The acquisition of an acoustic template is a fundamental component of vocal imitation learning, which is used to refine innate vocalizations and develop a species-specific song. In the absence of a model, birds fail to develop species typical songs. In zebra finches (Taeniopygia guttata), tutored birds produce songs with a stereotyped sequence of distinct acoustic elements, or notes, which form the song motif. Songs of untutored individuals feature atypical acoustic and temporal structure. Here we studied songs and associated respiratory patterns of tutored and untutored male zebra finches to investigate whether similar acoustic notes influence the sequence of song elements. A subgroup of animals developed songs with multiple acoustically similar notes that are produced with alike respiratory motor gestures. These birds also showed increased syntactic variability in their adult motif. Sequence variability tended to occur near song elements which showed high similarity in acoustic structure and underlying respiratory motor gestures. The duration and depth of the inspirations preceding the syllables where syntactic variation occurred did not allow prediction of the following sequence of notes, suggesting that the varying duration and air requirement of the following expiratory pulse is not predictively encoded in the motor program. This study provides a novel method for calculation of motor/acoustic similarity, and the results of this study suggest that the note is a fundamental acoustic unit in the organization of the motif and could play a role in the neural code for song syntax.

Rough-and-tumble play in juvenile rats and song in flocks of adult songbirds outside a breeding context (gregarious song) are two distinct forms of non-sexual social behavior. Both are believed to play roles in the development of sociomotor skills needed for later life-history events, including reproduction, providing opportunities for low-stakes practice. Additionally, both behaviors are thought to be intrinsically rewarded and are associated with a positive affective state. Given the functional similarities of these behaviors, this study used RNA-sequencing to identify commonalities in their underlying neurochemical systems within the medial preoptic area. This brain region is implicated in multiple social behaviors, including song and play, and is highly conserved across vertebrates. DESeq2 and rank-rank hypergeometric overlap analyses identified a shared neurotranscriptomic profile in adult European starlings singing high rates of gregarious song and juvenile rats playing at high rates. Transcript levels for several glutamatergic receptor genes, such as GRIN1, GRIN2A, and GRIA1, were consistently upregulated in highly gregarious (i.e., playful/high singing) animals. This study is the first to directly investigate shared neuromodulators of positive, non-sexual social behaviors across songbirds and mammals. It provides insight into a conserved brain region that may regulate similar behaviors across vertebrates.

In adult songbirds, new neurons are born in large numbers in the proliferative ventricular zone in the telencephalon and migrate to the adjacent song control region HVC (acronym used as proper name) [A. Reiner et al., J. Comp. Neurol. 473, 377-414 (2004)]. Many of these new neurons send long axonal projections to the robust nucleus of the arcopallium (RA). The HVC-RA circuit is essential for producing stereotyped learned song. The function of adult neurogenesis in this circuit has not been clear. A previous study suggested that it is important for the production of well-structured songs [R. E. Cohen, M. Macedo-Lima, K. E. Miller, E. A. Brenowitz, J. Neurosci. 36, 8947-8956 (2016)]. We tested this hypothesis by infusing the neuroblast migration inhibitor cyclopamine into HVC of male Gambel\'s white-crowned sparrows (Zonotrichia leucophrys gambelii) to block seasonal regeneration of the HVC-RA circuit. Decreasing the number of new neurons in HVC prevented both the increase in spontaneous electrical activity of RA neurons and the improved structure of songs that would normally occur as sparrows enter breeding condition. These results show that the incorporation of new neurons into the adult HVC is necessary for the recovery of both electrical activity and song behavior in breeding birds and demonstrate the value of the bird song system as a model for investigating adult neurogenesis at the level of long projection neural circuits.

Songbirds\' vocal mastery is impressive, but to what extent is it a result of practice? Can they, based on experienced mismatch with a known target, plan the necessary changes to recover the target in a practice-free manner without intermittently singing? In adult zebra finches, we drive the pitch of a song syllable away from its stable (baseline) variant acquired from a tutor, then we withdraw reinforcement and subsequently deprive them of singing experience by muting or deafening. In this deprived state, birds do not recover their baseline song. However, they revert their songs toward the target by about 1 standard deviation of their recent practice, provided the sensory feedback during the latter signaled a pitch mismatch with the target. Thus, targeted vocal plasticity does not require immediate sensory experience, showing that zebra finches are capable of goal-directed vocal planning.

Cultural and genetic inheritance combine to enable rapid changes in trait expression, but their relative importance in determining trait expression across generations is not clear. Birdsong is a socially learned cognitive trait that is subject to both cultural and genetic inheritance, as well as being affected by early developmental conditions. We sought to test whether early-life conditions in one generation can affect song acquisition in the next generation. We exposed one generation (F1) of nestlings to elevated corticosterone (CORT) levels, allowed them to breed freely as adults, and quantified their son\'s (F2) ability to copy the song of their social father. We also quantified the neurogenetic response to song playback through immediate early gene (IEG) expression in the auditory forebrain. F2 males with only one corticosterone-treated parent copied their social father\'s song less accurately than males with two control parents. Expression of ARC in caudomedial nidopallium (NCM) correlated with father-son song similarity, and patterns of expression levels of several IEGs in caudomedial mesopallium (CMM) in response to father song playback differed between control F2 sons and those with a CORT-treated father only. This is the first study to demonstrate that developmental conditions can affect social learning and neurogenetic responses in a subsequent generation.

Acoustic signalling is crucial in affecting movements and in social interactions. In species with dynamic social structures, such as multi-level societies, acoustic signals can provide a key mechanism allowing individuals to identify and find or avoid each other and to exchange information. Yet, if the spacing between individuals regularly exceeds the maximum signalling range, the relation between movements and signals becomes more complex. As the best-studied songbird in captivity, the zebra finch (Taeniopygia castanotis) is a species with individually distinct songs that are audible over just a few metres and a widely ranging dynamic multi-level social organization in the wild, raising questions on the actual role of its song in social cohesion and coordination. Here, we provide an overview of birdsong in social organizations (networks) and use the ecology of the zebra finch and male song to discuss how singing can facilitate social cohesion and coordination in species where the signal range is very short. We raise the question of the extent to which zebra finches are a representative species to understand the function of song in communication, and we broaden current views on the function of birdsong and its individual signature. This article is part of the theme issue \'The power of sound: unravelling how acoustic communication shapes group dynamics\'.

Vocalization is an important communication tool that can reflect many aspects of an individual\'s internal and external condition. This is especially true for birds. Previous research has shown that bird calls and songs change in response to a variety of potential stressors, although the extent and direction of the changes depend on the nature of the stressor and the environment. Circulating glucocorticoids, such as corticosterone, often increase in response to stressors and mediate some of the observed changes via alterations of the individual\'s physiological state. Acute elevations of corticosterone often occur as a physiological response to short-term stressors; however, the effects of this elevation on adult vocalizations have not been well documented. Here, we experimentally elevated corticosterone at two different levels using a noninvasive method and examined the effects on the vocal communication of male and female adult common mynas (Acridotheres tristis). Corticosterone elevation temporarily increased song output and some measures of song complexity, while call output decreased. These effects were dosage dependent (higher corticosterone levels had a stronger effect), most evident 40 min after ingestion, and some vocal changes were sex-specific. Future studies should investigate whether the changes in vocal performance due to elevated glucocorticoids have consequences for the birds\' behavior, reproductive success, and survival.

Communication needs to be complex enough to be functional while minimizing learning and production costs. Recent work suggests that the vocalizations and gestures of some songbirds, cetaceans and great apes may conform to linguistic laws that reflect this trade-off between efficiency and complexity. In studies of non-human communication, though, clustering signals into types cannot be done a priori, and decisions about the appropriate grain of analysis may affect statistical signals in the data. The aim of this study was to assess the evidence for language-like efficiency and structure in house finch (Haemorhous mexicanus) song across three levels of granularity in syllable clustering. The results show strong evidence for Zipf\'s rank-frequency law, Zipf\'s law of abbreviation and Menzerath\'s law. Additional analyses show that house finch songs have small-world structure, thought to reflect systematic structure in syntax, and the mutual information decay of sequences is consistent with a combination of Markovian and hierarchical processes. These statistical patterns are robust across three levels of granularity in syllable clustering, pointing to a limited form of scale invariance. In sum, it appears that house finch song has been shaped by pressure for efficiency, possibly to offset the costs of female preferences for complexity.

In sexually dimorphic zebra finches (Taeniopygia guttata), only males learn to sing their father\'s song, whereas females learn to recognize the songs of their father or mate but cannot sing themselves. Memory of learned songs is behaviorally expressed in females by preferring familiar songs over unfamiliar ones. Auditory association regions such as the caudomedial mesopallium (CMM; or caudal mesopallium) have been shown to be key nodes in a network that supports preferences for learned songs in adult females. However, much less is known about how song preferences develop during the sensitive period of learning in juvenile female zebra finches. In this study, we used blood-oxygen level-dependent (BOLD) functional magnetic resonance imaging (fMRI) to trace the development of a memory-based preference for the father\'s song in female zebra finches. Using BOLD fMRI, we found that only in adult female zebra finches with a preference for learned song over novel conspecific song, neural selectivity for the father\'s song was localized in the thalamus (dorsolateral nucleus of the medial thalamus; part of the anterior forebrain pathway, AFP) and in CMM. These brain regions also showed a selective response in juvenile female zebra finches, although activation was less prominent. These data reveal that neural responses in CMM, and perhaps also in the AFP, are shaped during development to support behavioral preferences for learned songs.

Migratory species inhabit many communities along their migratory routes. Across taxa, these species repeatedly move into and out of communities, interacting with each other and locally breeding species and competing for resources and niche space. However, their influence is rarely considered in analyses of ecological processes within the communities they temporarily occupy. Here, we explore the impact of migratory species on a breeding community using the framework of acoustic signal space, a limited resource in which sounds of species within communities co-exist. Migrating New World warblers (Parulidae, hereafter referred to as migrant species) often sing during refueling stops in areas and at times during which locally breeding warbler species (hereafter breeding species) are singing to establish territories and attract mates. We used eBird data to determine the co-occurrence of 19 migrant and 11 breeding warbler species across spring migration in SW Michigan, generated a signal space from song recordings of these species, and examined patterns of signaling overlap experienced by breeding species as migrants moved through the community. Migrant species were present for two-thirds of the breeding season of local species, including periods when breeding species established territories and attracted mates. Signaling niche overlap experienced by individual breeding species was idiosyncratic and varied over time, yet niche overlap between migrant and breeding species occurred more commonly than between breeding species or between migrant species. Nevertheless, the proportion of niche overlap between migrant and breeding warblers was similar to overlap among breeding species. Our findings showed that singing by migrant species overlapped the signals of many breeding species, suggesting that migrants could have unexplored impacts on communication in breeding species, potentially affecting song detection and song evolution. Our study contributes to a growing body of research documenting the impacts of migratory species on communities and ecosystems.