Ontogenetic changes in the pattern of androgen accumulation in song-control nuclei of male zebra finches

The present study examines the development of androgen accumulation in cells of two brain nuclei that are involved in controlling vocal behavior in zebra finches (Poephila guttata). HVc (caudal nucleus of the ventral hyperstriatum) is involved with vocal production in adult birds, and MAN (magnocellular nucleus of the anterior neostriatum) is involved with the initial ability to learn song. In both of these nuclei there is an increase in the proportion of cells that are labeled by systemic injections of tritiated dihydrotestosterone in juvenile male zebra finches during the time when production of song is becoming stereotyped (25-60 days). Within MAN there is an overall loss of cells during this time, such that the absolute number of androgen target cells in MAN remains at a constant level. However, it does not appear to be the case that unlabeled cells are selectively lost from MAN. Rather it appears that both labeled and unlabeled cells are lost, and the absolute number of labeled cells is maintained at a constant level via recruitment of additional labeled cells from the unlabeled population (i.e., some MAN cells that are unlabeled in young birds become labeled in older birds). In line with this hypothesis, there is a large increase in the density of labeling in individual MAN cells, indicating that these cells have an enhanced ability to concentrate androgen. In contrast to the situation in MAN, there is an increase in the overall number of cells within HVc during this time; this increase in total cell number combines with the increased proportion of labeled cells such that the absolute number of androgen target cells in HVc increases threefold. The ability of individual HVc cells to accumulate androgen remains constant. The relationship of these changes in the pattern of androgen accumulation to other aspects of neural and behavioral development related to song in zebra finches are discussed.     

Cell death during development of a forebrain nucleus involved with vocal learning in zebra finches

Lateral MAN (magnocellular nucleus of the anterior neostriatum) is a forebrain nucleus that is known to be importantly involved with vocal learning in juvenile male zebra finches only during a restricted period of the learning process: lesions of lMAN completely disrupt song behavior in zebra finches prior to 50 days of age but have little or no effect in older juvenile or adult birds. The development of lMAN, as of other song-control regions, is delayed until the time that song behavior is being learned. Lateral MAN undergoes a substantial loss of neurons between 25 and 55 days of age, a time that encompasses initial stages of vocal production as well as the interval during which lMAN lesions become ineffective. In this study, we measured both the time course of neuronal loss and the incidence of pyknotic cells within lMAN during the period of cell loss. There is a pronounced loss of neurons from lMAN between 20 and 35 days, after which the adult number of neurons is established. The incidence of pyknosis is greatest at 20 days, around the time when the loss of live cells is also most pronounced, suggesting that the loss of neurons from lMAN is attributable to cell death. The loss of neurons occurs well before lesions of lMAN become ineffective in disrupting vocal behavior. Thus the neurons remaining in lMAN after the period of cell loss apparently undergo a substantial change in function at the time lesions lose effectiveness (about 55-60 days).     

Changes in adult zebra finch song require a forebrain nucleus that is not necessary for song production

Male zebra finches normally crystallize song at approximately 90 days and do not show vocal plasticity as adults. However, changes to adult song do occur after unilateral tracheosyringeal (ts) nerve injury, which denervates one side of the vocal organ. We examined the effect of placing bilateral lesions in LMAN (a nucleus required for song development but not for song maintenance in adults) upon the song plasticity that is induced by ts nerve injury in adults. The songs of birds that received bilateral lesions within LMAN followed by right ts nerve injury silenced, on average, 0.25 syllables, and added 0.125 syllables (for an average turnover of 0.375 syllables), and changed neither the frequency with which individual syllables occurred within songs nor the motif types they used most often. In contrast, the songs of birds that received sham lesions followed by ts nerve injury lost, on average, 1.625 syllables, silenced 0.125 syllables, and added 0.75 syllables, turning over an average of 2.5 syllables. They also significantly changed both the frequency with which individual syllables were included in songs and the motif variants used. Thus, song plasticity induced in adult zebra finches with crystallized songs requires the presence of LMAN, a nucleus which had been thought to play a role in vocal production only during song learning. Although the changes to adult songs induced by nerve transection are more limited than those that arise during song development, the same circuitry appears to underlie both types of plasticity.     

HVC microlesions do not destabilize the vocal patterns of adult male zebra finches with prior ablation of LMAN

The songs of adult male zebra finches (Taeniopygia guttata) arise by an integration of activity from two neural pathways that emanate from the telencephalic nucleus HVC (proper name). One pathway descends directly from HVC to the vocal premotor nucleus RA (the robust nucleus of the arcopallium) whereas a second pathway descends from HVC into a basal ganglia circuit (the anterior forebrain pathway, AFP) that also terminates in RA. Although HVC neurons that project directly to RA outnumber those that contribute to the AFP, both populations are distributed throughout HVC. Thus, partial ablation (microlesion) of HVC should damage both pathways in a proportional manner. We report here that bilateral HVC microlesions in adult male zebra finches produce an immediate loss of song stereotypy from which birds recover, in some cases within 3 days. The contribution of the AFP to the onset of song destabilization was tested by ablating the output nucleus of this circuit (LMAN, the lateral magnocellular nucleus of the anterior nidopallium) prior to bilateral HVC microlesions. Song stereotypy was largely unaffected. Together, our findings suggest that adult vocal production involves nonproportional integration of two streams of neural activity with opposing effects on song--HVC's direct projection to RA underlies production of stereotyped song whereas the AFP seems to facilitate vocal variation. However, the rapid recovery of song in birds with HVC microlesions alone suggests the presence of dynamic corrective mechanisms that favor vocal stereotypy.     

A sensorimotor area in the songbird brain is required for production of vocalizations in the song learning period of development

Sensory feedback is essential for acquiring and maintaining complex motor behaviors, including birdsong. In zebra finches, auditory feedback reaches the song control circuits primarily through the nucleus interfacialis nidopalii (Nif), which provides excitatory input to HVC (proper name)—a premotor region essential for the production of learned vocalizations. Despite being one of the major inputs to the song control pathway, the role of Nif in generating vocalizations is not well understood. To address this, we transiently inactivated Nif in late juvenile zebra finches. Upon Nif inactivation (in both hemispheres or on one side only), birds went from singing stereotyped zebra finch song to uttering highly variable and unstructured vocalizations resembling sub‐song, an early juvenile song form driven by a basal ganglia circuit. Simultaneously inactivating Nif and LMAN (lateral magnocellular nucleus of the anterior nidopallium), the output nucleus of a basal ganglia circuit, inhibited song production altogether. These results suggest that Nif is required for generating the premotor drive for song. Permanent Nif lesions, in contrast, have only transient effects on vocal production, with song recovering within a day. The sensorimotor nucleus Nif thus produces a premotor drive to the motor pathway that is acutely required for generating learned vocalizations, but once permanently removed, the song system can compensate for its absence. © 2016 Wiley Periodicals, Inc. Develop Neurobiol 76: 1213–1225, 2016     

Distribution and developmental change in [3H]MK-801 binding within zebra finch song nuclei.

In many songbirds, vocal learning depends upon appropriate auditory experience during a sensitive period that coincides with the formation and reorganization of song-related neural pathways. Because some effects of early sensory experience on neural organization and early learning have been linked to activation of N-methyl-D-aspartate (NMDA) receptors, we measured binding to this receptor within the neural system controlling song behavior in zebra finches. Quantitative autoradiography was used to measure binding of the noncompetitive antagonist [3H]MK-801 (dizocilpine) in the brains of both adult and juvenile male zebra finches, focusing on four telencephalic regions implicated in song learning and production. Overall, the pattern of MK-801 binding in zebra finches was similar to the pattern found in rats (Monaghan and Cotman, 1985, J. Neurosci. 5:2909-2919; Sakurai, Cha, Penney, and Young, 1991, Neuroscience 40:533-543). That is, binding was highest in the telencephalon, intermediate in thalamic regions, and virtually absent from the brain stem and cerebellum. The telencephalic song areas exhibited intermediate levels of binding, and binding in the juveniles was not significantly different from adult levels in most song nuclei. However, in the lateral magnocellular nucleus of the anterior neostriatum (IMAN), binding at 30 days of age was significantly higher than binding in adults. Given the established role of NMDA receptors in other developing neural systems, both their presence in song control nuclei and their developmental regulation within a region implicated in song learning suggest that NMDA receptors play a role in mediating effects of auditory experience on the development of song behavior.     

Castration and antisteroid treatment impair vocal learning in male zebra finches.

Both song behavior and its neural substrate are hormone sensitive: castrated adult male zebra finches need replacement of gonadal steroids in order to restore normal levels of song production, and sex steroids are necessary to establish male-typical neural song-control circuits during early development. This pattern of results suggests that hormones may be required for normal development of learned song behavior, but evidence that steroids are necessary for normal neural and behavioral development during song learning has been lacking. We addressed this question by attempting to eliminate the effects of gonadal steroids in juvenile male zebra finches between the time of initial song production and adulthood. Males were castrated at 20 days of age and received systemic implants of either an antiandrogen (flutamide), an antiestrogen (tamoxifen), or both drugs. The songs of both flutamide- and tamoxifen-treated birds were extremely disrupted relative to normal controls in terms of the stereotypy and acoustic quality of individual note production, as well as stereotypy of the temporal structure of the song phrase. We did not discern any differences in the pattern of behavioral disruption between birds that were treated with either flutamide, tamoxifen, or a combination of both drugs. Flutamide treatment resulted in a reduced size of two forebrain nuclei that are known to play some role unique to early phases of song learning [lateral magnocellular nucleus of the anterior neostriatum (IMAN) and area X (X)], but did not affect the size of two song-control nuclei that are necessary for normal song production in adult birds [caudal nucleus of the ventral hyperstriatum (HVc) and robust nucleus of the archistriatum (RA)]. In contrast, treatment with tamoxifen did not result in any changes in the size of song-control nuclei relative to normal controls, and it blocked the effects of flutamide on the neural song-control system in birds that were treated with both drugs. Castration and antisteroid treatment exerted no deleterious effects on the quality of song behavior in adult birds, indicating that gonadal hormones are necessary for the development of normal song behavior during a sensitive period.     

Learning-Related Neuronal Activation in the Zebra Finch Song System Nucleus HVC in Response to the Bird's Own Song

Like many other songbird species, male zebra finches learn their song from a tutor early in life. Song learning in birds has strong parallels with speech acquisition in human infants at both the behavioral and neural levels. Forebrain nuclei in the 'song system' are important for the sensorimotor acquisition and production of song, while caudomedial pallial brain regions outside the song system are thought to contain the neural substrate of tutor song memory. Here, we exposed three groups of adult zebra finch males to either tutor song, to their own song, or to novel conspecific song. Expression of the immediate early gene protein product Zenk was measured in the song system nuclei HVC, robust nucleus of the arcopallium (RA) and Area X. There were no significant differences in overall Zenk expression between the three groups. However, Zenk expression in the HVC was significantly positively correlated with the strength of song learning only in the group that was exposed to the bird's own song, not in the other two groups. These results suggest that the song system nucleus HVC may contain a neural representation of a memory of the bird's own song. Such a representation may be formed during juvenile song learning and guide the bird's vocal output.     

Song learning in zebra finches: some effects of song model availability on what is learnt and when

This study indicates that captive male zebra finches (Taeniopygia guttata) normally learn their song during the juvenile period between independence from their parents and sexual maturity, from whatever suitable song model is available. Virtually nothing is learnt from the father before this time. Hybrid songs may develop if birds are removed from the father and given a new song model before song learning is complete. The absence of a song model during the juvenile stage appears to postpone the sensitive phase and abnormal song is produced until a suitable model becomes available. Normal song will then develop, even in a sexually mature adult. This indicates that experience and not age is the important factor determining the timing of the sensitive phase for song learning in zebra finches.     

Distribution and developmental change in [3H]MK-801 binding within zebra finch song nuclei

In many songbirds, vocal learning depends upon appropriate auditory experience during a sensitive period that coincides with the formation and reorganization of song-related neural pathways. Because some effects of early sensory experience on neural organization and early learning have been linked to activation of N-methyl-d-aspartate (NMDA) receptors, we measured binding to this receptor within the neural system controlling song behavior in zebra finches. Quantitative autoradiography was used to measure binding of the noncompetitive antagonist [³H]MK-801 (dizocilpine) in the brains of both adult and juvenile male zebra finches, focusing on four telencephalic regions implicated in song learning and production. Overall, the pattern of MK-801 binding in zebra finches was similar to the pattern found in rats (Monaghan and Cotman, 1985, J. Neurosci. 5:2909–2919; Sakurai, Cha, Penney, and Young, 1991, Neuroscience 40:533–543). That is, binding was highest in the telencephalon, intermediate in thalamic regions, and virtually absent from the brain stem and cerebellum. The telencephalic song areas exhibited intermediate levels of binding, and binding in the juveniles was not significantly different from adult levels in most song nuclei. However, in the lateral magnocellular nucleus of the anterior neostriatum (IMAN), binding at 30 days of age was significantly higher than binding in adults. Given the established role of NMDA receptors in other developing neural systems, both their presence in song control nuclei and their developmental regulation within a region implicated in song learning suggest that NMDA receptors play a role in mediating effects of auditory experience on the development of song behavior.     

At the interface of the auditory and vocal motor systems: NIf and its role in vocal processing,production and learning

Communication between auditory and vocal motor nuclei is essential for vocal learning. In songbirds, the nucleus interfacialis of the nidopallium (NIf) is part of a sensorimotor loop, along with auditory nucleus avalanche (Av) and song system nucleus HVC, that links the auditory and song systems. Most of the auditory information comes through this sensorimotor loop, with the projection from NIf to HVC representing the largest single source of auditory information to the song system. In addition to providing the majority of HVC’s auditory input, NIf is also the primary driver of spontaneous activity and premotor-like bursting during sleep in HVC. Like HVC and RA, two nuclei critical for song learning and production, NIf exhibits behavioral-state dependent auditory responses and strong motor bursts that precede song output. NIf also exhibits extended periods of fast gamma oscillations following vocal production. Based on the converging evidence from studies of physiology and functional connectivity it would be reasonable to expect NIf to play an important role in the learning, maintenance, and production of song. Surprisingly, however, lesions of NIf in adult zebra finches have no effect on song production or maintenance. Only the plastic song produced by juvenile zebra finches during the sensorimotor phase of song learning is affected by NIf lesions. In this review, we carefully examine what is known about NIf at the anatomical, physiological, and behavioral levels. We reexamine conclusions drawn from previous studies in the light of our current understanding of the song system, and establish what can be said with certainty about NIf’s involvement in song learning, maintenance, and production. Finally, we review recent theories of song learning integrating possible roles for NIf within these frameworks and suggest possible parallels between NIf and sensorimotor areas that form part of the neural circuitry for speech processing in humans.     

Representation of Early Sensory Experience in the Adult Auditory Midbrain: Implications for Vocal Learning

Vocal learning in songbirds and humans occurs by imitation of adult vocalizations. In both groups, vocal learning includes a perceptual phase during which juveniles birds and infants memorize adult vocalizations. Despite intensive research, the neural mechanisms supporting this auditory memory are still poorly understood. The present functional MRI study demonstrates that in adult zebra finches, the right auditory midbrain nucleus responds selectively to the copied vocalizations. The selective signal is distinct from selectivity for the bird''s own song and does not simply reflect acoustic differences between the stimuli. Furthermore, the amplitude of the selective signal is positively correlated with the strength of vocal learning, measured by the amount of song that experimental birds copied from the adult model. These results indicate that early sensory experience can generate a long-lasting memory trace in the auditory midbrain of songbirds that may support song learning.     

Castration and antisteroid treatment impari vocal learning in male zebra finches

Both song behavior and its neural substrate are hormone sensitive: Castrated adult male zebra finches need replacement of gonadal steroids in order to restore normal levels of song production, and sexsteroids are necessary to establish male-typical neural song-controlcircuits during early development. This pattern of results suggests that hormones may be required for normal development of learned songbehavior, but evidence that steroids are necessary for normal neuraland behavioral development during song learning has been lacking. Weaddressed this question by attempting to eliminate the effects of gonadal steroids in juvenile male zebra finches between the time of initial song production and adulthood. Males were castrated at 20 daysof age and received systemic implants of either an antiandrogen (flutamide). an antiestrogen (tamoxifen), or both drugs. The songs of both flutamide-and tamoxifen-treated birds were extremely disrupted relative to normal controls in terms of the stereotypy and acoustic quality of individual note production, as well as stereotypy of the temporal structure of the song phrase. We did not discern any differences in the pattern of behavioral disruption between birds that were treated with either flutamide, tamoxifen, or a combination of both drugs. Flutamide treatment resulted in a reduced size of two forebrain nuclei that are known to play some role unique to early phases of song learning [lateral magnocellular nucleus of the anterior neostriatum (IMAN) and area X (X)], but did not affect the size of two song-control nuclei that are necessary for normal song productionin adult birds [caudal nucleus of the ventral hyperstriatum (HVc) and robust nucleus of the archistriatum (RA)]. In contrast, treatment with tamoxifen did not result in any changes in the size of song-control nuclei relative to normal controls, and it blocked the effects of flutamide on the neural song-control system in birds that were treated with both drugs. Castration and antisteroid treatment exerted no deleterious effects on the quality of song behavior in adult birds, indicating that gonadal hormones are necessary for the development of normal song behavior during a sensitive period. © 1992 John Wiley & Sons, Inc.     

Matters of life and death in the songbird forebrain.

Male zebra finches learn a specific vocal pattern during a restricted period of development. They produce that song in stereotyped form throughout adulthood, and are unable to learn new song patterns. Development of the neural substrate for song learning and behavior is delayed relative to other brain regions, and neural song-control circuits undergo dramatic changes during the period of vocal learning due to both loss of neurons as well as incorporation of newly generated neurons. In contrast, canaries do learn new song patterns in adulthood and modify their vocal repertoires each breeding season. Adult canaries also maintain a large population of dividing cells in the ependymal zone of the telencephalon, and vast numbers of newly generated neurons migrate out to become incorporated into functional circuits and replace older neurons. We review the relationships between cellular and behavioral aspects of song learning in both zebra finches and canaries, as well as the role of gonadal hormones in regulating diverse aspects of the song-control system.     

Disconnection of a basal ganglia circuit in juvenile songbirds attenuates the spectral differentiation of song syllables

Similar to language acquisition by human infants, juvenile male zebra finches (Taeniopygia guttata) imitate an adult (tutor) song by transitioning from repetitive production of one or two undifferentiated protosyllables to the sequential production of a larger and spectrally heterogeneous set of syllables. The primary motor region that controls learned song is driven by a confluence of input from two premotor pathways: a posterior pathway that encodes the adult song syllables and an anterior pathway that includes a basal ganglia (BG)‐thalamo‐cortical circuit. Similar to mammalian motor‐learning systems, the songbird BG circuit is thought to be necessary for shaping juvenile vocal behaviour (undifferentiated protosyllables) toward specific targets (the tutor's song syllables). Here, we tested the hypothesis that anterior pathway activity contributes to the process of protosyllable differentiation. Bilateral ablation of lateral magnocellular nucleus of the anterior nidopallium (LMAN) was used to disconnect BG circuitry at ages before protosyllable production and differentiation. Comparison to surgical controls revealed that protosyllables fail to differentiate in birds that received juvenile LMAN ablation—the adult songs of birds with >80% bilateral LMAN ablation consisted of only one or two syllables produced with the repetitive form and spectral structure that characterizes undifferentiated protosyllables in normal juveniles. Our findings support a role for BG circuitry in shaping juvenile vocal behaviour toward the acoustic structure of the tutor song and suggest that posterior pathway function remains in an immature “default” state when developmental interaction with the anterior pathway is reduced or eliminated. © 2013 Wiley Periodicals, Inc. Develop Neurobiol 74: 574–590, 2014     

Behavioral discrimination of sexually dimorphic calls by male zebra finches requires an intact vocal motor pathway

Vocal communication between zebra finches includes the exchange of long calls (LCs) as well as song. By using this natural call behavior and quantifying the LCs emitted in response to playbacks of LCs of other birds, we have previously shown that adult male zebra finches have a categorical preference for the LCs of females over those of males. Female LCs are acoustically simpler than male LCs, which include complex acoustic features that are learned during development. Production of these male-typical features requires an intact nucleus RA, the sexually dimorphic source of the main telencephalic projection to brainstem vocal effectors. We have now made bilateral lesions of RA in 17 adult males and tested their discrimination behavior in the call response situation. Lesioned birds continue to call, but lose the male-typical preference for female LCs. The degree of loss is correlated with the extent of RA damage. Further, the simplified LCs of males with RA lesions have a variable duration that is correlated with stimulus features. In effect, the call response behavior of lesioned males becomes like that of females. Apparently, in the absence of RA, the remaining intact structures receive different call information than RA normally does, and/or process it differently. This suggests that the vocal motor nucleus RA could play a role in the transformation of a signal encoding the salience of stimulus parameters into a control signal that modulates the probability and strength of responding.     

Matters of life and death in the songbird forebrain

Male zebra finches learn a specific vocal pattern during a restricted period of development. They produce that song in stereotyped form throughout adulthood, and are unable to learn new song patterns. Development of the neural substrate for song learning and behavior is delayed relative to other brain regions, and neural song-control circuits undergo dramatic changes during the period of vocal learning due to both loss of neurons as well as incorporation of newly generated neurons. In contrast, canaries do learn new song patterns in adulthood and modify their vocal repertoires each breeding season. Adult canaries also maintain a large population of dividing cells in the ependymal zone of the telencephalon, and vast numbers of newly generated neurons migrate out to become incorporated into functional circuits and replace older neurons. We review the relationships between cellular and behavioral aspects of song learning in both zebra finches and canaries, as well as the role of gonadal hormones in regulating diverse aspects of the song-control system. © 1992 John Wiley & Sons, Inc.     

Manipulations of sensory experiences during development reveal mechanisms underlying vocal learning biases in zebra finches

Biological predispositions in learning can bias and constrain the cultural evolution of social and communicative behaviors (e.g., speech and birdsong), and lead to the emergence of behavioral and cultural “universals.” For example, surveys of laboratory and wild populations of zebra finches (Taeniopygia guttata) document consistent patterning of vocal elements (“syllables”) with respect to their acoustic properties (e.g., duration, mean frequency). Furthermore, such universal patterns are also produced by birds that are experimentally tutored with songs containing randomly sequenced syllables (“tutored birds”). Despite extensive demonstrations of learning biases, much remains to be uncovered about the nature of biological predispositions that bias song learning and production in songbirds. Here, we examined the degree to which “innate” auditory templates and/or biases in vocal motor production contribute to vocal learning biases and production in zebra finches. Such contributions can be revealed by examining acoustic patterns in the songs of birds raised without sensory exposure to song (“untutored birds”) or of birds that are unable to hear from early in development (“early‐deafened birds”). We observed that untutored zebra finches and early‐deafened zebra finches produce songs with positional variation in some acoustic features (e.g., mean frequency) that resemble universal patterns observed in tutored birds. Similar to tutored birds, early‐deafened birds also produced song motifs with alternation in acoustic features across adjacent syllables. That universal acoustic patterns are observed in the songs of both untutored and early‐deafened birds highlights the contribution motor production biases to the emergence of universals in culturally transmitted behaviors.     

The song must go on: resilience of the songbird vocal motor pathway

Stereotyped sequences of neural activity underlie learned vocal behavior in songbirds; principle neurons in the cortical motor nucleus HVC fire in stereotyped sequences with millisecond precision across multiple renditions of a song. The geometry of neural connections underlying these sequences is not known in detail though feed-forward chains are commonly assumed in theoretical models of sequential neural activity. In songbirds, a well-defined cortical-thalamic motor circuit exists but little is known the fine-grain structure of connections within each song nucleus. To examine whether the structure of song is critically dependent on long-range connections within HVC, we bilaterally transected the nucleus along the anterior-posterior axis in normal-hearing and deafened birds. The disruption leads to a slowing of song as well as an increase in acoustic variability. These effects are reversed on a time-scale of days even in deafened birds or in birds that are prevented from singing post-transection. The stereotyped song of zebra finches includes acoustic details that span from milliseconds to seconds--one of the most precise learned behaviors in the animal kingdom. This detailed motor pattern is resilient to disruption of connections at the cortical level, and the details of song variability and duration are maintained by offline homeostasis of the song circuit.     

Conspecific and heterospecific song discrimination in male zebra finches with lesions in the anterior forebrain pathway

Adult zebra finches can produce normal song in the absence of Area X, lMAN, or DLM, nuclei that constitute the anterior forebrain pathway of songbirds. Here, we address whether lesions involving Area X and lMAN affect adult male zebra finches' ability to discriminate between conspecific or heterospecific songs. Intact birds and lesioned birds were trained on an operant GO/NOGO conditioning paradigm to discriminate between hetero- or conspecific songs. Both lesioned and intact birds were able to learn all discriminations. Lesioned and intact birds performed equivalently on canary song discriminations. In contrast, discriminations involving bird's own song took significantly more trials to learn for lesioned birds than for intact birds. Discrimination between conspecific songs in general also took longer in the lesioned birds, but missed significance level. Birds with control lesions medial to Area X did not show any differences from intact animals. Our results suggest that an intact anterior forebrain pathway is not required to discriminate between heterospecific songs. In contrast, Area X and lMAN contribute to a male zebra finch's ability to discriminate between its own song and that of other zebra finches. © 1998 John Wiley & Sons, Inc. J Neurobiol 36: 81–90, 1998