Zaatar Et Al., 2024 Musica e Neuroplasticidade

Brain, Behavior, & Immunity - Health 35 (2024) 100716
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Brain, Behavior, & Immunity - Health

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The transformative power of music: Insights into neuroplasticity, health,

and disease
Muriel T. Zaatar *, Kenda Alhakim 1, Mohammad Enayeh 1, Ribal Tamer 1
Department of Biological and Physical Sciences, American University in Dubai, Dubai, United Arab Emirates
A R T I C L E I N F O A B S T R A C T
Keywords: Music is a universal language that can elicit profound emotional and cognitive responses. In this literature re
Neuroplasticity view, we explore the intricate relationship between music and the brain, from how it is decoded by the nervous
Music therapy system to its therapeutic potential in various disorders. Music engages a diverse network of brain regions and
Cognition
circuits, including sensory-motor processing, cognitive, memory, and emotional components. Music-induced
Sensory-motor processing
brain network oscillations occur in specific frequency bands, and listening to one’s preferred music can grant
easier access to these brain functions. Moreover, music training can bring about structural and functional
changes in the brain, and studies have shown its positive effects on social bonding, cognitive abilities, and
language processing. We also discuss how music therapy can be used to retrain impaired brain circuits in
different disorders. Understanding how music affects the brain can open up new avenues for music-based in
terventions in healthcare, education, and wellbeing.
1. Introduction in the brain. Proprioception and auditory feedbacks are crucial while
one learns to play an instrument, and long-term practice results in fewer
Music is an essential part of human life. Many of us have experienced neurons being recruited to execute the same movements (Gaser &
excitement and emotions induced by chords, rhythms, and melodies. Schlaug, 2003; James et al., 2014; Krings et al., 2000). Engaging in the
Long before music had been demonstrated to be a valuable tool for creative process of music production not only enhances musical skills
gaining an insight into various brain functions, the ancient Greeks but also fosters cognitive and motor skill development, contributing to
already knew about the importance of this art. Plato characterized music overall mental well-being.
as a guide toward goodness as it could touch the soul (Schoen-Nazzaro, Benefits of musical experiences begin already during pregnancy.
1978). Engaging with music by either performing it or simply listening to it
The process of deciphering sound involves several brain areas improves the mood and well-being during pregnancy, and strengthens
working in concert to grant the perception of a sound along with the the bond between a mother and her infant (Corbijn van Willenswaard
emotional valence for a specific melody. The music perception occurs et al., 2017; García González et al., 2018; Hepp et al., 2018; Wulff et al.,
through a series of events, starting with sound waves transforming into 2021). Early music experiences can improve the development of
an electric signal, which then progress through the auditory nerve to the cognitive, emotional, physical, and social domains (M. S. Barrett et al.,
brainstem and activates cortical areas, allowing the perception of a 2019; Cassidy et al., 2020; Ding et al., 2019; Fujioka et al., 2006; Sut
specific sound (Koelsch and Siebel, 2005; Moreno and Bidelman, 2014). cliffe et al., 2020; Swaminathan and Schellenberg, 2020). Among older
This intricate process not only contributes to our auditory experience adults, musical experiences contribute to their well-being, and are also
but also plays a crucial role in the potential mental health benefits associated with sustained brain volume and activation of networks
associated with music. involved in executive functions, memory, language processing and
Besides passive music listening, music involves playing and creating, emotions (Bugos et al., 2007; Chaddock-Heyman et al., 2021; Seinfeld
turning it into a unique experience (R. Zatorre, 2005). The acquisition of et al., 2013). These positive effects on well-being highlight the crucial
musical skills through repetitive training causes structural adaptations role of music in promoting mental health across the lifespan.
* Corresponding author. Department of Biological and Physical Sciences, P.O. Box: 28282, Dubai, United Arab Emirates.
E-mail address: [email protected] (M. T. Zaatar).
1
Authors contributed equally to this work (Co-first authors).
https://doi.org/10.1016/j.bbih.2023.100716
Received 7 October 2023; Received in revised form 4 December 2023; Accepted 8 December 2023
Available online 12 December 2023
2666-3546/© 2023 The Authors. Published by Elsevier Inc. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-
nc-nd/4.0/).
M. T. Zaatar et al. Brain, Behavior, & Immunity - Health 35 (2024) 100716
Music benefit individuals both in health and disease. Several studies et al., 2008). Whereas A1 co-activation with motor, pre-motor, insula,
show that many painful conditions and disorders can be alleviated by and cerebellum is related to the processing of emotional content of
music (Düzgün and Karadakovan, 2021; Feneberg et al., 2021; Gauba music (Pando-Naude et al., 2021). Another important functional loop
et al., 2021; Kim and Jeong, 2021; Monsalve-Duarte et al., 2021; C. between A1 and inferior frontal regions (particularly in the right
Wang and Tian, 2021). When a painful stimulus is applied to volunteers hemisphere) allows integration of working memory related to temporal
under control conditions while listening to their favorite songs, they dynamics of the sound. Since auditory events are not static, the brain’s
report lower pain rating scores as music modulates pain responses in ability to concatenate precise auditory information is crucial for the
cortical regions, brainstem and spinal cord (Dobek et al., 2014). This ability to maintain dynamic information for further processing (R. J.
pain modulation highlights one of the proposed mechanisms by which Zatorre and Salimpoor, 2013). Passive engagement with previously
music exerts its effects on mental health, acting as a distraction from unheard music activates subcallosal cingulate gyrus, prefrontal anterior
pain and modulating neural responses in regions associated with pain cingulate and retrosplenial cortices, hippocampus, anterior insula, and
perception. ventral striatum (S. Brown et al., 2004).
In addition, in a spontaneous and effortless way, music can trigger One of the most notable features of music is pitch. Lateral cortical
memories, awake emotions and intensify social bonding (Molnar-Sza regions to A1 are associated with pitch processing (Johnsrude et al.,
kacs and Overy, 2006). Improved attention and communication were 2000; McDermott and Oxenham, 2008). Studies using functional mag
observed in children with severe neurological impairments (Bringas netic resonance imaging (fMRI) scan indicate that pitch processing oc
et al., 2015) and a task-dependent cortical reorganization in stroke pa curs in a hierarchical manner, where more abstract sound properties are
tients occurs after piano and electronic drum pad lessons (Amengual encoded as one proceeds along with the sound analysis streams (R. J.
et al., 2013). Music interventions are also promising therapies for mood, Zatorre et al., 2007). Pitches unfold over time and are perceived as a
vigilance and the overall quality of life in patients with Parkinson’s melody. This dynamic perceptual process engages anterior and posterior
disease (Park and Kim, 2021; Pohl et al., 2020), Alzheimer disease and auditory pathways (Patterson et al., 2002).
other nervous system injuries (Gómez Gallego and Gómez García, 2017; Along with the pitch, music perception relies on the rhythm.
Martínez-Molina et al., 2021; Paprad et al., 2021; Pezzin et al., 2018). Anatomically, pitch and rhythm perception is separable as individuals
The primary goal of this literature review is to provide an insight into with brain injuries in a particular area can still perceive either pitch or
how music is decoded by the nervous system and its impacts on brain melody (depending on the area), but not both (Peretz and Coltheart,
function, as well as how it can unlock various brain states, engage 2003). In neuroimaging studies where subjects are listening only to
different brain circuits, and induce the release of neuromodulators. We rhythms, the cerebellum, the basal ganglia, premotor cortex, and sup
argue that given the fact that music imposes unique demands on the plementary motor area are activated (Grahn and Brett, 2007; Sakai
nervous system, by discussing differences between musicians’ and non- et al., 1999).
musicians’ neuronal network, we can better understand how music is
being utilized to retrain impaired brain circuits via music therapies for 2.2. Music performance: auditory-motor interaction (active involvement)
different disorders.
In contrary to passive music listening, music performance requires
2. The neural basis of the brain-music interaction (‘Brain on high precision control over motor execution and auditory perception
music’) systems. Meticulous motor control is required for the timing, sequence,
and spatial organization of the movement. These motor control com
2.1. Neurophysiology of the perceptual processing of music (passive ponents are associated with the architecture of musical rhythm, and the
involvement) tactile component of playing a musical instrument. The ability to
execute the movement at the precise timing has been linked to a neural
Music, as any auditory stimulus, is transmitted by a vibration of air, counter mechanism, which infers time through neuronal oscillations
which is then transduced into electrical impulses by the cochlea. The (Spencer et al., 2003; Wing, 2002), or a kinematic property of movement
cochlea is a spiral-shaped cavity in the inner ear that permits for diverse on its own (Penhune et al., 1998). The premotor and motor cortices,
frequencies to activate dedicated regions along the spiral (known as a cerebellum, and the basal ganglia contribute to the motor processes
tonotopic map). From the cochlea, sound waves-induced nerve impulses (timing, sequence) related to music performance (R. J. Zatorre et al.,
are carried to the brain for interpretation (Casale et al., 2021). A broad 2007). Activation of the supramarginal gyrus has been associated with
spectrum of sound features (e.g., pitch, timbre, sound intensity, inter tactile processes and limb positioning (Pando-Naude et al., 2021).
aural disparities) are encoded by diverse neural response properties and An auditory-motor interaction, which occurs when actively engaging
transformed to the auditory brainstem, namely superior olivary complex with music production can be defined as feedforward and feedback
and inferior colliculus (Langner and Ochse, 2006). From these brainstem communication. The motor system controls the fine movements when
regions, topographically organized auditory information travels to the playing an instrument. The produced sound is then processed by audi
auditory thalamus, specifically through the ventral geniculate body to tory circuits and adjust the motor output, if needed (R. J. Zatorre et al.,
the primary auditory cortex (A1) (Smith et al., 2012; Winer et al., 2002). 2007). The acquisition of auditory-motor sequences relies on cingulate
This pathway is known as the lemniscal pathway, and it represents the and cerebellar beta oscillations, which reflect the processing of auditory
major auditory signal processing pathway. The A1 receives diverse feedback-related adjustments during sensorimotor learning and perfor
temporal rate representations of acoustic signaling and uses them to mance (Herrojo Ruiz et al., 2017). Brain regions related to motor
encode for slowly changing sounds, and a neural firing rate-based rep planning and execution are co-activated (e.g., precentral, middle frontal
resentation to encode for fast varying sounds (X. Wang et al., 2008). The and supramarginal gyri). Along with motor function related to instru
internal representations of sound characteristics in the A1 no longer ment playing, these brain regions are also recruited during
reflect the original acoustic architecture. Sound structure alterations are sensory-motor coupling (Hanakawa et al., 2008).
imperative for the A1 to achieve sound segmentation, syntax processing, The emotional content during an instrument playing must be passed
and multisensorial integration (X. Wang et al., 2008). onto the listener. The anterior cingulate gyrus and insula are involved in
Since music is a multisensory stimulus, music listening activates A1 emotional information analysis and perceptual processing. The top-
along with motor and pre-motor regions, such as basal ganglia, primary down processing and control of emotional meaning, and bottom-up
motor areas, supplementary motor areas, and cerebellum (Pando-Naude analysis of the emotional content occurs in the anterior cingulate
et al., 2021). The A1 connectivity with the fronto-temporal-cerebellar gyrus solely (Pando-Naude et al., 2021). Interestingly, when profes
circuit yields perceptual processing during music listening (Chen sional composers are engaged with music creation, the integration of the
2
primary visual and motor areas is not required. These brain regions are pleasure when listening to music (Laeng et al., 2021). These experiments
instead involved in functional connectivity between the anterior suggest that the opioid system is not the primary mediator of
cingulate cortex (ACC) and the default mode network (DMN) to inte music-evoked pleasure and feeling of reward. However, music’s ability
grate sound with its emotional content (Lu et al., 2015). to stimulate opioid release is a powerful tool for pain-related therapeutic
conditions, thus further experiments on opioid system recruitment by
2.3. Neural circuits and neuromodulatory systems involved in the musical music are needed.
experience Along with the subjective increase of psychological wellbeing, music
exerts various physiological effects on the human body, mediated via the
Music can evoke a variety of emotions, feeling of pleasure/euphoria, autonomic nervous system (ANS). Music can induce changes in heart
increase in motivation and arousal. Music’s ability to tap into diverse rate, respiratory rate, blood pressure, electrodermal skin conductivity,
psychological and physiological brain states is mediated by the activa muscle tension, peripheral temperature (Blood and Zatorre, 2001; Fer
tion of the diverse neural circuits and neuromodulatory systems. In reri et al., 2019; Salimpoor et al., 2011). Chills evoked by highly plea
clinical settings, music acts as a non-pharmacological intervention that surable music are physiological markers of intense ANS activation (Mori
can attenuate various diseases, thus the mechanisms by which music and Iwanaga, 2017). The ANS function is primarily mediated by neu
exerts therapeutical effects are of great interest. romodulators noradrenaline, adrenaline, and acetylcholine (McCorry,
Listening to music, especially to subjectively preferred songs, en 2007).
gages brain pleasure pathways. A study using positron emission to In the brain, the primary source of noradrenaline (NA) is the locus
mography (PET) measured regional cerebral blood flow changes in coeruleus (LC), which activity has been associated with pupil dilation
response to the subject’s chosen highly pleasurable music, which would (Murphy et al., 2014), heart-rate variability (Mather et al., 2017),
evoke the experience of “chills” or “musical frisson” (Blood and Zatorre, emotional reactivity (Lerner et al., 2009). Pupil dilation increases in
2001). With increasing intensity of music-evoked pleasure, cerebral response to liked, predictable melodies (Bianco et al., 2019).
blood flow changes were registered in brain regions associated with Music-induced chills are correlated with greater pupil dilation, impli
reward, motivation, arousal, and emotions, namely ventral striatum, cating the central NA system in this phenomenon (Laeng et al., 2016).
midbrain, amygdala, orbitofrontal, and ventral medial prefrontal Another neuromodulator affected by auditory stimulation by music
cortices (Blood and Zatorre, 2001). These brain regions are similarly is serotonin (5-HT). Ascending from its source brainstem nucleus Dorsal
activated by highly rewarding stimuli such as food, drugs of abuse, sex Raphe, serotonin neurons project to parts of the auditory system where
(Berridge and Kringelbach, 2008; Oei et al., 2012). Although music does this monoamine neuromodulator acts as a prominent mechanism linking
not represent any biologically significant stimulus, it recruits the same external auditory processing with the internal state (Hurley and Hall,
brain circuits as the ones that are involved in pleasure and seeking 2011). Serotonin signaling has been mostly implicated in music
reward. Reward processes are known to recruit dopamine and opioid perception. In adult rats, exposure to classical music (Mozart’s sonata)
systems, as illustrated in studies in animal models (Peciña and Berridge, was associated with an increase of serotonin metabolites concentration
2013) and humans (Nummenmaa et al., 2018; Oei et al., 2012). The in caudate-putamen (CPu) (Moraes et al., 2018). Music’s effect on se
dopaminergic system related to reward mechanisms signals through the rotonin release in humans was studied by using the platelet model of
mesocorticolimbic pathway, which is comprised of the ventral serotonin content. According to the model, serotonin platelet content
tegmental area (VTA) (one of the two main nuclei of dopamine neurons reflects the serotonin content in neurons, such that decreased values for
in the brain) projecting to the ventral striatum, specifically nucleus the intracellular content reflect higher serotonin transmission history
accumbens (NAc) (Lammel et al., 2008). The dopamine release in (Evers and Suhr, 2000). In response to a pleasant music, the researchers
response to highly pleasurable music was measured by ligand-based PET measured an increase in serotonin platelets. In contrast, listening to
scanning. The study registered a release of dopamine in the dorsal and unpleasant music correlated with a decrease in serotonin platelets,
ventral striatum (NAc and the caudate putamen) at the peak of indicating an increased release of serotonin upon unpleasant music. The
emotional arousal when listening to one’s preferred song (Salimpoor results of the study suggest that serotonin release is modulated by the
et al., 2011). Furthermore, a causal link between dopamine release and music of different valence (Evers and Suhr, 2000). Serotonin’s role on
mediation of musical reward experience was shown in a study using music listening was further investigated in a study where participants
pharmacology. The study participants received (through oral adminis took lysergic acid diethylamide (LSD) (a psychedelic drug acting as
tration) either dopamine precursor (levodopa), D2-like dopamine re 5-HT2A receptor agonists) or LSD + ketanserin (5-HT2A receptor antag
ceptors inhibitors (risperidone), or placebo (lactose) while listening to onist) and their brain activity was assessed by collecting blood oxygen
music. The results revealed that dopamine receptor inhibition impaired level-dependent (BOLD) signal during music listening. The results show
participants’ ability to experience pleasure when listening to music, that serotonin receptor 5-HT2A signaling is implicated in neural response
whereas dopamine precursor enhanced musical pleasure (Ferreri et al., to music in brain regions related to higher-level musical processing, and
2019). supporting music-induced emotionality, meaningfulness, and connect
In animal models, reward associated with hedonic experiences in the edness (F. S. Barrett et al., 2018). Although evidence on serotonin
NAc is typically mediated by opioids, in contrast to dopamine trans modulation by listening to music is sparse, the comprehension about
mission shown in human studies exploring the music-associated feeling neuromodulatory underpinnings of musical experience is crucial for
of pleasure (Berridge and Kringelbach, 2008). According to animal health-relevant implications in daily life and therapeutical settings.
studies, music-evoked pleasure is not solely mediated by dopamine but Additionally, some neuropeptides, for example, oxytocin (Hansen
there is also a contribution of opioid signaling. This hypothesis has been and Keller, 2021; Keeler et al., 2015), are subjects to the modulation by
tested in a study in humans using naltrexone (NTX), a μ-opioid antag musical experience. The relation between oxytocin and music will be
onist, in a double-blind crossover study. Participants were administered discussed in the context of social bonding (part 3).
either NTX or a placebo and subjected to listen to pleasurable music
recordings of their choice, which would reliably produce intensely 2.4. Music-induced brain network oscillations
pleasurable feelings. NTX administration decreased the physiological
reactions associated with positive emotional experiences and Music engages an intricate set of brain regions and functional cir
self-reported measures of real-time pleasure (Mallik et al., 2017). A cuits, such as sensory-motor processing, cognitive, memory, and
similar study using NTX and placebo (d3 vitamin) found the opposite emotional components (Blood et al., 1999; Blood and Zatorre, 2001;
results. The μ-opioid antagonism did not change self-reported pleasure, Menon and Levitin, 2005). Moreover, music can modulate diverse brain
and it only dampened pupil response and decreased arousal during peak circuits and induce a release of various neuromodulators (Evers and
3
Suhr, 2000; Ferreri et al., 2019; Moraes et al., 2018). Due to afore act as an anxiolytic (Thaut et al., 2008). Listening to sad music,
mentioned reasons, music represents an ideal tool to investigate compared with happy music, is associated with stronger
multi-modal brain functioning and the integration of multisensory mind-wandering and greater transitions to the DMN (Taruffi et al.,
information. 2017). These results suggest that the emotional valence of the music can
Brain circuits recruited when listening to/playing music translate modulate the engagement of the DMN activity (Taruffi et al., 2017).
into large-scale brain oscillations that can largely impact neural popu
lation rhythms and thus, the general brain states. Neural population 3. Insights into brain function through musical experience
rhythms are cyclic changes in baseline neuronal activity that can be
observed in the local field potential (LFP), the electroencephalogram 3.1. Music and neuroplasticity
(EEG), and the magnetoencephalogram (MEG) (Slézia et al., 2011).
These neuronal population rhythms are usually evident in neocortical Neuroplasticity is brain’s ability to undertake functional and struc
and thalamic brain regions (Buzsáki and Draguhn, 2004; Slézia et al., tural modifications in response to experience or injury (von Bernhardi
2011). Thalamocortical oscillations occur in specific frequency bands of et al., 2017). These alterations might start as cellular changes and
delta (1–4 Hz), theta (4–8 Hz), alpha (8–12 Hz), beta (13–30 Hz), and progress to macro-modifications of the brain. Neuroplasticity can
gamma (30–70 Hz) (Buzsaki, 2006). Neural oscillations are the means of manifest as changes in neuron’s morphology, modifications of synaptic
communications between different nodes of the brain regions underly weight, synaptic pruning, cortical remapping (Olszewska et al., 2021;
ing cognitive processes or brain states (Buzsáki and Draguhn, 2004). Stegemöller, 2014; R. J. Zatorre, 2013). Music, as a multisensory stim
Connectivity dynamics of the neural circuits related to pleasantness ulus, has been shown to induce structural and functional changes in the
evoked by unfamiliar and familiar music has been addressed in a study brain, mostly due to continuous engagement of the brain regions related
using EEG. The authors observed an increase in right frontotemporal to music listening and/or performance.
theta synchronization along with higher reported pleasantness evoked Several studies have investigated the changes in musicians’ brains as
by previously unheard music. Significant theta synchronization a result of years of musical practice. The corpus callosum, a fiber tract
occurred between right temporal and left parietal signals (Ara and connecting the two cerebral hemispheres, is substantially larger in
Marco-Pallarés, 2021). Instead, inter-hemispheric temporoparietal theta musicians than in non-musicians, and the size of the corpus callosum
synchronization has been linked to pleasant feelings evoked by familiar positively correlated with the years of musical training (Reybrouck and
music. Together these results suggest that diverse theta connectivity Brattico, 2015). In multiple studies, it was confirmed that the corpus
patterns reflect the value assignment to music depending on whether the collosum exhibit a larger volume in musicians who started their practice
music is familiar to the listener or not (Ara and Marco-Pallarés, 2021). before the age of seven, as opposed to musicians who had a later onset
Theta and alpha frequency bands are observed when processing the (Reybrouck and Brattico, 2015; Schlaug et al., 2009; Wan & Schlaug,
music stimuli of different valence (consonant and dissonant chords). The 2010).
oscillation of these frequencies is driven by the amygdala and engages Notable music-induced changes in white and grain matter were also
the orbitofrontal and the auditory cortices (Omigie et al., 2015). reported. Bengtsson and colleagues (Bengtsson et al., 2005) studied the
Music listening evokes cortical activity in the high-gamma band in effects of long-term piano playing on the white matter in childhood,
the cortical regions ranging from the temporal lobe to the inferior frontal adolescence, and adulthood using diffusion tensor imaging. They
gyrus. Reversed temporal flow is observed during music recall. These discovered that pianists have more structured right posterior internal
mechanisms demonstrate bottom-up and top-down processes when capsule, than non-musicians. Training can cause plastic changes in the
listening to music and during the recall of a musical fragment (Ding white brain matter if training occurs during periods of fibre tracts
et al., 2019). Beta oscillations play a significant role in temporal pre maturation. Another study scientist looked at both grey and white
dictions to complex rhythms when listening to music, and enhance the matter in pianists and non-musicians (Han et al., 2009). In pianists,
processing of musical content (Doelling and Poeppel, 2015). Musical compared to non-musicians, grey matter density was higher in the left
improvisation and music creation is linked to feelings of ‘flow’ arising primary sensory-motor cortex and right cerebellum. Additionally, white
from a weak activity of the executive control network. The connectivity matter integrity was also higher in the right posterior internal capsule.
of sensorimotor and executive control networks does not differ when These findings suggest that long-term piano practice may cause grey and
imagining the performance or performing (Vergara et al., 2021). white matter adaptations in motor brain regions, which could affect the
Large-scale brain synchronization differs in musicians and non- number of synapses, glia volume, myelination, and axon diameter (Han
musicians. When listening to emotional sounds, musicians show et al., 2009).
greater activity in frontal theta and alpha bands (Nolden et al., 2017). As an evidence of music-related cortical remapping, Elbert and col
Professional musicians also exhibit more intense patterns of emotional leagues (Elbert et al., 1995) found that string players have the cerebral
activation and related brain synchronization when they listen to music representation augmented to the fingers of the hand that are used mostly
(Mikutta et al., 2014). In a study using EEG, mid-frontal theta and for playing the instrument. The more a given finger is used for playing an
posterior alpha-band activities during music perception were observed instrument, the larger the increase in cortical response is. Cortical
along with professional musicians’ more consistent ratings (compared to auditory representation in musicians is stronger compared to
non-musicians) of subjective arousal in response to classical music non-musicians as the magnitude of cortical activation for piano tones is
(Mikutta et al., 2014). by 25% greater in musicians. Both studies revealed a greater increase of
The brain state associated with music listening is the DMN. The DMN cortical representation in musicians who began musical training at
is related to specific brain functions, such as self-referential views, young age (Elbert et al., 1995; Pantev et al., 1998).
empathy, self-awareness, mind-wandering, imagining the future (Broyd As previously established by numerous studies, instrument playing
et al., 2009; Gusnard et al., 2001). This network is active when people engages the motor cortex and other structures involved in motor
listen to liked music, suggesting compatibility with the listener’s re movement coordination and initiation. Motor movements can be as
ported experiences of music-evoked introspection, mind-wandering, and simple as moving the musicians’ fingers, or as complex as coordination
self-referential thoughts (Wilkins et al., 2014). The activation of the of foot movement with hand movement. Music playing increases the size
DMN is related to the generation of innovative ideas, creation and in of grey matter in the primary motor cortex. The size of the right and left
ventiveness (Heuvel et al., 2009; Immordino-Yang et al., 2012). Thus, motor cortex in musicians and non-musicians was evaluated (Amunts
listening to one’s preferred music might grant the easier access to these et al., 1997). Both groups had leftward asymmetry but musicians had a
brain functions. This could have important therapeutical implications, smaller degree of asymmetry due to larger right motor cortex. The re
where music could improve executive functions and emotional states, searchers also discovered a negative relationship between the size of the
4
motor cortex in both hemispheres and the age at which musical practice verbal memory (Kraus and Chandrasekaran, 2010; Wan & Schlaug,
began (Amunts et al., 1997). The structural alterations in the motor 2010), faster neural response to speech (White-Schwoch et al., 2013),
cortex were a result of intensive and early hand skill instruction. Other and perform better in auditory tasks (Meyer et al., 2012) due to their
structures seen to be altered by music expertise are the cerebellum enhanced auditory attention (Kraus and Chandrasekaran, 2010). In the
(Olszewska et al., 2021; Reybrouck and Brattico, 2015) and precentral elderly individuals, volume and activity decrease in brain regions is a
gyrus, an area known for hand/finger movement integration (Wan & typical sign of aging. When compared to non-musicians, practicing
Schlaug, 2010). Schlaug group (Schlaug et al., 1995) measured the musicians have more grey matter volume in the left inferior frontal
volume of the cerebellum in musicians and non-musicians. They found gyrus. In contrast to non-musicians, musicians did not show substantial
that male musicians had a higher average cerebellar volume than age-related decline in overall brain volume or in brain areas such as the
non-musicians. These results showed that the cerebellum undergoes dorsolateral prefrontal cortex and the left inferior frontal gyrus. As a
microstructural adaptations in response to early initiation and result of their regular musical practices, musicians tend to be less sen
continued practice of complex finger sequences. sitive to age-related degenerations in the brain (Wan & Schlaug, 2010).
The auditory cortex is highly plastic in response to music as a year of Over 75-year-old participants were monitored for five years. When
music training can cause structural changes in the primary auditory compared to those who infrequently played a musical instrument, those
areas (Kraus and Chandrasekaran, 2010). Studies using diffusion tensor who regularly played a musical instrument were less likely to be diag
imaging have also shown that music exposure leads to stronger nosed for dementia. Playing music had a larger protective impact than
connection between the superior and middle parts of the gyrus (Meyer other cognitive tasks like reading, writing, or performing crossword
et al., 2012). Studies using fMRI scans have shown that all regions of the puzzles (Wan & Schlaug, 2010).
Heschl’s gyrus are activated stronger when exposed to music related
tasks involving rhythm and melody (Wan & Schlaug, 2010). The planum 3.2. Memory, emotions, and cognitive domains
temporal, a brain structure crucial in processing functions that allow for
the processing of music and speech, has been characterized by leftward As a research on music increases, researchers are drawn to explore
asymmetry and a greater size in musicians due to their ability to the effects of music on memory, emotions, and cognitive domains. A
perceive and process absolute pitch (Meyer et al., 2012; Reybrouck and study by Ferreri and Rodriguez-Fornells (Ferreri and
Brattico, 2015; Wan & Schlaug, 2010). Using voxel-based morphometry Rodriguez-Fornells, 2017) explored the effect of music-induced reward
analysis, Gaser & Schlaug (2003) discovered higher grey matter volume on episodic memory, revealing a correlation between pleasurable music,
in the motor, auditory, and visuospatial cerebral areas in musicians. The dopamine release, and enhanced memory (Ferreri and
findings point to structural brain adaptations in response to skill Rodriguez-Fornells, 2017). Brain activation during music listening or
acquisition and their long-term repetition. recall involves regions such as the temporal and frontal lobes, with
The hippocampus has also been found to have a greater grey matter anatomical differences observed in experienced musicians (Ding et al.,
volume in professional musicians in comparison to non-musicians (62). 2019; Ford et al., 2011; Groussard et al., 2010). Musicians, particularly,
Various studies were curious about the effect of musical process on the exhibit denser grey matter in the left hippocampus, resulting in more
connectivity between the NAc and other regions of the brain. The vivid and intense music-associated memories (Groussard et al., 2010).
findings showed that different sides of the NAc are connected with Verbal memory advantages in musicians, proven by studies (Franklin
different parts of the brain depending on whether the individual is a et al., 2008; Thaut et al., 2005), can be compromised by articulatory
musician or not. For instance, the NAc functional connectivity with the suppression, indicating enhanced verbal memory rehearsal mechanisms
hippocampus is stronger in non-musicians as opposed to the preferen (Franklin et al., 2008). Additionally, exposure to classical music during
tially stronger connectivity within the temporal pole and ventromedial study and sleep enhances short-term memory, especially in females (Gao
frontal areas in musicians (Reybrouck and Brattico, 2015). et al., 2020).
Musician’s engagement with music instruments is related to an The retrieval of memories through music can evoke emotions akin to
improvement of various music-related abilities. For instance, musicians those experienced during the initial events. Music-evoked emotions,
are better at pre-attentively extracting information out of musically conveyed as effectively as verbal language (Proverbio and Russo, 2021),
relevant stimuli, and demonstrate superior temporal integration induce powerful feelings, including chills or tears (Lundqvist et al.,
(Koelsch et al., 1999; Rüsseler et al., 2001). Another study demonstrated 2009; Mori and Iwanaga, 2017). Chills and tears represent happy or sad
that a year of high-intensity aural skills training in musicians improved emotions, respectively, with tears having a cathartic effect (Mori and
neural responses to temporal novelty in the hippocampus. This Iwanaga, 2017). Happy music induces more happiness, while sad music
improved sensitivity in the hippocampus positively correlated with increases empathy and pain processing (Lundqvist et al., 2009; Cheng
musical abilities of the participants (Li et al., 2018). Musacchia and et al., 2017). The limbic system, especially the amygdala, plays a crucial
colleagues (Musacchia et al., 2007) demonstrated that changes in role in music-evoked emotions (Koelsch et al., 2021). Furthermore,
functional organization also occurs within subcortical sensory structures music can influence visual information, affecting visual alertness and
in musicians compared to controls. Specifically, musicians have earlier attention shifts (Koelsch, 2014).
and larger auditory and audiovisual brainstem responses to speech and Music perception, composition, and imagery constitute basic
music stimuli. The magnitude of the brainstem response to speech contemporary music procedures engaging auditory cortices, sensori
stimuli positively correlated with the years of engagement in musical motor cortices, and the cerebellum. Imagery, particularly auditory
practice, suggesting that musical training enhances the representation of imagination, enhances pitch accuracy, while motor imagination aids
the pitch in the brain. Even though there are exceptional cases where coordination in large groups (Keller, 2012; R. Brown and Palmer, 2013;
musicians created beautiful pieces while deaf, like Beethoven, the ma Pando-Naude et al., 2021). Directed imagination exercises with music
jority of musicians rely heavily on their auditory perception to create influence vividness, sentiment, and perception, showcasing music’s
music as well as to perform it. Musicians are better at detecting and potential in therapeutic tools like Exposure Therapy and Imagery
processing micro-changes in essential aspects of music such as pitch, Rescripting (Herff et al., 2021). This suggests that music could be inte
timbre, and tempo (Kraus and Chandrasekaran, 2010; Meyer et al., grated into the standard therapeutic protocols, potentially enhancing
2012). Children also showed higher sensitivity to pitch patterns that are the effectiveness of these treatments and improving patient outcomes.
of their mother tongue (Kraus and Chandrasekaran, 2010). These ad Imagination is a powerful skill used by athletes and musicians to
vantages are due to more precise neuronal representations in their enhance their performance, impacting various cognitive and emotional
auditory cortex (Meyer et al., 2012). Individuals who were involved in aspects (Keller, 2012; Pando-Naude et al., 2021).
music whether on long-term or short-term basis have shown improved Multiple studies establish a positive correlation between music
5
playing and cognitive abilities (Grassi et al., 2017; Hars et al., 2014; 3.3. Social bonding
Okada and Slevc, 2018; Schneider et al., 2019). Musical experience in
fluences working memory processes (Okada and Slevc, 2018), and elder The attachment of memories to music does not only explain its as
musicians exhibit enhanced working memory and visual spatial per sociation with emotions, but it is also a reflection on music’s inherent
formance (Grassi et al., 2017). Weekly music-based multitask training social nature (Koelsch, 2014; Nummenmaa et al., 2021). In fact, the
reduces anxiety in older adults (Hars et al., 2014), and long-term music neural pathways activated by music processing are very close to those of
training improves both auditory and visual working memory in social processing (Nummenmaa et al., 2021). Thus, music can allow for
non-professional musicians (George and Coch, 2011; Putkinen et al., the attainment of social human needs (Koelsch, 2014). When groups of
2021). Music practice enhances attention, working memory, and people come together to sing or play instruments, stress and arousal
speech-in-noise perception, indicating improved cognitive factors levels are reduced due to an increase in adrenocorticotrophic hormone
(Kraus et al., 2012). This leads to musical training allowing one to hear levels after singing together (Keeler et al., 2015). Also, an increase in
speech in noise, due to the enhanced cognitive factors (Kraus et al., positivity, engagement, connectivity, and endorphin levels occurred
2012). Another study found a significant increase in verbal intelligence (Weinstein et al., 2016), all while negative emotions decreased and
among children in a music group that trained their music listening skills positive ones increased (Kreutz, 2014). Moreover, singing in large
(Moreno et al., 2011). During an executive-function task, verbal intel groups of unfamiliar people seems to have a more powerful effect when
ligence improvements were linked to functional brain plasticity changes compared to smaller more familiar groups (Weinstein et al., 2016). All in
(Moreno et al., 2011). The children also had better vocabulary knowl all, individual wellbeing and bonding increased dramatically when
edge, indicating an increase in verbal intelligence after music training placed in a musical group. All these positive effects could be due to the
exposure (Moreno et al., 2011). Thus, the effects of music training on increase in the levels of endorphins (Launay et al., 2016; Tarr et al.,
language and executive functions are related (Moreno et al., 2011). 2014; Weinstein et al., 2016), oxytocin (Keeler et al., 2015; Kreutz,
Interestingly, music, unlike other art forms, can affect brain processes 2014; Launay et al., 2016) and other hormones or neuromodulators
concerning auditory and other mechanisms (Moreno and Farzan, 2015). (Launay et al., 2016) when in musical groups. Synchronized human
Research has shown that children who had music training achieved activities, like group singing, were found to release endorphins (Launay
better results on second language acquisition and musical achievement, et al., 2016) and increase pain thresholds (Weinstein et al., 2016).
on the contrary to children who did not have any musical training (Yang Moreover, endorphins and the endogenous opioid system were found to
et al., 2014; Putkinen et al., 2021). However, development of cognitive assist in social bonding (Tarr et al., 2014). Meanwhile, oxytocin induces
abilities, such as mathematical skills, did not improve in the musician social bonding conditions like communication, cooperation, and eye
group (Yang et al., 2014). A sample of undergraduates showed positive contact (Keeler et al., 2015). In the context of diseases, these findings
relationships between musical capability and duration of music training, suggest that music-based interventions, particularly those involving
socioeconomic status, short-term memory, general cognitive skills, and group activities, have the potential to modulate hormonal and neuro
openness to new experiences (Swaminathan and Schellenberg, 2018). In modulator levels, providing a therapeutic avenue for conditions related
the combined analysis of these factors, musical competence was asso to stress, emotional well-being, and social bonding (Launay et al., 2016;
ciated with longer music training, better general cognitive skills, and Tarr et al., 2014; Weinstein et al., 2016; Keeler et al., 2015; Kreutz,
more openness (Swaminathan and Schellenberg, 2018). Moderation 2014). Dancing also seems to facilitate bonding and produce positive
analyses showed that participants who scored below the mean on the set emotions (Nummenmaa et al., 2021), and a stronger bond can form
measure of general cognitive skills showed the partial association be when there are similar musical preferences with others (Stupacher et al.,
tween musical proficiency and music training (Swaminathan and 2020). Finally, children with autism appeared to be able to communi
Schellenberg, 2018). Furthermore, general cognitive skills and openness cate better after being placed in music group therapies, with improve
were indirectly associated with musical capability since they predicted ments in many areas of their brain’s connectivity (Sharda et al., 2018).
music training, which was in turn related to musical competence
(Swaminathan and Schellenberg, 2018). There are many factors that 3.4. Music, speech and language
contribute to musical competence, including but not exclusive to music
training (Swaminathan and Schellenberg, 2018). Researcher looked at The separate concepts of music and language have existed for de
the N400 effect while examining arousal levels of participants during cades. However, it is becoming more apparent that both music listening/
reading comprehension with and without the music in the background. production and language share certain properties that make these two
The N400 effect was smaller in the group that was reading without processes interconnected (Peretz et al., 2015). This anatomical and
music, compared to the participants who were listening to background functional link occurs because both linguistic and musical syntax share
music, suggesting that background music increases the difficulty of se common syntactic processes for different domain-specific syntactic
mantic integration that occurs during reading comprehension (Du et al., representations, executed by the same brain regions (Patel, 2003).
2020). Experimental evidence shows that music reduces cognitive dis For instance, one study showed that people who were trained in
sonances, a discomfort caused by holding conflicting ideas simulta music showed better semantic processing than those who did not have
neously and inevitably leads to devaluation of conflicting ideas music training (Yu et al., 2017). However, phonological processing was
(Masataka and Perlovsky, 2012). Meditation involving singing or music not affected regardless of whether the person had musical training or not
listening has shown significant enhancement of subjective memory (Yu et al., 2017). Furthermore, semantic language processing and
function and objective cognitive function (Innes et al., 2017). Addi musical melodic analysis showed correlation at both the regional and
tionally, research indicates that both verbal and auditory working network levels, proving, that they are associated, and that there is a
memory rely on the ability to produce the instantly remembered sounds, neural basis link between them (Yu et al., 2017). In a study that
which suggests that sensorimotor representations are vital for the tem attempted to investigate the effects of music on word learning, musical
porary storage of auditory information in working memory (Schulze and stimuli, regardless of the degree of stimuli, were found to effectively
Koelsch, 2012). Music training results in significantly higher visual ease learning words and long-term retention of the learned words. Music
spatial ability, executive functioning, and naming skills compared to helped to encode the speech signal rather than recognition of the words
those who do not practice music in any way (Strong and Mast, 2019). (Ma et al., 2020). The study also showed that encoding was improved,
However, differences are not significant among those same groups particularly in terms of word association, but also possibly in terms of
regarding processing speed or episodic memory (Strong and Mast, word segmentation. The current findings suggest that including musical
2019). elements into speech improves both word learning and long-term
memory (Ma et al., 2020). This suggests that incorporating music into
6
therapeutic approaches for diseases involving language and memory non-musicians. In other words, encoding of these cues in native speakers
functions could potentially enhance word learning and retention, is enhanced (Intartaglia et al., 2017). A distinct finding was that the
providing a novel avenue for intervention strategies (Ma et al., 2020). neural responses to formant frequencies were the same for native
When listening to music, the pars orbitalis, a brain area involved in speakers and non-native musicians, suggesting that music training could
improving the linguistic structure of spoken and sign language, showed recouperate for the lack of language exposure by encoding crucial
activity related to focal activation (Levitin and Menon, 2003). Moreover, acoustic information (Intartaglia et al., 2017). The acquisition of lan
vocal or instrumental music was found to preferentially activate the guage and music seems to increase acoustic sensitivity in a functionally
superior temporal gyrus (STG), specifically bilaterally in the anterior relevant manner, such as phoneme discrimination (Intartaglia et al.,
planum polare (Whitehead and Armony, 2018). While on the other 2017). In one study, increasing syntactic complexity and distracting
hand, speech or singing engaged a great part of the superior temporal melodies caused the interference of one-timbre melodies with sentence
sulcus, suggesting that both areas integrate (Whitehead and Armony, recall (Fiveash et al., 2018). By contrast, alternating instruments in
2018). Several fronto-parietal areas, primarily the dorsolateral pre three-timbre melodies diminished interference on long-distance syn
frontal cortex, the supramarginal/angular gyrus and the precuneus, are tactic structure building, probably because they interrupted auditory
activated when listening to autobiographical music (familiar own streaming (Fiveash et al., 2018). Additionally, in contrast to
and/or other) (Castro et al., 2020). Also, familiar own music (compo three-timbre melodies, the participants had an easier time distinguish
sitions) activated brain regions including the medial prefrontal cortex, ing syntactically coherent one-timbre melodies (Fiveash et al., 2018).
visual imagery, and reward and emotion networks, whereas familiar Thus, results indicate syntactic analysis and auditory streaming may
other music (favorite) activated reward networks, such as ventral interact to influence recall of sentences (Fiveash et al., 2018). The beta
striatum (Castro et al., 2020). Accordingly, familiar music containing and gamma frequencies in EEG have been shown to have the greatest
self-related references (compositions) was associated with an enhanced significance for determining mental activities. The results of an analysis
activation of the autobiographical network during subsequent familiar of the energy distribution in EEG channels show a clear correlation
naming (in comparison to music that did not contain self-related refer between mental processes and the external world perception. Hence,
ences); the precuneus was strongly associated with such processing language skills may be affected by music training (Besedová et al.,
(Castro et al., 2020). 2019).
It is hypothesized that having a minor superiority in decoding speech
sounds may have led to the dominant role of the left hemisphere in many 4. Discussion
sophisticated linguistic functions (R. J. Zatorre et al., 2002). As a result,
the right hemisphere may have played a crucial role in aspects of Music is a complex stimulus that engages multiple brain regions and
musical perception, especially those related to tonal pitch, and may have circuits, influencing various domains of human functioning. The ability
been a result of, and a complement to, this specialization of language (R. of music to influence the brain is due to the brain’s neuroplasticity, the
J. Zatorre et al., 2002). Another hypothesis suggests that, syntax in ability of the brain to reorganize and adapt to new experiences. Studies
language and music is governed by hierarchical organization, as have shown that musical training can bring about structural and func
fundamental features of hierarchical control in the brain include func tional changes in the brain, resulting in increased grey and white matter
tional architecture of the frontal cortex, which maintains abstract rep density, larger corpus callosum, and greater cortical remapping in areas
resentations in anterior parts of the brain and more concrete related to music performance. Moreover, music can improve cognitive,
representations preferentially in posterior parts, dual control function (i. emotional, physical, and social domains, making it a valuable tool for
e. inhibition and selection) of the basal ganglia, and the influence of promoting health and wellbeing.
abstract temporally extended processes on concrete motor processes by One of the most remarkable aspects of music is its ability to tap into
means of cortico-basal-ganglia-thalamus circuits (Asano et al., 2021). different brain states, influencing diverse neural circuits and neuro
Evidence suggests that using neural networks to model cognitive modulatory systems. The emotional valence of music modulates the
tasks such as word learning could help provide a greater understanding engagement of brain regions involved in self-referential views, empathy,
of the neural dynamics underlying these tasks (Elmer and Jäncke, 2018). self-awareness, mind-wandering, and imagining the future. The activa
One study have shown that subjects with no special musical training tion of the default mode network (DMN) by music listening is also linked
who are naively presented with different musical excerpts associate the to the social and emotional functions of the brain. Music can trigger the
musical extracts with specific words similarly (Koelsch et al., 2004). release of hormones such as endorphins and oxytocin, which promote
Consequently, event-related potential (ERP) results, particularly the social bonding and alleviate pain. Music is a non-pharmacological
N400 priming effect, for the music indicated no differences in terms of intervention that can be used to treat various diseases, including
latency, scalp distribution, neural site, or amplitude between the music stroke, Parkinson’s disease, and dementia. Music therapy involves the
and language (Koelsch et al., 2004). Concurrent music processing use of music to address physical, cognitive, and emotional needs of
delayed N400 effects only when melodies are familiar, whereas double patients. Music therapy can improve motor control, speech, language,
violations of familiar melodies produce a sub-additive N400 effect and memory in stroke and Parkinson’s disease patients. In dementia
(Calma-Roddin and Drury, 2020). Additionally, both early negativity patients, music can trigger memories and improve mood, socialization,
effects (right anterior negativities RAN effects), which are associated and quality of life. Music therapy can also reduce anxiety, depression,
with musical syntax, as well as the music N400, were delayed in onset and pain in cancer patients.
for familiar melodies, and double violation cases associated with unfa The integration of music as a therapeutic tool prompts considerations
miliar melodies also caused RAN effects to be delayed (Calma-Roddin for its clinical implications and future research directions. To harness
and Drury, 2020, p. 400). Together, these patterns demonstrate the the full potential of music in healthcare, clinicians could explore tailored
existence of interference effects within these domains and add evidence music interventions for specific conditions, considering individual
regarding previously unreported types of interactions to the collection of preferences and cultural backgrounds. Additionally, interdisciplinary
findings important to determining if language and music share similar collaborations between neuroscientists, musicians, and healthcare pro
mechanisms (Calma-Roddin and Drury, 2020, p. 400). fessionals can advance our understanding of the underlying mechanisms
Experimental evidence shows that tonal music induces highly and optimize therapeutic applications. Moreover, investigating the long-
structured mental representations of musical pitch in both musicians term effects of sustained musical engagement and exploring innovative
and non-musicians (Patel, 2003). Language experience may contribute technologies, such as personalized music playlists, can enhance the ef
to an automatic encoding of subcortical electrophysiological responses ficacy of music-based interventions in diverse clinical settings. Future
to english syllables in native speakers as compared to non-native research should extend beyond music to examine the transferability of
7
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Brain, Behavior, & Immunity - Health 35 (2024) 100716

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Brain, Behavior, & Immunity - Health

The transformative power of music: Insights into neuroplasticity, health,

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