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The neuroscience of music and music-based interventions (MBIs) is a fascinating but challenging research field. While music is a ubiquitous component of every human society, MBIs may encompass listening to music, performing music, music-based movement, undergoing music education and training, or receiving treatment from music therapists. Unraveling the brain circuits activated and influenced by MBIs may help us gain better understanding of the therapeutic and educational values of MBIs by gathering strong research evidence. However, the complexity and variety of MBIs impose unique research challenges. This article reviews the recent endeavor led by the National Institutes of Health to support evidence-based research of MBIs and their impact on health and diseases. It also highlights fundamental challenges and strategies of MBI research with emphases on the utilization of animal models, human brain imaging and stimulation technologies, behavior and motion capturing tools, and computational approaches. It concludes with suggestions of basic requirements when studying MBIs and promising future directions to further strengthen evidence-based research on MBIs in connections with brain circuitry.SIGNIFICANCE STATEMENT Music and music-based interventions (MBI) engage a wide range of brain circuits and hold promising therapeutic potentials for a variety of health conditions. Comparative studies using animal models have helped in uncovering brain circuit activities involved in rhythm perception, while human imaging, brain stimulation, and motion capture technologies have enabled neural circuit analysis underlying the effects of MBIs on motor, affective/reward, and cognitive function. Combining computational analysis, such as prediction method, with mechanistic studies in animal models and humans may unravel the complexity of MBIs and their effects on health and disease.

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.

Neuronal tuning for spectral and temporal features has been studied extensively in the auditory system. In the auditory cortex, diverse combinations of spectral and temporal tuning have been found, but how specific feature tuning contributes to the perception of complex sounds remains unclear. Neurons in the avian auditory cortex are spatially organized in terms of spectral or temporal tuning widths, providing an opportunity for investigating the link between auditory tuning and perception. Here, using naturalistic conspecific vocalizations, we asked whether subregions of the auditory cortex that are tuned for broadband sounds are more important for discriminating tempo than pitch, due to the lower frequency selectivity. We found that bilateral inactivation of the broadband region impairs performance on both tempo and pitch discrimination. Our results do not support the hypothesis that the lateral, more broadband subregion of the songbird auditory cortex contributes more to processing temporal than spectral information. [ABSTRACT FROM AUTHOR]

Author(s): Mimi H. Kao (corresponding author) [1]; Allison J. Doupe [1]; Michael S. Brainard [1] Cortical-basal ganglia circuits have a critical role in motor control and motor learning [1]. In [...]

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 withinsyllable 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. [ABSTRACT FROM AUTHOR]

Song is a learned vocal behavior influenced by social interactions. Prior work has suggested that the anterior forebrain pathway (AFP), a specialized pallial- basal ganglia circuit critical for vocal plasticity, mediates the influence of social signals on song. Here, we investigate the signals the AFP sends to song motor areas and their dependence on social context by characterizing singing-related activity of single neurons in the AFP output nucleus LMAN (lateral magnocellular nucleus of the anterior nidopallium). We show that interaction with females causes marked, real-time changes in firing properties of individual LMAN neurons. When males sing to females ("directed"), LMAN neurons exhibit reliable firing of single spikes precisely locked to song. In contrast, when males sing alone ("undirected"), the same LMAN neurons exhibit prominent burst firing and trial-by-trial variability. Burst structure and timing vary substantially across repeated undirected trials. Despite context-dependent differences in firing statistics, the average pattern of song-locked firing for an individual neuron is similar across behavioral contexts, suggesting a common underlying signal. Different LMAN neurons in the same bird, however, exhibit distinct firing patterns, suggesting that subsets of neurons jointly encode song features. Together, our findings demonstrate that behavioral interactions reversibly transform the signaling mode of LMAN neurons. Such changes may contribute to rapid switching of motor activity between variable and precise states. More generally, our results suggest that pallial- basal ganglia circuits contribute to motor learning and production through multiple mechanisms: patterned signals could guide changes in motor output while state-dependent variability could subserve motor exploration. [ABSTRACT FROM AUTHOR]