Volume Of Song Control Nuclei Research Articles

Treatment with androgens plus estrogens cannot reverse sex differences in song and the song control nuclei in adult canaries

Adult treatments with testosterone (T) do not activate singing behavior nor promote growth of song control nuclei to the same extent in male and female canaries (Serinus canaria). Because T acts in part via aromatization into an estrogen and brain aromatase activity is lower in females than in males in many vertebrates, we hypothesized that this enzymatic difference might explain the sex differences seen even after exposure to the same amount of T. Three groups of castrated males and 3 groups of photoregressed females (i.e., with quiescent ovaries following exposure to short days) received either 2 empty 10 mm silastic implants, one empty implant and one implant filled with T or one implant filled with T plus one with estradiol (E2). Songs were recorded for 3 h each week for 6 weeks before brains were collected and song control nuclei volumes were measured in Nissl-stained sections. Multiple measures of song were still different in males and females following treatment with T. Co-administration of E2 did not improve these measures and even tended to inhibit some measures such as song rate and song duration. The volume of forebrain song control nuclei (HVC, RA, Area X) and the rate of neurogenesis in HVC was increased by the two steroid treatments, but remained significantly smaller in females than in males irrespective of the endocrine condition. These sex differences are thus not caused by a lower aromatization of the steroid; sex differences in canaries are probably organized either by early steroid action or by sex-specific gene regulation directly in the brain.

Photoperiodic control of singing behavior and reproductive physiology in male Fife fancy canaries

Temperate-zone birds display marked seasonal changes in reproductive behaviors and the underlying hormonal and neural mechanisms. These changes were extensively studied in canaries (Serinus canaria) but differ between strains. Fife fancy male canaries change their reproductive physiology in response to variations in day length but it remains unclear whether they become photorefractory (PR) when exposed to long days and what the consequences are for gonadal activity, singing behavior and the associated neural plasticity. Photosensitive (PS) male birds that had become reproductively competent (high song output, large testes) after being maintained on short days (SD, 8 L:16D) for 6 months were divided into two groups: control birds remained on SD (SD-PS group) and experimental birds were switched to long days (16 L:8D) and progressively developed photorefractoriness (LD-PR group). During the following 12 weeks, singing behavior (quantitatively analyzed for 3 × 2 hours every week) and gonadal size (repeatedly measured by CT X-ray scans) remained similar in both groups but there was an increase in plasma testosterone and trill numbers in the LD-PR group. Day length was then decreased back to 8 L:16D for LD-PR birds, which immediately induced a cessation of song, a decrease in plasma testosterone concentration, in the volume of song control nuclei (HVC, RA and Area X), in HVC neurogenesis and in aromatase expression in the medial preoptic area. These data demonstrate that Fife fancy canaries readily respond to changes in photoperiod and display a pattern of photorefractoriness following exposure to long days that is associated with marked changes in brain and behavior.

Rapid testosterone-induced growth of the medial preoptic nucleus in male canaries

Testosterone activates singing within days in castrated male songbirds but full song quality only develops after a few weeks. Lesions of the medial preoptic nucleus (POM) inhibit while stereotaxic testosterone implants into this nucleus increase singing rate suggesting that this site plays a key role in the regulation of singing motivation. Testosterone action in the song control system works in parallel to control song quality. Accordingly, systemic testosterone increases POM volume within 1–2 days in female canaries, while the increase in volume of song control nuclei takes at least 2 weeks. The current study tested whether testosterone action is associated with similar differences in latencies in males. Photosensitive castrated male canaries were implanted with testosterone-filled Silastic™ implants and control castrates received empty implants, while simultaneously the photoperiod was switched from short- to long-days. Brains were collected from all subjects two days later. Plasma testosterone was elevated in testosterone-treated but not in controls. HVC volumes were not affected, but testosterone significantly increased the POM volume as identified by the dense group of aromatase-immunoreactive neurons, the number and somal area of these neurons and the fractional area they cover in POM. Testosterone-treated females from a previous experiment had a smaller POM volume in similar conditions suggesting the existence of a stable sex difference potentially affecting singing behavior. Thus testosterone induces male POM growth and aromatase expression in this nucleus within two days without affecting HVC size, further supporting the notion that testosterone increases singing motivation via its action in POM.

Testosterone-induced neuroendocrine changes in the medial preoptic area precede song activation and plasticity in song control nuclei of female canaries.

Testosterone plays a key role in the control of seasonal changes in singing behavior and its underlying neural circuitry. After administration of exogenous testosterone, song quality and song control nuclei volumes change over the course of weeks, but song rate increases within days. The medial preoptic nucleus (POM) controls sexual motivation and testosterone action in POM increases sexually motivated singing. In this study, we investigated the time course of testosterone action in the song control nuclei and POM, at the gross anatomical and cellular level. Photosensitive female canaries were injected with BrdU to label newborn neurons. One day later they were transferred to a long-day photoperiod and implanted with testosterone-filled or empty implants. Brains and blood were collected 1, 2, 9 or 21days later. Testosterone increased POM volume within 1day, whereas the volume of song control nuclei increased significantly only on day 21 even if a trend was already observed for HVC on day 9. The density of newborn neurons in HVC, labeled by Bromodeoxyuridine (BrdU) and doublecortin, was increased by testosterone on days 9 and 21 although a trend was already detectable on day 2. In POM, testosterone increased the number and size of aromatase-immunoreactive neurons already after 1day. This rapid action of testosterone in POM supports its proposed role in controlling singing motivation. Although testosterone increased the number of newborn neurons in HVC rapidly (9, possibly 2days), it is unlikely that these new neurons affect singing behavior before they mature and integrate into functional circuits.

Reversing song behavior phenotype: Testosterone driven induction of singing and measures of song quality in adult male and female canaries (Serinus canaria)

In songbirds, such as canaries (Serinus canaria), the song control circuit has been shown to undergo a remarkable change in morphology in response to exogenous testosterone (T). It is also well established that HVC, a telencephalic nucleus involved in song production, is significantly larger in males than in females. T regulates seasonal changes in HVC volume in males, and exposure to exogenous T in adult females increases HVC volume and singing activity such that their song becomes more male-like in frequency and structure. However, whether there are sex differences in the ability of T to modulate changes in the song system and song behavior has not been investigated in canaries. In this study, we compared the effects of increasing doses of T on singing and song control nuclei volumes in adult male and female American Singer canaries exposed to identical environmental conditions. Males were castrated and all birds were placed on short days (8L:16D) for 8weeks. Males and females were implanted either with a 2, 6 or 12mm long Silastic™ implant filled with crystalline T or an empty 12mm implant as control. Birds were then housed individually in sound-attenuated chambers. Brains were collected from six birds from each group after 1week or 3weeks of treatment. Testosterone was not equally effective in increasing singing activity in both males and females. Changes in song quality and occurrence rate took place after a shorter latency in males than in females; however, females did undergo marked changes in a number of measures of song behavior if given sufficient time. Males responded with an increase in HVC volume at all three doses. In females, T-induced changes in HVC volume only had limited amplitude and these volumes never reached male-typical levels, suggesting that there are sex differences in the neural substrate that responds to T.

Early Exposure to 2,2′,4,4′,5-Pentabromodiphenyl Ether (BDE-99) Affects Mating Behavior of Zebra Finches

2,2',4,4',5-Pentabromodiphenyl ether (BDE-99) is a brominated flame retardant congener that has pervaded global food chains, being reported in avian egg and tissue samples throughout the world. Its effects on birds are not well known, but there is evidence in exposed mammals that it directly mediates and causes neurotoxicity, alters thyroid hormone homeostasis, and lowers sex steroid hormone concentrations. In birds, those processes could disrupt the song-control system and male mating behavior. In this study, the effects of nestling exposure to environmentally relevant levels of BDE-99 were assessed in a model songbird species, the zebra finch (Taeniopygia guttata). A tissue residue study in which zebra finch nestlings were orally exposed to 0, 2.5, 15.8, or 50.7 ng BDE-99/g body weight (bw) per day over the 21-day nesting period validated dosing methods and confirmed dose levels were environmentally relevant (332.7 ± 141.0 to 4450.2 ± 1396.2 ng/g plasma lipid). A full-scale study exposing nestlings to 0, 2.5, 15.8, 50.7, or 173.8 ng BDE-99/g bw/day was carried out to investigate long-term effects of BDE-99 on the adult song-control nuclei volumes, song quality, and male mating behavior. Early exposure to BDE-99 had significant effects on male mating behavior and the response of clean experienced females to exposed males. There was no effect on male song-control nuclei or song quality, and there were nondose-dependent effects on female song-control nuclei. The results demonstrate that early exposure to environmentally relevant levels of BDE-99 affects the behavior of zebra finches.

A dose–response study of estradiol's effects on the developing zebra finch song system

To gauge the sensitivity of the female zebra finch song system to estradiol (E2), we used subcutaneous implants to administer various doses of E2 to hatchling female zebra finches. Four different doses of E2 were administered: 50, 15, 5 and 0-μg via subcutaneous silicon “ropes” at hatching, and the brains were examined in adulthood. Further, we examined whether masculinization was all-or-none once a threshold was reached or if the morphology of the song system would show a graded response to the various doses of E2. Finally, we asked if the various dependent measures – volume of song nuclei, neuron size, and neuron number – would show differential sensitivity to E2. Fifteen micrograms was sufficient to masculinize many aspects of the song system and was often as effective as 50-μg, causing a dramatic difference relative to the 0-μg group. Different aspects of the song system seemed differentially sensitive to the effects of E2: volumes of song control nuclei, the size of RA neurons, and the number of HVC neurons were significantly masculinized by 15-μg E2, but the number of RA neurons and HVC and lMAN soma sizes required 50-μg. The results suggest that several developmental processes are influenced by E2, possibly because of multiple sites of action or multiple processes that respond to E2.

Photoperiodic induced changes in reproductive state of border canaries ( Serinus canaria) are associated with marked variation in hypothalamic gonadotropin-releasing hormone immunoreactivity and the volume of song control regions

In temperate zone songbirds, such as canaries ( Serinus canaria), seasonal variation in gonadal activity and behavior are associated with marked brain changes. These include gonadotropin-releasing hormone (GnRH) expression and the volume of brain areas controlling song production. Questions have been raised about the consistency of seasonal brain changes in canaries. Laboratory studies of the American singer strain raised doubts as to whether this strain exhibits a robust photoperiodic response along with changes in brain GnRH content, and studies of free-living canaries have failed to identify seasonal changes in volume of song control nuclei. We assessed differences in brain GnRH and the song control system associated with photoperiod-induced variation in reproductive state in Border canaries. We found that males and females maintained for 10 weeks on long days (14L:10D) regress their gonads, exhibit a decline in testosterone and initiate molt; a response consistent with the onset of absolute photorefractoriness (i.e., failed to respond to previously stimulating daylengths). All birds regained photosensitivity (i.e., exhibited gonadal response to stimulating daylengths) after experiencing short days (8L:16D) for 6 weeks. Furthermore, comparisons of birds in either a photosensitive, photostimulated, or photorefractory state revealed a marked increase in GnRH protein expression in the photosensitive and photostimulated birds over photorefractory birds. A similar variation was observed in the volume of key forebrain song nuclei. Thus, Border canaries demonstrate measurable neuroplasticity in response to photoperiodic manipulations. These data, along with previous work on other strains of canaries, indicate the presence of intra-specific variation in photoperiodically regulated neuroplasticity.

Auditory stimulation of reproductive function in male Rufous-winged Sparrows, Aimophila carpalis

Prolonged exposure to conspecific song stimulates gonadal function and reproductive hormone secretion in female birds but few studies have investigated the physiological effects of conspecific song exposure on males outside of short-term, aggressive interactions. We exposed male Rufous-winged Sparrows, Aimophila carpalis, either to conspecific song (CS Song), to heterospecific song (Black-throated Sparrow, Amphispiza bilineata; HS Song), or to no recorded song (No Song) for 59 consecutive days (two h per day). Birds were exposed to short days (8L:16D) for the first 21 days of treatment and were then transferred to long days (13L:11D) for the remaining 38 days. During long day exposure, CS Song birds experienced faster growth of testes than HS Song and No Song birds. HS Song birds also grew their testes faster than No Song birds. Plasma luteinizing hormone (LH) and testosterone did not differ between CS Song and No Song birds. However, plasma LH was higher in HS Song birds compared to other groups. There were no differences in hypothalamic immunocytochemical labeling for gonadotropin-releasing hormone, its precursor proGnRH, or gonadotropin-inhibitory hormone, nor were there differences in two song control nuclei volumes (HVC and RA) between CS Song and No Song treatment groups. Furthermore, we found no effect of heterospecific song on free-living Rufous-winged Sparrow aggressive behaviors. These data indicate that long-term exposure to auditory stimuli, such as song, can influence the reproductive system of male songbirds and different types of auditory stimuli can have differential effects on reproductive function.

Song and brain development in canaries raised under different conditions of acoustic and social isolation over two years

Early isolation experiments indicate that male songbirds learn their songs during an early sensitive period, although later work has shown that some open-ended learners modify songs in later years. Recent isolation experiments suggest that in some species song has a stronger genetic basis than previously thought. This study raised domestic canaries under different combinations of acoustic and social isolation and followed song development into the second year. Males raised alone in acoustic isolation developed songs with normal syllables, but larger repertoires and also produced syllables with lower repetition rates when compared to controls. The smallest repertoire occurred in males raised in a peer group. Isolate males had a smaller song control nucleus HVC than controls, but there was no effect on nucleus RA or on brain weight in general. In the second year, after introduction into a large normal colony, isolate and peer group males adjusted their syllable repertoire to normal size. In particular, the isolates reduced their repertoire even though the size of HVC showed a significant increase in volume. However, songs of isolate and peer group males still differ in repetition rate and number of single syllables in the common aviary. In contrast, control males showed low syllable turnover and no significant change in repertoire size. Nor did they show any significant change in the volumes of song control nuclei. It seems that complete isolation affects only some aspects of song and brain development, and later socialization corrects some but not all of these in the second year.

Social context affects testosterone‐induced singing and the volume of song control nuclei in male canaries (Serinus canaria)

The contribution of social factors to seasonal plasticity in singing behavior and forebrain nuclei controlling song, and their interplay with gonadal steroid hormones are still poorly understood. In many songbird species, testosterone (T) enhances singing behavior but elevated plasma T concentrations are not absolutely required for singing to occur. Singing is generally produced either to defend a territory or to attract a mate and it is therefore not surprising that singing rate can be influenced by the sex and behavior of the social partner. We investigated, based on two independent experiments, the effect of the presence of a male or female partner on the rate of song produced by male canaries. In the first experiment, song rate was measured in dyads composed of one male and one female (M-F) or two males (M-M). Birds were implanted with T-filled Silastic capsules or with empty capsules as control. The number of complete song bouts produced by all males was recorded during 240 min on week 1, 2, 4, and 8 after implantation. On the day following each recording session, brains from approximately one-fourth of the birds were collected and the volumes of the song control nuclei HVC and RA were measured. T increased the singing rate and volume of HVC and RA but these effects were affected by the social context. Singing rates were higher in the M-M than in the M-F dyads. Also, in the M-M dyads a dominance-subordination relationship soon became established and dominant males sang at higher rates than subordinates in T-treated but not in control pairs. The differences in song production were not reflected in the size of the song control nuclei: HVC was larger in M-F than in M-M males and within the M-M dyads, no difference in HVC or RA size could be detected between dominant and subordinate males. At the individual level, the song rate with was positively correlated with RA and to a lower degree HVC volume, but this relationship was observed only in M-M dyads, specifically in dominant males. A second experiment, carried out with castrated males that were all treated with T and exposed either to another T-treated castrate or to an estradiol-implanted female, confirmed that song rate was higher in the M-M than in the M-F condition and that HVC volume was larger in heterosexual than in same-sex dyads. The effects of T on singing rate and on the volume of the song control nuclei are thus modulated by the social environment, including the presence/absence of a potential mate and dominance status among males.

A test of absolute photorefractoriness and photo-induced neural plasticity of song-control regions in black-capped chickadees (Poecile atricapillus)

We tested whether male and female black-capped chickadees, Poecile atricapillus (L., 1766), were absolutely photorefractory according to Hamner's (1968) criteria of (i) spontaneous regression of gonads during prolonged long-day exposure and (ii) no subsequent recrudescence of gonads in response to constant light. We initially exposed black-capped chickadees to constant long-day photoperiods. Gonads regressed spontaneously, demonstrating that the birds met Hamner's first criterion for absolute photorefractoriness. Once their gonads fully regressed and the birds were in advanced prebasic moult, we exposed them to one of three photoperiods for an additional 2 weeks: constant light (24 h L), short days (8 h L), and controls (15 h L). Constant light challenge had no effect on gonadal condition or rate of moult, confirming that the birds met Hamner's second criterion for absolute refractoriness. We also compared volumes of song-control nuclei in the three groups and found that males overall had larger HVC, robust nucleus of arcopallium, and area X than females, but that longer days (24 h L) did not increase volumes and that shorter days (8 h L) did not decrease volumes compared with controls (15 h L). These data support the inference that black-capped chickadees do indeed become absolutely photorefractory, and that photorefractoriness precludes photo-induced plasticity of the song-control system.

Seasonal plasticity in the song control system: multiple brain sites of steroid hormone action and the importance of variation in song behavior.

Birdsong, in non-tropical species, is generally more common in spring and summer when males sing to attract mates and/or defend territories. Changes in the volumes of song control nuclei, such as HVC and the robust nucleus of the arcopallium (RA), are observed seasonally. Long photoperiods in spring stimulate the recrudescence of the testes and the release of testosterone. Androgen receptors, and at times estrogen receptors, are present in HVC and RA as are co-factors that facilitate the transcriptional activity of these receptors. Thus testosterone can act directly to induce changes in nucleus volume. However, dissociations have been identified at times among long photoperiods, maximal concentrations of testosterone, large song control nuclei, and high rates of song. One explanation of these dissociations is that song behavior itself can influence neural plasticity in the song system. Testosterone can act via brain-derived neurotrophic factor (BDNF) that is also released in HVC as a result of song activity. Testosterone could enhance song nucleus volume indirectly by acting in the preoptic area, a region regulating sexual behaviors, including song, that connects to the song system through catecholaminergic cells. Seasonal neuroplasticity in the song system involves an interplay among seasonal state, testosterone action, and behavioral activity.

Song activation by testosterone is associated with an increased catecholaminergic innervation of the song control system in female canaries

In canaries, singing and a large number of morphological features of the neural system that mediates the learning, perception and production of song exhibit marked sex differences. Although these differences have been mainly attributed to sex-specific patterns of the action of testosterone and its metabolites, the mechanisms by which sex steroids regulate brain and behavior are far from being completely understood. Given that the density of immunoreactive catecholaminergic fibers that innervate telencephalic song nuclei in canaries is higher in males, which sing, than in females, which usually do not sing, we hypothesized that some of the effects induced by testosterone on song behavior are mediated through the action of the steroid on the catecholaminergic neurons which innervate the song control nuclei. Therefore, we investigated in female canaries the effects of a treatment with exogenous testosterone on song production, on the volume of song control nuclei, and on the catecholaminergic innervation of these nuclei as assessed by immunocytochemical visualization of tyrosine hydroxylase. Testosterone induced male-like singing in all females and increased by about 80% the volume of two telencephalic song control nuclei, the high vocal center (HVC) and the nucleus robustus archistriatalis (RA). Testosterone also significantly increased the fractional area covered by tyrosine hydroxylase-immunoreactive structures (fibers and varicosities) in most telencephalic song control nuclei (HVC, the lateral and medial parts of the magnocellular nucleus of the anterior neostriatum, the nucleus interfacialis, and to a lesser extent RA). By contrast, testosterone did not affect the catecholaminergic innervation of the telencephalic areas adjacent to HVC and RA. Together these data demonstrate that, in parallel to its effects on song behavior and on the morphology of the song control system, testosterone also regulates the catecholaminergic innervation of most telencephalic song control nuclei in canaries. The endocrine regulation of singing may thus involve the neuromodulatory action of specialized dopaminergic and/or noradrenergic projections onto several key parts of the song control system.

The volumes of song control nuclei, HVC and lMAN, relate to differential behavioral responses of female European starlings to male songs produced within and outside of the breeding season

In seasonally breeding songbirds, such as European starlings ( Sturnus vulgaris), male song plays a direct role in immediate mate attraction during, but not outside of, the breeding season. The present experiment was designed to explore whether female starlings would display differential behavioral responses to male song recorded during (spring) and outside of (fall) the breeding season. Recordings of spring and fall male starling songs were broadcast to female starlings from nest boxes in an aviary setting. Females displayed marked elevations in general activity (landing, eating, and drinking) in response to spring compared to fall song, however as a group females did not preferentially land on nest boxes playing spring song. The volume of the song control nucleus HVC was largest in females that landed more often on nest boxes during playbacks of spring song compared to fall song, and the volume of the lateral portions of the magnocellular nucleus of the anterior neostriatum (lMAN) related positively to the extent to which females directed landings towards nest boxes playing spring rather than fall song. A possible relationship was also detected between female preferences for male spring compared to fall song and the volume of the robust nucleus of the archistriatum. Female responses to spring and fall song did not relate to the volumes of area X, or to estrogen concentrations. These data provide evidence that females attend to and respond behaviorally to seasonal changes in male starling song, and implicate song control nuclei, HVC and lMAN, in female perception and discrimination of male song.

Seasonal changes in the densities of alpha(2) noradrenergic receptors are inversely related to changes in testosterone and the volumes of song control nuclei in male European starlings.

The functions of song and the contextual cues that elicit song change seasonally in parallel with testosterone (T) concentrations in male European starlings. T is high in spring when at least one function of male song is that of immediate mate attraction, and low outside the context of breeding, when starlings primarily use song for dominance or flock maintenance. Several brain nuclei that control song contain high densities of alpha(2)adrenergic receptors. T can regulate the density of alpha(2)adrenergic receptors in the avian brain, indicating that the density of alpha(2) adrenergic receptors within the song system might change seasonally. Although the function of seasonal brain variation is not entirely clear, in many songbirds the volumes of song nuclei are largest when T is high and males sing most. Male starlings, however, sing both when T is high and when T is low. Therefore, exploring seasonal changes in T and the volumes of song nuclei could provide insight into the function of these changes. The present study was performed to explore the relationships among T, the volumes of song nuclei, and the densities of alpha(2) adrenergic receptors within the song system of male starlings. Song nuclei (the high vocal center [HVc], robust nucleus of the archistriatum [RA], and Area X) were largest, T was highest, and the density of alpha(2) adrenergic receptors (within HVc and RA) was lowest during the breeding season. The reverse pattern was observed outside of the breeding season. These results suggest that changes in T, volumes of song nuclei, and alpha(2) receptor densities might regulate seasonal changes in song behavior or the context that will elicit song in male starlings.

Seasonal Changes in Brain GnRH Immunoreactivity and Song-Control Nuclei Volumes in an Opportunistically Breeding Songbird

White-winged crossbills (Loxia leucoptera) are opportunistic breeders that can nest at almost any time of year if there is sufficient food. Other cardueline finches that have been shown to breed on a strictly seasonal schedule become absolutely refractory to the stimulatory effects of long-day photoperiod, dramatically down-regulate hypothalamic gonadotropin-releasing hormone (GnRH), and reduce the volume of several song-control nuclei in autumn. This study examined whether changes in photoperiod modify the GnRH and song-control systems in white-winged crossbills. Adult male and female crossbills were captured and held on a naturally changing photoperiod. Brains of male and female birds were collected in May, October, and January. GnRH content was assessed by immunocytochemistry and the volumes of Nissl-defined song-control nuclei (HVc, Area X, and the robust nucleus of the archistriatum) were reconstructed. In contrast to other cardueline finches, GnRH immunoreactivity was relatively stable across the year, exhibiting only modest seasonal variation. The song control system, on the other hand, exhibited large seasonal changes as well as sex differences. Thus, crossbills appear to maintain hypothalamic GnRH content year round, perhaps to facilitate a rapid response to favorable breeding conditions, even on short days. However, song control nuclei are dramatically affected by photoperiod. Future work should examine these systems in crossbills breeding on short days to compare photoperiod-dependent and -independent effects on neural plasticity.

TrkB-like immunoreactivity in the song system of developing zebra finches

The neural song system in zebra finches develops for approximately the first 2 months after hatching. During that time, male-biased sexual dimorphisms emerge in the volume of song control nuclei as well as in the number and size of neurons within them. Brain derived neurotrophic factor (BDNF) has been documented in song control nuclei at various stages of development. Its high affinity receptor (tyrosine kinase B; trkB) is also in the song system, at least at around 1 month of age. The present study was designed to more completely describe the timing and potential location of BDNF action by investigating trkB expression during sexual differentiation of the song control nuclei. The pattern of immunoreactivity to a trkB antibody was examined in male and female zebra finches at post-hatching days 3–60. Labeling in somata and neuropil appeared to define the telencephalic components of the motor pathway (high vocal center and robust nucleus of the archistriatum) for song production in males from days 30 to 60, and in females on days 45 and 60 (high vocal center). These results are consistent with the hypothesis that the receptor, and its ligand BDNF, play a role in processes related to song learning in both sexes, including perhaps the motor component exhibited by developing males.

Development of Topography within Song Control Circuitry of Zebra Finches during the Sensitive Period for Song Learning

Refinement of topographic maps during sensitive periods of development is a characteristic feature of diverse sensory and motor circuits in the nervous system. Within the neural system that controls vocal learning and behavior in zebra finches, axonal connections of the cortical nucleus lMAN demonstrate striking functional and morphological changes during vocal development in juvenile males. These circuits are uniquely important for song production during the sensitive period for vocal learning, and the overall size of these brain regions and their patterns of axonal connectivity undergo dramatic growth and regression during this time. Axonal connections to and from lMAN are topographically organized in adult males that have already learned song. We wondered whether the large-scale changes seen in lMAN circuitry during the time that vocal behavior is being learned and refined could be accompanied by the emergence of topographic mapping. However, results presented herein demonstrate that most of these song-control circuits show the same broad patterns of axonal connectivity between subregions of individual nuclei at the onset of song learning as seen in adult birds. Thus, coarse topographic organization is not dependent on the types of experience that are crucial for vocal learning. Furthermore, this maintenance of topographic organization throughout the period of song learning is clearly not achieved by maintenance of static axonal arbors. In fact, because the volumes of song-control nuclei are growing (or regressing), topography must be maintained by active remodeling of axonal arbors to adapt to the changes in overall size of postsynaptic targets. A salient exception to this pattern of conserved topography is the projection from lMAN to the motor cortical region RA: this pathway is diffusely organized at the onset of song learning but undergoes substantial refinement during early stages of song learning, suggesting that remodeling of axonal connections within this projection during the period of vocal learning may signify the production of increasingly refined vocal utterances.

Visual and song nuclei correlate with courtship skills in brown-headed cowbirds

We assessed courtship success in 14 adult male brown-headed cowbirds,Molothrus ater. Volumes of song control nuclei and visual nuclei were measured in Nissl stained tissue. Variation in courtship success was found to be related to variation in two areas of the avian brain: the song control nucleus, area X, and the thalamic visual area, nucleus rotundus. Volume of area X was negatively correlated with song potency, as assessed by female playback, and with rate of vocalizing. Volume of nucleus rotundus was positively correlated with song potency, vocalizing to females and courtship persistence. These data are the first to implicate a visual nucleus in the use of song. The data also complement previous findings with cowbirds suggesting that song learning involves visual attention to females. Taken as a whole, these results suggest that use of song depends on integration of auditory, vocal and visual information. In that female songbirds in many species assess multimodal performance of song, these findings with cowbirds suggest that future studies of brain and behaviour include a broader view of possible behavioural and neural correlates.