A neural circuit for vocal production responds to viscerosensory input in the songbird.

Motor performance is monitored continuously by specialized brain circuits and used adaptively to modify behavior on a moment-to-moment basis and over longer time periods. During vocal behaviors, such as singing in songbirds, internal evaluation of motor performance relies on sensory input from the auditory and vocal-respiratory systems. Sensory input from the auditory system to the motor system, often referred to as auditory feedback, has been well studied in singing zebra finches (), but little is known about how and where nonauditory sensory feedback is evaluated.

Female signal jamming in a socially monogamous brood parasite.

Acoustic signalling is vital to courtship in many animals, yet the role of female vocalizations is understudied. Here, we combine observational and experimental methods to assess the courtship function of the female chatter call in brown-headed cowbirds, . While the chatter call is likely multifunctional, it is frequently used in social interactions and overlapping duets with males during the breeding season.

Regulation of vocal precision by noradrenergic modulation of a motor nucleus.

Recent theories of norepinephrine (NE) function suggest that NE modulates the transition between stereotyped, goal-directed behavior and more variable, exploratory behaviors that facilitate learning and adaptation. We provide evidence for context-dependent switching by NE that is analogous to this explore/exploit strategy in the vocal system of the zebra finch (). Stimulation of the locus coeruleus, the major source of NE in the brain, decreases song trial-to-trial variability, transforming the variable, exploratory "undirected" song into song that resembles the more stereotyped, exploitative "directed" song that males sing to females.

3D Bird Reconstruction: a Dataset, Model, and Shape Recovery from a Single View.

Automated capture of animal pose is transforming how we study neuroscience and social behavior. Movements carry important social cues, but current methods are not able to robustly estimate pose and shape of animals, particularly for social animals such as birds, which are often occluded by each other and objects in the environment. To address this problem, we first introduce a model and multi-view optimization approach, which we use to capture the unique shape and pose space displayed by live birds.

Breathtaking Songs: Coordinating the Neural Circuits for Breathing and Singing.

The vocal behavior of birds is remarkable for its diversity, and songs can feature elaborate characteristics such as long duration, rapid temporal pattern, and broad frequency range. The respiratory system plays a central role in generating the complex song patterns that must be integrated with its life-sustaining functions. Here, we explore how precise coordination between the neural circuits for breathing and singing is fundamental to production of these remarkable behaviors.

Testing the evolutionary conservation of vocal motoneurons in vertebrates.

Medullary motoneurons drive vocalization in many vertebrate lineages including fish, amphibians, birds, and mammals. The developmental history of vocal motoneuron populations in each of these lineages remains largely unknown. The highly conserved transcription factor Paired-like Homeobox 2b (Phox2b) is presumed to be expressed in all vertebrate hindbrain branchial motoneurons, including laryngeal motoneurons essential for vocalization in humans.

The respiratory-vocal system of songbirds: anatomy, physiology, and neural control.

This wide-ranging review presents an overview of the respiratory-vocal system in songbirds, which are the only other vertebrate group known to display a degree of respiratory control during song rivalling that of humans during speech; this despite the fact that the peripheral components of both the respiratory and vocal systems differ substantially in the two groups. We first provide a brief description of these peripheral components in songbirds (lungs, air sacs and respiratory muscles, vocal organ (syrinx), upper vocal tract) and then proceed to a review of the organization of central respiratory-related neurons in the spinal cord and brainstem, the latter having an organization fundamentally similar to that of the ventral respiratory group of mammals. The second half of the review describes the nature of the motor commands generated in a specialized "cortical" song control circuit and how these might engage brainstem respiratory networks to shape the temporal structure of song.

Achieving perfection through variability: the basal ganglia helped me do it!

Motor variability can facilitate motor exploration necessary for learning. In this issue of Neuron, Woolley et al. (2014) record at different stages of the songbird basal ganglia and show that social-context modulation of motor variability first emerges in the pallidum.

Social brains in context: lesions targeted to the song control system in female cowbirds affect their social network.

Social experiences can organize physiological, neural, and reproductive function, but there are few experimental preparations that allow one to study the effect individuals have in structuring their social environment. We examined the connections between mechanisms underlying individual behavior and social dynamics in flocks of brown-headed cowbirds (Molothrus ater). We conducted targeted inactivations of the neural song control system in female subjects.

Characterization of respiratory neurons in the rostral ventrolateral medulla, an area critical for vocal production in songbirds.

Much is known about the neuronal cell types and circuitry of the mammalian respiratory brainstem and its role in normal, quiet breathing. Our understanding of the role of respiration in the context of vocal production, however, is very limited. Songbirds contain a well-defined neural circuit, known as the song system, which is necessary for song production and is strongly coupled to the respiratory system.

Linear and nonlinear auditory response properties of interneurons in a high-order avian vocal motor nucleus during wakefulness.

Motor-related forebrain areas in higher vertebrates also show responses to passively presented sensory stimuli. However, sensory tuning properties in these areas, especially during wakefulness, and their relation to perception, are poorly understood. In the avian song system, HVC (proper name) is a vocal-motor structure with auditory responses well defined under anesthesia but poorly characterized during wakefulness.

Breathing and vocal control: the respiratory system as both a driver and a target of telencephalic vocal motor circuits in songbirds.

The production of vocalizations is intimately linked to the respiratory system. Despite our understanding of neural circuits that generate normal respiratory patterns, very little is understood regarding how these pontomedullary circuits become engaged during vocal production. Songbirds offer a potentially powerful model system for addressing this relationship.

An IACUC perspective on songbirds and their use in neurobiological research.

Laboratory research using songbirds as a model system for investigating basic questions of neurobiological function has expanded rapidly and recently, with approximately 120 laboratories working with songbirds worldwide. In the United States alone, of the approximately 80 such laboratories nearly a third have been established in the past 10 years. Yet many animal facilities are not outfitted to manage these animals, and as a consequence laboratories often use alternative housing arrangements established by institutional animal care and use committees (IACUCs).

Sleep, off-line processing, and vocal learning.

The study of song learning and the neural song system has provided an important comparative model system for the study of speech and language acquisition. We describe some recent advances in the bird song system, focusing on the role of off-line processing including sleep in processing sensory information and in guiding developmental song learning. These observations motivate a new model of the organization and role of the sensory memories in vocal learning.

What birdsong can teach us about the central noradrenergic system.

Increasing evidence indicates that the noradrenergic system plays a key role in biasing the nervous system towards producing behaviors that help animals adapt to constantly changing environments. Most of the studies investigating noradrenergic function are performed in animals that have a limited repertoire of tractable natural behaviors. Songbirds, in contrast, with their rich set of precisely quantifiable vocal behaviors, provide a unique model system to study the noradrenergic system.

Using both sides of your brain: the case for rapid interhemispheric switching.

Individual brain hemispheres are often specialized for specific aspects of a behavior. How both sides of the brain coordinate their output to produce a perfectly seamless behavior is not known. Songbirds appear to achieve this by rapidly switching back and forth between hemispheres.

Bottom-up activation of the vocal motor forebrain by the respiratory brainstem.

Brainstem motor structures send output commands to the periphery and are dynamically modulated by telencephalic inputs. Little is known, however, about ascending brainstem control of forebrain motor structures. Here, we provide the first evidence for bottom-up activation of forebrain motor centers by the respiratory brainstem.

Hemispheric coordination is necessary for song production in adult birds: implications for a dual role for forebrain nuclei in vocal motor control.

Precise coordination across hemispheres is a critical feature of many complex motor circuits. In the avian song system the robust nucleus of the arcopallium (RA) plays a key role in such coordination. It is simultaneously the major output structure for the descending vocal motor pathway, and it also sends inputs to structures in the brain stem and thalamus that project bilaterally back to the forebrain.

Distributed and selective auditory representation of song repertoires in the avian song system.

For many songbirds, the vocal repertoire constitutes acoustically distinct songs that are flexibly used in various behavioral contexts. To investigate how these different vocalizations are represented in the song neural system, we presented multiple song stimuli while performing extracellular recording in nucleus HVC in adult male song sparrows Melospiza melodia, a species known for its complex vocal repertoire and territorial use of song. We observed robust auditory responses to natural song stimuli in both awake and anesthetized animals.

Brainstem and forebrain contributions to the generation of learned motor behaviors for song.

Brainstem nuclei have well established roles in generating nonlearned rhythmic behaviors or as output pathways for more complex, forebrain-generated behaviors. However, the role of the brainstem in providing information to the forebrain that is used to initiate or assist in the control of complex behaviors is poorly understood. In this study, we used electrical microstimulation in select nuclei of the avian song system combined with recordings of acoustic and respiratory output to examine how forebrain and brainstem nuclei interact in the generation of learned vocal motor sequences.

Sensorimotor nucleus NIf is necessary for auditory processing but not vocal motor output in the avian song system.

Sensorimotor integration in the avian song system is crucial for both learning and maintenance of song, a vocal motor behavior. Although a number of song system areas demonstrate both sensory and motor characteristics, their exact roles in auditory and premotor processing are unclear. In particular, it is unknown whether input from the forebrain nucleus interface of the nidopallium (NIf), which exhibits both sensory and premotor activity, is necessary for both auditory and premotor processing in its target, HVC.

Noradrenergic inputs mediate state dependence of auditory responses in the avian song system.

Norepinephrine (NE) plays a complex role in the behavioral state-dependent regulation of sensory processing. However, the role of forebrain NE action in modulating high-order sensory activity has not been directly addressed. In this study, we take advantage of the discrete, feedforward organization of the avian song system to identify a site and mechanism of NE action underlying state-dependent modulation of sensory processing.

Bilateral control and interhemispheric coordination in the avian song motor system.

Birdsong is a complex learned motor behavior controlled by an interconnected network of vocal control nuclei that are present in both cerebral hemispheres. Unilateral lesions of song nuclei in the left or the right hemisphere result in different effects on song structure, suggesting that normal song output results from the activation of two parallel but functionally different motor pathways. Because each syringeal half is innervated primarily by ipsilateral motor structures and activity in both halves is tightly coordinated during singing, motor commands originating from both hemispheres must be tightly coordinated to produce the appropriate vocal output.