous bird species showing a partial sex-role reversal, both sexes have a prominent frontal shield. We implanted both sexes with testosterone-filled implants to examine the effects of testosterone on shield characteristics and immune function. Shield size, thickness, and color were significantly increased by an experimental increase in testosterone concentrations in both males and females. Likewise, removal of the implants led to a rapid decrease in shield size and thickness in both males and females, suggesting that both sexes responded quickly to an increase or a decrease in testosterone. Moorhens implanted with testosterone had higher intensities of ectoparasite infestations than control birds, but other indirect measures of immunocompetence did not differ significantly between the two categories of birds.     

Does sexual dimorphism in human faces signal health?

Evolutionary psychologists suggest that a preference for sexually dimorphic traits in human faces is an adaptation for mate choice, because such traits reflect health during development. For male faces, this claim rests on the immunocompetence-handicap hypothesis, which states that the increased testosterone levels needed to develop large masculine traits stress the immune system. We examined whether masculine traits in adolescent male faces are associated with health during development, and also whether feminine traits in adolescent female faces signal health. Feminine traits are attractive, but it is less clear whether they should signal health. Rated masculinity in adolescent male faces correlated modestly with actual health, and was perceived as healthy, but not as attractive. Rated femininity in adolescent female faces did not correlate with actual health, although it was perceived as healthy and attractive. These results support the immunocompetence-handicap hypothesis for male faces in that masculine traits signalled health during adolescence. However, they suggest that any health-related evolutionary benefits obtained from preferences for attractive facial traits may be weak.     

Experimentally elevated testosterone levels enhance courtship behaviour and territoriality but depress acquired immune response in Red Bishops Euplectes orix

The immunocompetence handicap hypothesis (ICHH) posits that the hormone testosterone mediates a trade‐off between investment in reproduction and immunological condition. In this study, we tested the ICHH in the Red Bishop Euplectes orix, a polygynous weaverbird. Males of this strongly sexually dimorphic species show an elaborate courtship display to attract females and compete aggressively with other males for nesting sites in breeding colonies. We experimentally elevated testosterone levels in breeding male Red Bishops kept in an aviary with a subcutaneous implantation of testosterone‐releasing pellets. We then compared behaviour, development of territory size and immunological condition (as assessed through a white blood cell count) of the experimental group with a control group treated with placebos. In addition, we measured the primary and secondary response to phytohaemagglutinin (PHA) to investigate the effect of testosterone on both innate and acquired immunity. Males with elevated levels of testosterone enlarged their territories and conducted more courtship behaviour, while showing a decrease in health, expressed by an increased heterophil/lymphocyte ratio compared with the control group. Males of the control group showed an increase of the secondary response to PHA, as expected under the assumption that repeated exposure to an antigen enhanced the immune response due to acquired immunity. However, males with experimentally increased testosterone levels did not show such an enhanced immune response in the secondary PHA test (although sample size and power of the statistical tests were low), indicating that testosterone treatment might directly or indirectly suppress some component of the acquired immune response.     

Steroid hormones and immune function: experimental studies in wild and captive dark-eyed juncos (Junco hyemalis)

Monogamous and polygynous male songbirds generally differ in their breeding season profiles of circulating testosterone. Testosterone level spikes early in the breeding season of monogamists and then declines, but it remains high in polygynists. Male dark-eyed juncos (Junco hyemalis) are socially monogamous and exhibit the usual pattern, but experimental maintenance of high testosterone throughout the breeding season alters normal behavior and physiology and affects various components of annual reproductive success but not overall annual success. Because stabilizing selection predicts that alteration of naturally existing phenotypes should reduce lifetime reproductive success, we asked whether prolonged testosterone exposure might impair immune function and perhaps thereby reduce life span. We assessed immune function in captive and wild male juncos that we treated with either testosterone-filled or empty Silastic implants. Results indicate that prolonged elevation of testosterone suppresses antibody production in captive males and cell-mediated immunity in wild males. Together these results suggest that testosterone-treated males may be more susceptible to disease or parasitic infection. As earlier studies have shown, levels of corticosterone as well as testosterone are higher in testosterone-treated males, so it is unclear whether the immune suppression we observed is due to testosterone's direct effects on immunity or testosterone's influence on glucocorticoid production. We discuss results in the context of recent hypotheses regarding life-history theory and potential endocrine-immune interactions.     

Female choice and the benefits of mate guarding by male mallards

Pair formation and breeding in many species of waterfowl are separated both temporally and spatially. Most studies of female choice in this group have focused on male characteristics at the time of pairing, with less attention given to how mate choice affects breeding season outcomes. In this study I compared pairing success, male plasma testosterone level and mate-guarding ability of male mallards, Anas platyrhynchos, in two experiments. In the first experiment females and males were group housed with equal sex ratios, thus allowing all of these males to pair. At the same time, an equal number of males was housed in groups without access to females and remained unpaired. In this experiment testosterone levels of paired and unpaired males during autumn (baseline) and spring (breeding) did not differ, indicating that the process of pair formation and breeding does not cause elevated spring testosterone levels in males. However, testosterone did temporarily decrease in paired males during the winter (pair formation) season. In the second experiment groups were male biased, allowing only half of the males to pair. Here paired males had significantly higher testosterone levels than unpaired males during the breeding season, but not during the preceding autumn. Together the results of these experiments indicate that successful pair formation predicts but does not alter male testosterone level during the breeding season. I also found that females paired to males with high levels of testosterone were missing fewer feathers due to forced copulation attempts by nonmates, suggesting that females may choose males based on their mate-guarding abilities. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Effects of experimentally elevated testosterone on plasma glucocorticoids, body mass, and recapture rates in yellow-pine chipmunks, Tamias amoenus

In male yellow-pine chipmunks plasma levels of glucocorticoids (GCs) are low while plasma testosterone (T) levels peak during the mating season, suggesting that T suppresses GC levels. To test this hypothesis, free-living, post-reproductive males were implanted during summer with either a T-filled (T-males) or an empty silastic implant (controls or C-males). Body mass and plasma levels of corticosterone, cortisol, and T were measured immediately before and 1 month after implantation. Exogenous testosterone increased T to high physiological levels typical of reproductively active males. By 1 month after implantation, T-males decreased their mean body mass and plasma GC levels, while C-males maintained their mean body mass and GC levels. Even though T-males lost mass, recapture success 1 month after implantation for T-males (71%) was equal to that of C-males (71%). However, the overwinter recapture rate of C-males (83%) was significantly greater than that of T-males (20%). The results support the hypothesis that high plasma T of males during mating has a suppressive effect on plasma GC levels. Additionally, experimentally elevated T significantly reduced the rate of recapture during the following spring, and this may reflect a reduction in local overwinter survival. The suppression of adrenocortical activity by T may contribute to the reductions in prehibernation body mass and post-emergence recapture success. J. Exp. Zool. 287:378-383, 2000.     

The Effects of Experimentally Elevated Testosterone and Food Deprivation on Food Consumption and Prey Size Preferences in Male Dark-Eyed Juncos (Junco hyemalis, Emberizidae: Passeriformes)

Numerous studies have shown that the experimental elevation of circulating levels of testosterone reduces parental behaviour in male birds, particularly the provisioning of young. The mechanisms responsible for this change in behaviour are not fully understood. In this study, we examine the effects of elevated testosterone on food consumption and prey selection, both of which have potential consequences for nestling provisioning behaviour. We manipulated testosterone and performed two experiments on a captive, non-breeding population of male dark-eyed juncos ( Junco hyemalis ) on long day-lengths. In the first experiment, we subjected juncos to 3 h of food deprivation and compared food consumption and prey size selection by males with elevated testosterone (testosterone males) to that of control males. Testosterone males consumed more food than control males and showed a preference for larger prey. In a second experiment in which small prey were more abundant than large prey, food consumption and prey size preferences did not differ between testosterone and control males. We also manipulated the duration of food deprivation in the second experiment. Males of both treatments consumed more small prey under conditions of mild (1 h) or moderate (5 h) food deprivation and consumed more large prey under conditions of intermediate (3 h) food deprivation. We discuss our results and the effects that testosterone has on self-maintenance behaviour and male parental effort.     

Testosterone treatment is immunosuppressive in superb fairy-wrens, yet free-living males with high testosterone are more immunocompetent

The immunocompetence handicap hypothesis proposes that the immunosuppressive effect of testosterone enforces honesty of sexual signalling via a physiological trade-off between signal intensity and immunocompetence. However, evidence that testosterone is immunosuppressive is scant, particularly in birds. I studied the correlation between immunocompetence and testosterone in superb fairy-wrens (Malurus cyaneus), a species with intense intersexual selection. Males are seasonally dichromatic and testosterone increases during the moult from dull brown eclipse plumage into bright nuptial plumage. I determined the primary antibody response to immunization with sheep red blood cells (SRBCs) in (i) control and testosterone-implanted males in captivity, and (ii) a cross-section of free-living males with basal and elevated testosterone (in eclipse plumage, moulting and in nuptial plumage). Experimental treatment with testosterone decreased the likelihood of an antibody response to SRBCs in captive birds. In contrast, free-living males which had acquired the nuptial plumage and had naturally elevated testosterone were more likely to respond to SRBCs than males in eclipse plumage with basal testosterone levels. The association between higher immunocompetence and higher immunosuppressive testosterone could arise if both are positively correlated with male phenotypic quality In addition, the association could result if males compensate for potential immunosuppression by enhancing their humoral immune responses, particularly since high testosterone is linked to other demanding activities such as moulting and courtship displays.     

Testosterone levels of free-ranging resident mantled howler monkey males in relation to the number and density of solitary males: a test of the challenge hypothesis

The challenge hypothesis asserts that testosterone levels and aggression in male adult animals are closely associated with one another in the context of intense reproductive competition, particularly when males challenge one another for the access to females. For mantled howler monkeys (Alouatta palliata), the presence of a solitary male in the vicinity of a social group can be considered a threat for the reproductive success of the resident males, as group takeovers by solitary males are followed by a reestablishment of the access to females among group males and sometimes by infanticide. Therefore, these episodes should be accompanied by an increase in testosterone secretion according to the challenge hypothesis. In order to test our prediction, we relate different group and subpopulation variables to the testosterone levels measured in feces collected from 10 groups living in 6 forest fragments, at Los Tuxtlas, Mexico. Our results suggest that resident A. palliata males regard solitary individuals as potential contesters for their reproductive success and that they respond to interactions with them with an increase in testosterone levels, that is increasing their aggressiveness. Our results also suggest that in the studied groups all males have at least certain access to mates and that the entrance of a new male would affect their reproductive success negatively. Finally, the negative effects of chronically high testosterone levels may be negatively affecting the fitness of our study groups living in high population densities and small fragments.     

Physiological effects on demography: a long-term experimental study of testosterone's effects on fitness

Understanding physiological and behavioral mechanisms underlying the diversity of observed life-history strategies is challenging because of difficulties in obtaining long-term measures of fitness and in relating fitness to these mechanisms. We evaluated effects of experimentally elevated testosterone on male fitness in a population of dark-eyed juncos studied over nine breeding seasons using a demographic modeling approach. Elevated levels of testosterone decreased survival rates but increased success of producing extra-pair offspring. Higher overall fitness for testosterone-treated males was unexpected and led us to consider indirect effects of testosterone on offspring and females. Nest success was similar for testosterone-treated and control males, but testosterone-treated males produced smaller offspring, and smaller offspring had lower postfledging survival. Older, more experienced females preferred to mate with older males and realized higher reproductive success when they did so. Treatment of young males increased their ability to attract older females yet resulted in poor reproductive performance. The higher fitness of testosterone-treated males in the absence of a comparable natural phenotype suggests that the natural phenotype may be constrained. If this phenotype were to arise, the negative social effects on offspring and mates suggest that these effects might prevent high-testosterone phenotypes from spreading in the population.     

Effects of testosterone and corticosterone on immunocompetence in the zebra finch

The original immunocompetence handicap hypothesis (ICHH) suggested that testosterone has a handicapping effect in males by both promoting the development of sexual signals and suppressing immune function. A modified version, the stress-linked ICHH, has recently proposed that testosterone is immunosuppressive indirectly by increasing production of corticosterone. To test both the original and stress-mediated versions of the ICHH, we implanted male zebra finches taken from lines selected for divergent maximum stress-induced levels of corticosterone (high, low and control) with either empty or testosterone-filled implants. Their humoral and cell-mediated immune responses were then assessed by challenge with diphtheria:tetanus vaccine and phytohemagglutinin respectively. We found no effect of the hormone manipulations on either PHA or tetanus antibody responses, but found a significant interaction between titers of both testosterone and corticosterone on diphtheria secondary antibody response; antibody response was greatest in individuals with high levels of both hormones. There was also a significant interactive effect between testosterone treatment group and corticosterone titer on body mass; the body mass of males in the elevated testosterone treatment group decreased with increasing corticosterone titer. These results suggest that, contrary to the assumption of the stress-mediated version of the ICHH, high plasma levels of corticosterone are not immunosuppressive, but are in fact immuno-enhancing in the presence of high levels of plasma testosterone. Equally, the central assumption of the ICHH that testosterone is obligately immunosuppressive is also not supported. The same individuals with the highest levels of both hormones and consequently the most robust antibody response also possessed the lowest body mass.     

Adaptive immune responses are linked to the mating system of arvicoline rodents

ABSTRACT Males generally exhibit reduced immune responses and greater susceptibility to disease than females. The suppressive effect of testosterone on immune function is hypothesized to be one reason why males have lower immune responses than females. Presumably, this effect of testosterone should be more pronounced among polygynous than monogamous species because circulating testosterone is higher among polygynous than monogamous males. The present study examined the extent to which sex differences in specific humoral immunity are related to the endocrine status and mating system of two arvicoline rodents. Humoral immunity was evaluated among polygynous meadow voles (Microtus pennsylvanicus) and monogamous prairie voles (Microtus ochrogaster) by challenging them with the novel antigen keyhole limpet hemocyanin (KLH) and assessing specific immune responses 5, 10, and 15 d following immunization. Overall, meadow voles mounted higher anti-KLH IgM and IgG responses than prairie voles did. Sex differences were also apparent for anti-KLH IgM responses; male meadow voles mounted higher antibody responses than conspecific females, whereas female prairie voles mounted greater responses to KLH than did conspecific males. Male meadow voles had significantly higher testosterone concentrations and reproductive organ mass than male prairie voles did but had elevated immune responses, suggesting that testosterone may not be the primary factor involved in the observed sex and species differences in immune responses. Species and sex differences in corticosterone concentrations were also evident and may contribute to the observed differences in immune function. The influence of extrinsic factors on immune function is also discussed. Taken together, these data provide evidence that the mating system may influence endocrine-immune interactions.     

Male endocrine responses to females: Effects of social cues in cynomolgus macaques

This study tested the hypothesis that the hormonal responses of male macaques to sexually receptive females are mediated by specific socioenvironmental cues. Twelve, socially living male Macaca fascicularis were exposed to two-ovariectomized, estrogen-primed females under pair test, peer group, and peer group without dominant male conditions. Preexposure to postexposure changes in serum cortisol levels and testosterone levels were examined in relation to male dominance rank, age, and conditions of access to stimulus females. The males displayed significantly lower cortisol increases and greater testosterone increases with females under peer group than pair test conditions. Dominant males displayed greater testosterone increases than subordinate males under peer group conditions. The testosterone levels of subordinate males were greatly enhanced by removing the dominant male from each group. Adult males displayed greater cortisol responsivity to changes in conditions of access to females than did subadult males. Subadult males displayed greater testosterone responsivity to removal of the dominant male from the group than did adult males. It is concluded that specific social cues can greatly influence the endocrine responses of males to receptive females, and further, that there may be optimal socioenvironmental parameters for eliciting testosterone increases in male primates.     

Sexual selection for male mobility in a giant insect with female-biased size dimorphism

Female-biased size dimorphism, in which females are larger than males, is prevalent in many animals. Several hypotheses have been developed to explain this pattern of dimorphism. One of these hypotheses, the mobility hypothesis, suggests that female-biased size dimorphism arises because smaller males are favored in scramble competition for mates. Using radiotelemetry, we assessed the mobility hypothesis in the Cook Strait giant weta (Deinacrida rugosa), a species with strong female-biased size dimorphism, and tested the prediction that male traits promoting mobility (i.e., longer legs, smaller bodies) are useful in scramble competition for mates and thus promote reproductive success. Our predictions were supported: males with longer legs and smaller bodies exhibited greater mobility (daily linear displacement when not mating), and more mobile males had greater insemination success. No phenotypic traits predicted female mobility or insemination success. In species with female-biased size dimorphism, sexual selection on males is often considered to be weak compared to species in which males are large or possess weaponry. We found that male giant weta experience sexual selection intensities on par with males of a closely related harem-defending polygynous species, likely because of strong scramble competition with other males.     

Foreleg Ornaments Do Not Hinder Foraging Success in Brush-Legged Wolf Spiders

Secondary sexual traits are often assumed to increase reproductive success but can also carry costs for males bearing them. Here, we investigate potential foraging costs of foreleg ornamentation in wolf spiders in the genus Schizocosa. Given that the forelegs are involved in securing prey items and that ornaments may also inhibit general mobility, we hypothesized that the presence of ornamentation inhibits foraging success. To test this, we first took an ontogenetic approach. We conducted an experiment to examine variation in foraging behavior before and after ornament development in a population of Schizocosa that exhibits two male phentoypes: (a) brush-legged males that develop large brushes of hairs on their forelegs upon maturation and (b) non-ornamented males that lack ornamentation. When comparing penultimate versus mature foraging behavior between these two phenotypes, we predicted (i) reduced foraging intensity upon maturation for males of both groups (due to increased mate-searching behavior) and (ii) a greater decrease in foraging efficiency in brush-legged versus non-ornamented males (due to costs of ornamentation). As expected, we found that when exposed to prey, all males exhibited greater latencies to attack as matures than as penultimates, but unexpectedly, they differed in their patterns of attack frequency; brush-legged males decreased the number of attacks upon maturation while non-ornamented males increased. We found no support for our second prediction: neither male phenotype exhibited a change in foraging success upon maturation. Given that foraging behavior differed between ontogenetic stages, making it difficult to isolate the effects of ornamentation per se, we conducted a second experiment with a pure brush-legged population (S. ocreata) to directly test the influence of foreleg brushes on foraging behavior. We compared foraging behavior between mature males with brushes present versus mature males with brushes artificially removed and found no effect of the presence/absence of brushes on foraging success. In summary, using both ontogenetic and ornament manipulation approaches, we found no support for the hypothesis that ornamentation influences foraging success; however, we do document phenotype-dependent foraging behavior.     

Behavioral and physiological responses to experimentally elevated testosterone in female dark-eyed juncos (Junco hyemalis carolinensis)

Testosterone mediates the expression of many fitness-related traits in male vertebrates and is thought to account for numerous sex differences in trait expression. Testosterone is also secreted by females; however, far less is known regarding its effects on female physiology and behavior. Using a bird species in which the effects of testosterone on males are well characterized, the dark-eyed junco (Junco hyemalis), we tested whether an increase in exogenous testosterone in females would alter the phenotypic expression of a suite of behavioral and physiological traits. We found that increased testosterone levels in female dark-eyed juncos led to decreased cell-mediated immune function and increased intrasexual aggression, hypothalamo-pituitary-adrenal (HPA) axis responsiveness, baseline corticosterone and corticosterone-binding globulin (CBG) levels. Furthermore, immunosuppression following testosterone implantation was negatively correlated with total and free testosterone but did not appear to be related to either total or free corticosterone. These results demonstrate that the phenotypic impact of elevated testosterone is not confined to males in dark-eyed juncos, and that the impact in adults can be similar in males and females. We discuss these results in the context of potential endocrine-immune interactions and the evolution of sexual dimorphism.     

Testosterone treatment diminishes sickness behavior in male songbirds

Males of many vertebrate species are typically more prone to disease and infection than female conspecifics, and this sexual difference is partially influenced by the immunosuppressive properties of testosterone (T) in males. T-induced immunosuppression has traditionally been viewed as a pleiotropic handicap, rather than an adaptation. Recently, it has been hypothesized that suppression of sickness behavior, or the symptoms of infection, may have adaptive value if sickness interferes with the expression of T-mediated behaviors important for male reproductive success. We conduct a classic hormone replacement experiment to examine if T suppresses sickness behavior in a seasonally-breeding songbird, Gambel's white-crowned sparrow (Zonotrichia leucophrys gambelii). Triggered experimentally by bacterial lipopolysaccharide (LPS), sickness behavior includes decreased activity, anorexia, and weight loss. Gonadectomized (GDX) males that were treated with silastic implants filled with T exhibited suppression of behavioral and physiological responses to LPS compared to GDX and sham-GDX controls given empty implants. Sickness responses of control groups were statistically indistinguishable. T-implanted birds had significantly higher plasma T than control groups and levels were within the range associated with aggressive interactions during male-to-male contests. These findings imply that suppression of sickness behavior could occur when T is elevated to socially-modulated levels. Alternatively, it is possible that this suppressive effect is mediated through a stress-induced mechanism, as corticosterone levels were elevated in T-implanted subjects compared to controls. We propose that males wounded and infected during contests may gain a brief selective advantage by suppressing sickness responses that would otherwise impair competitive performance. The cost of immunosuppression would be manifested in males through an increased susceptibility to disease, which is presumably offset by capitalizing upon limited reproductive opportunities.     

Testosterone supplementation in subordinate, small male lizards: consequences for aggressiveness, color development, and parasite load

According to the immunocompetence hypothesis, testosterone stimulatesthe expression of male sexually selected traits while decreasingimmunocompetence. This proposed trade-off was studied by experimentalsupplementation of testosterone to small, subordinate, dull-coloredmale lizards, Psammodromus algirus. Experimental males showeda tendency to overlap their home range with fewer small malesthan did control males and tended to be more aggressive. However,control males were observed more frequently attending femalesthan experimental males. The area of patches of breeding coloration,the number of ticks, and the frequency of recoveries of testosterone-supplementedand control males did not differ significantly. The resultssuggest that small adult males with high levels of testosteronebehave more aggressively, which may be advantageous to securinga breeding territory in the next season. However, the hormonedid not apparently affect ornamentation or parasite load. Weargue that, whatever the mechanisms involved, blocking effectsof testosterone may be adaptive because being cryptic facilitatesa sneaking strategy, and low ectoparasite load may improve survival     

Androgens and the immunocompetence handicap hypothesis: unraveling direct and indirect pathways of immunosuppression in song sparrows

The immunocompetence handicap hypothesis proposes that testosterone (T)-dependent sexual signals are honest indicators of male health or genetic quality because only high-quality males are able to withstand the obligate effects of T-induced immunosuppression. In birds, the basic assumption that T suppresses immune function is equivocal, and the physiological mechanisms underlying T-induced immunosuppression remain to be investigated. We explored the proximate pathways of T-induced immunosuppression in song sparrows (Melospiza melodia) by treating captive nonbreeding males with different androgens and measuring several components of acquired immune function. Males implanted with T suppressed cell-mediated and humoral immune responses compared to males implanted with 5alpha-dihydrotestosterone (DHT), dehydroepiandrosterone, or control (empty) implants. Furthermore, T treatment increased plasma levels of corticosterone and decreased body mass and fat stores in relation to other treatments. The failure of DHT to depress immune function suggests that T-induced immunosuppression does not occur through a direct pathway because both T and DHT bind to androgen receptors on target cells. Instead, we outline indirect pathways that are likely responsible for suppression of the avian immune system that include stress-induced immunosuppression, aromatization to estrogen, and alterations in energy allocation that constrain expenditures toward immune system activation.