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A Brief Overview
DANIEL J. LEVITIN McGill University, Montreal, QC, Canada
ABSTRACT: Musical behaviors activate nearly every region of the music, and (3) effects of music on immune function and
brain that has so far been mapped. These neural structures are health.
reviewed in a domain-specific overview to include the stimulus-
response matrix for sensory activation, motor action planning, and Perceptual-Cognitive Processing of Music
cognitive and emotional reactions to music. Also reviewed are the Music consists of six fundamental attributes: pitch, dura-
effects of learning an instrument, similarities and differences in
tion, loudness, timbre, spatial location, and reverberant
speech versus musical communication, and the potential health
benefits of music. This review provides a framework and foundation environment (Levitin, 2010). A set of two or more pitches
for understanding the effects of music therapy and other interventions gives rise to contour and melody, and a set of two or more
nen, 2003; Warren Uppenkamp, Patterson, & Griffiths, similar, involving the forcing of air through the vocal folds and
2003).1 When two or more pitches combine to form intervals, motor movement of the vocal folds, tongue, and lips (Cook,
contour and melody activation is seen in the superior temporal 1999). Thus, music making and movement are intertwined.
gyrus (STG) and planum polare (Patterson et al., 2002). Listening to music may activate mirror neurons in the motor
The neural mechanisms for perceiving rhythm are intimate- cortex (Rizzolatti, Fadiga, Gallese, & Fogassi, 1996) that cause
ly bound with those for producing it, even in the absence of us to think (at least unconsciously) about those motor
physical movement (Grahn & Rowe, 2009). Sensorimotor movements that would be required to make the music.
coordination is facilitated by music (Altenmüller, Wiesen- Movement learning may be subserved by these mirror
danger, & Kesselring, 2006; Large, 2000), particularly music neurons, and eventually this activation could lead to new
with a strong beat (Grahn & Rowe, 2009) and tempo has been learned behaviors.
shown to mediate involvement of motor areas in beat When we move our bodies to music, there is neural
perception, specifically the premotor cortex, inferior frontal activation in the premotor cortex, motor cortex, the cerebel-
gyrus, superior temporal gyrus, and inferior parietal lobule lum and basal ganglia (Levitin, 2007; Mesulam, 1985) and
(McAuley, Henry, & Tkach, 2012). voluntary movements activate the precuneus as well, a region
Musical activities activate the sensory and motor cortices. of the parietal lobe (Brown & Parsons, 2008). Tapping in
Playing an instrument activates muscle regions of the synchrony to the pulse of a musical sequence (by humans)
(march) or triple (waltz) meter. The meter biased the violinists show increased responses to violin music, flutists to
perceptual representation and subsequent recall of the flute music, and so on (Margulis, Mlsna, Uppunda, Parrish, &
sequences. In effect, the movement itself created the (cross- Wong, 2009). Most North Americans are raised with exposure
modal) accented beat. This interactive process was found to be to a single musical system or culture (monomusical), and this
mediated by the vestibular system: although full body parallels development in a home where one language is
movement is the most effective in engendering the move- spoken (monolingual). Studies with bimusicals, individuals
ment-sound interaction, head movement alone is capable of who are exposed to different musical systems or cultures (such
producing it, while body movement alone is not (Phillips- as Indian and Western music), have shown that they use
Silver & Trainor, 2008). different parts of their brains to process music (Wong, Chan,
Roy, & Margulis, 2011). Monomusicals differentiated Indian
Effects of Practice and Training from Western music in auditory cortices, suggesting that the
Early instrumental musical training appears to train atten- primary distinction they made was sensory or perceptual.
tional networks in the brain as well as to train social and Bimusicals differentiated these same musics in limbic
interpersonal skills. Musically-trained children show improve- structures, suggesting the distinction was more they made
ments in nonverbal IQ, numeracy and spatial cognition was primarily on an emotional or affective basis.
(Neville et al., 2008). Children given music lessons showed Effective connectivity analysis, a mathematical technique to
suppressed in order to improvise well. Activation was rely on the hearing sense, and so there naturally exist
observed in neocortical sensory-motor areas that mediate overlapping circuits at the level of the ear, the cochlea, and
organization and execution of musical performance. This the auditory nerve. For music with lyrics, the lyrics are
pattern of activation and deactivation conforms to subjective generally processed in the same regions of the brain that
reports by musicians that improvisation relies on sub or process meaning in spoken or written language, semantic
preconscious processes that are outside the domain of association networks, and the internal mental lexicon that
conscious control and awareness. The lack of conscious include regions in the temporal lobes and hippocampus. The
control represents overlearned and automatic processes syntax of language is largely subserved by regions in the left
characteristic of experts in other fields of endeavor. frontal lobes, including Brodmann Areas 44, 45, and 47.
Thus, musical components are processed in distinctive and While it is not clear whether a true syntax of music exists
separable neural circuits. Musical activities engage a widely analogously to that of language, musical expectations and
dispersed network of brain regions, not just so-called auditory violations of those expectations are known to be processed in
regions. Musical training is typically associated with increases Brodmann Areas 44, 45, and 47 (Koelsch, 2005; Koelsch,
in gray matter volume and density, and helps to stave off age- Gunter, von Cramon, et al., 2002; Levitin & Menon, 2003).
related neural atrophy. Musical training is also typically When presented with a chord sequence that resolves in an
associated with a left-ward shift in processing, reflecting the unexpected fashion, an electrical signal (the evoked response
Perceptual-cognitive processing of music can lead to Many listeners report using music for mood regulation, and
emotional reactions, especially when lyrics are involved. may find comfort in sad music (Chamorro-Premuzic &
Researchers distinguish between the experience of musical Furnham, 2007). One might think that sad people would find
emotions and the mere recognition of them (Huron, 2011). happy music an antidote to their mood, but this is not always
That is, a listener can identify that a happy mood is intended in the case. Prolactin, a tranquilizing and consoling hormone, is
a piece of music without actually experiencing that happy produced by the anterior pituitary gland when we are sad
mood. What are the processes involved when we do (Huron, 2006; Panksepp, 2006). The evolutionary purpose of
experience music-evoked emotions? sorrow, Huron (2006) explains, is to aid in energy conserva-
tion and allow for reassessment of priorities for the future
Emotional Reactions to Music
following a traumatic event. Prolactin is released after orgasm,
Neural pathways carry the musical signal from the ear to the after birth, and during lactation in females.
inferior colliculi in the midbrain, and then to emotional A chemical analysis reveals that prolactin is not always
centers in the cerebellum, nucleus accumbens (NAc), ventral present in tears—it is not released in tears of lubrication of the
tegmental area (VTA), and amygdala; these regions are eye, or when the eye is irritated, or in tears of joy; it is only
activated when people listen to pleasurable music, and released in tears of sorrow. Huron speculates that sad music
dopamine levels are modulated as well (Menon & Levitin, allows us to ‘‘trick’’ our brains into releasing prolactin in
In contemporary society, music continues to be used to Kumar, & Kumar, 1997) and during stressful tasks (Khalfa,
promote health and well-being in clinical settings, such as for Dalla Bella, Roy, Peretz, & Lupien, 2003; Knight & Rickard,
pain management, relaxation, psychotherapy, and personal 2001). These results suggest that music can function effec-
growth (see Chanda & Levitin, 2013, and Kreutz, Quiroga tively to reduce stress across a variety of situations in the
Murcia & Bongard, 2012 for reviews). Although music therapy home, workplace, and in clinical settings.
interventions are based strongly on clinical-practice knowl- The mechanism for stress reduction likely involves a
edge, concepts from evidence-based practice and experiments reduction in central nervous system arousal levels. Because
are providing additional information about the specificity of some neuronal populations fire in synchrony with the tempo
these interventions. Indeed, the American Music Therapy of music, slow tempo music could lead to reductions in heart
Association (2011) defines its practices as relying on the and respiration rates. Music initiates reflexive brain stem
‘‘evidence-based use of music.’’ responses, mediating heart rate, pulse, blood pressure, body
For athletes, stamina, motivation (Becker et al., 1994; temperature, skin conductance, and muscle tension (Juslin &
Simpson & Karageorghis, 2006), and anaerobic power Västjäll, 2008; Lundqvist et al., 2009).
(Eliakim, Meckel, Nemet, & Eliakim, 2007) can be improved Music listening and performance appear to improve
by listening to music during sports activities and workouts. immune system function (Chanda & Levitin, 2013; Gangrade,
Indeed, the Ethiopian runner Haile Gebreselassie (who won 2012; Lane, 1994; Wachi et al., 2007). Specifically, increases
area, and the modulation of the neurochemical dopamine. Bidelman, G. M., Gandour, J. T., & Krishnan, A. (2011). Cross-domain effects of
Music has been shown to improve immune system function music and language experience on the representation of pitch in the human
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