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Background: Understanding the neural basis of behavior requires insight into how different brain systems coordinate with each other. Existing connectomes for various species have highlighted brain systems essential to various aspects of behavior, yet their application to complex learned behaviors remains limited. Research on vocal learning in songbirds has extensively focused on the vocal control network, though recent work implicates a variety of circuits in contributing to important aspects of vocal behavior. Thus, a more comprehensive understanding of brain-wide connectivity is essential to further assess the totality of circuitry underlying this complex learned behavior. Results: We present the Oscine Structural Connectome for Investigating NEural NETwork ORGanization (OSCINE-NET.ORG), the first interactive mesoscale connectome for any vocal learner. This comprehensive digital map includes all known connectivity data, covering major brain superstructures and functional networks. Our analysis reveals that the songbird brain exhibits small-world properties, with highly connected communities functionally designated as motor, visual, associative, vocal, social, and auditory. Moreover, there is a small set of significant connections across these communities, including from social and auditory sub-communities to vocal sub-communities, which highlight ethologically relevant facets of vocal learning and production. Notably, the vocal community contains the majority of the canonical vocal control network, as well as a variety of other nodes that are highly interconnected with it, meriting further evaluation for their inclusion in this network. A subset of nodes forms a "rich broker club," highly connected across the brain and forming a small circuit amongst themselves, indicating they may play a key role in information transfer broadly. Collectively, their bidirectional connectivity with multiple communities indicates they may act as liaisons across multiple functional circuits for a variety of complex behaviors. Conclusions: OSCINE-NET.ORG offers unprecedented access to detailed songbird connectivity data, promoting insight into the neural circuits underlying complex behaviors. This data emphasizes the importance of brain-wide integration in vocal learning, facilitating a potential reevaluation of the canonical vocal control network. Furthermore, we computationally identify a small, previously unidentified circuit—one which may play an impactful role in brain-wide coordination of multiple complex behaviors. [ABSTRACT FROM AUTHOR]

Zebra finches undergo a gradual refinement of their vocalizations, transitioning from variable juvenile songs to the stereotyped song of adulthood. To investigate the neural mechanisms underlying song crystallization—a critical phase in this developmental process—we performed intracellular recordings in HVC (a premotor nucleus essential for song learning and production) of juvenile birds. We then compared these recordings to previously published electrophysiological data from adult birds. We found that HVC projection neurons in juvenile zebra finches during the song crystallization phase exhibited more variable spiking patterns compared to the precise bursting observed in adult HVC projection neurons. Additionally, subthreshold membrane potential fluctuations in juvenile neurons exhibited longer durations and larger amplitude excitatory postsynaptic potentials. These distinct temporal dynamics in HVC during song crystallization likely play a crucial role in the fine-tuning processes that shape the precise timing and structure of the mature zebra finch song. [ABSTRACT FROM AUTHOR]

In this article, I present three main points that could benefit the "vocal learning and rhythmic synchronization hypothesis", encompassing neurogenetic mechanisms of gene expression transmission and single motor neuron function, classification of different behavioral motor phenotypes (e.g., spontaneous vs. voluntary), and other evolutionary considerations (i.e., the involvement of reward mechanisms). [ABSTRACT FROM AUTHOR]

Each commentary on my article raises important points and new ideas for research on rhythmic processing in humans and other species. Here I respond to points concerning the role of social factors in the ontogeny of beat synchronization, the neural connectivity underlying beat synchronization, the evolution of this connectivity, and the mechanisms by which evolutionary changes in the strength of one white matter tract (driven by natural selection) can have knock-on effects on the structure of an adjacent tract. [ABSTRACT FROM AUTHOR]

Dancing to music is ancient and widespread in human cultures. While dance shows great cultural diversity, it often involves nonvocal rhythmic movements synchronized to musical beats in a predictive and tempo-flexible manner. To date, the only nonhuman animals known to spontaneously move to music in this way are parrots. This paper proposes that human-parrot similarities in movement to music and in the neurobiology of advanced vocal learning hold clues to the evolutionary foundations of human dance. The proposal draws on recent research on the neurobiology of parrot vocal learning by Jarvis and colleagues and on a recent cortical model for speech motor control by Hickock and colleagues. These two lines of work are synthesized to suggest that gene regulation changes associated with the evolution of a dorsal laryngeal pitch control pathway in ancestral humans fortuitously strengthened auditory-parietal cortical connections that support beat-based rhythmic processing. More generally, the proposal aims to explain how and why the evolution of strong forebrain auditory-motor integration in the service of learned vocal control led to a capacity and proclivity to synchronize nonvocal movements to the beat. The proposal specifies cortical brain pathways implicated in the origins of human beat-based dancing and leads to testable predictions and suggestions for future research. [ABSTRACT FROM AUTHOR]

Abstract Dancing to music is ancient and widespread in human cultures. While dance shows great cultural diversity, it often involves nonvocal rhythmic movements synchronized to musical beats in a predictive and tempo-flexible manner. To date, the only nonhuman animals known to spontaneously move to music in this way are parrots. This paper proposes that human-parrot similarities in movement to music and in the neurobiology of advanced vocal learning hold clues to the evolutionary foundations of human dance. The proposal draws on recent research on the neurobiology of parrot vocal learning by Jarvis and colleagues and on a recent cortical model for speech motor control by Hickock and colleagues. These two lines of work are synthesized to suggest that gene regulation changes associated with the evolution of a dorsal laryngeal pitch control pathway in ancestral humans fortuitously strengthened auditory-parietal cortical connections that support beat-based rhythmic processing. More generally, the proposal aims to explain how and why the evolution of strong forebrain auditory-motor integration in the service of learned vocal control led to a capacity and proclivity to synchronize nonvocal movements to the beat. The proposal specifies cortical brain pathways implicated in the origins of human beat-based dancing and leads to testable predictions and suggestions for future research.

Abstract Each commentary on my article raises important points and new ideas for research on rhythmic processing in humans and other species. Here I respond to points concerning the role of social factors in the ontogeny of beat synchronization, the neural connectivity underlying beat synchronization, the evolution of this connectivity, and the mechanisms by which evolutionary changes in the strength of one white matter tract (driven by natural selection) can have knock-on effects on the structure of an adjacent tract.

Abstract In this article, I present three main points that could benefit the “vocal learning and rhythmic synchronization hypothesis”, encompassing neurogenetic mechanisms of gene expression transmission and single motor neuron function, classification of different behavioral motor phenotypes (e.g., spontaneous vs. voluntary), and other evolutionary considerations (i.e., the involvement of reward mechanisms).

Harbor seals (Phoca vitulina) are more solitary than many other pinnipeds. Yet, they are capable of vocal learning, a form of social learning. Most extant literature examines social animals when investigating social learning, despite sociality not being a prerequisite. Here, we report two formerly silent harbor seals who initiated vocalizations, after having repeatedly observed a conspecific receiving food rewards for vocalizing. Our observations suggest both social and vocal learning in a group of captive harbor seals, a species that lives semi-solitarily in the wild. We propose that, in this case, social learning acted as a shortcut to acquiring food rewards compared to the comparatively costly asocial learning.

Abstract Background Which mammals show vocal learning abilities, e.g., can learn new sounds, or learn to use sounds in new contexts? Vocal usage and comprehension learning are submodules of vocal learning. Specifically, vocal usage learning is the ability to learn to use a vocalization in a new context; vocal comprehension learning is the ability to comprehend a vocalization in a new context. Among mammals, harbor seals (Phoca vitulina) are good candidates to investigate vocal learning. Here, we test whether harbor seals are capable of vocal usage and comprehension learning. Results We trained two harbor seals to (i) switch contexts from a visual to an auditory cue. In particular, the seals first produced two vocalization types in response to two hand signs; they then transitioned to producing these two vocalization types upon the presentation of two distinct sets of playbacks of their own vocalizations. We then (ii) exposed the seals to a combination of trained and novel vocalization stimuli. In a final experiment, (iii) we broadcasted only novel vocalizations of the two vocalization types to test whether seals could generalize from the trained set of stimuli to only novel items of a given vocal category. Both seals learned all tasks and took ≤ 16 sessions to succeed across all experiments. In particular, the seals showed contextual learning through switching the context from former visual to novel auditory cues, vocal matching and generalization. Finally, by responding to the played-back vocalizations with distinct vocalizations, the animals showed vocal comprehension learning. Conclusions It has been suggested that harbor seals are vocal learners; however, to date, these observations had not been confirmed in controlled experiments. Here, through three experiments, we could show that harbor seals are capable of both vocal usage and comprehension learning.

Background: Which mammals show vocal learning abilities, e.g., can learn new sounds, or learn to use sounds in new contexts? Vocal usage and comprehension learning are submodules of vocal learning. Specifically, vocal usage learning is the ability to learn to use a vocalization in a new context; vocal comprehension learning is the ability to comprehend a vocalization in a new context. Among mammals, harbor seals (Phoca vitulina) are good candidates to investigate vocal learning. Here, we test whether harbor seals are capable of vocal usage and comprehension learning. Results: We trained two harbor seals to (i) switch contexts from a visual to an auditory cue. In particular, the seals first produced two vocalization types in response to two hand signs; they then transitioned to producing these two vocalization types upon the presentation of two distinct sets of playbacks of their own vocalizations. We then (ii) exposed the seals to a combination of trained and novel vocalization stimuli. In a final experiment, (iii) we broadcasted only novel vocalizations of the two vocalization types to test whether seals could generalize from the trained set of stimuli to only novel items of a given vocal category. Both seals learned all tasks and took ≤ 16 sessions to succeed across all experiments. In particular, the seals showed contextual learning through switching the context from former visual to novel auditory cues, vocal matching and generalization. Finally, by responding to the played-back vocalizations with distinct vocalizations, the animals showed vocal comprehension learning. Conclusions: It has been suggested that harbor seals are vocal learners; however, to date, these observations had not been confirmed in controlled experiments. Here, through three experiments, we could show that harbor seals are capable of both vocal usage and comprehension learning. [ABSTRACT FROM AUTHOR]

Abstract Background Most vocal learning species exhibit an early critical period during which their vocal control neural circuitry facilitates the acquisition of new vocalizations. Some taxa, most notably humans and parrots, retain some degree of neurobehavioral plasticity throughout adulthood, but both the extent of this plasticity and the neurogenetic mechanisms underlying it remain unclear. Differential expression of the transcription factor FoxP2 in both songbird and parrot vocal control nuclei has been identified previously as a key pattern facilitating vocal learning. We hypothesize that the resilience of vocal learning to cognitive decline in open-ended learners will be reflected in an absence of age-related changes in neural FoxP2 expression. We tested this hypothesis in the budgerigar (Melopsittacus undulatus), a small gregarious parrot in which adults converge on shared call types in response to shifts in group membership. We formed novel flocks of 4 previously unfamiliar males belonging to the same age class, either “young adult” (6 mo − 1 year) or “older adult” (≥ 3 year), and then collected audio-recordings over a 20-day learning period to assess vocal learning ability. Following behavioral recording, immunohistochemistry was performed on collected neural tissue to measure FoxP2 protein expression in a parrot vocal learning center, the magnocellular nucleus of the medial striatum (MMSt), and its adjacent striatum. Results Although older adults show lower vocal diversity (i.e. repertoire size) and higher absolute levels of FoxP2 in the MMSt than young adults, we find similarly persistent downregulation of FoxP2 and equivalent vocal plasticity and vocal convergence in the two age cohorts. No relationship between individual variation in vocal learning measures and FoxP2 expression was detected. Conclusions We find neural evidence to support persistent vocal learning in the budgerigar, suggesting resilience to aging in the open-ended learning program of this species. The lack of a significant relationship between FoxP2 expression and individual variability in vocal learning performance suggests that other neurogenetic mechanisms could also regulate this complex behavior.

Vocalizations coordinate social interactions between conspecifics by conveying information concerning the individual or group identity of the sender. Social accommodation is a form of vocal learning where social affinity is signalled by converging or diverging vocalizations with those of conspecifics. To investigate whether social accommodation is linked to the social lifestyle of the sender, we investigated sex-specific differences in social accommodation in a dispersed living primate, the grey mouse lemur (Microcebus murinus), where females form stable sleeping groups whereas males live solitarily. We used 482 trill calls of 36 individuals from our captive breeding colony to compare acoustic dissimilarity between individuals with genetic relatedness, social contact time and body weight. Our results showed that female trills become more similar the more time females spend with each other, independent of genetic relationship, suggesting vocal convergence. In contrast, male trills were affected more by genetic than social factors. However, focusing only on socialized males, increasing time as cage partners caused greater divergence in males' trills. Thus, grey mouse lemurs show the capacity for social accommodation, with females converging their trills to signal social closeness to sleeping group partners, whereas males do not adapt or diverge their trills to signal individual distinctiveness. This article is part of the theme issue 'The power of sound: unravelling how acoustic communication shapes group dynamics'. [ABSTRACT FROM AUTHOR]

Background: Most vocal learning species exhibit an early critical period during which their vocal control neural circuitry facilitates the acquisition of new vocalizations. Some taxa, most notably humans and parrots, retain some degree of neurobehavioral plasticity throughout adulthood, but both the extent of this plasticity and the neurogenetic mechanisms underlying it remain unclear. Differential expression of the transcription factor FoxP2 in both songbird and parrot vocal control nuclei has been identified previously as a key pattern facilitating vocal learning. We hypothesize that the resilience of vocal learning to cognitive decline in open-ended learners will be reflected in an absence of age-related changes in neural FoxP2 expression. We tested this hypothesis in the budgerigar (Melopsittacus undulatus), a small gregarious parrot in which adults converge on shared call types in response to shifts in group membership. We formed novel flocks of 4 previously unfamiliar males belonging to the same age class, either "young adult" (6 mo − 1 year) or "older adult" (≥ 3 year), and then collected audio-recordings over a 20-day learning period to assess vocal learning ability. Following behavioral recording, immunohistochemistry was performed on collected neural tissue to measure FoxP2 protein expression in a parrot vocal learning center, the magnocellular nucleus of the medial striatum (MMSt), and its adjacent striatum. Results: Although older adults show lower vocal diversity (i.e. repertoire size) and higher absolute levels of FoxP2 in the MMSt than young adults, we find similarly persistent downregulation of FoxP2 and equivalent vocal plasticity and vocal convergence in the two age cohorts. No relationship between individual variation in vocal learning measures and FoxP2 expression was detected. Conclusions: We find neural evidence to support persistent vocal learning in the budgerigar, suggesting resilience to aging in the open-ended learning program of this species. The lack of a significant relationship between FoxP2 expression and individual variability in vocal learning performance suggests that other neurogenetic mechanisms could also regulate this complex behavior. [ABSTRACT FROM AUTHOR]

A recent reshuffling in the avian phylogeny indicates that New Zealand wrens and songbirds share a close common ancestor with parrots – making New Zealand wrens an excellent group to test for vocal learning. The New Zealand wrens have previously been classified as vocal non-learners, but their vocal learning ability has never been experimentally tested. Here, we explore the potential presence of vocal learning in one species of New Zealand wren, the titipounamu (Acanthisitta chloris). We expose nestlings to synthetic playback stimuli that simulate adult feeding calls and determine whether the nestlings altered their calls after exposure to the playback stimuli. We found that experimental nestlings did not alter their calls towards the playback stimuli. While this indicates that no vocal imitation occurred during the nestling period, other developmental stages should be tested for vocal learning, particularly during the fledgling or first year adult phase. [ABSTRACT FROM AUTHOR]

Animals' learned vocalizations often show high levels of intraspecific variation, and different variants may play different roles in attracting mates and settling territorial disputes. In any animal population with variable vocalizations, certain acoustic variants may be common and others rare. The responses of animals to local versus nonlocal sounds have been well characterized in prior investigations, but few investigations have explored the reactions of animals to songs that are present at different frequencies of occurrence within a population. Using an experimental approach to investigate a breeding population of Savannah sparrows, Passerculus sandwichensis , we tested the responses of territorial male songbirds to playback of locally common songs, locally rare songs and heterospecific songs. Male sparrows showed strong responses to conspecific song but weak responses to heterospecific song. Males showed their strongest responses to locally common songs and weaker responses to locally rare songs. Birds did not appear to discriminate between playback songs based on whether they were of the same song type as their own. We suggest that male Savannah sparrows perceive locally rare songs as weaker threats than locally common songs, similar to birds' reactions to foreign dialects. This could be due to low familiarity with locally rare songs, reduced female preference for locally rare songs, decreased success with territory defence for males singing locally rare songs or other perceived differences between locally rare and locally common songs. Overall, our results suggest that different song types elicit different responses from territorial songbirds, even songs that are found within the same population with different frequencies of occurrence. • Many animals show small-scale geographical variation in vocalizations. • Responses to locally common or locally rare vocalizations are seldom explored. • We used playback to test birds' responses to locally common and locally rare songs. • Savannah sparrows responded more strongly to locally common songs. • We suggest locally common songs are perceived as a stronger threat. [ABSTRACT FROM AUTHOR]

Rare disruptions of the transcription factor FOXP1 are implicated in a human neurodevelopmental disorder characterized by autism and/or intellectual disability with prominent problems in speech and language abilities. Avian orthologues of this transcription factor are evolutionarily conserved and highly expressed in specific regions of songbird brains, including areas associated with vocal production learning and auditory perception. Here, we investigated possible contributions of FoxP1 to song discrimination and auditory perception in juvenile and adult female zebra finches. They received lentiviral knockdowns of FoxP1 in one of two brain areas involved in auditory stimulus processing, HVC (proper name) or CMM (caudomedial mesopallium). Ninety-six females, distributed over different experimental and control groups were trained to discriminate between two stimulus songs in an operant Go/Nogo paradigm and subsequently tested with an array of stimuli. This made it possible to assess how well they recognized and categorized altered versions of training stimuli and whether localized FoxP1 knockdowns affected the role of different features during discrimination and categorization of song. Although FoxP1 expression was significantly reduced by the knockdowns, neither discrimination of the stimulus songs nor categorization of songs modified in pitch, sequential order of syllables or by reversed playback were affected. Subsequently, we analyzed the full dataset to assess the impact of the different stimulus manipulations for cue weighing in song discrimination. Our findings show that zebra finches rely on multiple parameters for song discrimination, but with relatively more prominent roles for spectral parameters and syllable sequencing as cues for song discrimination. NEW & NOTEWORTHY: In humans, mutations of the transcription factor FoxP1 are implicated in speech and language problems. In songbirds, FoxP1 has been linked to male song learning and female preference strength. We found that FoxP1 knockdowns in female HVC and caudomedial mesopallium (CMM) did not alter song discrimination or categorization based on spectral and temporal information. However, this large dataset allowed to validate different cue weights for spectral over temporal information for song recognition. [ABSTRACT FROM AUTHOR]

Vocalizations often vary in structure within a species, from the individual to population level. Vocal differences among social groups and populations can provide insight into biological processes such as vocal learning and evolutionary divergence, with important conservation implications. As vocal learners of conservation concern, intraspecific vocal variation is of particular interest in elephants. We recorded calls from individuals in multiple, wild elephant social groups in two distinct Kenyan populations. We used machine learning to investigate vocal differentiation among individual callers, core groups, bond groups (collections of core groups) and populations. We found clear evidence for vocal distinctiveness at the individual and population level, and evidence for much subtler vocal differences among social groups. Social group membership was a better predictor of call similarity than genetic relatedness, suggesting that subtle vocal differences among social groups may be learned. Vocal divergence among populations and social groups has conservation implications for the effects of social disruption and translocation of elephants.

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.

Prelinguistic babbling is a critical phase in infant language development and is best understood in temperate songbirds where it occurs primarily in males at reproductive maturity and is modulated by sex steroids. Parrots of both sexes are icons of tropical vocal plasticity, but vocal babbling is unreported in this group and whether the endocrine system is involved is unknown. Here we show that vocal babbling is widespread in a wild parrot population in Venezuela, ensues in both sexes during the nestling stage, occurs amidst a captive audience of mixed-aged siblings, and is modulated by corticosteroids. Spectrographic analysis and machine learning found phoneme diversity and combinatorial capacity increased precipitously for the first week, thereafter, crystalizing into a smaller repertoire, consistent with the selective attrition model of language development. Corticosterone-treated nestlings differed from unmanipulated birds and sham controls in several acoustic properties and crystallized a larger repertoire post-treatment. Our findings indicate babbling occurs during an early life-history stage in which corticosteroids help catalyse the transition from a universal learning programme to one finely tuned for the prevailing ecological environment, a potentially convergent scenario in human prelinguistic development.

Acoustic signals are ubiquitous across mammalian taxa. They serve a myriad of functions related to the formation and maintenance of social bonds and can provide conspecifics information about caller condition, motivation and identity. Disentangling the relative importance of evolutionary mechanisms that shape vocal variation is difficult, and little is known about heritability of mammalian vocalizations. Duetting--coordinated vocalizations within male and female pairs--arose independently at least four times across the Primate Order. Primate duets contain individual- or pair-level signatures, but the mechanisms that shape this variation remain unclear. Here, we test for evidence of heritability in two call types (pulses and chirps) from the duets of captive coppery titi monkeys (Plecturocebus cupreus). We extracted four features--note rate, duration, minimum and maximum fundamental frequency--from spectrograms of pulses and chirps, and estimated heritability of the features. We also tested whether features varied with sex or body weight. We found evidence for moderate heritability in one of the features examined (chirp note rate), whereas inter-individual variance was the most important source of variance for the rest of the features. We did not find evidence for sex differences in any of the features, but we did find that body weight and fundamental frequency of chirp elements covaried. Kin recognition has been invoked as a possible explanation for heritability or kin signatures in mammalian vocalizations. Although the function of primate duets remains a topic of debate, the presence of moderate heritability in titi monkey chirp elements indicates duets may serve a kin recognition function.

The acoustic structure of nonhuman primate vocalizations can vary substantially within a call type and may converge between social partners. Examining which social partners share call structure can inform our understanding of the function of vocal learning and communication in nonhuman animals. We assessed vocal convergence of female gelada contact grunts within multiple levels of gelada society: small female-philopatric reproductive units that maintain close proximity and larger bands of units that preferentially associate while foraging. We also measured the extent to which grunt acoustic structure varied by caller identity and behavioural state at the time of the call, as well as genetic relatedness, when assessing acoustic similarity between females' grunts. The acoustic structure of female gelada grunts differed between individuals as well as between foraging, travelling and socializing behavioural states. Female gelada grunts were more similar between pairs living in the same band than between pairs living in different bands. This effect was not found for pairs living in the same unit compared to pairs living in different units within the same band. As genetic relatedness did not predict similarity in grunt acoustic structure, we propose that vocal convergence within bands is the result of vocal learning. Vocal convergence at the higher level of gelada society suggests this vocal learning is selective and not simply the result of auditory exposure to conspecific calls. Vocal convergence in geladas may function to maintain spatial cohesion, which is a more critical challenge for bands than for units. • Female gelada contact calls differed between callers and across behavioural states. • Contact grunts were more similar between females living in the same band. • Genetic relatedness did not predict vocal similarity. • Our results suggest learned vocal convergence. [ABSTRACT FROM AUTHOR]

Introduction: Vocal dialects are a taxonomically widespread phenomenon which are typically only studied in a portion of a species' range. Thus, it is difficult to infer whether a geographic pattern of vocal dialects observed in one part of a species' range are typical across the range or whether local conditions influence their presence or absence. We examined the yellow-naped amazon, Amazona auropalliata, a parrot species with remarkable vocal learning capabilities. Although this species' native range spans across Mesoamerica, only Costa Rican populations have been evaluated long-term. Previous studies have shown that these populations have geographically and temporally stable vocal dialect patterns. Without data on populations outside of Costa Rica, it is impossible to know whether vocal dialects are present in northern range populations, and whether they show similar geographic structure to southern range populations. Introduction: We recorded yellow-naped amazon contact calls at 47 different sites across the species' range between 2016 and 2019 and evaluated them for the presence of dialects. We visually classified 14 contact call types based on spectrographic similarity and used spectrographic cross-correlation, principal component analysis, and Mantel-based spatial autocorrelations to assess acoustic similarity; we also evaluated the robustness of our findings using simulated data. Results and Discussion: The results from our study show that the vocal patterns previously seen in Costa Rica are also present in northern populations, supporting our hypothesis that this species has vocal dialects throughout its Mesoamerican range. Call typeswere regionally specific (e.g., vocal dialects occurred) across the range, and no call types were repeated across multiple regions. We did, however, observe distinctive structural characteristics that are found inmultiple call types, suggesting that different call types stem from a common origin. Alternatively, similarity in the acoustic features of call typesmay also be a result of physiological and anatomical features that are common to allmembers of the species. Vocal dialects in this species are likely maintained through a tendency toward philopatry and matching call types to enhance social identification. [ABSTRACT FROM AUTHOR]

Vocal production learning and beat perception and synchronization (BPS) share some common characteristics, which makes the vocal learning and rhythmic synchronization hypothesis (VLH) a reasonable explanation for the evolution of the capability for rhythmic synchronization. However, even in vocal learners, it is rare to see non-human animals demonstrate BPS to human music. Therefore, the first objective of this article is to propose some possible reasons why we do not see BPS in budgerigars, an excellent vocal learning species, while presenting some of my own findings. The second objective of this article is to propose a seamless bridge to connect the capability for vocal learning and BPS in locomotion. For this purpose, I present my own findings, wherein cockatiels spontaneously sang in synchrony with a melody of human music. This behavior can be considered a vocal version of BPS. Therefore, it can establish a connection between these two capabilities. This article agrees with the possibility that some mechanisms other than the vocal learning system may enable BPS, contrary to the original idea of VLH. Nevertheless, it is still reasonable to connect the capability for vocal learning and that for BPS. At the very least, the capability for vocal learning may contribute to the evolution of BPS. From these arguments, this article also proposes a scenario which includes vocalizing in synchrony as a driving force for the evolution of BPS and the capability for music production. [ABSTRACT FROM AUTHOR]

Sight-singing and ear training is one of the essential introductory courses for music majors, and the effectiveness of sight-singing and ear training teaching directly affects students’ growth and progress in music. In this paper, we have combined relevant contents of sight-singing, ear training, and vocal learning to construct a flipped classroom teaching model for sight-singing and ear training using the flipped classroom teaching process. Secondly, taking the students of L Conservatory of Music as the research object, two different questionnaires were designed for students and teachers, and the teaching experiment comparison method was used to test the role of sight-singing and ear-training flipped classroom on vocal music learning skills. Finally, the parameters of the multiple linear regression model were estimated using the least squares method, and the model was used to analyze the specific effects of sight-singing and ear training on vocal learning skills. It was found that after the flipped classroom teaching of sight-singing and ear-training, the ear-training scores of students in the experimental class were 90.43±9.26 points, which were 17.06 points higher than those of students in the control class, and there was no difference in the pre- and post-test scores of sight-singing and ear-training. The regression coefficient of students’ musical expression was 0.306. Every 1 percentage point increase in musical expression would enhance vocal learning skills by 0.306 percentage points. Sight-singing and ear-training can help students better master the basic skills of vocal music, cultivate their sense of rhythm, and enhance their vocal music learning skills.

Animal songs can change within and between populations as the result of different evolutionary processes. When these processes include cultural transmission, the social learning of information or behaviours from conspecifics, songs can undergo rapid evolutions because cultural novelties can emerge more frequently than genetic mutations. Understanding these song variations over large temporal and spatial scales can provide insights into the patterns, drivers and limits of song evolution that can ultimately inform on the species’ capacity to adapt to rapidly changing acoustic environments. Here, we analysed changes in fin whale (Balaenoptera physalus) songs recorded over two decades across the central and eastern North Atlantic Ocean. We document a rapid replacement of song INIs (inter-note intervals) over just four singing seasons, that co-occurred with hybrid songs (with both INIs), and a clear geographic gradient in the occurrence of different song INIs during the transition period. We also found gradual changes in INIs and note frequencies over more than a decade with fin whales adopting song changes. These results provide evidence of vocal learning in fin whales and reveal patterns of song evolution that raise questions on the limits of song variation in this species.

In this article, I am going through my scientific and personal journey using my work on oxytocin as a compass. I recount how my scientific questions were shaped over the years, and how I studied them through the lens of different fields ranging from linguistics and neuroscience to comparative and population genomics in a wide range of vertebrate species. I explain how my evolutionary findings and proposal for a universal gene nomenclature in the oxytocin-vasotocin ligand and receptor families have impacted relevant fields, and how my studies in the oxytocin and vasotocin system in songbirds, humans and non-human primates have led me to now be testing intranasal oxytocin as a candidate treatment for speech deficits. I also discuss my projects on the neurobiology of dance and where oxytocin fits in the picture of studying speech and dance in parallel. Lastly, I briefly communicate the challenges I have been facing as a woman and an international scholar in science and academia, and my personal ways to overcome them.

Why do humans make music? Theories of the evolution of musicality have focused mainly on the value of music for specific adaptive contexts such as mate selection, parental care, coalition signaling, and group cohesion. Synthesizing and extending previous proposals, we argue that social bonding is an overarching function that unifies all of these theories, and that musicality enabled social bonding at larger scales than grooming and other bonding mechanisms available in ancestral primate societies. We combine cross-disciplinary evidence from archeology, anthropology, biology, musicology, psychology, and neuroscience into a unified framework that accounts for the biological and cultural evolution of music. We argue that the evolution of musicality involves gene-culture coevolution, through which proto-musical behaviors that initially arose and spread as cultural inventions had feedback effects on biological evolution because of their impact on social bonding. We emphasize the deep links between production, perception, prediction, and social reward arising from repetition, synchronization, and harmonization of rhythms and pitches, and summarize empirical evidence for these links at the levels of brain networks, physiological mechanisms, and behaviors across cultures and across species. Finally, we address potential criticisms and make testable predictions for future research, including neurobiological bases of musicality and relationships between human music, language, animal song, and other domains. The music and social bonding hypothesis provides the most comprehensive theory to date of the biological and cultural evolution of music.

The temporal organization of sounds used in social contexts can provide information about signal function and evoke varying responses in listeners (receivers). For example, music is a universal and learned human behavior that is characterized by different rhythms and tempos that can evoke disparate responses in listeners. Similarly, birdsong is a social behavior in songbirds that is learned during critical periods in development and used to evoke physiological and behavioral responses in receivers. Recent investigations have begun to reveal the breadth of universal patterns in birdsong and their similarities to common patterns in speech and music, but relatively little is known about the degree to which biological predispositions and developmental experiences interact to shape the temporal patterning of birdsong. Here, we investigated how biological predispositions modulate the acquisition and production of an important temporal feature of birdsong, namely the duration of silent pauses ("gaps") between vocal elements ("syllables"). Through analyses of semi‐naturally raised and experimentally tutored zebra finches, we observed that juvenile zebra finches imitate the durations of the silent gaps in their tutor's song. Further, when juveniles were experimentally tutored with stimuli containing a wide range of gap durations, we observed biases in the prevalence and stereotypy of gap durations. Together, these studies demonstrate how biological predispositions and developmental experiences differently affect distinct temporal features of birdsong and highlight similarities in developmental plasticity across birdsong, speech, and music. Research Highlights: The temporal organization of learned acoustic patterns can be similar across human cultures and across species, suggesting biological predispositions in acquisition.We studied how biological predispositions and developmental experiences affect an important temporal feature of birdsong, namely the duration of silent intervals between vocal elements ("gaps").Semi‐naturally and experimentally tutored zebra finches imitated the durations of gaps in their tutor's song and displayed some biases in the learning and production of gap durations and in gap variability.These findings in the zebra finch provide parallels with the acquisition of temporal features of speech and music in humans. [ABSTRACT FROM AUTHOR]

Why do humans make music? Theories of the evolution of musicality have focused mainly on the value of music for specific adaptive contexts such as mate selection, parental care, coalition signaling, and group cohesion. Synthesizing and extending previous proposals, we argue that social bonding is an overarching function that unifies all of these theories, and that musicality enabled social bonding at larger scales than grooming and other bonding mechanisms available in ancestral primate societies. We combine cross-disciplinary evidence from archeology, anthropology, biology, musicology, psychology, and neuroscience into a unified framework that accounts for the biological and cultural evolution of music. We argue that the evolution of musicality involves gene-culture coevolution, through which proto-musical behaviors that initially arose and spread as cultural inventions had feedback effects on biological evolution because of their impact on social bonding. We emphasize the deep links between production, perception, prediction, and social reward arising from repetition, synchronization, and harmonization of rhythms and pitches, and summarize empirical evidence for these links at the levels of brain networks, physiological mechanisms, and behaviors across cultures and across species. Finally, we address potential criticisms and make testable predictions for future research, including neurobiological bases of musicality and relationships between human music, language, animal song, and other domains. The music and social bonding hypothesis provides the most comprehensive theory to date of the biological and cultural evolution of music.

Human caregivers interacting with children typically modify their speech in ways that promote attention, bonding, and language acquisition. Although this "motherese," or child- directed communication (CDC), occurs in a variety of human cultures, evidence among nonhuman species is very rare. We looked for its occurrence in a nonhuman mammalian species with long- term mother--offspring bonds that is capable of vocal production learning, the bottlenose dolphin (Tursiops truncatus). Dolphin signature whistles provide a unique opportunity to test for CDC in nonhuman animals, because we are able to quantify changes in the same vocalizations produced in the presence or absence of calves. We analyzed recordings made during brief catch- and- release events of wild bottlenose dolphins in waters near Sarasota Bay, Florida, United States, and found that females produced signature whistles with significantly higher maximum frequencies and wider frequency ranges when they were recorded with their own dependent calves vs. not with them. These differences align with the higher fundamental frequencies and wider pitch ranges seen in human CDC. Our results provide evidence in a nonhuman mammal for changes in the same vocalizations when produced in the presence vs. absence of offspring, and thus strongly support convergent evolution of motherese, or CDC, in bottlenose dolphins. CDC may function to enhance attention, bonding, and vocal learning in dolphin calves, as it does in human children. Our data add to the growing body of evidence that dolphins provide a powerful animal model for studying the evolution of vocal learning and language. [ABSTRACT FROM AUTHOR]

Hummingbirds have been recognized, along with songbirds and parrots, as capable of learning songs. However, it is still unclear whether singing can be treated as a homologous trait within the family, analogous to songbirds. Therefore, we systematically compared the information about hummingbird vocalizations in the literature in a phylogenetic framework. In general, songs were emitted by perching males in a reproductive context, while calls were mainly in agonistic contexts. Singing was ancestral in most of the nine major hummingbird clades, but has been lost at least once in the mountain gem clade and twice in the bee clade. This evolutionary loss of singing might suggest heterogeneity of vocal-production learning. [ABSTRACT FROM AUTHOR]

The interactions between species-specific predispositions and cultural plasticity in the development of human musical behavior have recently become the rationale for a possible Baldwinian origin of human musicality. In the previously suggested Baldwinian scenarios of music origin, social bonding has been indicated as the crucial adaptive value that became the main cause of the co-evolutionary process that led to our musicality. However, the adaptive value of social bonding does not explain the cultural variability of musical expressions that enabled the Baldwinian evolution of musicality. The main aim of this article is to show that free rider recognition, along with social bonding and signaling commitment, could have been a possible adaptive function of hominin musical rituals. In the proposed scenario, free rider recognition became a "flywheel" of the arms race between deception and cooperation. As a result, the interplay between the canalization and plasticity of musical learning became a part of music evolution. This process created a cultural niche in which hominin vocal learning was specialized in the imitation of discrete pitch and rhythm. [ABSTRACT FROM AUTHOR]

In some species, the ability to acquire new vocalizations persists into adulthood and may be an important mediator of social interactions. While it is generally assumed that vocal learning persists undiminished throughout the lifespan of these open-ended learners, the stability of this trait remains largely unexplored. We hypothesize that vocal learning exhibits senescence, as is typical of complex cognitive traits, and that this decline relates to age-dependent changes in social behaviour. The budgerigar (Melopsittacus undulatus), an open-ended learner that develops new contact call types that are shared with social associates upon joining new flocks, provides a robust assay for measuring the effects of ageing on vocal learning ability. We formed captive flocks of 4 previously unfamiliar adult males of the same age class, either 'young adults' (6 mo−1 y) or 'older adults' (≥ 3 y), and concurrently tracked changes in contact call structure and social interactions over time. Older adults exhibited decreased vocal diversity, which may be related to sparser and weaker affiliative bonds observed in older adults. Older adults, however, displayed equivalent levels of vocal plasticity and vocal convergence compared to young adults, suggesting that many components of vocal learning are largely maintained into later adulthood in an open-ended learner. [ABSTRACT FROM AUTHOR]

Multimodal imitation of actions, gestures and vocal production is a hallmark of the evolution of human communication, as both, vocal learning and visual-gestural imitation, were crucial factors that facilitated the evolution of speech and singing. Comparative evidence has revealed that humans are an odd case in this respect, as the case for multimodal imitation is barely documented in nonhuman animals. While there is evidence of vocal learning in birds and in mammals like bats, elephants and marine mammals, evidence in both domains, vocal and gestural, exists for two Psittacine birds (budgerigars and grey parrots) and cetaceans only. Moreover, it draws attention to the apparent absence of vocal imitation (with just a few cases reported for vocal fold control in an orangutan and a gorilla and a prolonged development of vocal plasticity in marmosets) and even for imitation of intransitive actions (not object related) in monkeys and apes in the wild. Even after training, the evidence for productive or "true imitation" (copy of a novel behavior, i.e., not pre-existent in the observer's behavioral repertoire) in both domains is scarce. Here we review the evidence of multimodal imitation in cetaceans, one of the few living mammalian species that have been reported to display multimodal imitative learning besides humans, and their role in sociality, communication and group cultures. We propose that cetacean multimodal imitation was acquired in parallel with the evolution and development of behavioral synchrony and multimodal organization of sensorimotor information, supporting volitional motor control of their vocal system and audio-echoic-visual voices, body posture and movement integration. [ABSTRACT FROM AUTHOR]

The present review presents a new and feasible assumption in the comparative neurobiology of vocal production learning that underpins human speech. Vocal learning, the competence of modulating acoustic sounds or imitating novel sounds, is a requisite requirement of human spoken language and is considered a major innovation for the evolutionary origin of human verbal communication. Having numerous examples of impaired language in many neurodegenerative diseases caused by brain injury, until now our knowledge about the underlying neural procedure for verbal communication is inadequate. This capacity is restricted in a few distantly related groups of mammals and birds like humans, elephants, cetaceans, pinnipeds, bats, a few primates, and three genera of birds—parrots, songbirds, and hummingbirds. Astonishingly atypical neuronal type-spindle-shaped bipolar projection neurons-the von Economo neurons (VENs) and fork neurons are also detected in V layer of the anterior cingulate cortex (ACC) and frontoinsular cortex (FI) of identical groups of mammals and even in some telencephalic nuclei of bird species (parrots) that do not have a neocortex. Review evaluations speculate that capability of vocal imitation and learning is the shared feature of all animals containing VENs and probably fork neurons. The communication systems and their neural architecture in nonhuman animals have been reviewed and their significance has been discussed. The present review is intended to develop a new speculation regarding the neural mechanisms of vocalizition in nonhuman animals and human language. [ABSTRACT FROM AUTHOR]

Behaviors emerge via a combination of experience and innate predispositions. As the brain matures, it undergoes major changes in cellular, network, and functional properties that can be due to sensory experience as well as developmental processes. In normal birdsong learning, neural sequences emerge to control song syllables learned from a tutor. Here, we disambiguate the role of tutor experience and development in neural sequence formation by delaying exposure to a tutor. Using functional calcium imaging, we observe neural sequences in the absence of tutoring, demonstrating that tutor experience is not necessary for the formation of sequences. However, after exposure to a tutor, pre-existing sequences can become tightly associated with new song syllables. Since we delayed tutoring, only half our birds learned new syllables following tutor exposure. The birds that failed to learn were the birds in which pre-tutoring neural sequences were most ‘crystallized,’ that is, already tightly associated with their (untutored) song.

Despite much research on mimicry, little is known about the ecology of dynamic mimetic signals involving mimicry of multiple species. Some of the most conspicuous examples of phenotypically plastic mimicry are produced by oscine passerines, where vocal production learning enables some species to mimic multiple models and flexibly adjust what they mimic and when. While singing from a perch, male superb lyrebirds (Menura novaehollandiae) accurately imitate multiple songs and calls of over 20 species of bird. However, at key moments within their multimodal displays performed on display arenas on the forest floor, males mimic a small number of mobbing-alarm calls creating the acoustic illusion of a mixed-species mobbing flock ('D-song'). Using observations from camera footage and a field-based playback experiment, we tested six hypotheses for alarm call model selection within D-song. Mimicked species were remarkably invariant, with 79% of D-song made up of imitations of just three different bird species. Males did not mimic the most common species in their general environment, but neither did they mimic rare species. Instead, males imitated the mobbing-alarm calls of heterospecific birds that foraged on or near the forest floor. Indeed, males primarily mimicked the alarm calls of heterospecific species that foraged alongside lyrebirds and were likely to appear together in experimentally-induced, terrestrial mobbing flocks. These findings support the hypothesis that males mimic a cue of a terrestrial predatory threat to lyrebirds, most likely to exploit the antipredator behaviour of female lyrebirds. Our study illustrates the importance of investigating the drivers of model selection in dynamic multi-model mimicry. [ABSTRACT FROM AUTHOR]

Infants' prelinguistic vocalizations reliably organize vocal turn‐taking with social partners, creating opportunities for learning to produce the sound patterns of the ambient language. This social feedback loop supporting early vocal learning is well‐documented, but its developmental origins have yet to be addressed. When do infants learn that their non‐cry vocalizations influence others? To test developmental changes in infant vocal learning, we assessed the vocalizations of 2‐ and 5‐month‐old infants in a still‐face interaction with an unfamiliar adult. During the still‐face, infants who have learned the social efficacy of vocalizing increase their babbling rate. In addition, to assess the expectations for social responsiveness that infants build from their everyday experience, we recorded caregiver responsiveness to their infants' vocalizations during unstructured play. During the still‐face, only 5‐month‐old infants showed an increase in vocalizing (a vocal extinction burst) indicating that they had learned to expect adult responses to their vocalizations. Caregiver responsiveness predicted the magnitude of the vocal extinction burst for 5‐month‐olds. Because 5‐month‐olds show a vocal extinction burst with unfamiliar adults, they must have generalized the social efficacy of their vocalizations beyond their familiar caregiver. Caregiver responsiveness to infant vocalizations during unstructured play was similar for 2‐ and 5‐month‐olds. Infants thus learn the social efficacy of their vocalizations between 2 and 5 months of age. During this time, infants build associations between their own non‐cry sounds and the reactions of adults, which allows learning of the instrumental value of vocalizing. [ABSTRACT FROM AUTHOR]

Comparative analysis is a fundamental tool in biology. Comparative biolinguistics emphasizes that the foundations of cognition are shared between a wide range of species and provides insights into the nature of human speech, language, and social interaction by comparing the communication mechanisms of humans, animals, and hominins. This paper reviewed comparative biolinguistic studies in the modern population (between modern humans and hominins, and in model animals of vocal learning) to reveal the significance of comparative biology in the study of developmental language disorders and language evolution at the levels of genome, neurogenetics, and ethology.

Multimodal imitation of actions, gestures and vocal production is a hallmark of the evolution of human communication, as both, vocal learning and visual-gestural imitation, were crucial factors that facilitated the evolution of speech and singing. Comparative evidence has revealed that humans are an odd case in this respect, as the case for multimodal imitation is barely documented in non-human animals. While there is evidence of vocal learning in birds and in mammals like bats, elephants and marine mammals, evidence in both domains, vocal and gestural, exists for two Psittacine birds (budgerigars and grey parrots) and cetaceans only. Moreover, it draws attention to the apparent absence of vocal imitation (with just a few cases reported for vocal fold control in an orangutan and a gorilla and a prolonged development of vocal plasticity in marmosets) and even for imitation of intransitive actions (not object related) in monkeys and apes in the wild. Even after training, the evidence for productive or “true imitation” (copy of a novel behavior, i.e., not pre-existent in the observer’s behavioral repertoire) in both domains is scarce. Here we review the evidence of multimodal imitation in cetaceans, one of the few living mammalian species that have been reported to display multimodal imitative learning besides humans, and their role in sociality, communication and group cultures. We propose that cetacean multimodal imitation was acquired in parallel with the evolution and development of behavioral synchrony and multimodal organization of sensorimotor information, supporting volitional motor control of their vocal system and audio-echoic-visual voices, body posture and movement integration.

Vocal learning is a complex acquired social behavior that has been found only in very few animals. The process of animal vocal learning requires the participation of sensorimotor function. By accepting external auditory input and cooperating with repeated vocal imitation practice, a stable pattern of vocal information output is eventually formed. In parallel evolutionary branches, humans and songbirds share striking similarities in vocal learning behavior. For example, their vocal learning processes involve auditory feedback, complex syntactic structures, and sensitive periods. At the same time, they have evolved the hierarchical structure of special forebrain regions related to vocal motor control and vocal learning, which are organized and closely associated to the auditory cortex. By comparing the location, function, genome, and transcriptome of vocal learning-related brain regions, it was confirmed that songbird singing and human language-related neural control pathways have certain analogy. These common characteristics make songbirds an ideal animal model for studying the neural mechanisms of vocal learning behavior. The neural process of human language learning may be explained through similar neural mechanisms, and it can provide important insights for the treatment of language disorders.

The basal ganglia (BG) participate in aspects of reinforcement learning that require evaluation and selection of motor programs associated with improved performance. However, whether the BG additionally contribute to behavioral variation ("motor exploration") that forms the substrate for such learning remains unclear. In songbirds, a tractable system for studying BG-dependent skill learning, a role for the BG in generating exploratory variability, has been challenged by the finding that lesions of Area X, the song-specific component of the BG, have no lasting effects on several forms of vocal variability that have been studied. Here we demonstrate that lesions of Area X in adult male zebra finches (Taeniopygia gutatta) permanently eliminate rapid within-syllable variation in fundamental frequency (FF), which can act as motor exploration to enable reinforcement-driven song learning. In addition, we found that this within-syllable variation is elevated in juveniles and in adults singing alone, conditions that have been linked to enhanced song plasticity and elevated neural variability in Area X. Consistent with a model that variability is relayed from Area X, via its cortical target, the lateral magnocellular nucleus of the anterior nidopallium (LMAN), to influence song motor circuitry, we found that lesions of LMAN also eliminate within-syllable variability. Moreover, we found that electrical perturbation of LMAN can drive fluctuations in FF that mimic naturally occurring within-syllable variability. Together, these results demonstrate that the BG are a central source of rapid behavioral variation that can serve as motor exploration for vocal learning.SIGNIFICANCE STATEMENT Many complex motor skills, such as speech, are not innately programmed but are learned gradually through trial and error. Learning involves generating exploratory variability in action ("motor exploration") and evaluating subsequent performance to acquire motor programs that lead to improved performance. Although it is well established that the basal ganglia (BG) process signals relating to action evaluation and selection, whether and how the BG promote exploratory motor variability remain unclear. We investigated this question in songbirds, which learn to produce complex vocalizations through trial and error. In contrast with previous studies that did not find effects of BG lesions on vocal motor variability, we demonstrate that the BG are an essential source of rapid behavioral variation linked to vocal learning.

Vocal communication is the main mean of communication for cetaceans and some species developed vocal culture and traditions. They are maintained through both production and contextual learning, used to acquire two types of sounds: signature calls and songs. Signature calls are present only in species living in stable groups or fission-fusion society. They are used as an identification tool to maintain cohesion and contact with conspecifics. Songs are present in most baleen whales, but only humpback and bowhead whales change within and between seasons. They use vocal learning to conform to one type used by all individuals. Vocal learning is also employed during vocal development in ontogenesis, together with maturation. The existence of social groups in other species together with the lack of research suggest that traditions are more widespread among cetaceans than is currently known. [ABSTRACT FROM AUTHOR]