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 archaeology, 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 due to 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 (MSB) hypothesis provides the most comprehensive theory to date of the biological and cultural evolution of music.

Communicating with one's environment requires efficient neural interaction between action and perception. Neural substrates of sound perception and production are connected by the arcuate fasciculus (AF). Although AF is known to be involved in language, its roles in non-linguistic functions are unexplored. Here, we show that tone-deaf people, with impaired sound perception and production, have reduced AF connectivity. Diffusion tensor tractography and psychophysics were assessed in tone-deaf individuals and matched controls. Abnormally reduced AF connectivity was observed in the tone deaf. Furthermore, we observed relationships between AF and auditorymotor behavior: superior and inferior AF branches predict psychophysically assessed pitch discrimination and sound production perception abilities, respectively. This neural abnormality suggests that tone deafness leads to a reduction in connectivity resulting in pitch-related impairments. Results support a dual-stream anatomy of sound production and perception implicated in vocal communications. By identifying white matter differences and their psychophysical correlates, results contribute to our understanding of how neural connectivity subserves behavior.

Knowledge of musical rules and structures has been reliably demonstrated in humans of different ages, cultures, and levels of music training, and has been linked to our musical preferences. However, how humans acquire knowledge of and develop preferences for music remains unknown. The present study shows that humans rapidly develop knowledge and preferences when given limited exposure to a new musical system. Using a non-traditional, unfamiliar musical scale (Bohlen-Pierce scale), we created finite-state musical grammars from which we composed sets of melodies. After 25-30 min of passive exposure to the melodies, participants showed extensive learning as characterized by recognition, generalization, and sensitivity to the event frequencies in their given grammar, as well as increased preference for repeated melodies in the new musical system. Results provide evidence that a domain-general statistical learning mechanism may account for much of the human appreciation for music.

Structure and function of the human brain are affected by training in both linguistic and musical domains. Individuals with intensive vocal musical training provide a useful model for investigating neural adaptations of learning in the vocal–motor domain and can be compared with learning in a more general musical domain. Here we confirm general differences in macrostructure (tract volume) and microstructure (fractional anisotropy, FA) of the arcuate fasciculus (AF), a prominent white-matter tract connecting temporal and frontal brain regions, between singers, instrumentalists, and non-musicians. Both groups of musicians differed from non-musicians in having larger tract volume and higher FA values of the right and left AF. The AF was then subdivided in a dorsal (superior) branch connecting the superior temporal gyrus and the inferior frontal gyrus (STG ↔ IFG), and ventral (inferior) branch connecting the middle temporal gyrus and the inferior frontal gyrus (MTG ↔ IFG). Relative to instrumental musicians, singers had a larger tract volume but lower FA values in the left dorsal AF (STG ↔ IFG), and a similar trend in the left ventral AF (MTG ↔ IFG). This between-group comparison controls for the general effects of musical training, although FA was still higher in singers compared to non-musicians. Both musician groups had higher tract volumes in the right dorsal and ventral tracts compared to non-musicians, but did not show a significant difference between each other. Furthermore, in the singers’ group, FA in the left dorsal branch of the AF was inversely correlated with the number of years of participants’ vocal training. Our findings suggest that long-term vocal–motor training might lead to an increase in volume and microstructural complexity of specific white-matter tracts connecting regions that are fundamental to sound perception, production, and its feedforward and feedback control which can be differentiated from a more general musician effect.

The source of conscious experience has fueled scientific and philosophical debates for centuries. In auditory and motor domains, it is yet unknown how consciously and unconsciously obtained information may combine to enable the production and perception of speaking and singing. Both forms of vocalizations rely upon the interaction of brain networks responsible for perception and action. While perceptual experience and executed actions are usually well coupled, dissociations between perception and action can be informative of their underlying neural systems. Here we report a dissociation between production and perception: tone-deaf individuals, who cannot consciously perceive pitch differences, can paradoxically reproduce pitch intervals in correct directions. Our results suggest that multiple neural pathways have evolved for sound perception and production, so that pitch information sufficient for intact speech can be obtained separately from pathways necessary for conscious perception.

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Connectivity in the human brain has received increased scientific interest in recent years. Although connection disorders can affect perception, production, learning, and memory, few studies have associated brain connectivity with graded variations in human behavior, especially among normal individuals. One group of normal individuals who possess unique characteristics in both behavior and brain structure is absolute pitch (AP) musicians, who can name the appropriate pitch class of any given tone without a reference. Using diffusion tensor imaging and tractography, we observed hyperconnectivity in bilateral superior temporal lobe structures linked to AP possession. Furthermore, volume of tracts connecting left superior temporal gyrus to left middle temporal gyrus predicted AP performance. These findings extend previous reports of exaggerated temporal lobe asymmetry, may explain the higher incidence of AP in developmental disorders, and may provide a model for understanding the heightened connectivity that is thought to underlie savant skills and cases of exceptional creativity.

Humans uniquely appreciate aesthetics, experiencing pleasurable responses to complex stimuli that confer no clear intrinsic value for survival. However, substantial variability exists in the frequency and specificity of aesthetic responses. While pleasure from aesthetics is attributed to the neural circuitry for reward, what accounts for individual differences in aesthetic reward sensitivity remains unclear. Using a combination of survey data, behavioral and psychophysiological measures and diffusion tensor imaging, we found that white matter connectivity between sensory processing areas in the superior temporal gyrus and emotional and social processing areas in the insula and medial prefrontal cortex explains individual differences in reward sensitivity to music. Our findings provide the first evidence for a neural basis of individual differences in sensory access to the reward system, and suggest that social-emotional communication through the auditory channel may offer an evolutionary basis for music making as an aesthetically rewarding function in humans.