Male Starlings Research Articles

Dehydroepiandrosterone (DHEA) increases undirected singing behavior and alters dopaminergic regulation of undirected song in non-breeding male European starlings (Sturnus vulgaris).

It has been proposed that in species that defend territories across multiple life history stages, brain metabolism of adrenal dehydroepiandrosterone (DHEA) regulates aggressive behavior at times when gonadal androgen synthesis is low (i.e. the non-breeding season). To date, a role for DHEA in the regulation of other forms of social behavior that are expressed outside of the context of breeding remains unknown. In this experiment, we used the European starling (Sturnus vulgaris) model system to investigate a role for DHEA in the neuroendocrine regulation of singing behavior by males in non-breeding condition. Starling song in a non-breeding context is spontaneous, not directed towards conspecifics, and functions to maintain cohesion of overwintering flocks. Using within-subjects design, we found that DHEA implants significantly increase undirected singing behavior by non-breeding condition male starlings. Given that DHEA is known to modulate multiple neurotransmitter systems including dopamine (DA) and DA regulates undirected song, we subsequently used immunohistochemistry for phosphorylated tyrosine hydroxylase (pTH, the active form of the rate-limiting enzyme in DA synthesis) to investigate the effect of DHEA on dopaminergic regulation of singing behavior in a non-breeding context. Pearson correlation analysis revealed a positive linear association between undirected singing behavior and pTH immunoreactivity in the ventral tegmental area and midbrain central gray of DHEA-implanted, but not control-implanted, males. Taken together, these data suggest that undirected singing behavior by non-breeding starlings is modulated by effects of DHEA on dopaminergic neurotransmission. More broadly, these data expand the social behavior functions of DHEA beyond territorial aggression to include undirected, affiliative social communication.

Night-brooding behaviour in provisioning cavity-nesting birds is a trade-off between adult predation risk and nestling thermoregulation needs

Night-brooding of nestlings in cavity-nesting avian species carries predation risks to parents. Anecdotally, several species are known to shift from constant adult night-brooding behaviour to leaving nestlings unattended at night during offspring development but the timing, speed of change and sex-specific differences between parents, and the factors shaping this behaviour have rarely been described. Moreover, the location and nature of night roosts used by adults whilst provisioning nestlings has received little research attention. We studied breeding Common Starlings Sturnus vulgaris and hypothesised that, in such a cavity-nesting species, 1) nestlings would only be night-brooded until they achieved thermal independence, 2) since the species is frequently polygynous, female parents would most likely exclusively night-brood offspring despite provisioning by both sexes and 3) night-brooding would be more likely during nights with lowest temperatures. Nightly video recordings throughout the nestling phase at eight Starling nests together with data from 18 Starlings fitted with GPS-loggers during 26 nights provided support for hypotheses 1) and 2), while we found no support for hypothesis 3). All tagged male Starlings always roosted far from the nesting site (up to 8 km) independent of nestling age; all females brooded nestlings, usually up to the first seven-nine days after hatching (when the nestlings achieve thermoregulation), but roosted with males after day 10, when all tagged Starlings from the same nesting ‘colony’ roosted together. These results confirm differential sex-related parental effort in provisioning Starlings, suggesting females only night-brood until young achieve homeothermy.

Immunolabeling Provides Evidence for Subregions in the Songbird Nucleus Accumbens and Suggests a Context-Dependent Role in Song in Male European Starlings (Sturnus vulgaris)

Birdsong is well known for its role in mate attraction during the breeding season. However, many birds, including European starlings (Sturnus vulgaris), also sing outside the breeding season as part of large flocks. Song in a breeding context can be extrinsically rewarded by mate attraction; however, song in nonbreeding flocks, referred to here as gregarious song, results in no obvious extrinsic reward and is proposed to be intrinsically rewarded. The nucleus accumbens (NAC) is a brain region well known to mediate reward and motivation, which suggests it is an ideal candidate to regulate reward associated with gregarious song. The goal of this study was to provide new histochemical information on the songbird NAC and its subregions (rostral pole, core, and shell) and to begin to determine subregion-specific contributions to gregarious song in male starlings. We examined immunolabeling for tyrosine hydroxylase (TH), neurotensin, and enkephalin (ENK) in the NAC. We then examined the extent to which gregarious and sexually motivated song differentially correlated with immunolabeling for the immediate early genes FOS and ZENK in each subdivision of the NAC. We found that TH and ENK labeling within subregions of the starling NAC was generally similar to patterns seen in the core and shell of NACs in mammals and birds. Additionally, we found that gregarious song, but not sexually motivated song, positively correlated with FOS in all NAC subregions. Our observations provide further evidence for distinct subregions within the songbird NAC and suggest the NAC may play an important role in regulating gregarious song in songbirds.

Perinatal exposure to antibiotics reduces affiliative behavior after post-weaning in zebra finches (Taeniopygia guttata)

Social behavior is influenced by a host of factors, including the immune system; for example, song quality in male starlings predicts immunocompetence suggesting the development of the immune system is interconnected with aspects social development (Duffy and Ball, 2002). Treating birds with antibiotics during the perinatal period may alter this development, and thereby, social behaviors beyond song. We asked if antibiotic exposure during the perinatal period effected parenting and offspring social behavior (e.g. aggressive and affiliative behaviors) in zebra finches? We treated the drinking water of zebra finch parents and hatchlings from post-hatch day 5–14 with azithromycin or a vehicle control and monitored parenting/social behavior. After weaning, we transferred offspring from the breeding cage to group housing and monitored social behavior and integration into the colony by measuring aggressive and affiliative behaviors. For all treatments we saw a reduction in the number of songs performed by fathers, however, specifically for antibiotic treated offspring there was a reduction in affiliative behaviors relative to vehicle treated controls suggesting the immune system, perhaps via the guts microbiome, influences certain aspects of social behaviors in birds.

Beak coloration of starling (Sturnus unicolor) males depends on the length of their throat feathers

Abstract Within the context of complex sexual signaling, most research has focused on exploring the associations between several signals and/or their relationships with different proxies of individual quality. However, very few studies have focused on checking whether the expression of one signal is conditioned by the expression of the others. Here, by experimentally shortening the throat feathers of male spotless starlings (Sturnus unicolor), we evaluated the influence of this trait on the color expression of the beak base. In addition, we tested the relationship between these two sexually dimorphic characters with traits indicating individual qualities, such as body condition and color reflectance at the wavelength related to carotenes in the tip of the beak. Our results show that the coloration of the beak base in males, but not in females, is positively related to body condition and to the length of ornamental throat feathers. Moreover, the experimental shortening of throat feathers in males had a negative effect on the blue chroma intensity of their beak base 1 year after manipulation. These results support for the first time a causal link between the expression of two sexually dimorphic characters, which is essential to understand their functionality in a multiple signaling framework.

Exploration and sociability in a highly gregarious bird are repeatable across seasons and in the long term but are unrelated

Personality traits and behavioural syndromes are often assumed to relate to life history strategies and lifetime fitness variation and hence may be generally under selection. Key in this regard is the, often untested, assumption that individual differences in (correlated) behaviours are maintained across contexts and over an individual's lifetime. Here, we tested this assumption, using a population of 30 captive male starlings, Sturnus vulgaris, a highly gregarious avian species. We repeatedly assayed novel environment exploration and different aspects of sociability towards a female conspecific, across seasonal contexts (spring and autumn) and across a 2-year period, which represents a substantial portion of a starling's life span. We found that, regardless of plasticity at the population level, both exploration behaviour and sociability traits investigated were moderately repeatable across seasons and years, with no significant differences between repeatability estimates over different timescales. However, no evidence was found for significant between-individual correlations between the investigated traits, including different aspects of sociability. Taken together, our results provide empirical evidence that exploration and sociability are personality traits that are stable across seasons and in the long term but do not form behavioural syndromes. Given the recent evidence that personality traits are often heritable, the traits assessed in our study might have the potential to evolve independently under selection. This long-term consistency in exploration and sociability might have important implications for the social organization within complex social environments and influence a wide variety of ecologically relevant processes.

Prodynorphin and kappa opioid receptor mRNA expression in the brain relates to social status and behavior in male European starlings

Numerous animal species display behavioral changes in response to changes in social status or territory possession. For example, in male European starlings only males that acquire nesting sites display high rates of sexual and agonistic behavior. Past studies show that mu and delta opioid receptors regulate behaviors associated with social ascension or defeat. Opioids also act at kappa receptors, with dynorphin binding with the highest affinity; however, the role of these opioids in social behavior has not been well studied. We observed flocks of male starlings during the breeding season and ran quantitative real-time polymerase chain reaction (qPCR) to measure expression of kappa opioid receptors (OPRK1) and prodynorphin (PDYN) in brain regions involved in social behavior and motivation (ventral tegmental area [VTA], medial preoptic nucleus [mPOA]) and vocal behavior (Area X). Males with nesting territories displayed more sexual/agonistic behavior than males without nesting territories. They also had lower OPRK1 expression in VTA and mPOA. OPRK1 expression in VTA correlated negatively with sexual/agonistic behaviors, consistent with past studies showing kappa receptors in VTA to inhibit sociosexual behaviors. PDYN in mPOA correlated negatively with a measure of nesting behavior that may also reflect sexual motivation. PDYN in Area X related positively to song. Distinct patterns of OPRK1 and PDYN expression in VTA, mPOA, and Area X related to gonad volume, suggesting that breeding condition may modify (or be modified by) OPRK1 and PDYN expression. Studies are now needed to further characterize the role of OPRK1 and PDYN in status-appropriate social behaviors.

Contributions of testosterone and territory ownership to sexually-motivated behaviors and mRNA expression in the medial preoptic area of male European starlings

Animals integrate social information with their internal endocrine state to control the timing of behavior, but how these signals are integrated in the brain is not understood. The medial preoptic area (mPOA) may play an integrative role in the control of courtship behavior, as it receives projections from multiple sensory systems, and is central to the hormonal control of courtship behavior across vertebrates. Additionally, data from many species implicate opioid and dopaminergic systems in the mPOA in the control of male courtship behavior. We used European starlings to test the hypothesis that testosterone (T) and social status (in the form of territory possession) interact to control the timing of courtship behavior by modulating steroid hormone-, opioid- and dopaminergic-related gene expression in the mPOA. We found that only males given both T and a nesting territory produced high rates of courtship behavior in response to a female. T treatment altered patterns of gene expression in the mPOA by increasing androgen receptor, aromatase, mu-opioid receptor and preproenkephalin mRNA and decreasing tyrosine hydroxylase mRNA expression. Territory possession did not alter mRNA expression in the mPOA, despite the finding that only birds with both T and a nesting territory produced courtship behavior. We propose that T prepares the mPOA to respond to the presence of a female with high rates of courtship song by altering gene expression, but that activity in the mPOA is under a continuous (i.e. tonic) inhibition until a male starling obtains a nesting territory.

Neural endocannabinoid CB1 receptor expression, social status, and behavior in male European starlings

Many species modify behavior in response to changes in resource availability or social status; however, the neural mechanisms underlying these modifications are not well understood. Prior work in male starlings demonstrates that status-appropriate changes in behavior involve brain regions that regulate social behavior and vocal production. Endocannabinoids are ubiquitously distributed neuromodulators that are proposed to play a role in adjusting behavior to match social status. As an initial step to provide insight into this hypothesis we observed flocks of male starlings in outdoor aviaries during the breeding season. We used quantitative real-time PCR to measure expression of endocannabinoid CB1 receptors in brain regions involved in social behavior and motivation (lateral septum [LS], ventral tegmental area [VTA], medial preoptic nucleus [POM]) and vocal behavior (Area X and robust nucleus of the arcopallium; RA). Males with nesting sites sang to females and displaced other males more than males without nesting sites. They also had higher levels of CB1 receptor expression in LS and RA. CB1 expression in LS correlated positively with agonistic behaviors. CB1 expression in RA correlated positively with singing behavior. CB1 in VTA also correlated positively with singing when only singing birds were considered. These correlations nicely map onto the well-established role of LS in agonistic behavior and the known role of RA in song production and VTA in motivation and song production. Studies are now needed to precisely characterize the role of CB1 receptors in these regions in the production of status-appropriate social behaviors.

Reward associated with singing behavior correlates with opioid-related gene expression in the medial preoptic nucleus in male European starlings.

Birdsong consists of species-specific learned vocal sequences that are used primarily to attract mates and to repel competitors during the breeding season. However, many birds continue to sing at times when vocal production has no immediate or obvious impact on conspecific behavior. The mechanisms that ensure that animals produce important behaviors in contexts in which the function of these behaviors is not immediate or obvious are not known. One possibility is that animals engage in such behaviors because they are associated with pleasure. Here we examined the hypothesis that male European starlings sing outside of the breeding season in part because the act of singing in this context is facilitated and/or maintained by opioid-mediated reward. We measured song-associated reward using a conditioned place preference (CPP) test in male starlings producing fall, non-breeding season-typical song. We used quantitative real time PCR to measure expression of the enkephalin opioid precursor preproenkephalin (PENK) and mu opioid receptors (MOR) in the medial preoptic nucleus (POM; a region in which opioids are implicated in both reward and starling fall song) and additionally the song control region HVC as a control. Starlings developed a strong preference for a place that had been paired previously with the act of producing fall-typical song, indicating that fall song production was associated with a positive affective state. Both PENK and MOR mRNA expression in the POM, but not HVC, correlated positively with both individual reward state (as reflected in CPP) and undirected singing behavior. These results suggest that singing induces opioid receptor and enkephalin expression in the POM and consequent reward, and/or that opioid release in the POM induced by individual or environmental factors (e.g., the presence of food, safety of a flock or the absence of predators) induces a positive affective state which then facilitates singing behavior.

Status-appropriate singing behavior, testosterone and androgen receptor immunolabeling in male European starlings (Sturnus vulgaris)

Vocalizations convey information about an individual's motivational, internal, and social status. As circumstances change, individuals respond by adjusting vocal behavior accordingly. In European starlings, a male that acquires a nest site socially dominates other males and dramatically increases courtship song. Although circulating testosterone is associated with social status and vocal production it is possible that steroid receptors fine-tune status-appropriate changes in behavior. Here we explored a possible role for androgen receptors. Male starlings that acquired nest sites produced high rates of courtship song. For a subset of males this occurred even in the absence of elevated circulating testosterone. Immunolabeling for androgen receptors (ARir) was highest in the medial preoptic nucleus (POM) in males with both a nest site and elevated testosterone. For HVC, ARir was higher in dominant males with high testosterone (males that sang longer songs) than dominant males with low testosterone (males that sang shorter songs). ARir in the dorsal medial portion of the nucleus intercollicularis (DM) was elevated in males with high testosterone irrespective of dominance status. Song bout length related positively to ARir in POM, HVC and DM, and testosterone concentrations related positively to ARir in POM and DM. Results suggest that the role of testosterone in vocal behavior differs across brain regions and support the hypothesis that testosterone in POM underlies motivation, testosterone in HVC relates to song quality, and testosterone in DM stimulates vocalizations. Our data also suggest that singing may influence AR independent of testosterone and that alternative androgen-independent pathways regulate status-appropriate singing behavior.

Nest Composition and Yolk Hormones: Do Female European Starlings Adjust Yolk Androgens to Nest Quality?

The nest is a key element of avian reproductive fitness. It provides the developmental environment for the embryo and nestling thereby affecting their quality and survival. Nests are often constructed by the male and nest quality, elaboration, and ornamentation vary among males in a species. Therefore, male nest-building behaviour is likely under sexual selection and females may use male nest-building behaviour and nest features as cues in mate choice and their own reproductive investment. We tested this hypothesis in European starlings ( Sturnus vulgaris). Male starlings present green plants to females during their courtship display and furbish their nests with variable quantities of fresh green plants. The highest frequency of plant incorporation into the nest occurs during pair formation and oogenesis. We determined nest composition and measured yolk testosterone concentrations in eggs. Yolk testosterone concentrations were positively correlated with the amount of green plant material in nests. Because males use green nest material in courtship and green nest material positively affects nestlings’ development, we suggest that female European starlings use male nest-building effort, ritualised courtship display of green plants, and nest characteristics in mate choice, and adjust yolk testosterone levels in the eggs to the quality of the nest to optimise offspring growth.

Curvilinear relationships between mu-opioid receptor labeling and undirected song in male European starlings (Sturnus vulgaris)

Female-directed communication in male songbirds has been reasonably well studied; yet, relatively little is known about communication in other social contexts. Songbirds also produce song that is not clearly directed towards another individual (undirected song) when alone or in flocks. Although the precise functions of undirected song may differ across species, this type of song is considered important for flock maintenance, song learning or practice. Past studies show that undirected song is tightly coupled to analgesia and positive affective state, which are both mediated by opioid activity. Furthermore, labeling for the opioid met-enkephalin in the medial preoptic nucleus (POM) correlates positively with undirected song production. We propose that undirected song is facilitated and maintained by opioid receptor activity in the POM and other brain regions involved in affective state, analgesia, and social behavior. To provide insight into this hypothesis, we used immunohistochemistry to examine relationships between undirected song and mu-opioid receptors in male starlings. Polynomial regression analyses revealed significant inverted-U shaped relationships between measures of undirected song and mu-opioid receptor labeling in the POM, medial bed nucleus of the stria terminalis (BSTm), and periaqueductal gray (PAG). These results suggest that low rates of undirected song may stimulate and/or be maintained by mu-opioid receptor activity; however, it may be that sustained levels of mu-opioid receptor activity associated with high rates of undirected song cause mu-opioid receptor down-regulation. The results indicate that mu-opioid receptor activity in POM, BSTm, and PAG may underlie previous links identified between undirected song, analgesia, and affective state.

Male European Starlings Do Not Use Egg Spots as a Cue to Adjust Investment in Nestlings

The effects of post-laying egg spottiness on nestling condition and parental provisioning were investigated in a nest box-breeding population of European Starlings (Sturnus vulgaris). Our objectives were to ascertain whether egg spottiness was associated with the nest ectoparasite Carnus hemapterus, and to examine potential relationships between egg spottiness, presence of C. hemapterus, nestling condition, and parental provisioning effort in European Starlings. Spotted-egg clutches were present over all 3 years in our population, but the spots did not reflect C. hemapterus abundance. Nestlings from spotted-egg clutches did not have more C. hemapterus than those from unspotted-egg clutches. However, nestlings from spotted clutches were in better condition than those from unspotted clutches. Nestling condition was not associated with C. hemapterus abundance. Adult male and female provisioning rates to the offspring did not differ between spotted and unspotted clutches. Similarly, the proportion of provisioning visits by males did not differ significantly between spotted and unspotted clutches, indicating that parents in our population of European Starlings do not use egg spots as a cue to altering their provisioning effort. Further research is required to fully understand the cause and consequences of egg spots in European Starlings. Received 29 September 2011. Accepted 25 September 2012.

Functional changes between seasons in the male songbird auditory forebrain

Songbirds are an excellent model for investigating the perception of learned complex acoustic communication signals. Male European starlings (Sturnus vulgaris) sing throughout the year distinct types of song that bear either social or individual information. Although the relative importance of social and individual information changes seasonally, evidence of functional seasonal changes in neural response to these songs remains elusive. We thus decided to use in vivo functional magnetic resonance imaging (fMRI) to examine auditory responses of male starlings that were exposed to songs that convey different levels of information (species-specific and group identity or individual identity), both during (when mate recognition is particularly important) and outside the breeding season (when group recognition is particularly important). We report three main findings: (1) the auditory area caudomedial nidopallium (NCM), an auditory region that is analogous to the mammalian auditory cortex, is clearly involved in the processing/categorization of conspecific songs; (2) season-related change in differential song processing is limited to a caudal part of NCM; in the more rostral parts, songs bearing individual information induce higher BOLD responses than songs bearing species and group information, regardless of the season; (3) the differentiation between songs bearing species and group information and songs bearing individual information seems to be biased toward the right hemisphere. This study provides evidence that auditory processing of behaviorally-relevant (conspecific) communication signals changes seasonally, even when the spectro-temporal properties of these signals do not change.

Modulation of male song by naloxone in the medial preoptic nucleus.

Studies in songbirds implicate opioid neuropeptides in singing behavior; however, past results are contradictory. In starlings, the effect of opioid manipulations on sexually motivated courtship song differed in birds naturally singing at low compared to high rates, and mu-opioid receptors were denser in several regions, including the medial preoptic nucleus (POM) in low singing males. In the present study, we found that low singing male starlings also had significantly higher enkephalin (ENK) immunolabeling densities in the POM than high singers. We blocked opioid receptor activity in the POM with naloxone injections and found that this increased both song rate and song bout length in low singers, suggesting that high densities of mu receptors and ENK in the POM actively suppress song in these males. In contrast to its effects on low singers, naloxone in the POM of high singers dose dependently decreased song rate and tended to reduce song bout length. This suggests that at least some level of opioid activity in POM is necessary for song production. Our results are the first to demonstrate that direct administration of naloxone into the POM influences sexually motivated song, and that effects differ depending on an individual's initial rate of song and associated density of ENK. We suggest that differential effects seen in past studies of opioids and song may in part be explained by differences in the natural song rate of subjects and accompanying differences in ENK activity and neural substrate sensitivity to opioids in POM.

Context-Dependent Links between Song Production and Opioid-Mediated Analgesia in Male European Starlings (Sturnus vulgaris)

Little is known about the neural mechanisms that ensure appropriate vocal behaviors within specific social contexts. Male songbirds produce spontaneous (undirected) songs as well as female-directed courtship songs. Opioid neuropeptide activity in specific brain regions is rewarding, at least in mammals, and past studies suggest that the opioid met-enkephalin in such areas is more tightly linked to undirected than female-directed song. Recent data using a song-associated place preference paradigm further suggest that production of undirected but not directed song is tightly linked to intrinsic reward. Opioids have analgesic properties. Therefore, if production of undirected song is closely linked to opioid-mediated reward, the production of undirected but not directed song should be associated with analgesia. Consistent with this prediction, in male starlings we identified a positive correlation between analgesia (decreased reactivity to a hot water bath) and undirected song (in non-breeding season condition males in affiliative flocks) but not female-directed song (in breeding season condition males presented with females). When breeding condition males were divided according to social status, a negative correlation was found in subordinate males (i.e. males that failed to acquire a nest box). These data are consistent with the hypotheses 1) that the production of undirected song is facilitated or maintained by opioids (and/or other neuromodulators that also induce analgesia) and 2) that production of female-directed song is not linked in the same way to release of the same neuromodulators. Results also demonstrate a link between analgesia and song in subordinate individuals lacking a nesting territory within the breeding season. Overall, the findings indicate that distinct neural mechanisms regulate communication in different social contexts and support the working hypothesis that undirected but not directed song is tightly linked to opioid release.

Reward and vocal production: Song-associated place preference in songbirds

Vocal production is crucial for successful social interactions in multiple species. Reward can strongly influence behavior; however, the extent to which reward systems influence vocal behavior is unknown. In songbirds, singing occurs in different contexts. It can be spontaneous and undirected (e.g., song produced alone or as part of a large flock) or directed towards a conspecific (e.g., song used to attract a mate or influence a competitor). In this study, we developed a conditioned place preference paradigm to measure reward associated with different types of singing behavior in two songbird species. Both male zebra finches and European starlings developed a preference for a chamber associated with production of undirected song, suggesting that the production of undirected song is tightly coupled to intrinsic reward. In contrast, neither starlings nor zebra finches developed a place preference in association with directed song; however, male starlings singing directed song that failed to attract a female developed a place aversion. Unsuccessful contact calling behavior was also associated with a place aversion. These findings suggest that directed vocal behavior is not tightly linked to intrinsic reward but may be externally reinforced by social interactions. Data across two species thus support the hypothesis that the production of undirected but not directed song is tightly coupled to intrinsic reward. This study is the first to identify song-associated reward and suggests that reward associated with vocal production differs depending upon the context in which communication occurs. The findings have implications for understanding what motivates animals to engage in social behaviors and ways in which distinct reward mechanisms function to direct socially appropriate behaviors.

DHEA and estradiol levels in brain, gonads, adrenal glands, and plasma of developing male and female European starlings

Traditionally, sexual differentiation of the brain was thought to be driven by gonadal hormones, particularly testosterone (T). However, recent studies in songbirds suggest that other steroids may also be important. For example, dehydroepiandrosterone (DHEA) can be synthesized by the gonads, adrenal glands, and/or brain and locally metabolized into T and 17β-estradiol (E(2)). Here, we examined DHEA and E(2) levels in the brain, peripheral tissues, and plasma of wild European starlings (Sturnus vulgaris). In Study 1, samples were collected from males and females at P0 (day of hatch), P6, and P8. In Study 2, samples were collected at P4. At P0, DHEA levels in the diencephalon were higher in males than females. DHEA levels were generally high in the gonads and adrenals, and they were higher in testes than ovaries at P8. Further, E(2) levels were non-detectable in most brain samples, suggesting that DHEA was not metabolized to E(2) or that locally produced E(2) was rapidly inactivated. At P4, DHEA levels in telencephalic regions were lower in males than females. Taken together, these data suggest that sex differences in peripheral DHEA secretion and neural DHEA metabolism at specific ages during development might play a role in sexual differentiation of the songbird brain.

α 1-Noradrenegic receptor antagonism disrupts female songbird responses to male song

Female songbirds respond behaviorally to differences in male song structure. Past data suggest a complex role for norepinephrine in female responses to song. Here, we examined the effects of central infusions of the α 1-noradrenergic receptor antagonist prazosin on female European starling ( Sturnus vulgaris) responses to nest boxes broadcasting male song. Prazosin disrupted female preferential responses to male starling song over the less biologically relevant purple martin ( Progne subis) song. Prazosin decreased female responses to male starling song in a linear dose–response fashion; whereas, it affected responses to purple martin song in a U-shaped dose–response fashion. Results suggest that the role of norepinephrine in female responses to male song differs depending upon drug dose and the biological relevance of the song stimulus.