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Brain mechanisms that underlie music interventions in the exercise domain
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Brain mechanisms that underlie music interventions in the exercise domain Costas I. Karageorghis*,1, Marcelo Bigliassi*, Segole`ne M.R. Guerin†, Yvonne Delevoye-Turrell† *Brunel University London, Uxbridge, United Kingdom † University of Lille, Lille, France 1 Corresponding author: Tel.: +44 01895 266476, e-mail address: [email protected]
Abstract In this chapter we review recent work from the realms of neuroscience and neuropsychology to explore the brain mechanisms that underlie the effects of music on exercise. We begin with an examination of the technique of electroencephalography (EEG), which has proven popular with researchers in this domain. We go on to appraise work conducted with the use of functional magnetic resonance imaging (fMRI) and then, looking more toward the future, we consider the application of functional near-infrared spectroscopy (fNIRS) to study brain hemodynamics. The experimental findings expounded herein indicate that music has the potential to guide attention toward environmental sensory cues and prevent internal, fatigue-related signals from entering focal awareness. The brain mechanisms underlying such effects are primarily associated with the downregulation of theta waves across the cortex surface, reduction of communication among somatosensory regions, and increased activation of the left inferior frontal gyrus. Taken holistically, research in this subfield of exercise psychology demonstrates a vibrant and reflexive matrix of attentional, emotional, behavioral, physiological, and psychophysiological responses to music across a variety of exercise modalities and intensities. The emergent hypotheses that we propose can be used to frame future research efforts.
Keywords Brain, Exercise, Music, Neuroscience, Psychophysiology
1 INTRODUCTION Music making has been a staple of human civilizations since time immemorial. Anthropologists have argued that music is part of our genetic blueprint and an activity that almost defines the human species (Mithen, 2006). The celebrated neuropsychologist, Daniel Levitin, indicated how music is unusual among all human Progress in Brain Research, ISSN 0079-6123, https://doi.org/10.1016/bs.pbr.2018.09.004 © 2018 Elsevier B.V. All rights reserved.
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activities for both its ubiquity and its antiquity. In the modern era, music has become an essential element of the exercise experience, albeit the music-in-exercise phenomenon was predicated on scant empirical research (see Karageorghis and Terry, 1997). Following 200 or so experimental studies in the period since the early 1990s, it is only very recently that researchers have grappled with neurophysiological or brain mechanisms that underlie the application of music in the exercise domain.
1.1 THE MAIN EFFECTS OF MUSIC DURING EXERCISE In the exercise and physical activity context, the most common effects reported for when people are exposed to music fall under the categories of psychological, psychophysical, psychophysiological, and ergogenic (Terry and Karageorghis, 2011). The psychological effects pertain to how music can influence mood, affect (i.e., core feelings of pleasure displeasure), emotion, cognition, and behavior. As a close cousin of psychological effects, psychophysical effects pertain to the psychological perception of one’s physical state. In exercise science, this is most often operationalized through the rating of perceived exertion (RPE; e.g., Jones et al., 2017; Stork and Martin Ginis, 2017). The mainstay of the literature has focused on psychological and psychophysical effects in response to music during continuous exercise (see Clark et al., 2016; Karageorghis, 2016 for reviews). Given the consistency of findings regarding, for example, enhancements in affect across a range of exercise intensities and the lowering of perceived exertion at low-to-moderate intensities (Hutchinson and Karageorghis, 2013; Taylor et al., 2007), the underlying cerebral mechanisms hold particular interest for the exercise science community. Psychophysiological effects pertain to how music can influence aspects of physiological functioning and thus common measures employed by researchers include heart rate, blood pressure, and oxygen uptake (Bacon et al., 2012; Karageorghis et al., 2019). This is also the category under which EEG falls, and, as we will discover later in this chapter, this is a technique that has enabled us to shine a light on salient cerebral mechanisms in the music–exercise relationship. Ergogenic effects are evident when music elevates work output or causes higher-than-expected power output, endurance, or productivity (Terry and Karageorghis, 2011). On occasion, researchers have used brain-monitoring techniques in tandem with electromyography to investigate how the reorganization of electrical activity in the brain has a bearing on muscular activity (e.g., Bigliassi et al., 2018a).
1.2 APPLICATIONS OF MUSIC IN EXERCISE In the exercise context, music is applied in three primary ways and, as we will go on to detail, these can follow on sequentially from one another. There is the pretask application wherein music is used to prime an exerciser or get them in the right frame of mind to be active (see Smirmaul, 2017 for a review). Given the imperative to “lead” an individual from a sedate state into an active state, such music will often be at a moderate tempo and have inspirational or motivational lyrics. For example,
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Work From Home (100 bpm) by Fifth Harmony (feat. Ty Dolla $ign), which is an ideal precursor to a home workout. The in-task application of music has, by far and away, attracted the most research interest (see Karageorghis and Priest, 2012a,b). In this instance, music can be applied synchronously or asynchronously. The former entails a process of auditory-motor synchronization in which the music adopts a metronomic function to regulate movement and evenly distribute energy expenditure (see, e.g., Bacon et al., 2012; Karageorghis et al., 2009). The latter entails the absence of conscious synchronization and represents the way in which music is most commonly applied to individual exercise routines. For example, if an exerciser is engaged in a resistance training routine and music is piped into the gymnasium via a music-video channel, the strong likelihood is that the music will be used in the asynchronous mode. Hence, asynchronous music is akin to background or ambient music, although it can be played at a high intensity. In terms of the synchronous application of music, technological advancements have reduced the need for an exerciser to engage in conscious or active synchronization with a musical beat. Synchronization can be facilitated by means of an interface that uses accelerometry and algorithm-generated playlists to ensure coupling of musical rhythm with an individual’s work rate (see, e.g., Moens et al., 2014; D-Jogger). Such advancements have recently led to new definitions for the two main forms of auditory synchronization: (a) active synchronization entails a motor process in which an individual or group consciously synchronize their movement rate with the rhythmical qualities of music and (b) passive synchronization entails a motor process wherein a digital interface adapts the tempo of music in real-time or assigns a track at a tempo to match the movement rate of an individual or group (Karageorghis, 2019). The third main type of music application is post-task music (music used immediately after an exercise task) and similar to the synchronous application, two forms have recently been delineated. The first of these was coined respite music by Jones et al. (2017) and entails the use of music in between high-intensity intervals with a view to tempering the negative affective state that is typically elicited by such activity. Respite music also bears a positive influence on hemodynamic and cardiovascular recovery processes. The second is known as recuperative music (Terry and Karageorghis, 2011) and can be applied to both active (movement-based) and passive (static) forms of exercise recovery. Recuperative music playlists are typified by a progressively slower tempo (90 to 60 bpm), as the intention is to leave the exerciser with a sense of calmness and revitalization. Investigation of the brain mechanisms that underlie post-task music has yet to be initiated and thus provides tantalizing opportunities for future researchers. In this chapter, we will explore the brain mechanisms that underlie the use of in-task music in the exercise and physical activity domain. Some of the in-task mechanisms provide strong clues as to how music can take effect in the pretask phase (e.g., Bishop et al., 2014). We will begin with an examination of the technique of electroencephalography (EEG; i.e., measuring electrical activity in the brain), which
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has proven the most popular in this domain. Thereafter we will review work conducted with functional magnetic resonance imaging (fMRI)—a brain imaging technique that entails use of a scanner but is restricted to experimental tasks that involve no head movement. Finally, looking toward the future, and on the basis of a relatively small corpus of work, we have included a section on functional near-infrared spectroscopy (fNIRS) in which we present a hypothesis that could form the basis of future study. fNIRS is a newly-popularized technique that offers tremendous promise in terms of furthering our understanding of hemodynamic changes in the cortex that are caused by music during exercise.
2 EFFECTS OF MUSIC ON ELECTRICAL ACTIVITY IN THE BRAIN One of the first experiments to investigate the cerebral mechanisms underlying the effects of music on high-intensity exercise was conducted by Bigliassi et al. (2016). The authors employed an isometric, closed-kinetic chain exercise mode (an ankledorsiflexion task) and delivered the well-known track, Eye Of The Tiger by Survivor (109 bpm), in-task as a means by which to motivate participants. The exercise task was executed at 40% of each participant’s maximal voluntary contraction until the point of volitional exhaustion. Electrical activity in the brain was captured by means of a 64-channel EEG device, and a number of perceptual and affective responses were assessed. The results indicated that asynchronous music increased the use of dissociative thoughts and enhanced task performance to a greater degree than a no-music control condition. The authors also showed that music was sufficiently potent to downregulate theta waves (4–7 Hz) in the frontal, central, parietal, and occipital regions of the brain (see Fig. 1). This psychophysiological response appears to be associated with a mechanism pertaining to the suppression of fatigue-related symptoms (see Craig et al., 2012; Tanaka et al., 2012). Accordingly, Bigliassi et al. (2016) proposed a series of mechanisms in this study, suggesting that music-related interventions have the potential to reallocate attentional focus toward task-unrelated factors (e.g., exteroceptive signals), and ameliorate exertional responses to a greater degree than no-music conditions (Karageorghis et al., 2017). The upshot is that the processing of afferent feedback is reduced when individuals exercise with asynchronous music; a psychophysiological response that facilitates the execution of movements and optimizes the neural activation of working muscles (Bigliassi, 2015). Subsequently, the same group of researchers conducted a more complex experiment to identify the effects of music on motor unit recruitment and brain connectivity during whole-body exercise modes (Bigliassi et al., 2017). Participants exercised on a mechanically-braked cycle ergometer at light-to-moderate-intensities (i.e., at 10% below ventilatory threshold [VT]) for 12 min, while exposed to stimulative, asynchronous music. They also used another auditory stimulus—an audiobook—as a means by which to isolate the effects of musical components (e.g., melody and harmony) on perceptual, affective, and psychophysiological
ankle-dorsiflexion task. Psychophysiology 53, 1472–1483. https://doi.org/10.1111/psyp.12693. Copyright 2016, with permission from John Wiley & Sons.
Reprinted from Bigliassi, M., Karageorghis, C.I., Nowicky, A.V., Orgs, G., Wright, M.J., 2016. Cerebral mechanisms underlying the effects of music during a fatiguing isometric
Low-frequency components (theta waves) of the power spectrum presented for CO, MM, and MO. Note: The colored scale indicates the power of the band frequency (power [signal units^2/Hz*10^ {11}]); CO ¼ control condition; MM ¼ music condition; MO ¼ music-only condition.
FIG. 1
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responses. Time-frequency analyses were used to identify the synchronization– desynchronization–resynchronization cycle at the Cz electrode site. This site was chosen given that it provides a vista into the neural activity of lower limbs (i.e., the central area of the premotor cortex; Jain et al., 2013). The results of the Bigliassi et al. (2017) study indicate that music has the potential to inhibit alpha resynchronization and increase the number of motor units recruited to execute the task. The authors hypothesized that this psychophysiological mechanism could be representative of more autonomous control of working muscles. Put another way, asynchronous music appears to distract participants, guiding attention toward exteroceptive cues, and such distraction leads to a more efficient/autonomous neural activation. The results were supported by electromyographic (EMG) analysis, which indicated an increase in the number of motor units recruited in the quadriceps. Finally, the authors decided to run brain connectivity analysis to investigate whether the communication across somatosensory regions of the cortex was affected by the presence of auditory stimuli. Interestingly, music reduced the connectivity between Cz and both frontal (Fz) and central (C4) electrodes (see Fig. 2). The research team proposed that the reduction in communication could have been induced by a decrease in corollary discharges emitted from the central motor command to somatosensory regions of the cortex. This study demonstrated that music can lead to rearrangements of the electrical activity in the brain that are reliant upon exercise intensity, complexity, and mode. The results serve to illustrate that more robust methods such as timefrequency decomposition (e.g., wavelet transformations) and brain connectivity analysis (e.g., spectral coherence) might help to clarify underlying mechanisms that are not easily evident using traditional EEG procedures such as time-domain analysis (e.g., event-related potential). More recently, Bigliassi et al. (2018b) investigated the effects of asynchronous music on real-life physical activity. Participants were required to walk 400 m at a pace of their choosing on an outdoor running track. Electrical activity in the brain was recorded by use of a portable EEG device with active shielding technology. Such technology appears to function as a form of portable Faraday cage that protects the core components of cables against external artifacts during the execution of movements. The researchers also measured perceptual (state attention and perceived exertion) and affective (valence, arousal, and perceived enjoyment) outcomes immediately after each exercise bout. The results indicated that asynchronous music enhanced affective responses, increased the use of dissociative thoughts, and upregulated beta waves in the frontal and frontal-central regions of the brain to a greater degree than no-music control and podcast conditions (see Fig. 3). Based on previous findings (Bailey et al., 2008; Sayorwan et al., 2013), the researchers hypothesized that rearrangement of beta waves elicited by music serves to up/downregulate affective responses. Moreover, that this positive psychophysiological state could be capitalized upon during many forms of physical activity, as a means by which to render a given activity more pleasurable.
Physiol. Behav. 177, 135–147. https://doi.org/10.1016/j.physbeh.2017.04.023. Copyright 2017, with permission from Elsevier.
Reprinted from Bigliassi, M., Karageorghis, C.I., Wright, M.J., Orgs, G., Nowicky, A.V., 2017. Effects of auditory stimuli on electrical activity in the brain during cycle ergometry.
Group data alpha coherence values in CO (control condition), AB (audiobook condition), and MU (music condition). Reduced spectral coherence is manifest across central and frontal electrodes sites in MU when compared to CO and AB.
FIG. 2
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FIG. 3 Group data time-averaged band frequencies for CO, PO, and MU. Note: SMR, sensorimotor rhythm. The colored scale indicates the power of the band frequencies (signal^2/Hz*10^{10}); CO ¼ control condition; PO ¼ podcast condition; MU ¼ music condition. Reprinted from Bigliassi, M., Karageorghis, C.I., Hoy, G.K., Layne, G.S., 2018b. The Way You Make Me Feel: psychological and cerebral responses to music during real-life physical activity. Psychol. Sport Exerc. Advance online publication. https://doi.org/10.1016/j.psychsport.2018.01.010. Copyright 2018, with permission from Elsevier.
3 EFFECTS OF MUSIC ON SUBCORTICAL BRAIN REGIONS It is also necessary to examine the effects of music on exercise by use of high-spatial resolution techniques, which allow researchers to actually “see” the brain regions that activate in response to music and exercise; one such technique is functional magnetic resonance imaging (fMRI). This is oft-used in the field of neuroscience and cognitive psychology to detect changes in oxygenated hemoglobin that are coupled with neuronal activation. An fMRI scanner quantifies the energy emitted by protons and creates brain images based on the chemical structures of brain tissue. The scanner can also identify inhomogeneities in the brain that are caused by the deoxygenated/oxygenated blood ratio. This biophysical response is used to identify the speed of decay of the MRI signal, representative of neural activity, and referred to as the blood oxygenation level dependent (BOLD) response. Bishop et al. (2014) conducted one of the initial studies with fMRI and pretask music, albeit this was oriented more toward sport-related tasks. The researchers manipulated tempo and intensity of music (dBA) in order to identify changes in choice reaction-time. The hypotheses were predicated on the assumption that shorter reaction times occur at an optimal arousal state, and fast-tempo music was expected to
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upregulate arousal with corollary changes in brain activity. This study illustrates the nexus between musical components and neural systems, such as attention and motor control. The results supported the authors’ predictions that fast-tempo music would increase visuomotor activity through the elicitation of emotional responses. The subcallosal gyrus was significantly activated during the reaction-time task that followed fast-tempo, pretask music. Hence, fast-tempo music presumably elicited greater levels of arousal, which modulated reaction time by activating indirect pathways. In this case, arousal dictated participants’ level of readiness and voluntary/ involuntary attention. The only fMRI study designed to investigate the effects of in-task music on exercise was very recently conducted by Bigliassi et al. (2018c). The authors decided to use a simple exercise mode (handgrip) to investigate the brain regions that activate in response to exercise coupled with asynchronous music. The squeezing task was adopted in order to prevent head movements, which can easily compromise the fidelity of a scan. Participants were required to execute 30 exercise trials (i.e., 1 trial ¼ 10 s contraction + 10 s rest) at 30% of their maximal voluntary contraction. Each condition lasted for 10 min and brain activity was compared across conditions as a means by which to identify the interactive effects of music and exercise on the BOLD response. Psychological measures were also taken immediately after each exercise bout. The results indicated that the combined effects of exercise and music elicited increased activation in the left inferior frontal gyrus (Brodmann area 47; see Fig. 4). Moreover, a negative correlation between activity of this region and exertional responses was identified. The authors hypothesized that the presence of asynchronous music has the potential to reduce processing of interoceptive cues (e.g., muscle afferent feedback) by increasing activation in the left inferior frontal gyrus. They also proposed that this region could potentially represent a hub of sensory integration where internal and external sensory cues are processed during the execution of movements. In such instances, an increase in the influence of an external stimulus (e.g., the intensity of music) could, potentially, prevent interoceptive signals from entering focal awareness and thus ameliorate negative bodily sensations, such as breathlessness and limb discomfort.
4 SHINING SOME (INFRA-RED) LIGHT ON THE UNDERLYING MECHANISMS Functional near-infrared spectroscopy (fNIRS) is a noninvasive imaging method developed to quantify chromophore concentration resolved from near-infrared light. fNIRS devices are usually attached to the participant’s scalp, where optodes (optical sensor devices) emit near-infrared light (Leo´n-Carrio´n and Leo´n-Domı´nguez, 2012). Light in the spectrum of 700–900 nm penetrates the soft tissues and bone and is absorbed by oxy- and deoxy-hemoglobin. Accordingly, this near-infrared range represents the perfect absorption spectrum for hemoglobin. Light receivers positioned 2–3 cm from the optodes absorb the light emitted by the optode and quantify
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FIG. 4 Significant group-level, random-effects activation of the left inferior frontal gyrus during exercise and music (contrast: exercise and music > exercise > music). Note: The blue blob represents the activity of the left inferior frontal gyrus; S ¼ superior; I ¼ inferior; R ¼ right; L ¼ left; A ¼ anterior. Reproduced from Bigliassi, M., Karageorghis, C.I., Bishop, D.T., Nowicky, A.V., Wright, M.J., 2018c. Cerebral effects of music during isometric exercise: an fMRI study. Int. J. Psychophysiol. 133, 131–139. https://doi.org/10.1016/j.ijpsycho.2018.07.475. Copyright 2018, with permission from Elsevier.
the concentration of hemoglobin. This technology allows researchers to investigate the changes in oxyhemoglobin in particular brain regions, given that such changes are indicative of neural activity. Since its first application some 40 years ago (J€obsis, 1977), near-infrared spectroscopy has been an effective tool for the monitoring of local changes in cerebral oxygenation and hemodynamics during brain activation (Byun et al., 2014; Yanagisawa et al., 2010). The sum of the concentrations of oxy- and deoxyhemoglobin provides a measure of the local cerebral tissue blood volume, whereas the individual concentrations of the two forms of hemoglobin are the result of the interplay between physiological parameters such as regional blood volume, regional blood flow, and metabolic rate of oxygen. Therefore, fNIRS is a tool that holds particular promise for exercise-related protocols in which there is a focus on environmental manipulations (e.g., auditory or visual stimuli). It provides useful information on exercise metabolism and facilitates the study of cerebral mechanisms with an acceptable degree of spatial and temporal resolution. Using fNIRS, Bigliassi et al. (2015b) reported prefrontal hemodynamic changes during passive listening to different styles of music. Specifically, classical music
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(Beethoven’s Symphony No. 6 [Pastoral Symphony]) was sufficiently potent to increase the activity of the right and left dorsolateral prefrontal cortices and upregulate the parasympathetic activity of the autonomous nervous system to a greater degree than techno music (Cyber Trip Techno Shock by Techno Magnetiko). Accordingly, fNIRS appears to be a valuable technique with which to study complex psychophysiological responses and the interrelationship between sensory stimulation, brain activity, and peripheral physiological responses. Of particular interest in applying fNIRS to physical exercise is the fact that it provides an indirect measure of local changes in cerebral oxygenation, which reflects cortical activation—with brain measures resistant to head and body artifacts during dynamic exercise (see Ekkekakis, 2009). fNIRS has been used extensively with newborn babies and infants, a particularly suitable population owing to the porous nature of the head, which allows infrared light to penetrate the tissue with minimal biological noise caused by bones and hair. Since the mid-2000s, many studies have been published reporting significant effects of visual (e.g., Karen et al., 2008), olfactory (e.g., Bartocci et al., 2000), and auditory stimuli (e.g., Kotilahti et al., 2010) on the hemodynamic responses of babies’ brains. For example, the question of brain laterality has been addressed with reference to the processing of music: Homae et al. (2012) reported a left-dominant activity for language processing but no clear lateralization for pitch, melody, and rhythm in auditory stimuli of a various nature (e.g., tone sequences). The initial motor paradigms used in fNIRS research entailed finger-tapping tasks. Sato et al. (2007) instructed participants to place their thumbs on the tips of each finger of the same hand in serial order as quickly and precisely as they could. As expected, results revealed an increase in oxyhemoglobin with a concomitant decrease in deoxyhemoglobin, suggesting an increase in the functional activity of motor, somatosensory, and frontal cortices. Using a button-pressing task, research has shown that the degree of neural brain activation was modulated as a function of the frequency at which participants moved their index finger (Kuboyama et al., 2005). Similarly, task complexity (e.g., simple/complex unimanual and bimanual tasks) has also been identified as a salient factor (Holper et al., 2009). Such studies illustrated similar brain hemodynamic activity patterns to those reported in fMRI studies (Huppert et al., 2006). More importantly, they demonstrated the possible use of fNIRS to gain better understanding of brain hemodynamics during motor activities that require substantial head movement. In the exercise context, fNIRS techniques have been used as a means by which to further understanding of the neurophysiological and psychobiological changes that accompany exercise. For example, Tempest et al. (2014) examined the functional capacity of the prefrontal cortex and explored the changes in affective response that occur during exercise performed at various intensities. The authors identified that oxy-, deoxy-, and total hemoglobin were greater in mostly ventral than dorsal regions of the prefrontal cortex during exercises performed above the ventilatory threshold. Interestingly, negative correlations were observed between affective responses and oxygenation in all regions of the left hemisphere. The results indicate
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that changes in prefrontal cortex activity might moderate the affective responses experienced during exercise performed at very high intensities. A similar exercise protocol was employed by Chang et al. (2013) who investigated the effects of music and exercise on cognitive performance and brain activity through the application of fNIRS. The authors identified a positive effect of exercise on cognitive performance as well as increased activation in the prefrontal cortex. Nonetheless, the undisclosed piece of music that was used in the study was not sufficiently potent to improve cognitive performance or influence hemodynamic activity in the prefrontal cortex. We would speculate that the absence of cognitive and neurophysiological responses reported by Chang et al. could be attributed to the music choice and/or the use of an intensity of 100 dBA, which is considered to be harmful from an audiological perspective (Lindgren and Axelsson, 1988). While retaining a reasonable degree of temporal-resolution, the use of multichannel devices is necessary in order to identify brain activity in the subcortical regions. Only a few fNIRS devices provide such measures (e.g., Shimadzu and Artinis models). One of the rare studies including multichannel measures for whole-body exercises was conducted by Miyai et al. (2001), who reported that hemodynamic responses of both the medial primary sensory-motor cortices and the supplementary motor areas were incremental: increasingly greater levels of brain co-activations were observed for motor imagery, feet-flexion, and treadmill-walking. The EEG findings presented earlier in this chapter strongly suggest that music has the potential to delay the subjective experience of fatigue during exercise. Karageorghis et al. (2017) proposed that the effects of music on affective responses, perception of exertion, and exercise endurance are mediated by corresponding changes in the activity of the dorsolateral prefrontal cortex (cf. Bigliassi et al., 2015a). Notably, Ekkekakis (2009) showed that oxygenation of this region increases at moderate exercise intensity but drops to below baseline levels shortly before an individual reaches voluntary exhaustion. Therefore, it seems plausible that, during exercise, the presence of music can delay the increase in oxygenation. This is because moderate-intensity exercise is experienced as being more pleasant or less unpleasant when compared to engaging in exercise without auditory stimulation (Karageorghis et al., 2017). Along similar lines, it is also plausible that music-listening during exercise serves to shift the entire oxygenation curve toward higher levels of intensity. This causes a delay of the eventual decline in prefrontal oxygenation and prolongs exercise performance (i.e., engenders an ergogenic effect). Testing of these two hypotheses—music delaying the increase in oxygenation and/or shifting of the oxygenation curve—will enable future researchers to establish the physiological “efficacy zone” of musicrelated interventions during exercise. Specifically, the range of exercise intensity over which music might be expected to facilitate the cognitive control of unpleasant sensations. Such sensations markedly increase in severity at high exercise intensities, and thus render music progressively ineffectual (Bigliassi et al., 2016; Karageorghis et al., 2009). The reduced availability of oxygen in the brain might also influence the neural activation of working muscle and is thus a precursor to a sharp degradation in exercise performance (e.g., Oussaidene et al., 2013).
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5 SUMMARY AND RECOMMENDATIONS The cerebral mechanisms that underlie the psychophysical, psychological, and psychophysiological effects of music on exercise appear to be associated with the rearrangement of the brain electrical frequencies and increased activation of the left frontal cortex. Reallocation of attentional focus toward environmental sensory cues prevents internal signals from entering focal awareness (Hutchinson and Karageorghis, 2013; Hutchinson et al., 2015; Rejeski, 1985). This psychophysiological mechanism appears to lead to a sequence of perceptual, affective, and psychophysiological responses that have been frequently reported in the literature (e.g., Clark et al., 2016; Karageorghis and Priest, 2012a,b). It has also been hypothesized that low-frequency waves increase in amplitude as a means by which to slow the body down (Craig et al., 2012) and force individuals toward task disengagement. Accordingly, upregulation of theta waves appears to heighten exercise consciousness, forcing individuals to make decisions as to whether they should continue with a given activity (Marcora, 2016). Interestingly, musicrelated interventions have the potential to partially inhibit the upregulation of theta waves, reduce processing of interoceptive signals, and enhance task performance (Bigliassi et al., 2016). We provided insight from studies that have examined the exercise-related hemodynamics and, on the basis of extant evidence, proposed two hypotheses that can be tested by future researchers: (a) music delays the increase in brain oxygenation and (b) music shifts the entire oxygenation curve toward higher levels of exercise intensity (Karageorghis et al., 2017). The series of mechanisms proposed in the EEG, fMRI, and fNIRS studies reviewed herein can be reliant upon exercise mode, complexity, and intensity. It is notable that relatively few exercise modes have been investigated to date—mainly simple, repetitive motor tasks, such as cycle ergometry—while complex forms of activity such as resistance training or calisthenics have thus far evaded researchers’ attention. Accordingly, there remains ample scope for further work to elucidate the brain mechanisms that underlie the effects of music during exercise. In particular, the mechanisms that underpin the recently-reported benefits of both respite and recuperative music represent investigative territory that is entirely uncharted (Jones et al., 2017; Karageorghis et al., 2019).
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Brain mechanisms that underlie music interventions in the exercise domain Costas I. Karageorghis*,1, Marcelo Bigliassi*, Segole`ne M.R. Guerin†, Yvonne Delevoye-Turrell† *Brunel University London, Uxbridge, United Kingdom † University of Lille, Lille, France 1 Corresponding author: Tel.: +44 01895 266476, e-mail address: [email protected]
Abstract In this chapter we review recent work from the realms of neuroscience and neuropsychology to explore the brain mechanisms that underlie the effects of music on exercise. We begin with an examination of the technique of electroencephalography (EEG), which has proven popular with researchers in this domain. We go on to appraise work conducted with the use of functional magnetic resonance imaging (fMRI) and then, looking more toward the future, we consider the application of functional near-infrared spectroscopy (fNIRS) to study brain hemodynamics. The experimental findings expounded herein indicate that music has the potential to guide attention toward environmental sensory cues and prevent internal, fatigue-related signals from entering focal awareness. The brain mechanisms underlying such effects are primarily associated with the downregulation of theta waves across the cortex surface, reduction of communication among somatosensory regions, and increased activation of the left inferior frontal gyrus. Taken holistically, research in this subfield of exercise psychology demonstrates a vibrant and reflexive matrix of attentional, emotional, behavioral, physiological, and psychophysiological responses to music across a variety of exercise modalities and intensities. The emergent hypotheses that we propose can be used to frame future research efforts.
Keywords Brain, Exercise, Music, Neuroscience, Psychophysiology
1 INTRODUCTION Music making has been a staple of human civilizations since time immemorial. Anthropologists have argued that music is part of our genetic blueprint and an activity that almost defines the human species (Mithen, 2006). The celebrated neuropsychologist, Daniel Levitin, indicated how music is unusual among all human Progress in Brain Research, ISSN 0079-6123, https://doi.org/10.1016/bs.pbr.2018.09.004 © 2018 Elsevier B.V. All rights reserved.
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activities for both its ubiquity and its antiquity. In the modern era, music has become an essential element of the exercise experience, albeit the music-in-exercise phenomenon was predicated on scant empirical research (see Karageorghis and Terry, 1997). Following 200 or so experimental studies in the period since the early 1990s, it is only very recently that researchers have grappled with neurophysiological or brain mechanisms that underlie the application of music in the exercise domain.
1.1 THE MAIN EFFECTS OF MUSIC DURING EXERCISE In the exercise and physical activity context, the most common effects reported for when people are exposed to music fall under the categories of psychological, psychophysical, psychophysiological, and ergogenic (Terry and Karageorghis, 2011). The psychological effects pertain to how music can influence mood, affect (i.e., core feelings of pleasure displeasure), emotion, cognition, and behavior. As a close cousin of psychological effects, psychophysical effects pertain to the psychological perception of one’s physical state. In exercise science, this is most often operationalized through the rating of perceived exertion (RPE; e.g., Jones et al., 2017; Stork and Martin Ginis, 2017). The mainstay of the literature has focused on psychological and psychophysical effects in response to music during continuous exercise (see Clark et al., 2016; Karageorghis, 2016 for reviews). Given the consistency of findings regarding, for example, enhancements in affect across a range of exercise intensities and the lowering of perceived exertion at low-to-moderate intensities (Hutchinson and Karageorghis, 2013; Taylor et al., 2007), the underlying cerebral mechanisms hold particular interest for the exercise science community. Psychophysiological effects pertain to how music can influence aspects of physiological functioning and thus common measures employed by researchers include heart rate, blood pressure, and oxygen uptake (Bacon et al., 2012; Karageorghis et al., 2019). This is also the category under which EEG falls, and, as we will discover later in this chapter, this is a technique that has enabled us to shine a light on salient cerebral mechanisms in the music–exercise relationship. Ergogenic effects are evident when music elevates work output or causes higher-than-expected power output, endurance, or productivity (Terry and Karageorghis, 2011). On occasion, researchers have used brain-monitoring techniques in tandem with electromyography to investigate how the reorganization of electrical activity in the brain has a bearing on muscular activity (e.g., Bigliassi et al., 2018a).
1.2 APPLICATIONS OF MUSIC IN EXERCISE In the exercise context, music is applied in three primary ways and, as we will go on to detail, these can follow on sequentially from one another. There is the pretask application wherein music is used to prime an exerciser or get them in the right frame of mind to be active (see Smirmaul, 2017 for a review). Given the imperative to “lead” an individual from a sedate state into an active state, such music will often be at a moderate tempo and have inspirational or motivational lyrics. For example,
ARTICLE IN PRESS 1 Introduction
Work From Home (100 bpm) by Fifth Harmony (feat. Ty Dolla $ign), which is an ideal precursor to a home workout. The in-task application of music has, by far and away, attracted the most research interest (see Karageorghis and Priest, 2012a,b). In this instance, music can be applied synchronously or asynchronously. The former entails a process of auditory-motor synchronization in which the music adopts a metronomic function to regulate movement and evenly distribute energy expenditure (see, e.g., Bacon et al., 2012; Karageorghis et al., 2009). The latter entails the absence of conscious synchronization and represents the way in which music is most commonly applied to individual exercise routines. For example, if an exerciser is engaged in a resistance training routine and music is piped into the gymnasium via a music-video channel, the strong likelihood is that the music will be used in the asynchronous mode. Hence, asynchronous music is akin to background or ambient music, although it can be played at a high intensity. In terms of the synchronous application of music, technological advancements have reduced the need for an exerciser to engage in conscious or active synchronization with a musical beat. Synchronization can be facilitated by means of an interface that uses accelerometry and algorithm-generated playlists to ensure coupling of musical rhythm with an individual’s work rate (see, e.g., Moens et al., 2014; D-Jogger). Such advancements have recently led to new definitions for the two main forms of auditory synchronization: (a) active synchronization entails a motor process in which an individual or group consciously synchronize their movement rate with the rhythmical qualities of music and (b) passive synchronization entails a motor process wherein a digital interface adapts the tempo of music in real-time or assigns a track at a tempo to match the movement rate of an individual or group (Karageorghis, 2019). The third main type of music application is post-task music (music used immediately after an exercise task) and similar to the synchronous application, two forms have recently been delineated. The first of these was coined respite music by Jones et al. (2017) and entails the use of music in between high-intensity intervals with a view to tempering the negative affective state that is typically elicited by such activity. Respite music also bears a positive influence on hemodynamic and cardiovascular recovery processes. The second is known as recuperative music (Terry and Karageorghis, 2011) and can be applied to both active (movement-based) and passive (static) forms of exercise recovery. Recuperative music playlists are typified by a progressively slower tempo (90 to 60 bpm), as the intention is to leave the exerciser with a sense of calmness and revitalization. Investigation of the brain mechanisms that underlie post-task music has yet to be initiated and thus provides tantalizing opportunities for future researchers. In this chapter, we will explore the brain mechanisms that underlie the use of in-task music in the exercise and physical activity domain. Some of the in-task mechanisms provide strong clues as to how music can take effect in the pretask phase (e.g., Bishop et al., 2014). We will begin with an examination of the technique of electroencephalography (EEG; i.e., measuring electrical activity in the brain), which
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has proven the most popular in this domain. Thereafter we will review work conducted with functional magnetic resonance imaging (fMRI)—a brain imaging technique that entails use of a scanner but is restricted to experimental tasks that involve no head movement. Finally, looking toward the future, and on the basis of a relatively small corpus of work, we have included a section on functional near-infrared spectroscopy (fNIRS) in which we present a hypothesis that could form the basis of future study. fNIRS is a newly-popularized technique that offers tremendous promise in terms of furthering our understanding of hemodynamic changes in the cortex that are caused by music during exercise.
2 EFFECTS OF MUSIC ON ELECTRICAL ACTIVITY IN THE BRAIN One of the first experiments to investigate the cerebral mechanisms underlying the effects of music on high-intensity exercise was conducted by Bigliassi et al. (2016). The authors employed an isometric, closed-kinetic chain exercise mode (an ankledorsiflexion task) and delivered the well-known track, Eye Of The Tiger by Survivor (109 bpm), in-task as a means by which to motivate participants. The exercise task was executed at 40% of each participant’s maximal voluntary contraction until the point of volitional exhaustion. Electrical activity in the brain was captured by means of a 64-channel EEG device, and a number of perceptual and affective responses were assessed. The results indicated that asynchronous music increased the use of dissociative thoughts and enhanced task performance to a greater degree than a no-music control condition. The authors also showed that music was sufficiently potent to downregulate theta waves (4–7 Hz) in the frontal, central, parietal, and occipital regions of the brain (see Fig. 1). This psychophysiological response appears to be associated with a mechanism pertaining to the suppression of fatigue-related symptoms (see Craig et al., 2012; Tanaka et al., 2012). Accordingly, Bigliassi et al. (2016) proposed a series of mechanisms in this study, suggesting that music-related interventions have the potential to reallocate attentional focus toward task-unrelated factors (e.g., exteroceptive signals), and ameliorate exertional responses to a greater degree than no-music conditions (Karageorghis et al., 2017). The upshot is that the processing of afferent feedback is reduced when individuals exercise with asynchronous music; a psychophysiological response that facilitates the execution of movements and optimizes the neural activation of working muscles (Bigliassi, 2015). Subsequently, the same group of researchers conducted a more complex experiment to identify the effects of music on motor unit recruitment and brain connectivity during whole-body exercise modes (Bigliassi et al., 2017). Participants exercised on a mechanically-braked cycle ergometer at light-to-moderate-intensities (i.e., at 10% below ventilatory threshold [VT]) for 12 min, while exposed to stimulative, asynchronous music. They also used another auditory stimulus—an audiobook—as a means by which to isolate the effects of musical components (e.g., melody and harmony) on perceptual, affective, and psychophysiological
ankle-dorsiflexion task. Psychophysiology 53, 1472–1483. https://doi.org/10.1111/psyp.12693. Copyright 2016, with permission from John Wiley & Sons.
Reprinted from Bigliassi, M., Karageorghis, C.I., Nowicky, A.V., Orgs, G., Wright, M.J., 2016. Cerebral mechanisms underlying the effects of music during a fatiguing isometric
Low-frequency components (theta waves) of the power spectrum presented for CO, MM, and MO. Note: The colored scale indicates the power of the band frequency (power [signal units^2/Hz*10^ {11}]); CO ¼ control condition; MM ¼ music condition; MO ¼ music-only condition.
FIG. 1
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responses. Time-frequency analyses were used to identify the synchronization– desynchronization–resynchronization cycle at the Cz electrode site. This site was chosen given that it provides a vista into the neural activity of lower limbs (i.e., the central area of the premotor cortex; Jain et al., 2013). The results of the Bigliassi et al. (2017) study indicate that music has the potential to inhibit alpha resynchronization and increase the number of motor units recruited to execute the task. The authors hypothesized that this psychophysiological mechanism could be representative of more autonomous control of working muscles. Put another way, asynchronous music appears to distract participants, guiding attention toward exteroceptive cues, and such distraction leads to a more efficient/autonomous neural activation. The results were supported by electromyographic (EMG) analysis, which indicated an increase in the number of motor units recruited in the quadriceps. Finally, the authors decided to run brain connectivity analysis to investigate whether the communication across somatosensory regions of the cortex was affected by the presence of auditory stimuli. Interestingly, music reduced the connectivity between Cz and both frontal (Fz) and central (C4) electrodes (see Fig. 2). The research team proposed that the reduction in communication could have been induced by a decrease in corollary discharges emitted from the central motor command to somatosensory regions of the cortex. This study demonstrated that music can lead to rearrangements of the electrical activity in the brain that are reliant upon exercise intensity, complexity, and mode. The results serve to illustrate that more robust methods such as timefrequency decomposition (e.g., wavelet transformations) and brain connectivity analysis (e.g., spectral coherence) might help to clarify underlying mechanisms that are not easily evident using traditional EEG procedures such as time-domain analysis (e.g., event-related potential). More recently, Bigliassi et al. (2018b) investigated the effects of asynchronous music on real-life physical activity. Participants were required to walk 400 m at a pace of their choosing on an outdoor running track. Electrical activity in the brain was recorded by use of a portable EEG device with active shielding technology. Such technology appears to function as a form of portable Faraday cage that protects the core components of cables against external artifacts during the execution of movements. The researchers also measured perceptual (state attention and perceived exertion) and affective (valence, arousal, and perceived enjoyment) outcomes immediately after each exercise bout. The results indicated that asynchronous music enhanced affective responses, increased the use of dissociative thoughts, and upregulated beta waves in the frontal and frontal-central regions of the brain to a greater degree than no-music control and podcast conditions (see Fig. 3). Based on previous findings (Bailey et al., 2008; Sayorwan et al., 2013), the researchers hypothesized that rearrangement of beta waves elicited by music serves to up/downregulate affective responses. Moreover, that this positive psychophysiological state could be capitalized upon during many forms of physical activity, as a means by which to render a given activity more pleasurable.
Physiol. Behav. 177, 135–147. https://doi.org/10.1016/j.physbeh.2017.04.023. Copyright 2017, with permission from Elsevier.
Reprinted from Bigliassi, M., Karageorghis, C.I., Wright, M.J., Orgs, G., Nowicky, A.V., 2017. Effects of auditory stimuli on electrical activity in the brain during cycle ergometry.
Group data alpha coherence values in CO (control condition), AB (audiobook condition), and MU (music condition). Reduced spectral coherence is manifest across central and frontal electrodes sites in MU when compared to CO and AB.
FIG. 2
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FIG. 3 Group data time-averaged band frequencies for CO, PO, and MU. Note: SMR, sensorimotor rhythm. The colored scale indicates the power of the band frequencies (signal^2/Hz*10^{10}); CO ¼ control condition; PO ¼ podcast condition; MU ¼ music condition. Reprinted from Bigliassi, M., Karageorghis, C.I., Hoy, G.K., Layne, G.S., 2018b. The Way You Make Me Feel: psychological and cerebral responses to music during real-life physical activity. Psychol. Sport Exerc. Advance online publication. https://doi.org/10.1016/j.psychsport.2018.01.010. Copyright 2018, with permission from Elsevier.
3 EFFECTS OF MUSIC ON SUBCORTICAL BRAIN REGIONS It is also necessary to examine the effects of music on exercise by use of high-spatial resolution techniques, which allow researchers to actually “see” the brain regions that activate in response to music and exercise; one such technique is functional magnetic resonance imaging (fMRI). This is oft-used in the field of neuroscience and cognitive psychology to detect changes in oxygenated hemoglobin that are coupled with neuronal activation. An fMRI scanner quantifies the energy emitted by protons and creates brain images based on the chemical structures of brain tissue. The scanner can also identify inhomogeneities in the brain that are caused by the deoxygenated/oxygenated blood ratio. This biophysical response is used to identify the speed of decay of the MRI signal, representative of neural activity, and referred to as the blood oxygenation level dependent (BOLD) response. Bishop et al. (2014) conducted one of the initial studies with fMRI and pretask music, albeit this was oriented more toward sport-related tasks. The researchers manipulated tempo and intensity of music (dBA) in order to identify changes in choice reaction-time. The hypotheses were predicated on the assumption that shorter reaction times occur at an optimal arousal state, and fast-tempo music was expected to
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upregulate arousal with corollary changes in brain activity. This study illustrates the nexus between musical components and neural systems, such as attention and motor control. The results supported the authors’ predictions that fast-tempo music would increase visuomotor activity through the elicitation of emotional responses. The subcallosal gyrus was significantly activated during the reaction-time task that followed fast-tempo, pretask music. Hence, fast-tempo music presumably elicited greater levels of arousal, which modulated reaction time by activating indirect pathways. In this case, arousal dictated participants’ level of readiness and voluntary/ involuntary attention. The only fMRI study designed to investigate the effects of in-task music on exercise was very recently conducted by Bigliassi et al. (2018c). The authors decided to use a simple exercise mode (handgrip) to investigate the brain regions that activate in response to exercise coupled with asynchronous music. The squeezing task was adopted in order to prevent head movements, which can easily compromise the fidelity of a scan. Participants were required to execute 30 exercise trials (i.e., 1 trial ¼ 10 s contraction + 10 s rest) at 30% of their maximal voluntary contraction. Each condition lasted for 10 min and brain activity was compared across conditions as a means by which to identify the interactive effects of music and exercise on the BOLD response. Psychological measures were also taken immediately after each exercise bout. The results indicated that the combined effects of exercise and music elicited increased activation in the left inferior frontal gyrus (Brodmann area 47; see Fig. 4). Moreover, a negative correlation between activity of this region and exertional responses was identified. The authors hypothesized that the presence of asynchronous music has the potential to reduce processing of interoceptive cues (e.g., muscle afferent feedback) by increasing activation in the left inferior frontal gyrus. They also proposed that this region could potentially represent a hub of sensory integration where internal and external sensory cues are processed during the execution of movements. In such instances, an increase in the influence of an external stimulus (e.g., the intensity of music) could, potentially, prevent interoceptive signals from entering focal awareness and thus ameliorate negative bodily sensations, such as breathlessness and limb discomfort.
4 SHINING SOME (INFRA-RED) LIGHT ON THE UNDERLYING MECHANISMS Functional near-infrared spectroscopy (fNIRS) is a noninvasive imaging method developed to quantify chromophore concentration resolved from near-infrared light. fNIRS devices are usually attached to the participant’s scalp, where optodes (optical sensor devices) emit near-infrared light (Leo´n-Carrio´n and Leo´n-Domı´nguez, 2012). Light in the spectrum of 700–900 nm penetrates the soft tissues and bone and is absorbed by oxy- and deoxy-hemoglobin. Accordingly, this near-infrared range represents the perfect absorption spectrum for hemoglobin. Light receivers positioned 2–3 cm from the optodes absorb the light emitted by the optode and quantify
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FIG. 4 Significant group-level, random-effects activation of the left inferior frontal gyrus during exercise and music (contrast: exercise and music > exercise > music). Note: The blue blob represents the activity of the left inferior frontal gyrus; S ¼ superior; I ¼ inferior; R ¼ right; L ¼ left; A ¼ anterior. Reproduced from Bigliassi, M., Karageorghis, C.I., Bishop, D.T., Nowicky, A.V., Wright, M.J., 2018c. Cerebral effects of music during isometric exercise: an fMRI study. Int. J. Psychophysiol. 133, 131–139. https://doi.org/10.1016/j.ijpsycho.2018.07.475. Copyright 2018, with permission from Elsevier.
the concentration of hemoglobin. This technology allows researchers to investigate the changes in oxyhemoglobin in particular brain regions, given that such changes are indicative of neural activity. Since its first application some 40 years ago (J€obsis, 1977), near-infrared spectroscopy has been an effective tool for the monitoring of local changes in cerebral oxygenation and hemodynamics during brain activation (Byun et al., 2014; Yanagisawa et al., 2010). The sum of the concentrations of oxy- and deoxyhemoglobin provides a measure of the local cerebral tissue blood volume, whereas the individual concentrations of the two forms of hemoglobin are the result of the interplay between physiological parameters such as regional blood volume, regional blood flow, and metabolic rate of oxygen. Therefore, fNIRS is a tool that holds particular promise for exercise-related protocols in which there is a focus on environmental manipulations (e.g., auditory or visual stimuli). It provides useful information on exercise metabolism and facilitates the study of cerebral mechanisms with an acceptable degree of spatial and temporal resolution. Using fNIRS, Bigliassi et al. (2015b) reported prefrontal hemodynamic changes during passive listening to different styles of music. Specifically, classical music
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(Beethoven’s Symphony No. 6 [Pastoral Symphony]) was sufficiently potent to increase the activity of the right and left dorsolateral prefrontal cortices and upregulate the parasympathetic activity of the autonomous nervous system to a greater degree than techno music (Cyber Trip Techno Shock by Techno Magnetiko). Accordingly, fNIRS appears to be a valuable technique with which to study complex psychophysiological responses and the interrelationship between sensory stimulation, brain activity, and peripheral physiological responses. Of particular interest in applying fNIRS to physical exercise is the fact that it provides an indirect measure of local changes in cerebral oxygenation, which reflects cortical activation—with brain measures resistant to head and body artifacts during dynamic exercise (see Ekkekakis, 2009). fNIRS has been used extensively with newborn babies and infants, a particularly suitable population owing to the porous nature of the head, which allows infrared light to penetrate the tissue with minimal biological noise caused by bones and hair. Since the mid-2000s, many studies have been published reporting significant effects of visual (e.g., Karen et al., 2008), olfactory (e.g., Bartocci et al., 2000), and auditory stimuli (e.g., Kotilahti et al., 2010) on the hemodynamic responses of babies’ brains. For example, the question of brain laterality has been addressed with reference to the processing of music: Homae et al. (2012) reported a left-dominant activity for language processing but no clear lateralization for pitch, melody, and rhythm in auditory stimuli of a various nature (e.g., tone sequences). The initial motor paradigms used in fNIRS research entailed finger-tapping tasks. Sato et al. (2007) instructed participants to place their thumbs on the tips of each finger of the same hand in serial order as quickly and precisely as they could. As expected, results revealed an increase in oxyhemoglobin with a concomitant decrease in deoxyhemoglobin, suggesting an increase in the functional activity of motor, somatosensory, and frontal cortices. Using a button-pressing task, research has shown that the degree of neural brain activation was modulated as a function of the frequency at which participants moved their index finger (Kuboyama et al., 2005). Similarly, task complexity (e.g., simple/complex unimanual and bimanual tasks) has also been identified as a salient factor (Holper et al., 2009). Such studies illustrated similar brain hemodynamic activity patterns to those reported in fMRI studies (Huppert et al., 2006). More importantly, they demonstrated the possible use of fNIRS to gain better understanding of brain hemodynamics during motor activities that require substantial head movement. In the exercise context, fNIRS techniques have been used as a means by which to further understanding of the neurophysiological and psychobiological changes that accompany exercise. For example, Tempest et al. (2014) examined the functional capacity of the prefrontal cortex and explored the changes in affective response that occur during exercise performed at various intensities. The authors identified that oxy-, deoxy-, and total hemoglobin were greater in mostly ventral than dorsal regions of the prefrontal cortex during exercises performed above the ventilatory threshold. Interestingly, negative correlations were observed between affective responses and oxygenation in all regions of the left hemisphere. The results indicate
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that changes in prefrontal cortex activity might moderate the affective responses experienced during exercise performed at very high intensities. A similar exercise protocol was employed by Chang et al. (2013) who investigated the effects of music and exercise on cognitive performance and brain activity through the application of fNIRS. The authors identified a positive effect of exercise on cognitive performance as well as increased activation in the prefrontal cortex. Nonetheless, the undisclosed piece of music that was used in the study was not sufficiently potent to improve cognitive performance or influence hemodynamic activity in the prefrontal cortex. We would speculate that the absence of cognitive and neurophysiological responses reported by Chang et al. could be attributed to the music choice and/or the use of an intensity of 100 dBA, which is considered to be harmful from an audiological perspective (Lindgren and Axelsson, 1988). While retaining a reasonable degree of temporal-resolution, the use of multichannel devices is necessary in order to identify brain activity in the subcortical regions. Only a few fNIRS devices provide such measures (e.g., Shimadzu and Artinis models). One of the rare studies including multichannel measures for whole-body exercises was conducted by Miyai et al. (2001), who reported that hemodynamic responses of both the medial primary sensory-motor cortices and the supplementary motor areas were incremental: increasingly greater levels of brain co-activations were observed for motor imagery, feet-flexion, and treadmill-walking. The EEG findings presented earlier in this chapter strongly suggest that music has the potential to delay the subjective experience of fatigue during exercise. Karageorghis et al. (2017) proposed that the effects of music on affective responses, perception of exertion, and exercise endurance are mediated by corresponding changes in the activity of the dorsolateral prefrontal cortex (cf. Bigliassi et al., 2015a). Notably, Ekkekakis (2009) showed that oxygenation of this region increases at moderate exercise intensity but drops to below baseline levels shortly before an individual reaches voluntary exhaustion. Therefore, it seems plausible that, during exercise, the presence of music can delay the increase in oxygenation. This is because moderate-intensity exercise is experienced as being more pleasant or less unpleasant when compared to engaging in exercise without auditory stimulation (Karageorghis et al., 2017). Along similar lines, it is also plausible that music-listening during exercise serves to shift the entire oxygenation curve toward higher levels of intensity. This causes a delay of the eventual decline in prefrontal oxygenation and prolongs exercise performance (i.e., engenders an ergogenic effect). Testing of these two hypotheses—music delaying the increase in oxygenation and/or shifting of the oxygenation curve—will enable future researchers to establish the physiological “efficacy zone” of musicrelated interventions during exercise. Specifically, the range of exercise intensity over which music might be expected to facilitate the cognitive control of unpleasant sensations. Such sensations markedly increase in severity at high exercise intensities, and thus render music progressively ineffectual (Bigliassi et al., 2016; Karageorghis et al., 2009). The reduced availability of oxygen in the brain might also influence the neural activation of working muscle and is thus a precursor to a sharp degradation in exercise performance (e.g., Oussaidene et al., 2013).
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5 SUMMARY AND RECOMMENDATIONS The cerebral mechanisms that underlie the psychophysical, psychological, and psychophysiological effects of music on exercise appear to be associated with the rearrangement of the brain electrical frequencies and increased activation of the left frontal cortex. Reallocation of attentional focus toward environmental sensory cues prevents internal signals from entering focal awareness (Hutchinson and Karageorghis, 2013; Hutchinson et al., 2015; Rejeski, 1985). This psychophysiological mechanism appears to lead to a sequence of perceptual, affective, and psychophysiological responses that have been frequently reported in the literature (e.g., Clark et al., 2016; Karageorghis and Priest, 2012a,b). It has also been hypothesized that low-frequency waves increase in amplitude as a means by which to slow the body down (Craig et al., 2012) and force individuals toward task disengagement. Accordingly, upregulation of theta waves appears to heighten exercise consciousness, forcing individuals to make decisions as to whether they should continue with a given activity (Marcora, 2016). Interestingly, musicrelated interventions have the potential to partially inhibit the upregulation of theta waves, reduce processing of interoceptive signals, and enhance task performance (Bigliassi et al., 2016). We provided insight from studies that have examined the exercise-related hemodynamics and, on the basis of extant evidence, proposed two hypotheses that can be tested by future researchers: (a) music delays the increase in brain oxygenation and (b) music shifts the entire oxygenation curve toward higher levels of exercise intensity (Karageorghis et al., 2017). The series of mechanisms proposed in the EEG, fMRI, and fNIRS studies reviewed herein can be reliant upon exercise mode, complexity, and intensity. It is notable that relatively few exercise modes have been investigated to date—mainly simple, repetitive motor tasks, such as cycle ergometry—while complex forms of activity such as resistance training or calisthenics have thus far evaded researchers’ attention. Accordingly, there remains ample scope for further work to elucidate the brain mechanisms that underlie the effects of music during exercise. In particular, the mechanisms that underpin the recently-reported benefits of both respite and recuperative music represent investigative territory that is entirely uncharted (Jones et al., 2017; Karageorghis et al., 2019).
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