CHAPTER

Reflections on music, affect,

and sociality

Matthew Sachs, Assal Habibi, Hanna Damasio1

8
Brain and Creativity Institute, University of Southern California, Los Angeles, CA, United States
1
Corresponding author: Tel.: +1-213-740-3462, e-mail address: [email protected]

Abstract
Music is an important facet of and practice in human cultures, significantly related to its
capacity to induce a range of intense and complex emotions. Studying the psychological
and neurophysiological responses to music allows us to examine and uncover the neural mech-
anisms underlying the emotional impact of music. We provide an overview of different aspects
of current research on how music listening produces emotions and the corresponding feelings,
and consider the underlying neurophysiological mechanisms. We conclude with evidence to
suggest that musical training may influence the ability to recognize the emotions of others.

Keywords
Music, Affect, Pleasure, Functional magnetic resonance imaging, Diffusion tensor imaging

1 INTRODUCTION
Music is certainly but so far inexplicably tied to pleasure. While enjoyment associ-
ated with the procurement of food, sex, social interactions, and monetary goods can
be explained because they directly promote homeostasis, music appears to be intrin-
sically rewarding without serving a clear physiological purpose. Music does not di-
rectly nourish our bodies. It is not a tangible commodity. Throughout history, there
have never been threats of music shortage, nor cases of persons becoming addicted or
overdosing on music. And yet, the enjoyment of music making and music listening
has remained an integral aspect of human cultures across time and place.
Because of its special status, music has become a tool in the study of human emo-
tions and feelings. Indeed, many scholars contend that the rewarding aspects of
music derive from its emotive power. Music can make us smile and move us to tears,
excite us and calm us, and connect us to our feelings and to the feelings of others.
This ability is all the more baffling when one considers that music is in essence
abstract, an art form without an objective, agreed upon meaning.
Progress in Brain Research, Volume 237, ISSN 0079-6123, https://doi.org/10.1016/bs.pbr.2018.03.009
© 2018 Elsevier B.V. All rights reserved.
153
154 CHAPTER 8 Reflections on music, affect, and sociality

In this chapter, we discuss findings from psychological and neuroimaging studies

that attempt to describe the neurobiological mechanisms that may support music’s
extraordinary hold over us. We begin with a review of the literature on the neural
responses associated with music-evoked emotions, highlighting some of our own
work in which we used advanced analytical techniques of neuroimaging data to
elucidate the nature of affective experiences caused by music listening. We then
explore cases in which music induces intense emotional responses, pleasurable or
not, underlining how such extremes further our understanding of neural structures
involved in musical experiences. We conclude with a discussion of musical training,
addressing how the development of musical skills through the process of learning to
play an instrument may influence the ability to process emotions as well as the ability
to understand, connect with, and bond with others.

2 NEUROBIOLOGY OF MUSIC-EVOKED EMOTIONS

2.1 EMOTIONS AND FEELINGS
To begin the discussion of how music is able to co-opt human affective processes, it is
useful to summarize what is known about affect in more general terms. Emotions are
often conceptualized as physiological action programs triggered by changes in the
internal or external environment, and executed by a partnership of brain and body.
The changes that result from the action programs may serve to draw attention to a par-
ticular aspect of the environment that is relevant for survival or to motivate a behavior
that promotes or executes homeostatic regulation (Phillips et al., 2003). Emotions are
therefore adaptive and evolutionary and are significant because they aim at optimal life
regulation (Damasio, 1999). It is generally accepted that several “basic” emotions,
those that are universal and utilitarian, are seen across many species. These basic emo-
tions include happiness, sadness, disgust, fear, surprise, and anger (Ekman, 1992).
Each category is associated with qualitatively distinct bodily state changes which serve
unique functions in terms of regulating homeostasis (James, 1890).
In addition to emotions, many species also have emotional feelings, phenomena
that specifically refer to the mental experience of the bodily state changes associ-
ated with the emotion (Damasio and Carvalho, 2013). Feelings serve to mark a par-
ticular situation or circumstance in a way that signifies whether it should, in future
occasions, be pursued or avoided (Berridge and Kringelbach, 2008). Situations
marked by pleasant feelings, often labeled as having a “positive valence,” are to
be approached in the future. Situations marked by unpleasant feelings, having a
“negative valence,” are to be avoided. We can experience a range of feelings, which
lie somewhere on a valence spectrum, from negative to positive. Emotions and feel-
ings can also be categorized along a spectrum corresponding to how activating or
energizing they are (Russell, 1980). Often termed arousal, this component is what
separates the positively valenced emotion of joy from another positively valenced
emotion such as peacefulness, for example.
 2 Neurobiology of music-evoked emotions 155

Capitalizing on the spatial resolution that functional magnetic resonance imaging

(fMRI) provides, neuroscientists have been using an array of affective stimuli, includ-
ing pictures, faces, smells, sounds, films, music, and autobiographical memories, to
induce discrete emotional states, while measuring neural activity. The available results
suggest that emotional experiences of varying degrees of valence and arousal involve
the cooperative functioning of a distributed network of brain regions associated with
basic psychological processes such as reward processing, salience detection, learning,
and memory (Lindquist et al., 2012). There is no one-to-one match between the func-
tion of a singular brain region and a particular emotional state. Instead, emotional
experience engages brain networks such as the limbic system—including the amyg-
dala, ventral striatum (including the nucleus accumbens), anterior cingulate cortex
(ACC), thalamus, hypothalamus, and hippocampus—as well as the related paralimbic
system—including the insula, orbitofrontal cortex (OFC), and medial prefrontal cortex
(MPFC; Kober et al., 2008).

2.2 BRAIN REGIONS INVOLVED IN EMOTIONAL RESPONSES TO MUSIC

How does music convey emotions and how does the human brain process them? It
has been shown that basic emotions, e.g., happiness, sadness, fear, and anger, can be
detected aurally through the acoustical aspects and structure of music (Balkwill and
Thompson, 1999; Fritz et al., 2009). Music, additionally, has the ability to induce
those emotions and respective feelings in the listener, as well as a wide range of
more complex feelings that are rarely experienced outside of an aesthetic context,
such as transcendence, wonder, and nostalgia (Zentner et al., 2008). There is evi-
dence that the same psychophysiological (Lundqvist et al., 2008) and cognitive
changes (Vuoskoski and Eerola, 2012) associated with basic, everyday emotions
have also been found in association with music-evoked emotions, such as changes
in facial muscular activity, changes to internal body temperature, responses in the
sympathetic nervous system, and biases in perception and judgment.
Similarly, results from both functional neuroimaging and lesion studies suggest
that the same brain networks collectively involved in processing everyday basic emo-
tions are also involved in processing emotions conveyed through music (Habibi and
Damasio, 2014). Music selected to evoke a wide array of emotions (happiness, sad-
ness, joy, fear, or pleasantness/unpleasantness) engages regions in the paralimbic
and limbic systems, such as the ventral striatum, insula, ACC, parahippocampal
gyrus, hippocampus, amygdala, and OFC (Brattico et al., 2011; Koelsch, 2014;
Mitterschiffthaler et al., 2007).
The exact role that each of these regions plays in processing emotions in music is
not entirely clear, but in conjunction with findings from neuroimaging studies on
emotional responses to more basic stimuli, it is possible to advance some hypotheses.
The amygdala, hippocampus, and parahippocampal gyrus, three closely related brain
regions located in the medial temporal lobe, may be collectively involved in inducing
early emotional reactions to sounds, specifically those related to fear and surprise
uhholz et al., 2014). The amygdala is believed to trigger positive and negative
(Fr€
156 CHAPTER 8 Reflections on music, affect, and sociality

reactions to potentially salient stimuli and to induce the appropriate approach or

avoidance behaviors (Lindquist et al., 2012). The hippocampus is primarily known
for its role in learning and memory and therefore is an ideal structure to be involved
in connecting past experiences and previous associations with the musical stimuli
with current behavioral responses to music (Juslin, 2013). The parahippocampal gy-
rus may be particularly associated with processing music with negative valence,
given that patients with damage in either the left or right region exhibited reduced
responses to dissonant music (Gosselin et al., 2006).
The insula and ACC, two brain regions that are functionally and structurally con-
nected (Medford and Critchley, 2010), appear to be involved in numerous cognitive
processes. Regarding affect, these regions are likely responsible for mapping the
body-state changes that constitute emotional responses (Craig, 2009) and are the
basis for the respective feelings (Damasio et al., 2000). Both the insula and ACC
were found to be activated in functional neuroimaging studies in which participants
listened to pleasant musical stimuli, rather than to neutral musical stimuli (Blood and
Zatorre, 2001; Brown et al., 2004; Trost et al., 2012).
The ventral striatum (nucleus accumbens) and the OFC are part of the reward
circuitry of the brain (Leknes and Tracey, 2008). Both regions have been found to
be activated in conjunction with feelings of liking (Brattico et al., 2013) and are also
activated in response to intensely pleasurable musical stimuli. The OFC has been
shown to be involved in numerous decision-making tasks and, therefore, seems to
be a good candidate for initiating the process of making aesthetic judgments about
beauty (Brown et al., 2011). The nucleus accumbens is believed to be the central site
involved in both the anticipation and consummation of a reward (Berridge and
Kringelbach, 2008). Its activation is also correlated with peak pleasurable experi-
ences with music (Salimpoor et al., 2013).
What we provided so far is a brief overview of brain regions associated with
emotional experiences caused by music. We believe these regions form a complex
network that is involved in numerous aspects of the emotional experiences to music,
and they certainly are not to be seen as “centers” for that experience.

2.3 ADVANTAGES OF NEWER ANALYTICAL TECHNIQUES

Machine learning algorithms, used primarily in the field of computer science, have
more recently been applied to the analysis of neuroimaging data. The goal with such
methods is to identify patterns within the recorded fMRI signal and relate these pat-
terns to the cognitive states being experienced by the participant. Termed multivoxel
pattern analysis (MVPA; Norman et al., 2006), this method evaluates the fMRI signal
recorded from multiple voxels at once, which can provide a more nuanced picture of
how information is spatially encoded/distributed across the brain. By classifying
cognitive states based on the spatially distributed patterns of neural signal associated
with them, this type of analysis does not identify brain regions that are “activated” by
a stimulus, but, rather, brain regions that carry key information regarding some iden-
tifying property of that stimulus (Mitchell et al., 2004).
 2 Neurobiology of music-evoked emotions 157

MVPA has been used to classify emotional states induced by music based on
patterns of fMRI data. Kragel and LaBar (2015) found that distinct patterns of activ-
ity within several cortical and subcortical brain regions could predict one of seven
discrete emotional states induced by music. These included the precuneus, cingu-
late, insula, thalamus, amygdala, and prefrontal cortex. Another study using music
with either positive or negative valence corroborated those findings, reporting that
patterns of activity within the precuneus, cingulate, thalamus, and prefrontal cor-
tex could successfully predict the valence of the musical piece that was presented
(Kim et al., 2017).
Recently, in our own work (Sachs, Habibi, Damasio, & Kaplan, 2018), we
showed that emotional categories of musical sounds expressing happiness, sadness,
and fear, produced by a violin and clarinet, could be reliably distinguished in fMRI
data, based on the patterns of the neural signal detected in bilateral auditory cortices,
insulae, parietal opercula, post- and precentral gyri, inferior frontal gyri, and right
medial prefrontal cortex. Neural patterns within the primary and secondary auditory
cortices, insulae, and parietal opercula could also reliably distinguish the same three
emotions when training the classifier on data using a different set of nonmusical,
nonverbal vocalizations, e.g., the sound of a person crying, screaming, or laughing
(see Belin et al., 2008), suggesting that these patterns of activity are not unique to the
processing of emotions conveyed through musical instruments. In combination, the
MVPA results appear to support the conclusions drawn from fMRI studies with uni-
variate data, namely that feelings induced by music evoke responses similar to those
associated with feelings induced by other stimuli (Koelsch, 2014).
In addition to MVPA, other model-free methods have recently been employed
with neuroimaging to assess brain responses to more naturalistic, ecologically valid
stimuli. One such approach involves recording neural signal continuously (using ei-
ther EEG or fMRI) during the presentation of a full-length piece of music. This type
of approach does not rely on a predefined model to evaluate the changes in neural
signal. Instead, it involves calculating correlations between the neural signal in dif-
ferent brain regions or between the neural signal in the same brain region across dif-
ferent participants. Changes in these correlations over time can then be related to
specific changes in the music. In this way, the fluctuating patterns of brain syn-
chrony can be linked to an emotional experience, shared among different subjects,
in response to the music presented. Studies using such an technique have shown that
continuous ratings of valence and arousal while listening to a music piece were
associated with synchronized activity in the amygdala, insula, ACC, and caudate
nucleus across participants (Trost et al., 2014). Furthermore, within-network ana-
lyses showed that synchrony of activity within regions of the limbic system was
correlated with changes in valence ratings during a listening period (Singer
et al., 2016). While these methods have only been used with musical stimuli re-
cently and sparsely, the early findings provide further support for the hypothesis
that music-evoked feelings involve the continuous interaction of multiple brain re-
gions over time, highlighting the importance of capturing the temporal parameters
of emotional responses to music.
158 CHAPTER 8 Reflections on music, affect, and sociality

2.4 MECHANISMS OF MUSIC-EVOKED EMOTIONS

Based on self-reports, psychophysiological evidence, and neuroimaging results, we
have established that music can induce a range of emotional states. But what are the
actual mechanisms which allow music to do so? Some theorize that it is the detection
of certain acoustical features in music, which have come to be associated with spe-
cific emotions, that actually trigger an affective response, e.g., faster tempos, major
tonalities, increased volume, higher fundamental frequencies, and faster tonal at-
tacks tend to characterize happy music (Juslin and Laukka, 2003). But it is difficult
to accept that acoustic properties of music alone could account for the range and in-
tensity of possible feelings. Often, the emotions evoked by music are neither discrete
nor immutable; rather, they are multifaceted and mercurial, continually influenced
by one’s auditory abilities, previous experiences, personality, and current mood.
In an effort to systematize the breadth of complex factors at play, Juslin (2013)
proposed a list of eight mechanisms by which music is able to induce an emotion,
each characterized by a unique set of brain regions and networks. These mechanisms
can work independently or in tandem, which may result in multiple distinct feelings
in response to the same piece of music. One way that emotions can be induced by
music, according to the model, is through the process of evaluative conditioning, i.e.,
the pairing of a stimulus with an affective marker. This can occur when the sounds of
a given piece of music trigger a memory that, through previous experience, was as-
sociated with a certain emotion. For example, a piece of music played repeatedly at a
joyous occasion can eventually trigger joy, even when removed from that event. Or,
as described earlier, it could also be that the sounds in and of themselves are asso-
ciated with positive or negative feelings because, over time, and through the process
of learning, these sounds have acquired the affective significance of the objects or
events that typically produced them (Damasio, 2018). The sound of the blaring trum-
pet, for example, may induce unpleasant feelings because these acoustic properties
commonly accompany fearful or distressing events. This process of pairing sounds
with affective labels likely involves the amygdala. On the other hand, an emotion to
the same piece of music could be induced through a separate mechanism, such as that
of assessing the aesthetic qualities of the piece, in which case it is likely that other
brain regions in the prefrontal cortex might be involved. This is not the place to
review in detail all eight mechanisms proposed by Juslin; but we would like to
acknowledge that, while further research is needed to corroborate the hypotheses
presented in Juslin’s model, the framework provides a systematized way to under-
stand and evaluate the vast array of emotional experiences and neural activity that
we see associated with listening to music.

3 INTENSE MUSICAL PLEASURE AND ITS ABSENCE

One aspect of music listening that warrants its own discussion is its strong connection
to feelings of pleasure. Engaging with music is often reported as a highly enjoyable
activity in many people’s lives and there is empirical evidence to suggest that such
 3 Intense musical pleasure and its absence 159

engagement can trigger the same biological and psychological responses associated
with other highly rewarding, and yet transparently valuable stimuli, such as food,
sex, and money (Salimpoor et al., 2013). Psychophysiological, highly pleasurable
experiences with music were associated with increases in heart rate and electroder-
mal activity (Salimpoor et al., 2009). Regions in the brain involved in hedonic
responses and reward, such as the ventral striatum, insula, ACC, and OFC, are
activated when listening to pleasant or self-selected pleasurable music (Brown
et al., 2004; Koelsch, 2014; Menon and Levitin, 2005; Trost et al., 2012). Activity
in the ventral striatum in particular was shown to be positively correlated with sub-
jective ratings of how much a person liked a piece of music (Salimpoor et al., 2013)
and activity in the insula and amygdala was shown to be correlated with physiolog-
ical measures of arousal during music listening (Blood and Zatorre, 2001).
How do feelings of pleasure relate to the previously discussed emotional
responses to music? Neuroimaging and behavioral evidence suggest that feelings
of pleasure and conscious enjoyment of a piece of music emerge later in time,
after the initial emotional response. In a recent report, Brattico et al. (2013) drew
on the findings from previous fMRI and EEG studies to attempt to outline the tem-
poral order of the neural events involved in pleasurable responses to music. These
authors hypothesized that after the initial processing of the acoustical properties
of the sounds in the brainstem, and subsequently the auditory cortex, the recognition
of the valence and arousal level of the music is processed in association with acti-
vation of the amygdala, hippocampus, and parahippocampal gyrus. Subsequently,
feelings associated with valence and arousal emerge in association with activity
in the insula and ACC. The aesthetic judgment of beauty occurs in association with
activity in the OFC (Brown et al., 2011), which may or may not result in feelings of
pleasure and enjoyment. If the product of all of these complex processes does result
in a pleasurable response, Brattico and colleagues assume that the ventral striatum
and nucleus accumbens are most likely involved in such experiences (Brattico
et al., 2013).

3.1 EXPERIENCE OF CHILLS AND OTHER “STRONG EXPERIENCES

OF MUSIC”
While many people find music pleasurable, there are certain sensations associated
with music listening that go beyond the everyday emotional experience in terms
of intensity and are rarely felt outside of an aesthetic domain. These “strong expe-
riences of music,” sometimes referred to as “aesthetic emotions,” include sensations
such as awe, feeling of a lump in the throat, feeling moved, and experiencing chills
(Lamont, 2011). Chills are one of the most widely documented strong emotional
experiences to music. While the nature of the sensation can vary markedly across
individuals, chills are most frequently reported as “shivers down the spine,” or scalp,
or back of the neck are often accompanied by “goose bumps” and “hair standing on
end”, and tend to be associated with moments of peak pleasure and enjoyment
(Nusbaum et al., 2014). A variety of different terms have been used throughout
160 CHAPTER 8 Reflections on music, affect, and sociality

the literature to describe this phenomenon when it occurs in response to the arts,
including “aesthetic chills,” “thrills,” and “frisson” (Nusbaum and Silvia, 2011).
Why we would experience sensations typically associated with coldness or fear
in response to music, a stimulus that poses no clear threat to our well-being, remains
largely unclear. However, recent explorations, in particular several neuroimaging
studies, have begun to provide some answers as to how such a link may have
emerged.
Research on the frequency and likelihood of experiencing aesthetic chills has gar-
nered several possible explanations for their existence. Early findings that music
expressing sadness is more likely to induce chills in women lead researchers to pos-
tulate that the sensation was the product of the music’s resemblance to separation
calls, sounds emitted from offspring that inform their parents of their whereabouts
(Panksepp, 1995). Such sounds are designed to induce deep, biological responses
that encourage social bonding behaviors and thus motivate a parent to find and pro-
tect her progeny (Panksepp and Bernatzky, 2002). This explanation, however, does
not account for chills that occur in response to other modalities, such as to nonmu-
sical works of art, and subsequent studies have failed to replicate these gender dif-
ferences (Silvia and Nusbaum, 2011).
Huron (2006) proposed that chills are the body’s programmatic response to
stimuli that are surprising and therefore considered to be potentially threatening at
first, but that are later reappraised as nonthreatening. In this account, the pleasurable
feelings that accompany chills would derive from this mismatch between the initial
negative physiological response and its subsequent cognitive appraisal as some-
thing positive (Huron, 2006). Chills may also be the physiological by-product
of feelings of awe (Silvia et al., 2015), an intense emotional mixture of joy and fear
that arises when one experiences something unfathomably vast, novel, and/or
beautiful (Konečni, 2005). Music is a frequently reported inducer of feelings of
awe, suggesting that chills to music may in fact represent a deeply rooted emotional
reaction designed to help process startlingly novel information of any kind (Silvia
et al., 2015).
Intriguingly, not everyone experiences chills in response to music and the actual
rates of frequency vary wildly from study to study. In one particular sample, 90% of
participants reported experiencing chills in response to music (Sloboda, 1991),
whereas in a study in which participants listened and responded to musical stimuli
in a laboratory setting, less than 25% felt chills (Grewe et al., 2005). A more recent
study had participants reported on their activities, environment, and feelings in real
time at various moments throughout the day and found that participants experienced
chills 14% of the time while listening to music (Nusbaum et al., 2014).
The frequency and likelihood of experiencing chills to music, as well as several
other intense aesthetic experiences, have been shown to be associated with one of the
Big Five personality traits called openness to experience, a trait marked by intellectual
curiosity, and a tendency to examine one’s own feelings (McCrae, 2007). Knowledge
about, or engagement with, the arts and music is also found to be associated with the
experience of chills by the same authors. In a follow-up assessment, the authors
 3 Intense musical pleasure and its absence 161

suggested that one’s musical experience and the frequency of engagement with
music (e.g., listening to music, going to concerts, seeking out new bands) seemed
to mediate the interaction between personality and chills (Nusbaum and Silvia,
2011). No specific acoustical, harmonic, or lyrical features have been found to
be a direct trigger of chills; however, there are certain music structures and themes
that are commonly associated with the experience. People often report feeling chills
during passages that contain an entrance of a new voice or instrument, rapid or large
changes in tempo or dynamics, or unexpected harmonic shifts (Grewe et al., 2007;
Guhn et al., 2007). Individuals were also more likely to experience chills to pieces
that were more familiar; even so, the frequency of chills did not increase with an
increase in familiarity (Grewe et al., 2007). Taken together, these findings suggest
that the induction of chills in response to music is likely caused by an interaction
between a subject’s personality and the previous musical experience, mood at the
time of the experience, level of attention, and the structural features of the music in
question (Grewe et al., 2007).
Because chills can be highly variable and hard to define, recent attempts have
been made to identify more objective, neurophysiological markers that accompany
this experience. Increases in psychophysiological measures, including electrodermal
activity, heart, and respiration rates, have been observed during self-reported expe-
riences of chills in response to music stimuli (Grewe et al., 2009; Sachs et al., 2016;
Salimpoor et al., 2009). These increases generally indicate intensified feelings of
arousal (Fowles, 1980). Importantly, similar increases were not observed in those
subjects who did not report experiencing chills or pleasurable responses while listen-
ing to the same pieces of music, suggesting that the autonomic nervous system
response could not be solely accounted for by changes in acoustical elements of
the music, such as increases in volume or tempo (Salimpoor et al., 2009).
Neuroimaging techniques have also been used to further our understanding of
chills induced by music. Several fMRI studies have shown increases in brain activity
in the ventral striatum, insula, ACC, thalamus, right OFC, and dorsomedial prefron-
tal cortex when participants experienced chills; decreases in neural signal during the
same episodes were observed in the amygdala and hippocampus (Blood and Zatorre,
2001). Within the ventral striatum in particular, dopamine appears to be released
during both the anticipation and culmination of a peak pleasurable response to
self-selected pieces of music (Salimpoor et al., 2011). Other neuroimaging studies
exploring brain responses to highly pleasurable music, but not the experience of
chills per se, have found similar patterns of brain activity, primarily in reward
and emotion processing regions, including the ventral striatum, ACC, anterior
insula, and hippocampus (Brown et al., 2004; Koelsch et al., 2006; Menon and
Levitin, 2005).
Individual variation in the experience of chills may additionally be a function
of structural differences in the human brain. Our own work with diffusion tensor
imaging, a neuroimaging tool that allows the visualization and quantification
of white matter connections between various regions of the brain, has shown that
those who frequently and reliably experience chills while listening to music have
162 CHAPTER 8 Reflections on music, affect, and sociality

greater white matter volume in the tracks connecting the auditory cortex, insula, and
prefrontal cortex as compared to people who rarely, if ever, experience them (Sachs
et al., 2016). This suggests that the capacity for music to trigger intense emotional
responses may be related to this more robust structural connection between auditory
and emotion processing areas of the brain.

3.2 MUSIC ANHEDONIA

What do we make of individuals who do not experience chills? Individual variation
in ability to experience strong emotional responses to music is such that there are
people who report being unable to derive any pleasure from music, despite normal
responses to other rewarding stimuli, such as those associated with money, sex, and
drugs (Mas-Herrero et al., 2014). Even more surprising is that these individuals show
normal perceptual abilities when it comes to both processing structural features of
music and identifying the intended emotion that music conveys (Zatorre, 2015). This
incapacity to derive pleasure specifically from music has been called “musical
anhedonia” and recent investigations found that roughly 5% of a large sample of
individuals demonstrated such a condition (Mas-Herrero et al., 2014).
In addition to these behavioral differences, while listening to pieces of music
rated as highly pleasing by an independent group of participants, people with musical
anhedonia displayed attenuated psychophysiological responses (in both heart rate
and electrodermal activity) compared to a group of participants who experienced
typical emotional responses to music (Mas-Herrero et al., 2014). Importantly, no dif-
ferences in psychophysiological responses were found between musical anhedonics
and controls when the experiment involved monetary rewards, confirming that the
lack of pleasure experienced in the subjects with music anhedonia was specific to
musical stimuli. Recently, this experiment was replicated with fMRI. Both the
music-listening task and the monetary reward task activated the nucleus accumbens;
however, the music anhedonia group showed significantly less activation in the nu-
cleus accumbens during music listening compared to the control group, a difference
that was not seen during the monetary reward task (Martı́nez-Molina et al., 2016).
Furthermore, during the music-listening task, the patterns of activity in the ventral
striatum and auditory cortex were less correlated in the music anhedonia group, sug-
gesting that the origin of this special form of anhedonia could result from abnormal
communication between the reward and auditory processing regions of the brain
(Martı́nez-Molina et al., 2016).
In another study [in which one of us (Sachs) was involved], this hypothesis was
tested by obtaining structural brain images of an individual with musical anhedonia
using diffusion imaging and probabilistic tractography. When compared to a group
of adult participants, who demonstrated normal sensitivity to musical rewards
(though were not matched in terms of age or gender), the music anhedonic participant
showed less volume in the white matter track connecting the left auditory cortex and
the left nucleus accumbens as well as that connecting the left anterior insula to left
nucleus accumbens. Furthermore, in the comparison group, sensitivity to musical
 4 The case of sad music 163

rewards was directly related to volume in these white matter pathways. Specifically,
people who were more likely to experience pleasurable and rewarding responses to
music had larger tracks connecting the left auditory cortex and left anterior insula
(Loui et al., 2017). The results from this study and the functional imaging results
referred to above suggest that the lack of reward from music may be linked to aber-
rant connections between regions of the brain involved in processing sounds and re-
gions of the brain involved in processing pleasure, rather than malfunctioning within
one or several brain regions.

4 THE CASE OF SAD MUSIC

The relationship between emotions and pleasure, affect and reward, is clearly
entwined, though the exact relation can become rather complex and muddled when
it comes to music. For example, there are certain situations in which an intended
emotion can be recognized within a music piece without the listener feeling the said
emotion (Gabrielsson, 2001). Even more intriguing, the valence of the felt emotion
might be different from, or even the opposite of, the emotion that is identified as the
intended one. Such a mismatch is commonly observed in the case of sad music, when
people often report experiencing positive feelings, such as tenderness and enjoy-
ment, in response to what they would classify as sad music (Sachs et al., 2015). This
attraction to sad music has been labeled the “tragedy paradox” and is particularly
intriguing from a psychological standpoint because sadness is generally conceptual-
ized as a negatively valenced emotion, an unpleasant feeling that is to be avoided.
The fact that so many people report enjoying music meant to convey sadness to such
an extent that they explicitly seek it out is therefore strange and worthy of empirical
investigation.
To better understand this paradox, several researchers have attempted to both
qualify and quantify the exact nature of the emotional experience typically reported
in response to sad music. Some have suggested that because music does not pose a
tangible threat, any corresponding sensations are devoid of real-life consequences
and therefore must be “vicarious” by nature, the implication being that the emotion
of sadness can be recognized in the music, but it need not be experienced as unpleas-
ant or pleasant (Kawakami et al., 2013). Instead, people can actually feel pleasure as
a result of the beauty of the music or the music-listening experience itself, regardless
of the emotional intent of the piece (Kawakami et al., 2013; Schubert, 1996).
On the other hand, there is psychological and neurophysiological evidence
suggesting that full-fledged, “genuine” feelings of sadness in response to music that
conveys sadness, feelings as intense and visceral as sadness felt in response to a real
or perceived loss, can be felt in response to music that conveys sadness. Psychophys-
iological differences were found in participants when listening to sad music vs happy
music, including decreased skin conductance, higher finger temperature, and de-
creased activity in the muscles of the face (Lundqvist et al., 2008). Listening to sad
music also had subsequent effects on performance during a word recall task and a
164 CHAPTER 8 Reflections on music, affect, and sociality

picture judgment task, and these effects were similar to those caused by sadness in-
duced through autobiographical recall (Vuoskoski and Eerola, 2012), suggesting that
affective responses to sad music can alter perception and judgment in a similar way as
sadness induced in other situations. In neuroimaging studies, sad music activated some
of the same regions associated with sad affective states triggered by other means, and
these included the hippocampus, amygdala, caudate nucleus, and the thalamus
(Brattico et al., 2011; Mitterschiffthaler et al., 2003; Vytal and Hamann, 2010).
In the case of attraction to sad music, these genuine feelings of sadness are likely
experienced in tandem with other, more positive emotions. As pointed out in Juslin’s
model described earlier, more than one mechanisms of music-evoked emotions can
be engaged during music listening and at times, these different mechanisms can cul-
minate in two different emotional experiences (Juslin, 2013). In this account, when
listening to enjoyable, sad music, a person may feel sadness through one mecha-
nism, by the process of recognizing and internalizing the negative emotion being
conveyed, for instance, while also feel positive emotions through a separate mech-
anism, by the process of assessing the aesthetic qualities of the music, for example
(Juslin, 2013).
Whether people who enjoy listening to sad music actual feel sad or, rather, only
recognize sadness, but feel some other, more positive emotion most likely depends
on a combination of factors. Previous experiences and associations with the music
may play a role. It has been suggested that sad music may become linked to certain
psychological benefits that are less commonly associated with music and that convey
more positive valence. When participants were asked about their motivations for en-
gaging with self-identified sad music over happy music, they reported that sad music
allowed them to understand, savor, and ultimately resolve their own intense feelings
without any real-life implications (Taruffi and Koelsch, 2014). Sad music may also
be enjoyed because it has the ability to trigger specific memories, to connect people
with others, to distract them from current problems or situations, and to regulate or
enhance mood, or purely because it has become associated with something that is
beautiful and aesthetically pleasing (Eerola and Peltola, 2016).
Whether a person can experience these psychological benefits from sad music
may depend on personality. Empathy, for instance, has been shown to be correlated
with the liking of sad music (Kawakami and Katahira, 2015). It has been proposed
that people with higher empathic abilities may be more likely to gain the psycholog-
ical benefits from listening to sad music because they can more easily resonate with
and vicariously feel the intense emotions being conveyed (Sachs et al., 2015). Open-
ness to experience was also correlated with liking of sad music (Ladinig and
Schellenberg, 2012; Vuoskoski et al., 2011), a relation that was also found with
experiencing chills, suggesting that both experiences may be related, in that they
are likely to occur with music that is moving and in people that like to be moved.
Stemming from the findings that rumination is also found to be positively correlated
with enjoyment of sad music, other researchers have concluded that some individuals
have a maladaptive attraction to sad music, and enjoy it because of the negative feel-
ings associated with it (Garrido and Schubert, 2011). Musical expertise might
 5 Music training and emotion processing 165

additionally play a role, as some studies have found that individuals with more music
training tend to report feeling joy and pleasure in response to sad music (Eerola and
Peltola, 2016).
The social context surrounding the music-listening experience is also important.
People report choosing to listen to sad music more often when they are alone, when
they are in contact with nature, when they are in distress or feeling lonely, or when
they are in reflective or introspective moods (Taruffi and Koelsch, 2014). Positive
feelings toward sad-sounding music were also shown to increase when played after
a sad-mood induction paradigm (Hunter et al., 2011). However, there is also evi-
dence that some people select sad music when they are not feeling sad. In one study,
people who exhibited a subtype of empathy called perspective taking, referring to the
tendency to cognitively adopt the viewpoint of an observed other, were more likely to
listen to sad music when in a positive mood; on the other hand, people who exhibited
another subtype of empathy called fantasy proneness, referring to the tendency to
become transported into the feelings of characters when engaging with works of
fiction (Davis, 1983), were more likely to listen to sad music when they were in a
negative mood (Taruffi and Koelsch, 2014).
Preliminary survey data collected in our laboratory may help explain how the
different subtypes of empathy can impact emotional responses to sad music. We col-
lected responses in a survey in which people answered questions regarding various
reasons why they might engage with sad music. We found that people who scored
high on the perspective taking reported enjoying sad music because it was associated
with certain cognitive benefits, such as allowing them to better understand their sit-
uation and emotional state. In this survey, we also assessed the situations in which
people were likely to engage with sad music and found that people who scored higher
on the fantasy component of empathy chose to listen to sad music when in a negative
mood, such as after a breakup or when feeling lonely, because, paradoxically,
they were more likely to feel positive, rather than negative, emotions in response
to the music.

5 MUSIC TRAINING AND EMOTION PROCESSING

So far, we have discussed varied ways in which music is systematically tied to emo-
tions and feelings. But can this connection be tightened or altered with music train-
ing? Learning to play a musical instrument has been associated with improvement in
auditory skills (Kraus and Chandrasekaran, 2010; Schellenberg and Moreno, 2010;
Sch€on et al., 2004), yet, it is unclear whether music training is associated with better
social or emotional abilities. With this in mind, we have been investigating the im-
pact of music training on child development, specifically whether music instruction
in childhood is predictive of improvements in recognizing and understanding the
emotions of others and whether this improvement can be detected at the neural level.
For this study, young musicians were recruited from music schools and studios in
the greater Los Angeles area. Participants were between the ages of 8 and 13 and had
166 CHAPTER 8 Reflections on music, affect, and sociality

been playing a string instrument continuously for at least 2 years. These children
completed two emotion perception tasks while inside an MRI scanner: one musical
(auditory) and one nonmusical (visual). Using both a musical and nonmusical task of
emotion perception would allow us to inquire if music training influences the pro-
cessing of emotions when presented in the auditory domain, and if such influence
extends to a nonauditory domain with nonmusical stimulus.
For the auditory (musical) task, we developed a set of auditory stimuli by recording
an adult violinist performing two musical pieces (Jules Massenet’s Meditation from
Tha€ıs and Sergei Rachmaninoff’s Rhapsody on a Theme of Paganini, Variation 18)
in four different emotional styles: happy, sad, peaceful, and angry. During the task,
children listened to the recordings and were instructed to respond to the question
“How do you think the musician is feeling during the performance” using an MRI-
compatible button box with four answer choices. Given that the same musical piece
appeared in every video clip, this task required that the children pay close attention to
the subtle differences of timing and expression in the performance in order to deter-
mine which emotion was being conveyed.
For the visual (nonmusical) task, the children were presented with a series of
full color emotional faces, taken from the NimStim dataset (Tottenham et al.,
2009), corresponding to one of three basic emotions: sadness, fear, or anger. The
faces were subsequently morphed, to varying degrees, with neutral, expressionless
faces to create images that ranged in intensity of expression. In one condition, chil-
dren had to judge the intensity of the emotional expression, independently of the type
of emotion being expressed. Like the auditory (musical) emotion perception task,
accurate performance on this condition requires attention to specific details of the
stimulus. As a control condition, children were asked to identify which emotion
was being conveyed. Responses were again recorded via the MRI-compatible button
box with four possible choices. We analyzed correlations between years of music
training and (1) the behavioral responses to the two tasks as well as (2) the degree
of neural activation.
Behaviorally, we found that during the auditory task, years of musical training
were positively correlated, after controlling for age, with correct identification of
the emotion that the musician was attempting to convey. For the visual task, no
correlation was found between years of music training and correct identification
of the intensity of the emotion being conveyed by the face nor between music train-
ing and correct identification of the emotion.
For the fMRI analysis of the auditory task, we contrasted the positively valenced
audio stimuli (happy and peaceful) with the negatively valenced audio stimuli
(sad and angry) in order to quantify brain activity related to perceiving differences
in the emotional style of the performances. We then correlated these signal differ-
ences with years of musical training and found significant positive relationships:
children with more musical training showed increased activity for this contrast in
the right superior parietal lobule, postcentral gyrus, and middle and superior frontal
gyri. These regions are proposed to be part of the action–observation network,
i.e., a collection of brain areas involved in both performing an action and observing
 References 167

that same action in others (Caspers et al., 2010; Grezes and Decety, 2001;
Meyer et al., 2011) and, relatedly, are activated during various tasks that involve
socioemotional experiences (McLellan et al., 2012; Schulte-R€uther et al., 2007).
Our neuroimaging finding might therefore indicate that children with more music
training may be better at mentally simulating the actions involved in expressing emo-
tions through music, which may help to correctly identifying the intended emotion.
In the visual (nonmusical) task, the judgment of emotional intensity is a more
sensitive measure of emotional perception than simply labeling an emotion and, pre-
dictably, produced greater brain activation. However, neural activations from this
contrast did not correlate with years of music training. Thus, in the visual task neither
behavior nor brain activation was related to musical training.
These findings suggest that learning to play a musical instrument can enhance the
ability to perceive emotions conveyed by musical stimuli. This improvement in the
behavior may be related to maturation of brain functions, as our study shows that music
training was associated with heightened activity in the neural systems that under-
lie emotion perception. However, given that the number of years of music training
is highly correlated with age, it is not clear, in the absence of a control group of
same-age children without music training, if the effect we detected might or might
not be related to age. Furthermore, our results did not provide evidence that the emo-
tion perception benefits of musical training transfer to the visual, nonmusical domain.
Nevertheless, we should note that there is evidence that adult musicians demonstrate
an enhanced perception of emotion in nonmusical auditory stimuli, such as speech,
suggesting that music training can provide perceptual benefits to nonmusical stimuli
presented to the auditory system (Lima and Castro, 2011; Strait et al., 2009). It remains
to be determined if the possible influence of musical training on the emotion processes
in a nonauditory domain would be evident with a different task or in a larger sample.

6 CONCLUDING REMARKS
We have provided an overview of current research on the ways in which music and
human affective experiences are connected and on the brain mechanisms that support
this connection. Future neuroimaging and psychological explorations are likely to
provide a clearer picture of how music and reward processes interact and continue
to reveal the ways in which music plays a role in so many aspects of human cultures.

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