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Aggression is a behavioral strategy for securing limited resources and its expression is strongly influenced by their presence and value. In particular, males are generally thought to guard females after mating to ward off other males, but the underlying control mechanisms are unknown. Here, we investigated the role of amines on male courtship behavior and its subsequent effect on male-male aggression in crickets (Gryllus bimaculatus). Contrary to the guarding hypothesis, female presence alone had no immediate effect on male-male aggression. Furthermore, confirming studies on other species, prior female contact, but not necessarily courtship or copulation, promoted subsequent male-male aggression in subordinate, but not socially naive crickets. This promoting effect of female contact is transient and slowly wanes after her removal. Selective aminergic receptor antagonists revealed that the promoting effect of prior female contact on male-male aggression is mediated by octopamine (OA), as well as by serotonin (5HT) acting most likely via 5HT1 and/or 5HT7 like receptors. This contrasts the role of 5HT2-like receptors in maintaining reduced aggressiveness after social defeat. Furthermore, while dopamine (DA) is necessary for the recovery of aggression in subordinates after defeat, it appears to play no part in female induced aggression. Male courtship, on the other hand, is selectively promoted by DA and 5HT, again most likely via 5HT1 and/or 5HT7 like receptors, but not by OA. We conclude that OA, DA and 5HT each differentially modulate different aspects of courtship and aggressive behavior in a context specific fashion.

Performing appropriate mating behaviors is crucial to male reproductive success, especially in species where mating is predominantly via female mate choice. Mating behaviors are hormonally regulated and may be sexually selected traits: courtship displays are selected via mate choice, while forced copulations and aggressive behaviors are selected for via intrasexual competition. Endocrine disrupting compounds interfere with proper hormonal functioning in exposed animals. Exposures during developmentally crucial life stages can have irreversible effects lasting through adulthood. I tested the effects of prenatal exposure to environmentally relevant doses of a commonly used herbicide, atrazine (1 and 13.5μg/L) on mating behaviors in male guppies. Guppies were used as a model organism to test the effects of atrazine exposure on wildlife reproductive health. Adult female guppies were mated and exposed to the treatments throughout the gestation period, and offspring born to them were raised without further treatment. At adulthood, the males were tested for the effects of prenatal exposure on their mating behaviors such as courtship displays, gonopodium swings, forced copulatory attempts, and competitive and aggressive behaviors towards rivals who were not exposed to atrazine. I also tested female preference for treated males compared to control males. Atrazine-exposed males were less likely to perform the mating behaviors, and performed them less frequently, than control males. Atrazine exposure also made males less aggressive towards rivals. Females preferred untreated males over atrazine-treated males. In all cases, a non-monotonic pattern was seen, highlighting the significance of low-dose exposures.

Water dragons (Intellegama [Physignathus] lesueurii) are large (to >1m) agamid lizards from eastern Australia. Males are fiercely combative; holding a territory requires incessant displays and aggression against other males. If a dominant male is absent, injured or fatigued, another male soon takes over his territory. Our sampling of blood from free-ranging adult males showed that baseline levels of both testosterone and corticosterone were not related to a male's social tactic (territorial versus non-territorial), or his frequency of advertisement display, aggression, or courtship behavior. Even when we elicited intense aggression by non-territorial males (by temporarily removing territory owners), testosterone did not increase with the higher levels of aggression that ensued. Indeed, testosterone levels decreased in males that won contests. In contrast, male corticosterone levels increased with the heightened aggression during unsettled conditions, and were higher in males that won contests. High chronic male-male competition in this dense population may favor high testosterone levels in all adult males to facilitate advertisement and patrol activities required for territory maintenance (by dominant animals), and to maintain readiness for territory take-overs (in non-territorial animals). Corticosterone levels increased in response to intense aggression during socially unstable conditions, and were higher in contest winners than losers. A positive correlation between the two hormones during socially unstable conditions suggests that the high stress of contests decreased androgen production. The persistent intense competition in this population appears to exact a high physiological cost, which together with our observation that males sometimes lose their territories to challengers may indicate cycling between these two tactics to manage long-term energetic costs.

Hormones are critical for the development, maturation, and maintenance of physiological systems; therefore, understanding their involvement during maturation of the brain is important for the elucidation of mechanisms by which adults become behaviorally competent. Changes in exogenous and endogenous factors encountered during sexual maturation can have long lasting effects in mature adults. In this study, we investigated the role of the gonadotropic hormone, juvenile hormone (JH), in the modulation of adult behaviors in Drosophila. Here we utilized methoprene (a synthetic JH analog) and precocene (a JH synthesis inhibitor) to manipulate levels of JH in sexually immature male and female Drosophila with or without decreased synthesis of neuronal dopamine (DA). Locomotion and courtship behavior were assayed once the animals had grown to sexual maturity. The results demonstrate a sexually dimorphic role for JH in the modulation of these centrally controlled behaviors in mature animals that is dependent on the age of the animals assayed, and present DA as a candidate neuronal factor that differentially interacts with JH depending on the sex of the animal. The data also suggest that JH modulates these behaviors through an indirect mechanism. Since gonadotropic hormones and DA interact in mammals to affect brain development and later function, our results suggest that this mechanism for the development of adult behavioral competence may be evolutionarily conserved.

Androgens, specifically 11-ketotestosterone, are hypothesized to be important in the expression of pre-spawning behaviors such as courtship and aggression in many teleost species. This experiment attempted to elucidate the roles of androgens in the expression of pre-spawning courtship and aggression in male convict cichlids (Amatitlania nigrofasciata). In a laboratory experiment, males were treated with either the androgen receptor antagonist flutamide or blank control and subsequently exposed to social conditions to stimulate inter-sexual courtship or intra-sexual aggression. Males treated with flutamide expressed significantly fewer courtship behaviors than control males but did not differ from control males in pre-spawning intra-sexual aggression. In a field experiment, males treated with flutamide expressed significantly less courtship behavior than males given blank capsules or unmanipulated control males, but did not differ from either set of control males in aggression towards conspecifics or overall aggression to con- and heterospecifics. These data suggest that androgens mediate pre-spawning courtship behavior but not pre-spawning aggression in this species.

Reproductive behavior in amphibians, as in other vertebrate animals, is controlled by multiple hormones. A neurosteroid, 7α-hydroxypregnenolone, has recently been found to enhance locomotor activity in the male newt, Cynops pyrrhogaster. Here, we show that this neurosteroid is also involved in enhancing the expression of courtship behavior. Intracerebroventricular (ICV) injection of 7α-hydroxypregnenolone enhanced courtship behavior dose-dependently in the sexually undeveloped males that had been pretreated with prolactin and gonadotropin, which is known to bring the males to a sexually developed state. But, unlike the case in the locomotion activity, 7α-hydroxypregnenolone did not elicit the behavior in males receiving no prior injections of these hormones. ICV administration of ketoconazole, an inhibitor of the steroidogenic enzyme cytochrome P450s, suppressed the spontaneously occurring courtship behavior in sexually active males. Supplementation with 7α-hydroxypregnenolone reversed the effect of ketoconazole in these animals. It was also demonstrated that the effect of the neurosteroid on the courtship behavior was blocked by a dopamine D2-like, but not by a D1-like, receptor antagonist. These results indicate that endogenous 7α-hydroxypregnenolone enhances the expression of the male courtship behavior through a dopaminergic system mediated by a D2-like receptor in the brain.

Social environment can affect the expression of sex-typical behavior in both males and females. Males of the African cichlid species Astatotilapia burtoni have long served as a model system to study the neural, endocrine, and molecular basis of socially plastic dominance behavior. Here we show that in all-female communities of A. burtoni, some individuals acquire a male-typical dominance phenotype, including aggressive territorial defense, distinctive color patterns, and courtship behavior. Furthermore, dominant females have higher levels of circulating androgens than either subordinate females or females in mixed-sex communities. These male-typical traits do not involve sex change, nor do the social phenotypes in all-female communities differ in relative ovarian size, suggesting that factors other than gonadal physiology underlie much of the observed variation. In contrast to the well-studied situation in males, dominant and subordinate females do not differ in the rate of somatic growth. Dominant females are not any more likely than subordinates to spawn with an introduced male, although they do so sooner. These results extend the well known extraordinary behavioral plasticity of A. burtoni to the females of this species and provide a foundation for uncovering the neural and molecular basis of social dominance behavior while controlling for factors such as sex, gonadal state and growth.

Due to the differential cost of reproduction in promiscuous species, like domesticated goat(s) (Capra hircus), it is expected that females should mate with higher quality males, while males should mate with a greater number of females. Females may use conspicuous secondary sexual characteristics of males such as courtship display to distinguish among high and low quality males. Testosterone (T) controls a large suite of secondary sexual characteristics and variation in T concentrations may account of differences in courtship rates. Two studies were conducted to examine the relationship between T concentrations and courtship rate and its role in mammalian female mate choice. Experiment 1 utilized bucks (intact males) and Experiment 2 utilized T-replaced wethers (castrated pre-pubertally). During the first year of Experiment 2, T-replaced wethers received either vehicle control (CON), 25mg or 100mg testosterone propionate (TP). During the second year of Experiment 2, T-replaced wethers were treated with either 5mg, 15 mg or 25mg TP. For all experiments, mean courtship rates and circulating T concentrations were measured, as well as female preference for males displaying different courtship frequencies. T concentrations and courtship rate were positively correlated for bucks and estrous females preferred high courting bucks. Males receiving 25mg and 100mg TP courted females at a similar rate, but both were significantly higher than courtship rates of the CON wethers. Courtship rates of the 25mg, 15 mg and 5mg TP-treated males were all significantly different. Females did not show a preference between the 100mg or 25mg TP-treated wethers, however both were preferred in comparison to the CON wethers. Females did not show preference between the 25mg or 15 mg TP-treated wethers, however both were preferred in comparison to the 5mg TP-treated wethers. Taken together, these studies suggest that courtship rate is T-dependent. Further, females can use courtship rate to distinguish among males.

Male sexual behavior depends on gonadal androgens in species of all major vertebrate lineages, including reptiles. However, male sexual behavior includes distinct appetitive and consummatory phases, typically denoted as courtship and mounting, with potentially different hormonal control. Different proximate controls of courtship versus mounting could enable disconnected evolutionary losses and gains of various aspects of male sexual behavior. Male courtship display, which is activated by testosterone (T) in many species, is an ancestral trait in the lizard family Eublepharidae. However, Coleonyx elegans (Yucatan Banded Gecko) lost the courtship display, while retaining a highly simplified male sexual behavior that involves only mounting for copulation. We performed surgical manipulations (castration with and without T replacement in adult males; implantation of adult females with exogenous T) to investigate hormonal mechanisms involved in this evolutionary novelty. Our results indicate that the expression of simplified sexual behavior in C. elegans does not require elevated circulating levels of T, a finding that is previously unreported in lizards. In females, however, exogenous T induced male-like mounting. Thus, the mounting phase of sexual behavior is not activated by T in the traditional sense of this term but probably requires post-natal, maturational organization (if not periodic reorganization) by androgens. We conclude that the simplification of male sexual behavior and its independence from elevated levels of circulating androgens in C. elegans evolved via 1) evolutionary loss of the androgen-activated courtship display and 2) retention of the mounting phase, which has a longer "functional memory" for the effects of androgenic steroids.

Previous studies have shown that situations relevant for human mating can affect the levels of many hormones. This study focused on the hypothalamus-pituitary-adrenal axis by measuring salivary cortisol levels in 84 young men prior to and after a period of short social contact with a woman or man. Results showed that after contact with another man the cortisol levels of the participants declined according to the circadian release pattern of cortisol. However, cortisol levels in men declined less when they had contact with a woman. Furthermore, cortisol levels of men increased when they perceived the woman with whom they had contact as attractive. Our findings provide indirect evidence for the role of the hypothalamus-pituitary-adrenal axis in human courtship. During social contact with attractive women, moderate increases in cortisol levels may reflect apprehension over an opportunity for courtship. (C) 2010 Elsevier Inc. All rights reserved.

Ghrelin is a peptide hormone released by the stomach that stimulates hunger. Ghrelin also suppresses reproductive physiology by inhibiting the HPG axis. However, to our knowledge, our results are the first to demonstrate ghrelin's quick suppression of sex-hormone-regulated behaviors. In experiment 1, 2 orexigenic i.p. ghrelin injections (0.165 mg/kg and 0.33 mg/kg) suppressed male courtship behavior (ultrasonic calling to a female) and intermale aggression (latency to attack a stimulus male) 20 min following administration. Experiment 2 (examining only the 0.33 mg/kg dose ) replicated ghrelin's suppression of ultrasonic calling and intermale aggression; however, a third behavior, preference for volatile female odors (20 min following administration), was not significantly inhibited. In experiment 2, ghrelin treatment did not affect general locomotor activity (distance traveled 20 min following injection) or seminal vesicle weight (measured 5 days after completing ghrelin injections). We hypothesize that ghrelin's quick suppression of male aggression and ultrasonic mating calls was mediated through its effects on the brain (rather than indirectly through inhibition of the HPG axis).

In the monogamous and biparental California mouse (Peromyscus californicus), paternal care is critical for maximal offspring survival. Animals form pair bonds and do not engage in extrapair matings, and thus female evaluation of paternal quality during courtship is likely to be advantageous. We hypothesized that male endocrine or behavioral response to courtship interactions would be predictive of future paternal behavior. To test this hypothesis, we formed 20 pairs of California mice, and evaluated their behavior during the first hour of courtship interactions and again following the birth of young. We also collected blood from males at baseline, 1-hr after pairing, 3-weeks paired, and when young were four days old to measure testosterone (T). We found that male T-response to courtship interactions predicted future paternal behavior, specifically the amount of time he huddled over young when challenged by the temporary removal of his mate. Males that mounted T increases at courtship also approached pups more quickly during this challenge than males who had a significant decrease in T at courtship. Proximity of the male and female during courtship predicted paternal huddling during a 1-hr observation, and a multiple regression analysis revealed that courtship behavior was also predictive of birth latency. We speculate that male T-response to a female in P. californicus is an honest indicator of paternal quality, and if detectable by females could provide a basis for evaluation during mate choice.

In most bird species, male courtship behavior is controlled by testosterone (T) and its metabolites. In species breeding in temperate and arctic regions T circulates at high levels during a relatively short courtship period because high levels of T can be costly in terms of immunocompetence and parental care. Few studies have investigated androgen modulation of courtship behavior in tropical birds. Male golden-collared manakins (Manacus vitellinus) aggregate in leks for several months and perform spectacular, acrobatic courtship displays. Here we examined whether T is elevated in golden-collared manakins during the displaying period and if courtship behavior is modulated by androgen action on androgen receptors. We measured T levels in displaying males at the beginning of the breeding season and again, one month later. In addition, both wild and captive males were treated with the anti-androgen, flutamide, and their courtship behavior was recorded for several weeks. T levels were relatively high shortly after leks were established but decreased substantially a month later, even though the amount of courtship did not change. Flutamide reduced male courtship activity for one week, but display behavior then increased after two weeks of flutamide treatment. Our studies show that androgens modulate male manakin courtship, but the amount of courtship is not directly correlated with the concentration of circulating T. These results suggest that the relationships between androgen and courtship might differ between tropical and temperate birds.

Male golden-collared manakins gather on leks and perform an acrobatic display to attract females. In temperate breeding species, testosterone (T) activation of courtship displays has been well studied. Few studies have examined T activation of displays in tropical species; even fewer have explored the activational role of T in elaborate courtship displays such as in the manakin. In some tropical species, including manakins, territorial aggression or song behavior are uncoupled from T. We have previously shown that T activates display behavior in manakin males when endogenous T levels are low in the non-courtship season. To understand how T functions in breeding birds, we examined T levels in a large group of manakins sampled during the courtship and non-courtship season. In addition, during the courtship season, we gave T implants to adult males, juvenile males, and females. We found that T levels were low during the non-courtship season and comparatively higher on average during the courtship season. However, T levels were low in many adult males during the courtship season, especially when compared to temperate breeding species. Regardless of initial endogenous T levels during the courtship season, T implants did not increase the display frequency of adult males. T-treated females and juvenile males did display under similar conditions. Our data suggest that the effects of T on manakin display vary with season, sex, and age and that high T is not necessary for display.

In order to attract females, male golden-collared manakins gather in leks and perform a complex display consisting of acrobatics accompanied by loud "wingsnapping". During this display, males show off their yellow beard and yellow, black, and green plumage that is striking in comparison to the dull green plumage of young males and females. We investigated the role of testosterone (T) in activating the display of manakins and in stimulating the growth of the adult male plumage. T regulates song, copulation, and territorial aggression in temperate species. In tropical species, however, T levels can be relatively low year round, which has raised questions about the involvement of T in courtship display and male aggression in these species. In neither temperate nor tropical species has the role of hormones in the shift from juvenile to adult plumage been well studied. Therefore, we implanted green-plumaged birds and adult males with either a T pellet or an inert pellet (controls) and observed the display behaviors of these birds in the field and in captivity. In captive birds, we also plucked feathers from sexually dimorphic regions and observed color and regeneration rate of new feathers. We found that birds implanted with T increased several display behaviors compared to controls. All plucked feathers grew back the same color as prior to treatment; however, we observed some differences in feather growth rate between T-treated birds and controls.

It is well known that androgens play a critical role in mediating the reproductive behavior of males. However, many laboratory experiments that examined the effects of testosterone in male songbirds typically limited their investigations to the early phase of breeding. We sought to determine the influence of testosterone on social behavior, pair bonding, nesting, and use of space in captive zebra finch (Taeniopygia guttata) males as a function of breeding stage (pre-laying, incubation, and nestling phases). Fourteen males were released into an aviary with 14 females and allowed to breed for 7 weeks. Half of the males were given testosterone implants and half were given control implants. During the pre-laying phase, testosterone-implanted males spent significantly more time in nesting activities than control birds and more time elapsed from starting to build a nest to when their mates initiated egg-laying. During the incubation phase, testosterone-implanted subjects spent significantly more time in female-directed and undirected singing. Use of space varied between hormone conditions depending upon breeding phase: there was no difference during pre-laying, but during the incubation and nestling phases, testosterone-implanted subjects used significantly more space. This significant increase in "home range" during the latter phase of the breeding cycle coincides with results from field studies on other species. These results underscore the importance of considering breeding phase in assessing the behavioral sensitivity to hormones.

Despite having detailed information on mechanisms mediating sex-typical behavior in many species, we have little understanding of whether the same mechanisms regulate these behaviors when they are performed in the same species under different social contexts. In the five field experiments of this study of bluehead wrasses (Thalassoma bifasciatum), a sex-changing fish, we examined the roles of arginine vasotocin (AVT) and the potent teleost androgen 11-ketotestosterone (11KT) in mediating sexual and aggressive behaviors typical of dominant males. We demonstrated that AVT appears necessary for the assumption of dominant territorial status in males and females, but is sufficient only in the socially dominant terminal phase (TP) male phenotype. Specifically, an AVP V(1) receptor antagonist prevented both TP males and females from gaining dominance over recently vacated territories. However, unlike TP males in a previous study, neither females nor initial phase males responded to AVT treatment with increases in display of TP male typical behaviors when under social conditions that inhibit sex change. Treating females with 11KT did not alter responsiveness to AVT, but did induce male coloration and courtship behavior that was not observed in oil-treated females. Combined with the results of a previous study, these results indicate that the ability of AVT to induce male-typical behavior differs among sexual phenotypes and that this differential responsiveness appears to be dependent on social context and not directly on exposure to 11KT. Furthermore, since 11KT can induce courtship behavior in females that is not affected by AVT, there may be different hormonal mechanisms mediating courtship behavior under different social contexts.

Zebra finches evolved in arid areas of Australia. Their reproduction is stimulated by water availability, which is unpredictable. Cheng (Poult. Sci. Rev. 5 (1993) 37) hypothesized that the primary mechanism controlling reproduction in species relying on unpredictable cues should be inhibitory. The onset of stimulatory environmental conditions terminates the inhibition, allowing rapid initiation of reproduction. As the primary hormone regulating water balance in birds, arginine vasotocin (AVT) appears a likely candidate to modulate reproduction in finches. Drought conditions cause sustained AVT release, which in other species inhibits androgen production. To determine whether increased AVT inhibits reproductive behavior, intact males were tested with females and divided into three groups matched for courtship behavior. Osmotic minipumps containing (a) saline, (b) 264 ng AVT, or (c) 1320 ng AVT in saline were implanted subcutaneously and males tested 48 h later. AVT-treated males socialized with females, but the high dose significantly reduced singing and courtship displays. To determine whether AVT acted by depressing androgen secretion, additional males were given subcutaneous androgen implants and divided into two groups matched for courtship behavior. Males were then implanted with minipumps containing (a) saline or (b) the high AVT dose. Males treated with AVT plus androgen showed no deficits in courtship behavior. These data suggest that AVT secretion during periods of drought may inhibit reproduction by inhibiting androgen production. Inhibition of reproductive behavior by AVT may be a more general phenomenon. Large quantities of AVT or, in mammals the closely-related peptide vasopressin (VP), are released when animals are stressed, and high levels of AVT/VP may inhibit reproductive behavior. The extremely short half-life of these peptides means that once proximal factors become more favorable, the gonads should rapidly be released from the peptides' inhibitory actions.

Previous studies have found that vasotocin (AVT) administration to male roughskin newts (Taricha granulosa) enhances courtship clasping as well as appetitive responses to specific sexual stimuli and that treating female newts with androgens plus AVT induces the expression of male-typical courtship clasping (the selective clasping of females). However, the unique and/or interactive effects of sex steroids and AVT on appetitive responses to specific sexual stimuli have not yet been determined. To first identify male-typical, sexually dimorphic appetitive responses to female sexual stimuli, we tested intact newts during the breeding season and found that males, but not females, are attracted to female visual and pheromonal sexual stimuli. We then used ovariectomized (ovx) females implanted with empty silastic capsules (Blk) or with capsules containing testosterone (T), dihydrotestosterone (DHT), or estradiol (E2) and then injected with either saline or AVT to determine the effects of steroids and AVT, alone or in combination with each other, on male-typical behavioral responses to those stimuli. E2 treatment depressed responses toward female visual stimuli independently of AVT. On the other hand, only T-implanted, AVT-injected females displayed male-typical behavioral responses toward female olfactory stimuli, preferring to spend more time in proximity to female-scented than unscented newt models and selectively clasping the female-scented models. Together, these results support the conclusion that sex steroids and AVT influence behavioral responses to sexual stimuli via sensory-specific mechanisms. Furthermore, they suggest that T and AVT interact within the brain to influence sensorimotor processing in the pathways that integrate olfactory sexual stimuli into male-typical courtship behaviors.

The ability of an animal to respond with appropriate defensive behaviors when confronted with an immediate threat can affect its survival and reproductive success. In the roughskin newt (Taricha granulosa), exogenous corticosterone (CORT) rapidly blocks and vasotocin (VT) enhances reproductive behaviors (mainly clasping behavior). Electrophysiological studies have shown that pretreatment of male Taricha with VT counteracts the inhibitory effects of CORT on neuronal activity in the medulla. To test whether similar interactions between VT and CORT influence reproductive behaviors in Taricha, we recorded the time spent and incidence of clasping in males injected with VT or vehicle at 60 min and then CORT or vehicle at 5 min before presentation of a female. This study found that clasping behavior is suppressed in males that received vehicle and then CORT, but is not suppressed in males that received VT and then CORT. Considering these results and the possibility that the performance of clasping behaviors might cause increases in endogenous VT activity, we tested whether the suppressive effects of CORT administration on clasping behavior would occur in males that had recently clasped females. The study found that, in contrast to males that had been isolated from females, CORT administration did not suppress clasping behavior in males that had been allowed to clasp females for 60 min prior to the hormone injection. Our results suggest that, at least in this amphibian and perhaps in other animals, the neuroendocrine regulation of alternative behavioral responses to threats involves functional interactions between corticosteroids and VT-like peptides.

Most seasonally breeding songbirds display dramatic seasonal fluctuations in plasma testosterone (T) levels and mate attraction behaviors, including song. However, males of some songbird species, such as the European starling (Sturnus vulgaris), continue to sing at high levels after the breeding season, when T levels are basal. In male starlings song during the breeding season functions mainly to attract mates, whereas song during the nonbreeding season appears unrelated to reproduction. This suggests that song produced in a context unrelated to female courtship, unlike song directed toward females, is not regulated by plasma T. In captive males housed in large outdoor aviaries we explored the relationship between plasma T and song produced during the breeding season within and outside a courtship context. This was achieved by determining the effects of castration and subsequent T treatment on song and mate attraction behaviors in both the presence and the absence of a female. Compared to intact males, castrated males did not show reduced song activity in the absence of a female for at least 6 months after the operation, strongly suggesting that the expression of noncourtship song is not regulated by plasma T. Likewise, we found that experimentally elevating T levels in castrated males did not affect noncourtship song rates. However, control castrated males receiving empty implants tended to show reduced noncourtship song rates after implantation. This may have been due to a suppressive effect caused by the presence of the T-implanted castrated males in the same aviary. In contrast, courtship singing was clearly controlled by plasma T: it was abolished by castration and restored by subsequent T replacement when males were housed both individually and in a group situation. High plasma levels of T also appeared necessary for the activation of three other behavioral traits critical for mate attraction, namely, nesthole occupancy, spending time (singing) in a nesthole, and carrying green nesting material into a nesthole.

Courtship declines and ceases while parental care increases in the presence of developing eggs during the nesting cycle of the male three-spined stickleback, Gasterosteus aculeatus. Furthermore, circulating 11-ketotestosterone (11KT) levels are higher during the initial “courtship phase” than during the later “parental phase,” similar to that found in other paternal fishes. This study aimed to investigate a possible functional relationship between changes in 11KT levels and changes in reproductive behavior during the nesting cycle. To this end, groups of nonspawned and spawned male sticklebacks were sham-operated, castrated, or castrated and treated with 11-ketoandrostenedione (11KA), and the effects of the treatments on courtship and parental care were studied. Castration removed circulating 11KT, while 11KA replacement prevented the natural decline in 11KT during the parental phase (11KA converts to 11KT extratesticularly), as assessed by radioimmunoassay. Regardless of treatment, parental care remained low and courtship was present in all nonspawned males, even at the end of the experiment. However, courtship did eventually decline in castrated nonspawned males compared to the other two nonspawned groups. In all treatments of spawned males there was a drastic decline in courtship and an increase in parental care. In castrated spawned males, however, the decline in courtship came earlier than in the other two spawned groups. 11KA treatment did not prevent the natural decline in courtship/increase in parental care in spawned males, indicating that the natural decline in 11KT is not responsible for the main portion of the rapid changes in these behaviors over the stickleback's nesting cycle. The limited effects of castration also exclude other gonadal hormones from being responsible for most of these changes.

In male vertebrates the display of courtship behavior depends on the presence of testicular androgens. However, social experiences in adulthood can alter the hormonal dependence of courtship behavior in a variety of species, and we have previously proposed that these behavioral changes are linked to changes in neural metabolic capacity (cytochrome oxidase activity). Here we investigated the effects of prior social experience (housing with females vs housing in isolation) on the retention of courtship behavior following gonadectomy and on cytochrome oxidase (CO) activity in male little striped whiptail lizards, Cnemidophorus inornatus. In Experiment 1, we found that males that were previously housed with females (HWF males) continued to display courtship behavior longer after castration than males previously housed in isolation (ISOLATE males). This is similar to the behavioral plasticity found in rodents and cats. On the other hand, courtship behavior while gonadally intact was indistinguishable between HWF and ISOLATE males. Because all males were housed individually following castration, the difference is due to different social experiences prior to castration. In Experiment 2, we found that gonadally intact HWF males had significantly elevated CO activity in the preoptic area, amygdala, and anterior and ventromedial hypothalamic areas relative to intact ISOLATE males. No significant differences in metabolism were found in the lateral septum, lateral hypothalamus, and habenula or in hindlimb muscle, suggesting that the increase in metabolism is specific to brain nuclei involved in courtship behavior. Altogether, this demonstrates that elevations in metabolic capacity correlate with experience-dependent increases in robustness to castration.

Previous work on lizards has shown that many sexually dimorphic traits depend on testosterone (T), but the details of this control can vary among species. Here, we tested the role of T on the expression of morphological, physiological, and behavioral traits in Lichtenfelder's gecko (Goniurosaurus lichtenfelderi), from the lizard family Eublepharidae notable for interspecific variation in sexually dimorphic traits and the mode of sex determination. Experiments included three groups of males (intact control, surgically castrated, castrated with T replacement) and two groups of females (intact control, T supplemented). In males, castration caused reductions in 1) the size of hemipenes, 2) offensive aggression, 3) male sexual behavior in a neutral arena, 4) activity of precloacal glands, and 5) loss of male chemical cues for sex recognition. These reductions were not observed in castrated males with T replacement. Interestingly, castrated males performed sexual behavior in their home cages, which shows that the effect of T depends on the environmental context. Notably, tail vibration, previously reported as a courtship behavior in other eublepharids, is displayed by males of G. lichtenfelderi during interactions with conspecifics of both sexes, suggesting an evolutionary shift in the meaning of this signal. In females, T induced growth of hemipenes and male-typical courtship but did not induce precloacal pore activity, aggression, or mounting. In comparison to previous reports on Eublepharis macularius, our results indicate that effects of T do not depend on the mode of sex determination. Further, our results extend our understanding of the complexity of control of male traits and illustrate how lability in the effects of T can be a general mechanism causing evolutionary changes in the components of suites of functionally correlated traits.

Preoptic dopamine release is integral to the display of copulatory behaviors in male mammals and birds. However, while the anatomical distributions of the dopamine synthesizing enzyme tyrosine hydroxylase are similar among vertebrates, evolutionary changes in the functional role of dopamine are poorly understood. In this study, we tested whether a dopamine D1 receptor agonist would facilitate the display of courtship and copulatory behaviors in two related Cnemidophorine lizards (Cnemidophorus inornatus and Cnemidophorus uniparens). Cnemidophorus lizards offer a unique system to study evolutionary changes in functionality because ancestral (e.g., C. inornatus) and descendant (e.g., C. uniparens) species can be studied in parallel. Cnemidophorus uniparens is an all-female, parthenogenetic species and is the triploid descendant of the sexual and diploid species C. inornatus. Here we report that in castrated male C. inornatus and ovariectomized C. uniparens a dopamine D1 agonist increased the proportion of individuals mounting and decreased the latency to mount. Moreover, there was a species difference in sensitivity to the agonist: Mounting was elicited at a lower dose in C. uniparens than in C. inornatus. One possible explanation for this heightened sensitivity in the triploid parthenogen is that, by virtue of the increased ploidy, the parthenogen has elevated levels of D1 receptor in limbic brain areas modulating courtship behavior.

In many species of birds, males perform complex visual and acoustic courtship displays to attract and stimulate females. Some of these displays involve considerable use of the wings and legs, suggesting that they may be controlled by sexually dimorphic spinal motoneurons and their target muscles. Sex steroid hormones are known to organize and activate many sexually dimorphic phenotypes, so these neuromuscular systems may also be steroid sensitive. To test these ideas, we have begun studies of wild golden-collared manakins ( Manacus vitellinus ) in Central America. Males of this species establish a courtship arena in the forest, where they perform an elaborate dance that includes use of their wings to generate loud snapping sounds. Here we describe male golden-collared manakin courtship behavior, including the various “wingsnaps.” We also review our studies, and those of others, showing sexually dimorphic properties of manakin wings, the wing musculature, and sex steroid accumulation in the spinal cord. These data suggest that manakins are useful models for evaluating steroid control of complex peripheral neuromuscular systems.

We examined the proximate causes of delayed breeding and alloparental behavior in the cooperatively breeding red-cockaded woodpecker by measuring plasma concentrations of testosterone (T) and prolactin (PRL) in female breeders, male breeders, and male helpers during different stages of the reproductive cycle. Male breeders and helpers have low T during the prebreeding period, highest T during copulation, and low concentrations of T during the egg-laying/incubation and nestling provisioning stages. Helpers appear physiologically capable of reproducing; their T concentrations equal that of male breeders. Helpers unrelated to the breeding female have higher T than helpers sharing a territory with their mother. Sexual inactivity by male helpers might be explained by behavioral suppression resulting from interactions of male helpers with the breeding pair that somehow differ in accordance with the helper's relatedness to the breeding female (e.g., female breeders are generally unreceptive to courtship from male helpers and least receptive to related helpers). Female breeder, male breeder, and male helper PRL levels did not differ and increased from the prebreeding stage through the copulation and egg-laying/incubation stages. During the nestling provisioning stage, male breeder and helper PRL declined, while female PRL continued to increase. Based on these results, we conclude that the physiological bases of alloparental behavior have not diverged from those that mediate parental behavior in this species.

During the breeding season, testosterone in male birds is often linked to some secondary sexual ornaments, courtship behaviors, and intrasexual aggression. I examined the effect of castration on plumage expression in Gambel's Quail (Callipepla gambelii), a species in which males are highly ornate, and in Scaled Quail (C. squamata), an unornamented species. Using male pairs, each consisting of a castrate and a control, I also assessed whether castration affected (1) the behavior of males, (2) the mating decisions of females, or (3) the outcome of male-male competition. Castration did not alter the plumage of male Gambel's or Scaled Quail. In these species, and some other members of the avian order Galliformes, production of ornate plumage appears to be independent of testosterone. In contrast, castration reduced or eliminated courtship behaviors. Females almost never preferred castrated individuals. During male-male competition, castrates also exhibited lower rates of threat behaviors, which appear to be identical to those used during courtship. Castration did not, however, influence the outcome of male-male competition. Castrates of both species exhibited overt aggression (pecks, chases, displacement) and frequently won male contests. Such results suggest that certain types of aggressive behavior may be testosterone-independent. In both Gambel's and Scaled Quail, male body size correlated positively with dominant individuals.

In male starlings (Sturnus vulgaris) courtship song plays a critical role in mate attraction. During the breeding season courtship song occurs prior to copulation and appears to reflect male sexual arousal. Outside the breeding season starlings sing, but song appears unrelated to reproduction. The aromatization of testosterone (T), likely within the medial preoptic nucleus (POM), is critical for the expression of male sexual arousal. The present study was performed to determine whether seasonal changes in the POM might relate to seasonal changes in courtship singing behavior in male starlings. T concentrations, the volume of the POM, and aromatase within the POM were examined both during and outside of the breeding season in male starlings. Song was also recorded at these times both with and without a female present. The POM was largest and contained dense aromatase immunostaining only during the spring breeding season, when T concentrations were highest and males responded to a female with an increase in courtship song. Outside the breeding season the volume of the POM was small, T concentrations were low, and males displayed no changes in song expression in response to female conspecifics. Song bout length was positively related to POM volume, and males sang longer songs in spring. Only males with nestboxes in spring responded to a female, and the POM tended to be larger in these males, suggesting that nestbox possession might influence neuroplasticity within the POM. Overall, the findings suggest that T-dependent plasticity and aromatase activity within the POM might regulate courtship singing in a wild songbird.

The aromatization of testosterone (T) in the medial preoptic nucleus (POM) is known to regulate male courtship and sexual behaviors expressed prior to, and in anticipation of, copulation. Singing in male European starlings is used to attract mates prior to physical sexual contact, suggesting that the POM might be involved. The present study was performed to examine the effects of lesions targeting the POM on singing and courtship behavior in reproductively active male starlings. A significant decrease in song output and the gathering of green nest materials was observed in males with lesions to the POM compared to males with damage to brain areas outside of the POM. Lesions did not affect a male's tendency to remain near a female or to occupy a nestbox, suggesting that the effects of POM lesions were specific to courtship behaviors. Behavioral differences were not related to testis mass or volume, and GnRH immunoreactivity was observed within the hypothalamus and median eminence for each male, suggesting that the effects of POM lesions were related specifically to POM involvement in song expression rather than to a disruption of the GnRH axis. These results suggest a general role for the POM in the expression of behaviors related to sexual arousal or anticipation, including song.

This study reports the gender and seasonal specificity of hormonal, behavioral, and coloration responses displayed by "resident" male lizards (Sceloporus undulatus) exposed to male or female "intruders" during staged encounters in outdoor enclosures. Resident males were engaged in staged encounters with males or females for 1 h per day on 9 consecutive days during the breeding and postbreeding seasons. Male-specific responses occurred during the breeding but not the postbreeding season. These included (1) a transient increase in plasma testosterone (T) that was evident on Day 4 and had subsided by Day 10, (2) behavioral displays of aggression (full shows and chases), and (3) a lightening of dorsal integumental color. Female-specific behavioral responses (nod sets) were displayed in both seasons. Season-specific responses consisted only of a transient increase in plasma corticosterone (B) during the breeding season that was evident on Day 4 and had subsided by Day 10. Pushups were displayed in response to both genders during both seasons, although the frequency of pushups was significantly higher in response to females than to males during the postbreeding season. The coloration of residents did not change in response to male intruders during the postbreeding season or to females during either season. These results define the gender and seasonal specificity of hormonal, behavioral, and coloration responses of resident male S. undulatus in social interactions with conspecifics. Thus, our results clarify the biological significance of these responses in terms of potentially aggressive versus courtship interactions and breeding versus postbreeding contexts.

In cricket frogs, Acris crepitans, sexually active males can switch between calling and noncalling (satellite) mating strategies and injections of the neuropeptide arginine vasotocin (AVT) stimulate calling behavior. We report here that this behavioral variation of animals under field conditions is associated with variations in AVT-immunoreactive (AVT-ir) staining in distinct brain nuclei. In both calling and satellite males, one AVT-ir brain region was found in a continuous string of cells between the medial amygdala and the nucleus accumbens (ACC). Satellite males possessed significantly more AVT-ir staining in the brain (cells and fibers) than calling males at the level of the ACC, although not in the medial amygdala. This difference in AVT-ir staining in the ACC can, in part, be explained by differences in the density of staining within the cells and in cell size. In addition, satellite males had significantly higher AVT-ir staining in the fibers medial to the ACC than calling males. Because other studies have demonstrated that AVT stimulates calling behavior, a plausible hypothesis is that calling males are releasing more AVT from neurons in the ACC, depleting reserves within the cells, and that the released AVT elicits calling behavior. AVT immunoreactivity levels are also higher in the ACC in both calling and satellite males than in female cricket frogs, which do not call. Satellite males may therefore have AVT reserves that might allow them to call depending on the social conditions.

Recent studies have demonstrated that physiological doses of progesterone may facilitate the androgen-dependent display of male sexual behavior in laboratory rats and three species of lizard. We used mice with a targeted disruption of the progesterone receptor to investigate whether such interactions exist in male mice and whether they may be modified by sexual experience. We found that naive intact male progesterone receptor knockout (PRKO) mice exhibit reduced mount frequencies compared to wild-type (WT) mice. Also unlike WT mice, sexually experienced PRKO males show profound losses in many measures of sexual behavior following castration. In a second experiment, we tested whether male mice heterozygous for a null mutation at the progesterone receptor locus were responsive to testosterone and progesterone treatment. We found that heterozygous males showed a reduced response to testosterone. The data are consistent with experiments indicating that the progesterone receptor is able to facilitate male-typical sex behaviors in other species and suggest novel mechanisms underlying the interaction of androgens and experience.

Adult males of African weakly discharging electric fish (family: Mormyridae) are distinguished from juveniles and adult females by a dorsally directed indentation of the posterior ventral body wall and by massive bone expansion of the bases of a select number of anal-fin rays. These sexually dimorphic structures seem to facilitate the anal-fin reflex that is displayed during courtship when the male envelopes its anal fin around the female's to form a common spawning pouch. Expanded bone could provide additional surface for muscle attachment and thus assist in part with the courtship sequence. Based on the fact that the expression of the male sexually dimorphic electric organ discharge (EOD) is under androgen control, and that the female EOD can be masculinized through testosterone administration, we hypothesized that androgens should also drive anal-fin ray bone expansion in male mormyrids and equally effect male-like changes in treated juveniles and adult females. Exogenous androgen treatment (17alpha-methyltestosterone) of adult female Brienomyrus niger resulted in a male-like EOD, and male-typical structural transformations (body wall indentation and anal-fin ray bone expansion). Some of these changes were immediate and receded following hormone withdrawal (EOD), while others developed more slowly and were apparently permanent (indentation and bone formation). 17alpha-Methyltestosterone administration affected only those targets in females that are normally involved in the male's reproductive behavior, i.e., its courtship signal (EOD) and two morphological features (body-wall indentation and bone expansion). Rays of the dorsal or caudal fins were never affected.

Both arginine vasotocin (AVT) and gonadotropin-releasing hormone (GnRH) are known to influence sexual behavior in many vertebrate taxa. We investigated the effects of both of these peptides on two different sexual behaviors, calling and amplexus, in the Great Plains toad (Bufo cognatus). AVT, at a dosage of 100 μg/100 g toad, significantly increased both the amount of calling behavior per individual and the probability that an individual would call. GnRH, however, had no effect on calling behavior. There was a reciprocal effect of these peptides on amplexus: AVT did not induce amplexus, while GnRH significantly induced this behavior. Furthermore, AVT-induced calling could be inhibited by Manning compound (an arginine vasopressin receptor antagonist). This is the first report of GnRH influencing sexual behavior in an anuran amphibian. These results suggest that specific sexual behaviors inB. cognatusmay be under the regulation of independent peptide control.

As in many mammalian and avian species, testicular androgens or their metabolites activate courtship and copulatory behaviors in adult male zebra finches. However, studies of sexual differentiation of these behaviors and related anatomical structures provide conflicting results. For example, posthatching estradiol can both masculinize courtship and the neural structures involved in song in females and inhibit the development of masculine copulation in males. These and other results have led to the hypotheses that (1) testicular androgens are converted to estradiol in the brain of developing males, and estradiol serves to masculinize the song system, whereas (2) estradiol secretion by the female ovary allows feminine rather than masculine copulatory behavior to develop. Treating embryonic zebra finches with the estrogen synthesis inhibitor fadrozole causes functional testicular tissue to develop in genetic females. The present study investigated the effects of such treatment on the development of singing and copulatory behavior as well as song system anatomy in males and females. While exogenous testosterone facilitated the display of sexual behaviors in adult males, the testicular tissue in females had no masculinizing effect on the production of audible courtship song or copulation. Their song control nuclei were also not masculinized, even in individuals lacking ovarian tissue. In contrast, embryonic inhibition of estrogen synthesis in males significantly stimulated song production. These results suggest that while manipulations of steroid hormone exposure can influence the display of sexual behaviors, gonadal secretions may not be required for normal sexual differentiation of the song system in zebra finches.

The relationship between courtship ultrasound emission rates and the volume of a discrete, sex-related, hypothalamic nucleus, the sexually dimorphic area, pars compacta (SDApc), in male and neonatally androgenized female gerbils is lateralized. Unbiased stereological estimates of neuron number and nuclear and neuropil volume are also laterally asymmetric in male SDApcs. In this study sexual differentiation and lateral asymmetry of stereologically assessed cytoarchitectural SDApc components, and their relationship to male-typical behaviors, including vocal emission, were examined in masculinized females. Female neonates received a single injection of testosterone propionate (TP) or control vehicle (Control) and were then implanted with silastic cannulae of testosterone at 65 days of age. Total SDApc volume, neuron number, nuclear volume, and neuropil volume had significantly greater values in TP compared to Control females. Neuron number was laterally asymmetric in TP females, since the left SDApc contained a greater number of smaller neurons, possibly interneurons, than the right. Courtship vocal emission and two other behaviors were masculinized in TP females. Left SDApc total volume and, most significantly, left neuron number, were correlated with vocal rates. No other lateralized correlations between behaviors and stereological estimates were found. It was concluded that various stereological parameters and the lateralization of vocal behavior and brain asymmetry depend on the early sexually differentiating effects of androgens. It is suggested that in gerbils, androgens have a role in the survival of interneurons in a laterally asymmetric hypothalamic nucleus which is an index of vocal control.

Previous studies found that pinealectomy of male Canadian red-sided garter snakes (Thamnophis sirtalis parietalis) in the autumn, before prolonged exposure to low temperatures (hibernation), significantly impaired the expression of courtship behavior upon emergence in the spring. Additionally, pinealectomized animals with a disrupted diel cycle of plasma melatonin did not court while those exhibiting a more typical diel pattern did. These results suggested that the pineal gland functions in the transduction of a temperature cue which stimulates courtship. To test this hypothesis, we pinealectomized males in the spring after they had undergone a normal hibernation but were still courting. Pinealectomy of courting males in the spring, in each of the 3 years of study, had no effect on courtship. This result suggests that once the cue is transduced, the pineal gland no longer has a modulatory effect on courtship behavior. Finally, we took advantage of the fact that, in the laboratory, there is always a small percentage of males that do not court upon emergence. Pinealectomy of these noncourters greatly increased the percent of males expressing courtship behavior in each of the study years. Plasma melatonin levels of unmanipulated courting and noncourting males was measured after emergence in successive years. In both years, courters had a typical pattern of melatonin secretion (low in the photophase, high in the scotophase) while persistent noncourters displayed the opposite pattern.

Aggressive and sexual behavior in the adult leopard gecko (Eublepharis macularius), a species with temperature-dependent sex determination (TSD), is influenced by the temperature experienced as an egg, as well as by prenatal and perinatal hormones. This study focused on the effects of hormonal manipulation of adult female leopard geckos from different incubation temperatures. Following ovariectomy, females from both all-female (26 degrees C) and male-biased (32.5 degrees C) incubation temperatures exhibited a significant decrease in high-posture (HP) aggression toward male and female stimulus animals. Testosterone treatment attenuated this decrease in HP aggression toward female but not toward male stimulus animals. Ovariectomy also resulted in a loss in attractiveness in both groups of females. Following treatment with testosterone, over 50% of the females were attacked by male stimulus animals, suggesting a change in the pheromonal cues normally secreted by females. Unmanipulated females never exhibit tail vibrations, a male-typical courtship behavior. However, following ovariectomy with testosterone treatment, half of the females from both incubation temperatures exhibited this behavior, indicating an activational effect of testosterone. An effect of incubation temperature on aggression was evident with females from the male-biased incubation temperature exhibiting a greater likelihood of aggression compared to females from the all-female incubation temperature. This effect continued to be detected after hormone manipulation. Ovariectomized females from the all-female incubation temperature were less aggressive even with testosterone treatment toward males, whereas females from the male-biased incubation temperature showed no significant decline in aggression following testosterone treatment, suggesting that individuals from different incubation temperatures may have different sensitivities to hormones.

Testosterone, 17 beta-estradiol, prostaglandin E2 (PGE2), and prostaglandin F2 alpha (PGF2alpha) were monitored in plasma and in vitro brain incubations during the main phases of male Triturus carnifex courtship. Plasma and in vitro results were similar: testosterone was highest in inactive males, while 17 beta-estradiol and PGF2 alpha were highest and PGE2 lowest during approach. PGF2 alpha in vitro treatments decreased testosterone and increased 17 beta-estradiol, while 9-ketoreductase inhibitor in vitro treatments increased testosterone and decreased estradiol. In addition, in vitro aromatase and PGE2-9-ketoreductase activities were highest during approach. PGF2 alpha in vitro treatments increased aromatase, while 9-ketoreductase inhibitor decreased aromatase. These results suggest that 9-ketoreductase and PGF2 alpha enhance aromatase activity to trigger male Triturus carnifex courtship.

Blood samples were taken from free-living male and female mockingbirds to determine correlations between plasma steroids and breeding behavior. Plasma levels of testosterone (T), estradiol (E2), progesterone (P), and corticosterone (B) were assessed during pre-breeding courtship, while males built nests, while females incubated, while parents fed young, and while males built new nests as pairs continued to care for dependent fledglings, Samples were also taken from unmated males holding spring territories. In males, T and P fluctuated with breeding phase, and in all groups, male T levels exceeded basal values measured in autumn. But, when mated males built nests in the absence of young, T levels were higher than in all other conditions, though only for early broods. In females too, T and P levels changed with breeding phase; T levels rose during courtship and while males built first nests and were intermediate during nest construction for later broods. Male and female P levels rose during courtship and when males built nests with young present. While feeding young in the nest, males showed significantly higher corticosterone levels than females, though in females, both P and B levels increased during incubation for replacement broods. These findings strengthen an earlier finding that T may activate male nest construction, and they suggest that breeding behavior may be related to T in females as well. Progesterone data raise the possibility that P too, may be linked to breeding behavior in male and female mockingbirds. Sex differences in corticosterone levels during the nestling period may reveal subtle costs associated with male mockingbirds' highly aggressive defense against predators.

Plasma levels of 11-ketotestosterone, 11-beta-OH-testosterone, testosterone, and 17-beta-estradiol were measured in reproductive Porichthys notatus, a teleost fish with two male morphs and alternative reproductive tactics. The two male types had contrasting androgen profiles. 11-Ketotestosterone was the predominant androgen in the Type I male morph which acoustically courts females, excavates nests, and guards eggs. Yet testosterone was predominant in the plasma of the Type II male morph which neither courts females nor nests, but instead parasitizes Type I males with sneak or satellite spawning tactics. The Type I-Type II male divergence in reproductive tactics and androgen levels is paralleled by dimorphisms in the vocal system, in body size, and in relative testis size. A review of endocrine data from six different species with male dimorphism shows consistent differences between morphotypes and a striking pattern: 11-ketotestosterone levels are uniformly elevated in each "courting male" morphotype relative to its "noncourting" conspecific. This cross-species pattern may reflect the behavioral, gonadal, or morphological differences which characterize the two morphotypes. At this point, the morphological interpretation is favored.

To clarify the role of corticosterone in the male and female crested newt, Triturus carnifex, and the possible involvement of this hormone in the reproductive processes, we monitored the seasonal and daily pattern of this corticosteroid; in addition, corticosterone, testosterone, and 17 beta-estradiol plasma levels were evaluated during the various phases of courtship. In both sexes, the seasonal pattern of corticosterone in plasma showed two peaks: one in winter, at the beginning of the reproduction, and the other one in summer, when the newts leave the pond. The corticosterone daily pattern showed a peak at 05:00 PM. During courtship, corticosterone was lower and testosterone higher in "inactive" male newts than in the courting animals, while estradiol was highest during the approach; in the females corticosterone was lower in "nonreceptive" animals than in "receptive" ones, and sex steroids did not show any difference. These data suggest that corticosterone is involved in the reproductive processes in male and female crested newts.

Endocrine disrupting chemicals (EDCs) are a widely studied group of chemicals that interfere with the endocrinology of organisms. So far, few studies have demonstrated the effect of EDCs on the reproductive behavior of aquatic wildlife. Here we show that sand goby males' ( Pomatoschistus minutus ) success in mating competition greatly decreases after an exposure for 7 to 24 days to 17α-ethinyl estradiol (EE2, measured concentration 4 ng L − 1 ). The sand goby exhibits a polygynous mating system with male parental care, in which males compete for nest sites and females. The aim of this study was to test how EE2 exposure affects the ability of males to compete for breeding resources, i.e. nest sites and mates. First, EE2 exposed males competed over a nest site against a non-exposed, control male of the same size. Secondly, we examined male courtship behavior and female mate preferences for EE2 exposed males and similar-sized non-exposed, control males. In addition to the behavioral experiments we determined the zona radiata protein (Zrp) mRNA gene expression and measured morphometric indicators of sexual maturation. Our study revealed that EE2 treated males were not able to acquire or defend a nest site. Additionally, EE2 treated males spent significantly less time in active courtship and nest leading behavior than control males. As a result, females clearly preferred to mate with control males. However, we found no significant differences in Zrp mRNA expression or the morphometric indicators between treatments. Our study illustrates that exposure to this EDC can greatly reduce the chances of an individual reproducing successfully. Moreover, it demonstrates that severe behavioral effects can be seen before any effects are detectable at the molecular or morphometric level.

A sequence of behaviours which we call courtship initiates reproduction in a large number of species. In vertebrates, as a component of male sexual behaviour courtship is strongly influenced by testicular androgen. Here I will review some salient issues about the regulation of courtship by testosterone in birds. The first section will briefly summarize the first 100 years of research on this topic. The specific role of testosterone or its oestrogenic metabolites in the control of different components of courtship will be the subject of the second section. Then, I will discuss how behavioural patterns can be recruited into courtship and modified in their structure by testosterone action. In the following section, the role of sexual selection and female choice in shaping the link between testosterone and courtship will be addressed. The problematic nature of the quantitative relationships between testosterone and behaviour will be topic of the fifth section. Finally, I will discuss how courtship traits that are activated by testosterone can be apparently independent of hormone blood concentrations. These issues will be examined in an evolutionary perspective, in an attempt to understand how natural and sexual selection have shaped the links between the hormone and the behaviour.

Crested auklets emit a citrus-like odorant that is seasonally modulated, suggesting that it is a secondary sexual trait. We hypothesized that expression of the chemical odorant is facilitated by steroid hormones, similar other secondary sexual traits in birds. Therefore we examined variation in concentrations of hormones in blood plasma and odor production during incubation and early chick rearing. A novel method was used to obtain and measure chemical emissions of crested auklets. Blood plasma samples were analyzed by radioimmunoassay. Progesterone was detected in all birds, and it varied during the breeding season. Octanal emissions covaried with progesterone levels in males but not in females. No seasonal patterns were detected in testosterone, estrogen or DHT, and these hormones were not detected in all breeding adults. Covariance of progesterone and octanal emissions in males suggests there could be at least an indirect relationship between odor emissions and steroid hormones in this species. Thus expression of the citrus-like odorant of crested auklets, like other secondary sexual traits in birds, could be regulated by steroid hormones.

Energy consumption is critical for the energetically expensive processes related to reproduction, and thus, mechanisms that increase ingestive behavior are directly linked to reproductive success. Similarly, the mechanisms that inhibit hunger and ingestive behavior might be most adaptive when these mechanisms cause individuals to stop foraging, hoarding and eating in order to find and court potential mates. In the laboratory, ingestive behaviors are typically studied separately from reproductive behaviors even though it is likely that these behaviors evolved under conditions in which both food and mates were available. We examined the choice between paracopulatory and ingestive behaviors in a semi-natural environment in which both food and potential mates were available. Intact female Syrian hamsters showed a high preference for males on days 3 and 4 (day 4 being the day of ovulation and estrous behavior), and a 48-h period of food deprivation significantly decreased preference for sex and increased preference for eating and food hoarding on day 3 in 89% of the hamsters, although none became anestrous. The same period of food deprivation significantly decreased the level of vaginal marking without significant effects on plasma estradiol concentrations. Next, hamsters were either food deprived (FD) or fed ad libitum, and half of each group was treated with vehicle or the adipocyte hormone leptin. The percentage of females with a low preference for sex was significantly greater in the FD compared to the ad libitum-fed groups, and leptin treatment prevented this effect. Metabolic fuels, possibly acting through leptin and other hormones, might influence sensitivity to estradiol or enhance the downstream effects of estradiol, thereby increasing motivation for sex and decreasing the relative motivation to forage, hoard and eat food.

In virtually every mammalian species examined, some males exhibit reflexive testosterone release upon encountering a novel female (or female-related stimulus). At the same time, not every individual male (or every published study) provides evidence for reflexive testosterone release. Four experiments using house mice (Mus musculus) examined the hypothesis that both the male's genotype and his degree of sexual arousal (as indexed by ultrasonic mating calls) are related to such variability. In Experiment 1, CF-1 males exhibited reflexive testosterone elevations 30 min after encountering female urine. CK males, on the other hand, did not exhibit testosterone elevations 20, 30, 50, 60, or 80 min after encountering female urine (Experiments 1 and 2) suggesting this strain incapable of reflexive release. In Experiment 3, we measured both mating calls and reflexive testosterone release in response to female urine in CF-1 and CK males. Most males of both strains called vigorously to female urine but not to water. But, only CF-1 males exhibited significant testosterone elevations to female urine. In Experiment 4, DBA/2J males called vigorously to females followed by testosterone elevations 30 min later. The first 3 experiments support the hypothesis that male genotype is an important variable underlying mammalian reflexive testosterone release. Statistically significant correlations between mating calls in the first minute after stimulus exposure and testosterone elevations 30 min later (Experiments 3 and 4) support the hypothesis that, in capable males, reflexive testosterone release is related to the male's initial sexual arousal.

The purpose of this study was to test whether sex steroid actions are necessary for courtship and pairing in socially monogamous birds. We examined the effects of an aromatase inhibitor, 1,4,6-androstatriene-3,17-dione (ATD), combined with an anti-androgen, flutamide (F), on the behavior and pairing status of initially unpaired male and female zebra finches (Taeniopygia guttata). In the first experiment, 24 adult males were implanted with either a combination of ATD and flutamide or empty implants. Two weeks after implantation, birds were housed in aviaries containing 3 ATD + F males, 3 control males, and 3 females and allowed 2 weeks to pair, with observations 7 times during the 2-week period. A second experiment tested the effects of these same treatments in females. During the first 4 days of testing, ATD + F males were less likely to attack conspecifics than were control males. ATD + F males were also less likely to "greet," or approach, females than were control males, but other courtship behaviors, including directed singing, were unaffected. ATD + F females did not differ from control females on any courtship behavior measured. Furthermore, these treatments did not affect pairing behaviors (time spent clumping or in a nest box together) or the likelihood of pairing with a partner of the opposite sex. ATD + F treatments in females did, however, increase the likelihood of same-sex pairing. This suggests that, although sex steroids may regulate some courtship behaviors in males, they do not regulate pairing behaviors and have little effect on the likelihood that a male or female will be chosen as a mate by a bird of the opposite sex.