Early life nutritional stress affects song learning but not underlying neural circuitry in zebra finches.

Early life stress negatively impacts behavior and underlying neural circuitry across species. The present study investigated the effects of nutritional stress (NS), which increases parental foraging, on song quality in males, song preferences in females, and the size and number of cells in song and auditory regions of the zebra finch brain. We hypothesized that NS would decrease song quality in males and decrease preference for high quality song in females. Furthermore, we predicted that NS would reduce the size of song nuclei and decrease the number of cells in song and auditory regions. In males, NS decreased the number of syllable types (a measure of song complexity), decreased the number of high notes and flat notes, and increased the number of noisy notes, but had no effect on song rate or song length. In females, NS reduced preferences for high quality song. Despite these behavioral effects, there were no effects of NS on song and auditory nuclei, although there were effects of age and sex. These results suggest that NS affects behavior in both males and females, but these effects are not attributable to the number of cells or size of song and auditory regions. Such findings add to our understanding of the effects of early life stress on behavior and cognition. (PsycInfo Database Record (c) 2020 APA, all rights reserved).

Acute exogenous corticosterone treatments have few effects on courtship and pair bonding in zebra finches.

Stress impacts social relationships. In turn, social relationships buffer the stress response in some species. Studies that have investigated the role of corticosterone (CORT) on courtship, mate choice, mating, and pairing have found mixed results. We therefore tested the role of CORT in these steps of the pairing process in the monogamous zebra finch. Male and female zebra finches received either one of 2 doses of corticosterone (CORT, 10 µg and 20 µg, referred to as low and high dose) or a vehicle control (peanut oil). Subjects were then given the opportunity to pair in mixed sex aviaries. Courtship and pair bonding behaviors were observed over 3 days. Overall, zebra finches of both sexes were equally likely to pair or not pair regardless of treatment, although a high dose of CORT increased the latency to form a pair bond. There were no effects of CORT on courtship behavior in either sex, though the low dose increased undirected (non-courtship) singing in males relative to the high dose. Animals treated with CORT, regardless of dose, engaged in fewer copulations than did control animals. When we examined pairing behaviors, we found a decrease in co-nesting in low dose animals. Our results suggest that acute CORT has few effects on pair bonding, suggesting species-specific effects of CORT on behavior.

Pairing Increases Activation of V1aR, but not OTR, in Auditory Regions of Zebra Finches: The Importance of Signal Modality in Nonapeptide-Social Behavior Relationships.

Social relationships are complex, involving the production and comprehension of signals, individual recognition, and close coordination of behavior between two or more individuals. The nonapeptides oxytocin and vasopressin are widely believed to regulate social relationships. These findings come largely from prairie voles, in which nonapeptide receptors in olfactory neural circuits drive pair bonding. This research is assumed to apply to all species. Previous reviews have offered two competing hypotheses. The work of Sarah Newman has implicated a common neural network across species, the Social Behavior Network. In contrast, others have suggested that there are signal modality-specific networks that regulate social behavior. Our research focuses on evaluating these two competing hypotheses in the zebra finch, a species that relies heavily on vocal/auditory signals for communication, specifically the neural circuits underlying singing in males and song perception in females. We have demonstrated that the quality of vocal interactions is highly important for the formation of long-term monogamous bonds in zebra finches. Qualitative evidence at first suggests that nonapeptide receptor distributions are very different between monogamous rodents (olfactory species) and monogamous birds (vocal/auditory species). However, we have demonstrated that social bonding behaviors are not only correlated with activation of nonapeptide receptors in vocal and auditory circuits, but also involve regions of the common Social Behavior Network. Here, we show increased Vasopressin 1a receptor, but not oxytocin receptor, activation in two auditory regions following formation of a pair bond. To our knowledge, this is the first study to suggest a role of nonapeptides in the auditory circuit in pair bonding. Thus, we highlight converging mechanisms of social relationships and also point to the importance of studying multiple species to understand mechanisms of behavior.

Early life stress increases testosterone and corticosterone and alters stress physiology in zebra finches.

Early life stress has enduring effects on behavior and physiology. However, the effects on hormones and stress physiology remain poorly understood. In the present study, parents of zebra finches of both sexes were exposed to an increased foraging paradigm from 3 to 33days post hatching. Plasma and brains were collected from chicks at 3 developmental time points: post hatching days 25, 60 and adulthood. Plasma was assayed for testosterone (T), estradiol (E2), and corticosterone (CORT). The paraventricular nucleus of the hypothalamus was assessed for corticotrophin releasing factor (CRH) and glucocorticoid receptor (GR) expression. As expected, body mass was lower in nutritionally stressed animals compared to controls at multiple ages. Nutritionally stressed animals overall had higher levels of CORT than did control and this was particularly apparent in females at post hatching day 25. Nutritionally stressed animals also had a higher number of cells expressing CRH and GR in the paraventricular nucleus of the hypothalamus than did controls. There was an interaction, such that both measures were higher in control animals at PHD 25, but higher in NS animals by adulthood. Females, regardless of treatment, had higher circulating CORT and a higher number of cells expressing CRH than did males. Nutritionally stressed animals also had higher levels of T than did control animals, and this difference was greatest for males at post hatching day 60. There were no effects of nutritional stress on E2. These findings suggest that nutritional stress during development has long-lasting effects on testosterone and stress physiology.

Sex and pairing status explain variations in the activation of nonapeptide receptors in song and motivation regions.

The nonapeptides oxytocin and vasopressin have been implicated in a variety of social behaviors. In zebra finches, oxytocin antagonists decrease pairing in both sexes, and pairing, in turn, increases expression of both mesotocin (the avian homologue of oxytocin) and vasotocin (the avian homologue of vasopressin). Increases in mesotocin and vasotocin mRNA are correlated with the amount of directed singing by males. Thus, in the present study, we examined the hypothesis that activation of cells containing nonapeptide receptors in song-related regions (ventral tegmental area, lateral septum, and medial preoptic nucleus) would also be correlated with directed singing in males. To rule out the possibility that these regions are involved in general pairing motivation, we also included females as subjects. In the ventral tegmental area, males had higher ZENK and V1aR than females and paired animals (regardless of sex) had higher ZENK and V1aR than did unpaired animals. In the medial preoptic nucleus, paired animals had higher ZENK than did unpaired animals, and there were no sex or pairing effects in the lateral septum. Only ZENK + V1aR in the medial preoptic nucleus was correlated with singing in males. These findings suggest that pairing is associated activation of nonapeptide receptors in the ventral tegmental area and the medial preoptic nucleus, but there is only partial evidence that courtship singing accounts for these findings. (PsycINFO Database Record

Early Life Manipulations of the Nonapeptide System Alter Pair Maintenance Behaviors and Neural Activity in Adult Male Zebra Finches.

Adult zebra finches (T. guttata) form socially monogamous pair bonds characterized by proximity, vocal communication, and contact behaviors. In this experiment, we tested whether manipulations of the nonapeptide hormone arginine vasotocin (AVT, avian homolog of vasopressin) and the V1a receptor (V1aR) early in life altered species-typical pairing behavior in adult zebra finches of both sexes. Although there was no effect of treatment on the tendency to pair in either sex, males in different treatments exhibited profoundly different profiles of pair maintenance behavior. Following a brief separation, AVT-treated males were highly affiliative with their female partner but sang very little compared to Controls. In contrast, males treated with a V1aR antagonist sang significantly less than Controls, but did not differ in affiliation. These effects on behavior in males were also reflected in changes in the expression of V1aR and immediate early gene activity in three brain regions known to be involved in pairing behavior in birds: the medial amygdala, medial bed nucleus of the stria terminalis, and the lateral septum. AVT males had higher V1aR expression in the medial amygdala than both Control and antagonist-treated males and immediate early gene activity of V1aR neurons in the medial amygdala was positively correlated with affiliation. Antagonist treated males showed decreased activity in the medial amygdala. In addition, there was a negative correlation between the activity of V1aR cells in the medial bed nucleus of the stria terminalis and singing. Treatment also affected the expression of V1aR and activity in the lateral septum, but this was not correlated with any behaviors measured. These results provide evidence that AVT and V1aR play developmental roles in specific pair maintenance behaviors and the neural substrate underlying these behaviors in a bird.

Temporary inactivation of NCM, an auditory region, increases social interaction and decreases song perception in female zebra finches.

The caudomedial nidopallium (NCM) is an important site for the storage of auditory memories, particularly song, in passerines. In zebra finches, males sing and females do not, but females use song to choose mates. The extent to which the NCM is necessary for female mate choice is not well understood. To investigate the role of NCM in partner preferences, adult female zebra finches were bilaterally implanted with chronic cannulae directed at the NCM. Lidocaine, a sodium channel blocker, or saline (control) was infused into the NCM of females using a repeated measures design. Females were then tested in 3 separate paradigms: song preference, sexual partner preference, and pairing behavior/partner preference. We hypothesized that lidocaine would increase interactions with males by decreasing song discrimination and that this would be further evident in the song discrimination task. Indeed, females, when treated with lidocaine, had no preference for males singing unaltered song over males singing distorted song. These same females, when treated with saline, demonstrated a significant preference for males singing normal song. Furthermore, females affiliated with males more after receiving lidocaine than after receiving saline in the pairing paradigm, although neither treatment led to the formation of a partner preference. Our results support the hypothesis that NCM plays an important role not only in song discrimination, but also affiliation with a male.

Same-sex partner preference in zebra finches: pairing flexibility and choice.

This study examined flexibility and choice in same-sex pair-bonding behavior in adult zebra finches (Taeniopygia guttata). Zebra finches form life-long monogamous relationships and extra pair behavior is very low, making them an ideal species in which to study same-sex pairing. We examined same-sex behaviors using both semi-naturalistic choice paradigms and skewed sex ratios. In the first experiment, we allowed zebra finches to pair in aviaries with equal sex ratios as part of multiple experiments. On average, 6.4% (N = 78) of unmanipulated pairs were same-sex: all but one was female-female. In a second experiment, we identified pairs from same-sex cages and selected 20 total same-sex pairs (10 of each sex). We then gave pairs a chance to court and pair with members of the opposite sex and observed their behavior for three days. Females did not retain their partner, but most paired with males. In contrast, some males did retain their partner. Similarly, females were more likely to engage in pairing behaviors with males than with their partners or other females whereas males were equally likely to engage in same-sex and opposite-sex pairing behaviors. These findings suggest that same-sex partnerships in zebra finches can be facultative, based on the sex ratio of the group in which they live, but can also be a choice, when opportunities to pair with opposite-sex individuals are possible. Furthermore, it is possible that females are more flexible in this choice of same-sex partnerships than are males.

N-methyl-D-aspartate receptor agonism and antagonism within the amygdaloid central nucleus suppresses pain affect: differential contribution of the ventrolateral periaqueductal gray.

UNLABELLED: The amygdala contributes to the generation of pain affect, and the amygdaloid central nucleus (CeA) receives nociceptive input that is mediated by glutamatergic neurotransmission. The present study compared the contribution of N-methyl-d-aspartate (NMDA) receptor agonism and antagonism in the CeA to generation of the affective response of rats to an acute noxious stimulus. Vocalizations that occur following a brief tail shock (vocalization afterdischarges) are a validated rodent model of pain affect and were preferentially suppressed, in a dose-dependent manner, by bilateral injection into the CeA of NMDA (.1, .25, .5, or 1 µg/side) or the NMDA receptor antagonist d-(-)-2-amino-5-phosphopentanoic acid (AP5; 1, 2, or 4 µg/side). Vocalizations that occur during tail shock were suppressed to a lesser degree, whereas spinal motor reflexes (tail flick and hind limb movements) were unaffected by injection of NMDA or AP5 into the CeA. Injection of NMDA, but not AP5, into the CeA increased c-Fos immunoreactivity in the ventrolateral periaqueductal gray, and unilateral injection of the µ-opiate receptor antagonist H-d-Phe-Cys-Tyr-d-Trp-Arg-Thr-Pen-Thr-NH2 (CTAP; .25 µg) into ventrolateral periaqueductal gray prevented the antinociception generated by injection of NMDA into the CeA. These findings demonstrate that although NMDA receptor agonism and antagonism in the CeA produce similar suppression of pain behaviors, they do so via different neurobiologic mechanisms. PERSPECTIVE: The amygdala contributes to production of the emotional dimension of pain. NMDA receptor agonism and antagonism within the CeA suppressed rats' emotional response to acute painful stimulation. Understanding the neurobiology underlying emotional responses to pain will provide insights into new treatments for pain and its associated affective disorders.

Effects of acute corticosterone treatment on partner preferences in male and female zebra finches (Taeniopygia guttata).

Stress alters physiology and behavior across species. Most research on the effects of stress on behavior uses chronic stressors, and most are correlational. The effects of acute stressors on physiology and behavior have been mixed. Here, we use zebra finches, a highly gregarious species that forms long-term pair bonds, to test the effects of an acute corticosterone (CORT) on opposite-sex partner preferences over a same-sex individual or a group (the latter is a highly appealing option). We had two competing hypotheses. First, we predicted that acute CORT would alter preferences for the opposite sex bird in both conditions in both sexes. However, since there is a sex difference in the effects of CORT on partner preferences in voles, these effects may be more pronounced in males than in females. To test our hypotheses, we administered 2 doses of CORT (10µg and 20µg) or vehicle (control) using a repeated measures design. In the male vs. female test, there was a significant Sex by Treatment interaction, such that in males, 10µg CORT increased preferences for a female over the male compared to when these same males were treated with saline at baseline. There were no effects of treatment in females. In the opposite-sex vs. group condition, there was an overall effect of Treatment, such that the 10µg dose increased preference for the opposite-sex individual over both saline treatments, regardless of sex. These findings further our understanding of the effects of an acute stressor on sexual partner preferences.

Repeated toluene exposure increases c-Fos in catecholaminergic cells of the nucleus accumbens shell.

Toluene is a frequently abused solvent. Previous studies have suggested that toluene acts like other drugs of abuse, specifically on the dopaminergic system in the nucleus accumbens (NAc) and ventral tegmental area (VTA) of the mesolimbic pathway. Although changes in dopamine (DA) levels and c-Fos have been observed in both acute and repeated exposure paradigms, the extent to which c-Fos is localized to catecholaminergic cells is unknown. The present study tested the effects of repeated toluene exposure (1000-4000ppm) on locomotor activity and cells containing c-Fos, tyrosine hydroxylase (TH), or both in the core and shell of the NAc, as well as the anterior and posterior VTA. We focused our study on adolescents, since adolescence is a time of great neural change and a time when individuals tend to be more susceptible to drug abuse. In early tests, toluene dose-dependently increased locomotor activity. Repeated exposure to the highest concentration of toluene resulted in sensitization to toluene's effects on locomotor activity. Although the number of cells immunopositive for c-Fos or TH did not significantly differ across groups, cells immunopositive for TH+c-Fos were higher in the NAc shell of animals exposed to 4000ppm than in animals exposed to air (control) or 1000ppm. Taken together, these findings demonstrate that repeated high dose toluene exposure increases locomotor activity as well as activation of catecholaminergic cells in the shell of the NAc.

DSP-4, a noradrenergic neurotoxin, produces sex-specific effects on pairing and courtship behavior in zebra finches.

Norepinephrine (NE) is involved in a variety of behaviors across vertebrate species. In songbirds, NE is involved in singing and auditory perception, fundamental components of pair formation. Mechanisms of pairing remain poorly understood in avian species. NE is likely involved given its role in vocal communication and perception. Here, we tested the hypothesis that DSP-4 treatments (a noradrenergic neurotoxin that decreases NE) decreases singing in males, song perception in females and pairing in both sexes using a naturalistic paradigm. Females were tested for preferences of either control or DSP-4 males in a two-choice paradigm using live males. Both sexes were then tested for courtship and pair formation in aviaries. In the two-choice paradigm, control females showed a significant preference for control males over DSP-4 males, whereas DSP-4 females showed no such preference. In the aviary tests, DSP-4 males engaged in less courtship behavior, showed decreased pairing behaviors and increased pair latencies compared to control males. In females, DSP-4 treatments did not alter courtship or pairing behavior. Lower neural densities of noradrenergic fibers in song, auditory, and affiliative regions were observed in DSP-4 animals of both sexes. Furthermore, DBH-ir densities in these regions explained variations in courtship and pairing behaviors, as well as pairing status. Our results extend previous findings to naturalistic contexts, provide evidence that DBH-ir densities in specific regions correlate with pairing-related behaviors, and inform us of sex differences in the role of NE in pairing.

17ß-Hydroxysteroid dehydrogenase type IV, a Z-linked gene, is higher in females than in males in visual and auditory regions of developing zebra finches.

One of the most important decisions in a monogamous animal's life is the choice of a partner (partner preference), but the process by which this occurs remains poorly understood. The present study tests the hypothesis that hormones and genes play a role in sexual differentiation of partner preferences, as in the song system. We focused on a Z-linked gene, 17ß-hydroxysteroid dehydrogenase type IV (HSD17B4), coding for a steroidogenic enzyme that converts estradiol (E2) into an inactive metabolite. HSD17B4 mRNA is expressed more in the song regions of males compared to females throughout development, suggesting that regulation of E2 is important for male-typical song development. Here, we focused on four regions associated with sexual partner preferences. Females had significantly higher levels of HSD17B4 mRNA in auditory (caudomedial nidopallium) and visual (hyperpallium apicale) regions than did males at day 25. HSD17B4 was expressed in the hippocampus and caudolateral nidopallium, but there were no sex differences. In a second experiment, animals of both sexes were treated with E2 and HSD17B4 and androgen receptor (AR) mRNA were measured, since masculinization of the song system is, in part, accomplished by AR. AR was low across the four regions and was not sexually differentiated. E2 treatments increased HSD17B4 mRNA in the auditory region of males, which is contrary to findings in the song system. Our research suggests that different behaviors may be guided by the same genes and hormones, but that the exact nature of the gene-hormone relationships may differ according to brain region and behavior.

Behavioral effects of progesterone on pair bonding and partner preference in the female zebra finch (Taeniopygia guttata).

Progesterone is a sex steroid known to be involved in reproduction, but its role in pair relationships is not well understood. This study explored the effects of exogenous progesterone (P4) on courtship and pairing behaviors in female zebra finches (Taeniopygia guttata) in two separate experiments: the first focused on courtship and initial pair formation and the second examined the effects on pair maintenance. In these experiments, we tested the hypothesis that P4 increases pairing behaviors and consequently influences their partner preference. In Experiment 1, animals engaged in significantly more pairing behaviors when they were treated with P4 than when they received the vehicle. However, this effect was not partner-specific, since the association index (a marker for female partner preference) did not differ between treatment conditions. In Experiment 2, females were given two weeks to form a pair and then injected with P4 or vehicle. Pairs were observed that day and the subsequent day to determine if P4 caused a decrease in mate-directed behavior and an increase in extra pair behavior. P4 did not affect the quality of the pair relationship and did not increase extra pair behavior. These results suggest that P4 influences the overall quantity of initial pairing behaviors and may slightly increase the likelihood of partner preference formation over short time courses. However, P4 does not alter a previously established bond, suggesting there are likely separate mechanisms for initial pairing behaviors and pair maintenance.

Sex-dependent species discrimination in auditory forebrain of naturally hybridizing birds.

Pairs of individuals breed together only if they recognize each other as the same species, but the process of recognizing conspecifics can depend on flexible criteria even when species-specific signals are innate and fixed. This study examines species recognition in naturally hybridizing sister species, California and Gambel's quail (Callipepla californica and Callipepla gambelii), that have vocalizations which are not learned. Specifically, this study tests whether being raised in a vocalizing mixed-species cohort affects neural activity in the adult auditory forebrain in response to heterospecific and conspecific calls. After hatching, quail chicks were raised either with their own kind or with both species. Once reaching reproductive condition, each adult was played a recording that was one of three types: Gambel's quail opposite-sex contact calls; California quail opposite-sex contact calls; or synthetic tones. Brains were collected following playback and assessed for neuronal activity by quantifying expression of the protein of the immediate early gene, ZENK, in two brain regions, the caudomedial nidopallium (NCM) and the caudomedial mesopallium (CMM). ZENK levels were greater in NCM of males than females, but female NCM cells responded differentially to conspecific compared to heterospecific calls. Namely, females had more immuno-positive NCM cells when they heard conspecific calls rather than heterospecific male calls. Early experience with heterospecific broodmates did not alter neural responses in the NCM or CMM to heterospecific vocalizations. This study suggests that the NCM plays a role in species discrimination but that rearing condition does not alter the response in these non-vocal-learning species.

Sexual differentiation of the zebra finch song system: potential roles for sex chromosome genes.

BACKGROUND: Recent evidence suggests that some sex differences in brain and behavior might result from direct genetic effects, and not solely the result of the organizational effects of steroid hormones. The present study examined the potential role for sex-biased gene expression during development of sexually dimorphic singing behavior and associated song nuclei in juvenile zebra finches. RESULTS: A microarray screen revealed more than 2400 putative genes (with a false discovery rate less than 0.05) exhibiting sex differences in the telencephalon of developing zebra finches. Increased expression in males was confirmed in 12 of 20 by qPCR using cDNA from the whole telencephalon; all of these appeared to be located on the Z sex chromosome. Six of the genes also showed increased expression in one or more of the song control nuclei of males at post-hatching day 25. Although the function of half of the genes is presently unknown, we have identified three as: 17-beta-hydroxysteroid dehydrogenase type IV, methylcrotonyl-CoA carboxylase, and sorting nexin 2. CONCLUSION: The data suggest potential influences of these genes in song learning and/or masculinization of song system morphology, both of which are occurring at this developmental stage.

Sexually dimorphic SCAMP1 expression in the forebrain motor pathway for song production of juvenile zebra finches.

Mechanisms regulating sexual differentiation of the zebra finch song system are not well understood. The present study was designed to more fully characterize secretory carrier membrane protein 1 (SCAMP1), which was identified in a cDNA microarray screen as showing increased expression in the forebrains of developing male compared with female zebra finches. We completed the sequence of the open reading frame and used in situ hybridization to compare mRNA in song control regions of juvenile (25-day-old) individuals. Expression was significantly greater in the HVC (used as a proper name) and robust nucleus of the arcopallium (RA) in males than in females. Immunohistochemistry revealed that SCAMP1 protein is also expressed in these two brain regions, and qualitatively appears greater in males. Western analysis confirmed that the protein is increased in the telencephalon of males when compared with females at 25 days of age. These results are consistent with the idea that SCAMP1 is involved in masculinization of these brain areas, perhaps facilitating the survival of cells within them.

Immediate early gene (ZENK) responses to song in juvenile female and male zebra finches: effects of rearing environment.

Accurate song perception is likely to be as important for female songbirds as it is for male songbirds. Male zebra finches (Taeniopygia guttata) show differential ZENK expression to conspecific and heterospecific songs by day 30 posthatch in auditory perceptual brain regions such as the caudomedial nidopallium (NCM) and the caudomedial mesopallium (CMM). The current study examined ZENK expression in response to songs of different qualities at day 45 posthatch in both sexes. Normally reared juvenile zebra finches showed higher densities of immunopositive nuclei in both the dorsal and ventral areas of NCM and CMM (formerly cmHV), but not HA, a visual area, in response to normal song over untutored song or silence. Male and female patterns of ZENK expression did not differ. We next compared responses of birds reared without exposure to normal song (untutored) to those of normally reared birds. Untutored birds did not show higher responses to normal song than to untutored song in the three song perception areas. Furthermore, untutored birds of both sexes showed lower densities of immunopositive nuclei in all four areas than did normally reared birds. In addition, ZENK expression was greater in untutored females than in males in the dorsal portion of NCM and in CMM. Our findings suggest that at least some neural mechanisms of song perception are in place in socially reared female and male finches at an early age. Furthermore, early exposure to song tutors affects responses to song stimuli.

The role of sex steroids in courtship, pairing and pairing behaviors in the socially monogamous zebra finch.

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.

Sex differences in juvenile rhesus macaque (Macaca mulatta) agonistic screams: life history differences and effects of prenatal androgens.

This study investigated sex differences in juvenile rhesus macaque (Macaca mulatta) vocal behavior during agonistic contexts, and the effects of prenatal androgens on these differences. A total of 59 subjects (5-8 per treatment group) received exogenous androgen (testosterone enanthate), an anti-androgen (flutamide) or vehicle injections (DMSO) for 30 or 35 days during the second (early) or third (late) trimester of pregnancy. An additional 19 unmanipulated controls were included in the analysis. Screams by juvenile males and females between the ages of 1 and 3 years were compared to the screams of adult female exemplars using a discriminant function analysis. Juvenile females produced more adult-female like screams than did juvenile males. Females exposed to androgen treatment late in gestation produced a more masculine pattern of screams. Flutamide treatment in males either early or late in gestation did not significantly affect scream production. Flutamide treatments in females late in gestation, however, masculinized scream production. Androgen treatments administered late in gestation hyper-masculinized male scream production. No sex differences in the contextual usage of screams emerged. These findings suggest that both life history differences and the early hormone environment contribute to sex differences in juvenile rhesus macaque vocal production.