The Neurobiology of Aesthetic Chills: How Bodily Sensations Shape Emotional Experiences

Abstract

The phenomenon of aesthetic chills—shivers and goosebumps associated with either rewarding or threatening stimuli—offers a unique window into the brain basis of conscious reward due to their universal nature and simultaneous subjective and physical counterparts. Elucidating the neural mechanisms underlying aesthetic chills can reveal fundamental insights about emotion, consciousness, and the embodied mind. What is the precise timing and mechanism of bodily feedback in emotional experience? How are conscious feelings and motivations generated from interoceptive predictions? What is the role of uncertainty and precision signaling in shaping emotions? How does the brain distinguish and balance processing of rewards versus threats? Here we review neuroimaging evidence and highlight key questions for understanding how bodily sensations shape conscious feelings of pleasure and meaning. This research stands to advance models of brain-body interactions shaping affect and may lead to novel non-pharmacological interventions for disorders of motivation and pleasure.

Full text

Vol.:(0123456789)
Cognitive, Affective, & Behavioral Neuroscience (2024) 24:617–630
https://doi.org/10.3758/s13415-024-01168-x
THEORETICAL REVIEW
The neurobiology ofaesthetic chills: How bodily sensations shape
emotional experiences
FelixSchoeller1,2· AbhinandanJain2· DiegoA.Pizzagalli3· NiccoReggente1
Accepted: 21 January 2024 / Published online: 21 February 2024
© The Author(s) 2024
Abstract
The phenomenon of aesthetic chills—shivers and goosebumps associated with either rewarding or threatening stimuli—offers
a unique window into the brain basis of conscious reward because of their universal nature and simultaneous subjective
and physical counterparts. Elucidating the neural mechanisms underlying aesthetic chills can reveal fundamental insights
about emotion, consciousness, and the embodied mind. What is the precise timing and mechanism of bodily feedback in
emotional experience? How are conscious feelings and motivations generated from interoceptive predictions? What is the
role of uncertainty and precision signaling in shaping emotions? How does the brain distinguish and balance processing of
rewards versus threats? We review neuroimaging evidence and highlight key questions for understanding how bodily sensa-
tions shape conscious feelings. This research stands to advance models of brain-body interactions shaping affect and may
lead to novel nonpharmacological interventions for disorders of motivation and pleasure.
Keywords Chills· Dopamine· Precision· Reward· Learning· Music· Film· Emotional· Valence· Arousal
Introduction
Aesthetic chills (AC) are a strong emotional response to
specific stimuli (chills stimuli [CS]), such as music, films,
or speech (Schoeller etal., 2022, 2023a, b, c, d), with a
characteristic neural signature involving the mesocortical
“reward” pathway (Blood & Zatorre, 2001; Ferreri etal.,
2019; Salimpoor etal., 2011). Crucially, AC is associated
with strong, discrete physiological response of shivering
and/or goosebumps (Benedek & Kaernbach, 2011), which
ordinarily regulate body temperature (hence AC are perhaps
best characterized as “psychogenic shivers”), thereby pro-
viding a unique opportunity to study how brain and body
interact during consummatory pleasure and reward learning
(Contreras-Huerta etal., 2023; Paulus & Stewart, 2014).
Interestingly, there seems to be two symmetrical types of
AC: 1) positive chills, tied to high rewards (Blood & Zatorre,
2001), and 2) negative chills, associated with high risks
(Zald & Pardo, 2002). Both activate the extended amygdala
(AMG) and are connected to the processing of uncertainty,
whether unexpectedly higher or lower than anticipated (Sch-
oeller & Perlovsky, 2016). The evidence that AC engage
both appetitive and aversive neural systems is a reminder
of philosophical discussions concerning the nature of awe
and the sublime as a mixture of positive and negative emo-
tions (Burke, 1757; Kant, 1951; Schiller; 1967; Longinus,
2022) and work on threat and reward processing in the brain
(Murty etal., 2023). The dynamics of AC offer a distinc-
tive window into the emergence of conscious feelings—e.g.,
how the latency between an unexpected interoceptive signal
(i.e., shivers and downstream effects, such as goosebumps)
and conscious awareness impacts the valuation of exterocep-
tive cues (Craig, 2008; Damasio & Damasio, 2023; Seth,
2013). Hence, in addition to providing novel insights into
how pleasure comes about in the mind and brain, an under-
standing of the neural process supporting AC could lead to
nonpharmacological substitutes for dopaminergic-related ill-
nesses, as suggested by the recent findings suggesting that
CS seem to improve reward learning (Jain etal., 2023a, b)
and maladaptive cognitions in anhedonic depression (Sch-
oeller etal., 2023a, b, c, d). CS include music (Blood &
* Felix Schoeller
felixsch@mit.edu
1 Institute forAdvanced Consciousness Studies,
SantaMonica, CA, USA
2 Media Lab, Massachusetts Institute ofTechnology,
Cambridge, MA, USA
3 McLean Hospital, Harvard Medical School, Belmont, MA,
USA

618 Cognitive, Affective, & Behavioral Neuroscience (2024) 24:617–630
Zatorre, 2001; de Fleurian & Pearce, 2021), films (Schoe-
ller etal., 2022; Schoeller, Eskinazi etal., 2018a; Schoeller
& Perlovsky, 2016), stories (Schoeller & Perlovsky, 2015),
poetry or speech (Wassiliwizky etal., 2017; Wassiliwizky
& Menninghaus, 2021), scientific practice (Schoeller,
2015a), and religious and secular rituals (Schoeller, 2015a).
In this short perspective, we review the neural mechanisms
of chills and reward (Section "Neural mechanisms of aes-
thetic chills and reward") and relate them to the notion of
precision encoding in the framework of predictive coding
(section "Dopamine and precision encoding in aesthetic
chills"). We discuss the notion that chills may be related to
the overall predictability of events given previous expecta-
tions (a.k.a., precision encoding) and conclude by exploring
how AC could influence dysfunctional precision-weighting
in psychopathology.
Neural mechanisms ofaesthetic chills
andreward
Evidence to date suggests that AC engage a distinct brain
network where neurons in the ventral tegmental area (VTA)
of the midbrain disperse dopaminergic projections through-
out the mesocorticolimbic system to exert neuromodula-
tory actions critical for a range of reward and motivation
processes, including hedonics, reward-related learning,
and behavioral adaptation (Blood & Zatorre, 2001; Chabin
etal., 2020; Ferreri etal., 2021; Witt etal., 2023; Salim-
poor etal., 2011; Zald & Pardo, 2002). As we discuss in
the following section, dopamine is thought to encode preci-
sion—statistically, the inverse of variance—of predictions,
signaling when neural representations are more reliable
(Friston etal., 2012)—although see alternative interpreta-
tions in Jeong etal. (2022). The relevant circuits are usually
broadly characterized as the salience and reward networks,
and as such include limbic (amygdala [AMG] and nucleus
accumbens [NAcc]) and frontal (orbitofrontal [OFC] and
ventromedial frontal cortex [vmPFC]) regions (Seeley,
2019; Sesack & Grace, 2010). As of this article, only seven
studies have examined the neural correlates of AC (Blood
& Zatorre, 2001; Chabin etal., 2020; Ferreri etal., 2019;
Witt etal., 2023; Sachs etal., 2016; Salimpoor etal., 2011;
Zald & Pardo, 2002). Unfortunately, the neuroimaging work
conducted so far has focused exclusively on music as an
eliciting stimulus (but see Wassiliwizky etal., 2017). We
simply cannot conclude anything yet regarding AC induced
by CS, such as films, speeches, stories, science, and rituals
at large. A recently validated, “gold standard” database of
CS, extending beyond music, has been established by Sch-
oeller etal. (2022) later validated on 3000+ participants
(Schoeller etal., 2023a, b, c, d), marking the initial steps in
uncovering the underlying neural mechanisms beyond mere
musical pleasure. We review some of the neural correlates
associated with musical chills as a springboard (Fig.1), with
all the limitations entailed.
Blood & Zatorre (2001) used PET imaging to investigate
musical chills, reporting increased blood flow to the ventral
striatum, midbrain, insula, right orbitofrontal cortex, thala-
mus, anterior cingulate, SMA, and cerebellum. The decrease
in blood flow was noted in areas, such as the amygdala, left
hippocampus, and vmPFC. The authors observed that this
profile is similar to neural responses in cocaine-dependent
subjects when administered cocaine (Breiter etal., 1997),
suggesting that musical chills may activate pathways akin
to those engaged by euphorogenic substances (Biederman
& Vessel, 2006; Nguyen etal., 2021). Given the causal
relationship between dopamine and the pleasure derived
from cocaine (Volkow etal., 1999), it raises the question of
whether the sensation of chills may engage a dopaminergic
response analogous to that which is classically observed dur-
ing “incentive salience” (Berridge, 2007). Early studies by
Goldstein suggest that chills can be inhibited by an opioid-
antagonist (Goldstein, 1980), but these results have failed
to replicate (Laeng etal., 2021). In the study by Laeng etal.
(2021), compared with placebo, the opioid antagonist (50
mg, naltrexone) did not alter self-reported pleasure associ-
ated to the CS, but it did specifically reduce pupil size dur-
ing AC. This suggests that while the endogenous μ-opioid
system is not essential for the subjective pleasure of AC,
blocking opioids induce a reduction in arousal in response
to the CS.
The influence of dopamine on various functions—such as
sensory pleasure (also known as hedonic impact), increased
motivation, and learning—is evident in overlapping brain
regions, including the ventral striatum (NAcc), midbrain
(VTA, Periaqueductal Gray [PAG], Pedunculopontine
Nucleus), amygdala, hippocampus, mPFC (Berridge & Rob-
inson, 1998; Terry etal., 1995). Consistent with this line
of reasoning, Salimpoor etal. (2011, 2013) subsequently
used PET to demonstrate that the pleasure derived from
music is associated with dopamine release in the dorsal and
ventral striatum, specifically the NAcc. This neural activity
was found to correlate with how much participants would
pay for pleasurable songs, connecting the experience of
musical pleasure with reward-based decision making. Fur-
thering confidence in this relationship, Mas-Herrero etal.
(2018) demonstrated a causal link between neural activity
and music appreciation, where excitatory TMS to the left
DLPFC increased pleasure, arousal, and monetary valuation
of music (i.e., the subject’s willingness to spend money on
songs), while inhibitory TMS had the opposite effect. Ferreri
etal. (2019) also showed that the administration of dopa-
mine precursors, such as levodopa upregulates the experi-
ence of pleasure during music listening (but not valence and
arousal), and drives a significant increase in chills incidence,

619Cognitive, Affective, & Behavioral Neuroscience (2024) 24:617–630
whereas dopamine antagonists reduced these effects com-
pared with placebo (Ferreri etal., 2019).
These results suggest that aesthetic pleasure may be
rooted in the interplay between reward/valuation systems
(striatal-limbic-paralimbic) and more phylogenetically
advanced perception/prediction systems (temporofrontal).
Goupil & Aucouturier (2019) proposed models that high-
light two distinct, yet possibly, interacting frameworks: (A)
a corticostriatal model connected with NAcc for musical
pleasures, and (B) a model for musical emotion involving
the thalamus, amygdala, and DLPFC. The orchestration of
these responses raises profound questions about whether A
and B operate independently, if A provides first-order input
to the construction of emotion, or if A evaluates B, thereby
illuminating the complex neural interactions that underpin
the experience of pleasure and emotion.
Interestingly, the brain correlates of chills map with the
first phase of the reward cycle (a.k.a., the “Wanting” phase),
characterized by midbrain dopamine projections to forebrain
targets, such as NAcc and other parts of striatum (Berridge
etal., 2009) (Fig.1). This contrasts with the dynamics of
AC in narrative films, where AC generally arise during the
culmination of the film—not beforehand (Fig.2). In other
words, people do not experience chills in anticipation of
the movie; rather, these sensations tend to manifest during
the film itself (although an exception could be made of film
“trailers” as evidenced in Schoeller etal., 2023a, b, c, d).
This is when the narrative's uncertainties are resolved, and
a cohesive meaning emerges from the tapestry of events that
unfold throughout the film—i.e., the film is over (Schoeller
& Perlovsky, 2015). This pattern would instead suggest that
AC correspond to a peak in consummatory pleasure (a.k.a.,
the “Liking” phase), kicking off the satiation process (a.k.a.,
the “Learning” phase) (Schoeller & Perlovsky, 2016), where
mesolimbic circuitry is responsible for generating incentive
salience (Berridge, 2012). Note that the desire for rewards
also is involved in generating fearful salience (Berridge &
Kringelbach, 2015), consistently with previous research on
negative AC by Zald & Pardo and the “reversed role” of the
amygdala in positive and negative AC during aversive audi-
tory stimulation (Zald & Pardo, 2002). To reconcile this,
we consider the neurobiological underpinnings of pleas-
ure and satisfaction, acknowledging that the neural circuits
involved—namely the Nucleus Accumbens (NAc), Ventral
Tegmental Area (VTA), and striatum—do not necessarily
exhibit exclusive activity in distinct phases of appetitive,
consummatory, and satiety processes. Instead, these regions
play a pivotal role in transitioning from consumption to
Fig. 1 Neural mechanisms of chills and reward. Green areas indicate
increased activation during the chills response. Red areas are deacti-
vated. Key regions involved include the ventral tegmental area (VTA)
projecting to the ventral striatum (nucleus accumbens) and the hip-
pocampus. The amygdala, orbital, and ventromedial prefrontal cortex
show deactivation during chills (Blood & Zatorre, 2001). The VTA,
nucleus accumbens, and striatum are part of the brain's reward sys-
tem, associated with pleasure, reward, and compulsive behavior. The
right orbitofrontal cortex (OFC) plays a role in sensory processing,
reward, and expected outcomes. Note that electroencephalographic
recording of chills showed theta activity with activation in the orbit-
ofrontal cortex, in correlation with emotional ratings (Chabin et al.,
2020)

620 Cognitive, Affective, & Behavioral Neuroscience (2024) 24:617–630
satiety, as evidenced by studies showing NAc’s role in ini-
tiating and maintaining feeding behaviors, and its potential
'unpausing' during the transition to satiety (Krause etal.,
2010). Furthermore, VTA lesions have been associated with
reduced overconsumption (Shimura etal., 2002), suggest-
ing its involvement in peak consumption and the subsequent
shift to satiety. This is further supported by evidence of
dopamine release in the striatum during peak consumma-
tory pleasure (Small etal., 2003). Therefore, we propose that
AC may not solely be an artifact of the “wanting” phase but
also could represent a neurobiological marker of the transi-
tion from consumption to satiety, characterized by a rela-
tive increase in neural activity (e.g., from pause to unpause
in NAc activity). This perspective aligns with the observed
overlap in neural correlates and the temporal dynamics of
AC. Additionally, we acknowledge that the temporal reso-
lution of fMRI may not be sufficiently sensitive to capture
these transient processes, especially considering the chal-
lenges in perfectly timing the self-reporting of chills with
neural activity.
Schoeller & Perlovksy suggested a computational models
for AC (Schoeller & Perlovsky, 2016) as a peak value in
uncertainty reduction at higher levels of cortical informa-
tion encoding—represented by the equation: L = ∏n [∑m
l(X(n) | M(m))], where l denotes the conditional similar-
ity of sensory signals given available models (see technical
details in Schoeller Perlovsky, etal., 2018b). Importantly,
the local peak value (i.e., modelled as a null derivative) is
independent of valence, accounting for the fact that chills
can be experienced in response to both positive (appetitive)
and negative (aversive) stimuli (Fig.3). In line with recent
models of affect as second order predictions about learning,
i.e., expected prediction errors (Joffily & Coricelli, 2013;
Kenett etal., 2023; Miller & Clark, 2018; Perlovsky & Sch-
oeller, 2019; Sarasso etal., 2020; Van de Cruys & Bervoets,
2022), this suggests that chills may correspond to a null
learning rate, or zero derivatives, signaling a local maxi-
mum (extremum) in learning (Schoeller, Perlovsky etal.,
2018b; Schoeller & Perlovsky, 2016). In other words, AC
occur when the individual reaches the limits of their ability
to learn more about their surroundings, and their learning
rate slows down, terminating the Wanting phase and kicking
off the Satiation phase, effectively corresponding to what
has been described in the field as a temporary satiation of
curiosity (Biederman & Vessel, 2006; Kenett etal., 2023;
Sarasso etal., 2020; Schoeller, 2016). In machine learning,
momentum-based gradient optimization algorithms utilize
a similar concept of momentum to expedite the convergence
process toward minima in the error landscape, enhancing the
efficiency of learning in areas with consistent error reduction
(Ruder, 2016). These methods mirror the adaptive modula-
tion of reward cycles, observed in both biological (Soltani
& Koechlin, 2022) and artificial (Piao etal., 2023) intel-
ligent, self-organizing systems. In both cases, the dynamic,
adaptive recalibration of reward mechanisms is pivotal for
optimizing learning trajectories. The mammalian brain, par-
ticularly in the prefrontal cortex, is equipped with computa-
tional abilities to form and evaluate task sets based on inter-
nal models associating stimuli, actions, and outcomes. This
configuration facilitates adaptive behavior through selective,
Fig. 2 Three phases of pleasure and chills as peak consummatory
pleasure. The "Wanting" phase represents the initial anticipation
and desire for rewards. The "Liking" phase corresponds to the peak
of consummatory pleasure, characterized by the experience of aes-
thetic chills (AC). Following the chills, the “Learning” phase begins,
involving the encoding of crucial information about the film. This
phase is associated with the consolidation of the CS meaning. In
other words, AC marks the onset of the satiation process, where the
viewer's curiosity is temporarily satisfied

621Cognitive, Affective, & Behavioral Neuroscience (2024) 24:617–630
predictive, and contextual models, integrating rewards, sta-
tistical inferences, and dynamic environmental cues. Addi-
tionally, the activation of motor regions of the cortex, such
as the cerebellum or the supplementary motor area, further
supports the hypothesis that chills may be related to action
orientation and readiness (Sarasso etal., 2019, 2020, 2021,
Sarasso, Francesetti etal., 2022a, Sarasso, Neppi-Modona
etal., 2022b). Sarasso and colleagues accumulated exten-
sive data on the role of aesthetic emotions in learning (and
their downstream effects in terms of perceptual and memory
improvement) and suggested the hypothesis that they may be
related specifically to the inhibition of action for the purpose
of seeking and acquiring knowledge (Sarasso etal., 2019,
2021, Sarasso, Neppi-Modona etal., 2022b), in line with the
hypothesis that AC may correspond to a satiation of curios-
ity (Schoeller, 2015b, 2016).
Activity in the insular cortex during the chills response
hints towards the role of interoception (the perception of
bodily sensations) and peripheral signals in the emotional
experience of chills (Craig, 2008; Damasio, 1996; Seth,
2013). Recent evidence suggests that lesions impairing
structural connectivity of the left insula in stroke patients
modulated AC (Witt etal., 2023). Specifically, a large left
hemispheric lesion including the left insula impaired the
bodily response of chill experience (objective chill response)
but left the cognitive aspects of chill processing (subjective
chill response) unaffected (Grunkina etal., 2017). The study
by Witt etal. (2023) examined stroke patients who primarily
had damage to the insula. The study found that these patients
experienced chills in response to auditory stimuli (such as
music or harsh sounds) at a frequency similar to that of peo-
ple without brain damage. However, there was a disconnect
between their emotional arousal and their physical responses
that correlated with structural connectivity between the left
anterior insula and the temporal pole. Additionally, fMRI
analyses revealed that during chills stimuli, reduced skin
conductance responses correlated with lower activation
of the temporal pole. These findings highlight a role for
the insula and temporal pole in integrating physiological
responses to arousing experiences like chills. This role of
interoceptive signals in chills also is evident in data show-
ing that physical manipulation of the somatic markers of
the emotion by means of a wearable prosthesis enhancing
the sensation of cold can enhance the feeling of pleasure
and some of the downstream effect of chills (Haar etal.,
2020; Ishikawa etal., 2023; Jain etal., 2020; Schoeller etal.,
2019). Interoceptive signals provide essential information
to the brain about the body’s internal state, contributing to
the shaping of emotional responses and the anticipation of
rewarding outcomes (Contreras-Huerta etal., 2023; Paulus
& Stewart, 2014). Positive stimuli, whether they are social
interactions, delicious food, or soothing experiences, such
as the warmth of sunlight, activate neural pathways that are
closely intertwined with interoception. Conversely, negative
experiences or threats can trigger aversive responses that
also involve interoceptive processing, contributing to our
understanding of interoception's role in the broader spec-
trum of emotions. Notably, amygdala and insula connectivity
modulate the effect of input arousal on effective connections,
with the amygdala as the main hub driving arousal input
(Wang etal., 2023).
Elaborating on the contributions of insular-cortical con-
nectivity to the uptake of arousing stimuli, Sachs etal.
(2016) accounted for individual differences in the trait
frequency of experiencing musical chills with white-mat-
ter tract volume (an indicator of structural connectedness
(Mollin etal., 2016; Seehaus etal., 2013, 2015)) within
a three-node network of the posterior superior temporal
gyrus (auditory association area) with the anterior insula
(interoception and emotion processing (Zaki etal., 2012))
Fig. 3 Learning (blue) and its rate of change (red), as the fit between
ascending signals and available models. Learning is represented by
the equation: L = ∏n [∑m l(X(n) | M(m))], where l denotes the con-
ditional similarity of data given conditional models (Schoeller etal.,
2018a, b). This sinusoidal representation is a gross oversimplifica-
tion for the sake of readability. The blue area at the peak of the curve
defines conscious aesthetic emotions (when the rate of change tends
toward zero), and the squared area describes a descending learning
slope corresponding to a negative derivative. The graph highlights the
interplay between learning and emotions, emphasizing how insights
(e.g., positive chills) and traumas (e.g., negative chills) shape an indi-
vidual’s overall mood and well-being. Adapted from Schoeller & Per-
lovsky, 2016

622 Cognitive, Affective, & Behavioral Neuroscience (2024) 24:617–630
and medial prefrontal cortex (reward processing (Knutson
etal., 2003; Tzschentke, 2000; Pastor and Medina, 2021)).
These findings, which illustrate that increased sensory
access (i.e., increased structural connectedness) to reward
systems accounts for increased sensitivity to chills (Mori
& Iwanaga, 2015), suggest a potential evolutionary foun-
dation for the aesthetically rewarding function of mean-
ing-making in humans. Given the evidence that practice
induces microstructural white-matter changes across several
domains and time scales (Sagi etal., 2012; Engvig etal.,
2011; Reid etal., 2017), it stands to reason that these effects
could be interpreted in reverse: frequent chills might lead
to enhanced structural connectivity between these regions.
Future research should investigate the structural effects of
repeated exposure to chills to better understand any causal
relationships involved.
Individual differences in the propensity to experience aes-
thetic chills also have been linked to both personality traits,
such as absorption and openness to experience (Silvia &
Nusbaum, 2011, Williams etal., 2018, Williams etal., 2023,
Silvia etal., 2015, Johnson etal., 2023, Schoeller etal.,
2023a; McCrae, 2007), as well as biological factors, such as
gene polymorphisms affecting neurotransmitter function. A
recent twin study found that approximately 36% of variance
in aesthetic chills experiences can be attributed to genetic
factors (Bignardi etal., 2022). This further supports a role
for underlying biological pathways in facilitating intense
emotional responses to aesthetic stimuli. Given evidence that
serotonin and dopamine systems interact, with dopamine D2
and serotonin 5-HT1a receptors showing functional opposi-
tion (Howell & Cunningham, 2015), an interesting question
is whether altering serotonin function could inhibit AC by
shifting this balance. For example, increased serotonin sign-
aling through selective serotonin reuptake inhibitors (SSRI)
antidepressants might conceivably dampen chill experiences
mediated by dopamine release. Testing effects of pharma-
cological manipulations on aesthetic chills could elucidate
neuromodulatory mechanisms underlying individual differ-
ences in emotional reactivity.
Dopamine andprecision encoding
inaesthetic chills
Within a view of the brain as an organ of prediction and
learning (Kveraga etal., 2007; Downing, 2009), these
findings and our knowledge of the brain structures marry
well with the growing amount of evidence that dopamin-
ergic neurons and pathways play an essential role in learn-
ing by encoding the precision of prediction errors (i.e.,
their confidence or reliability) along the cortical hierar-
chy (Diederen etal., 2017; Jeong etal., 2022). What’s
more, the peculiarity of the chills phenomenon make it an
excellent object of experimental study, insofar as chills can
be 1) consciously reported by subjects, 2) are, in principle,
measurable physically in terms of heat dispersal, and 3)
seem universal across cultures (McCrae, 2007). Precision
encoding is a process by which the brain continuously esti-
mates the reliability of the sensory inputs (i.e., the predic-
tion errors) encountered and the reliability of its models
of the world (Keshvari etal., 2012). Crucially, mounting
evidence indicates that dopaminergic pathways mediate
this process, where dopamine release occurs when reward
is greater than expected, which helps the brain update pre-
dictions and improve learning trajectories (Schultz, 2016;
Kiverstein etal., 2017). Precision encoding may be related
to the enhancement of memory and attention processes
observed during AC experiences (Ferreri etal., 2019,
2021). Recent evidence by Kathios etal. shows that when
people are exposed to novel musical melodies based on
an unfamiliar scale, liking ratings increase for frequently
presented melodies adhering to an implicitly learned struc-
ture, supporting predictive coding models whereby new
musical sounds become rewarding through learned predic-
tions (Kathios etal., 2023). By encoding the precision of
prediction errors, dopamine allows the brain to consolidate
and retain information, leading to improved memory and
attention of salient events (Adcock etal., 2006; Ferreri
etal., 2019, 2021; Sarasso etal., 2021).
In a series of experiments, it was repeatedly shown that
priming subjects with incoherent stimuli inhibits the chills
(Schoeller & Perlovsky, 2016; Schoeller & Eskinazi, 2019;
Schoeller etal., 2018b), most likely as incoherence (i.e., sur-
prisal) interferes with the precision system (Schoeller & Per-
lovsky, 2016). CS activate dopamine projections, aiding to
the formation and consolidation of emotional memory (Rip-
olles etal., 2016; Ferreri & Rodriguez-Fornells, 2017, 2022;
Ferreri etal., 2021, 2013; Sarasso etal., 2019, 2020, 2021,
Sarasso, Francesetti etal., 2022a, Sarasso, Neppi-Modona
etal., 2022b). Sarasso and colleagues accumulated evidence
that chills and aesthetic emotions induce an enhancement of
memory and attention processes, corroborating their hypoth-
esis and confirming previous studies by Laura Ferreri on
reward-driven music memory consolidation and the role
of dopamine in the process (Ferreri & Rodriguez-Fornells,
2017, 2022; Ferreri etal., 2021, 2013). Neurobiologically,
this makes sense insofar as VTA dopaminergic projections
to the hippocampus, as observed in the chills phenomenon,
are known to play a crucial role in encoding emotional mem-
ories (Ripolles etal., 2016; Shohamy & Adcock, 2010). The
extended amygdala’s involvement calls for studies on the
roles of acetylcholine and its combined effects with dopa-
mine in supporting aesthetic chills and their connection
to learning. That is because acetylcholine, acting through
nicotinic receptors, interacts with dopamine release and may
contribute to precision signaling (Matityahu etal., 2023).

623Cognitive, Affective, & Behavioral Neuroscience (2024) 24:617–630
Along similar lines, musical pleasure stems from expecta-
tions generated by learned musical patterns and the reward-
ing violations of those expectations (Krumhansl & Agres,
2008; Koelsch etal., 2019, Kraus, 2020).
As we engage with CS, our brain continually predicts
upcoming events based on implicit knowledge (Zatorre &
Salimpoor, 2013). Dopamine signals violations of expecta-
tions, or prediction errors, driving learning to update expec-
tations (Pessiglione etal., 2006; Egermann etal., 2013).
High precision tunes attention—enhancing memory and
learning—and the sudden drop in precision induced by an
incoherent prime interrupts this process (Schoeller & Per-
lovsky, 2016; Schoeller & Eskinazi, 2019; Schoeller etal.,
2018b). Precision tuning focuses attention and learning on
the most reliable predictions. Musical or narrative tension
builds uncertain predictions, engaging a cascade of stimulus-
driven expectations until resolution ultimately satisfies the
predictions, eliciting pleasure (Lehne etal., 2014, Schoeller
& Perlovsky, 2016—see also Deterding etal., 2022 in the
context of video games). Kathios etal. (2023) demonstrated
that functional MRI activity in auditory cortical regions
reflected prediction errors, while connectivity between audi-
tory and medial prefrontal cortex tracked both exposure and
prediction error signals, further implicating predictive cod-
ing in the context of musical reward. Across nine studies (n
= 1,185), statistical learning of sequential patterns was suf-
ficient to drive rewarding responses to expectation violations
for novel musical systems (Kathios etal., 2023).
CS engage a cyclic interplay between cortical systems
generating expectations, dopamine signaling of prediction
precision and errors to update expectations, and subcorti-
cal systems mediating emotion, reward, and memory. The
arousal system plays a significant role in this process, includ-
ing dopaminergic pathways, regulating sensory processing
and emotional responses (Pfaff & Banavar, 2007). Eleva-
tions in dopamine release within mesolimbic, mesocortical,
and nigrostriatal target sites coincide with arousal (Horvitz,
2000). Notably, the level of arousal before CS exposure is a
major predictor of whether the subject will experience chills
subsequently (Schoeller etal., 2023b); chills participants
report arousal levels twice as high as nonchills participants
on average before the CS exposure. This aligns with theo-
ries proposing primal brain areas as central neural corre-
lates of consciousness (Solms, 2021; Safron, 2021; Schoeller
etal., 2023a, b, c, d). AC and aesthetic emotions may ini-
tially originate from basic autonomic emotional responses
rather than higher-order cognitive processes, which would
explain why chills may feel surprising when they finally
enter consciousness.
Notably, dysfunctional precision encoding of prediction
errors by dopamine is implicated across psychiatric ill-
nesses, including schizophrenia, depression, and addiction.
For example, in schizophrenia, the precision of perceptual
priors driving inference is pathologically low, leading to hal-
lucinations and delusions. Dopamine has been suggested to
play a role in the precision weighting of unsigned prediction
errors, which signal the degree of surprise without indicat-
ing valence (Haarsma etal., 2021). Hence, dysfunctional
precision-weighting of unsigned prediction errors may be a
pivotal contributor to the pathogenesis of psychosis (Krys-
tal etal., 2017; Adams etal., 2013; Heinz etal., 2019). In
contrast, high precision in depression (especially concern-
ing social cues) may reflect impaired reward sensitivity and
learned helplessness (Smith etal., 2023). Hence, identi-
fying the relationship between aesthetic chills, the preci-
sion-weighting of prediction errors, and its dopaminergic
substrate could help to understand the role of dopamine in
these conditions. Using a recently constituted database of
CS, some preliminary studies have investigated the effects of
aesthetic chills on subjects diagnosed with Major Depressive
Disorder (Jain etal., 2023a; Schoeller etal., 2023c). Prelimi-
nary data suggest that chills have an effect in mood disorders
(Jain etal., 2023a, b; Schoeller etal., 2023c). A first study
(Jain etal., 2023b) investigated whether experiencing aes-
thetic chills could improve reward learning in 103 people
with depressive symptoms, especially anhedonia, using vid-
eos known to elicit chills and the Probabilistic Reward Task
(PRT) (Pizzagalli etal., 2008). Anhedonic participants who
experienced chills showed a significant increase in reward
bias on the PRT compared with those who did not experi-
ence chills (p = 0.004), suggesting temporary mitigation of
blunted reward learning; however, no difference was seen in
non-anhedonic depressed participants (Fig.4E).
Another study on depression investigated whether expe-
riencing aesthetic chills could shift maladaptive self-beliefs
(Schoeller etal., 2023c). The findings revealed that chills
positively alter (depressogenic) negative self-schemas
(e.g., related to shame and self-acceptance) (Schoeller
etal., 2023c). Crucially, the phenomenology associated
to the belief change induced by CS closely resemble the
effects observed in psychedelic-assisted therapy, includ-
ing the characteristic associated phenomenology of emo-
tional breakthrough (EBI) (Roseman etal., 2019; Schoeller
etal., 2023c), psychological insight (PIS) (Peill etal., 2022;
Schoeller etal., 2023d), and ego dissolution (Nour etal.,
2016; Christov-Moore etal., 2023) (Fig.4). The surpris-
ing, involuntary nature of chills, their emergence without
conscious learning, and their underlying neurobiology
suggest a strong similarity to so-called primary states of
consciousness (Schoeller etal., 2023c), which are consid-
ered phylogenetically older states of consciousness often
associated with profound changes in beliefs, perceptions,
and behavior (Schoeller, 2016; Carhart-Harris etal., 2014).
Psychedelic experiences, such as those induced by psilo-
cybin, which stimulates 5-HT2a receptors, often result in
similar transcendent experiences, with shared characteristics

624 Cognitive, Affective, & Behavioral Neuroscience (2024) 24:617–630
Fig. 4 A and B. Individuals who experience aesthetic chills display
higher psychological insight (PIS) and emotional breakthrough (EBI)
compared with control (Schoeller etal.,2023c; 2023d). C and D. Chills
intensity is positively correlated with emotional breakthrough (EBI)
and emotional awareness (MAIA). E. AC was associated with a sig-
nificant change in reward bias (PRT) pre- and post-exposure to stimuli
in subjects compared to control (Jain etal., 2023b). F. Experience of
aesthetic chills was reliably associated with patterns of ego dissolution,
connectedness, and moral elevation (Christov-Moore etal., 2023)

625Cognitive, Affective, & Behavioral Neuroscience (2024) 24:617–630
to aesthetic chills (Schoeller etal., 2023c, d; Christov-Moore
etal., 2023). However, currently no direct evidence links
serotonin 5-HT2a receptors to AC.
Perhaps the most robust effect found across these studies
was AC effects on emotional valence, reliably generating
an “emotional drift,” i.e., predictable change in their emo-
tional state, including in participants with anhedonic symp-
toms (results summarized in Fig.5). In all of these studies,
participants are invited to report their level of emotional
valence and arousal before and after the experience (Jain
etal., 2023a, b; Schoeller etal., 2022). Participants who
experienced chills reported a larger shift in valence scores
from pre- to post-assessment across all groups compared
with those who did not. Interestingly, in the anhedonic study
(Jain etal., 2023b), the endpoint arousal and valence scores
for anhedonic patients after experiencing chills were almost
the same as those for non-anhedonic, depressed individuals,
who did not experience chills. This suggests that experienc-
ing chills brings the emotional state of anhedonic patients
closer to that of depressed patients without anhedonia. Addi-
tionally, there was no difference in endpoint affective scores
for the depressed group (non-anhedonic) compared with the
control group. This indicates that the emotional state of non-
anhedonic depressed individuals was similar to that of the
healthy control group by the end of the study.
Conclusions
Aesthetic chills offer a unique opportunity to study reward
learning in both healthy and clinical populations because
of its universal nature across cultures and its ability to be
consciously reported and physically measured. AC activate
a specific brain network involving the VTA and its dopa-
minergic projections to the mesocorticolimbic system,
crucial for reward and motivation processes. This network
includes NAcc, AMG, and frontal regions, such as the OFC
and VMPFC, primarily studied in the context of musical
chills. In line with behavioral evidence showing high sur-
prisal inhibits the AC process, dopamine plays a key role in
AC, encoding the precision of predictions and enhancing
sensory pleasure, motivation, and learning. AC are linked to
Fig. 5 Circumplex model depicts participants’ self-reported arousal
and valence before (empty dots) and after chills exposure (full dots).
Participants with high anhedonia (HA) are shown in red, low anhedo-
nia (LA) in orange, and healthy controls in blue. Solid lines indicate
participants who experienced chills, while dotted lines show those
who did not. Empty dots represent pre-exposure ratings, while filled
dots show post-exposure ratings. HA participants who experienced
chills shifted toward LA participants without chills and control pre-
exposure levels. This suggests chills exposure increased arousal and
valence in high anhedonia to approach levels seen in low anhedonia
and controls

626 Cognitive, Affective, & Behavioral Neuroscience (2024) 24:617–630
the reward cycle’s phases: wanting, liking, and learning, rep-
resenting a peak in consummatory pleasure and the onset of
the learning phase. The insular cortex’s role in interoception
significantly affects the emotional experience of chills, with
individual differences in AC propensity linked to personal-
ity traits, structural brain connectivity, and genetic factors.
Understanding AC’s neural underpinnings has implications
for psychiatric conditions like schizophrenia, depression,
and addiction, where dopamine’s role in precision encod-
ing of prediction errors is crucial. Preliminary studies sug-
gest AC can positively affect mood disorders, potentially
improving reward learning in anhedonia and altering nega-
tive self-beliefs.
Acknowledgment All authors contributed to the manuscript. This work
was partially funded by Tiny Blue Dot Foundation and a Joy Ventures
Research Grant. DAP was partially supported by R37MH068376 and
P50 MH119467. The content is solely the responsibility of the authors
and does not necessarily represent the official views of the National
Institutes of Health. The authors thank two reviewers for taking the
time to engage with the manuscript and their insightful remarks that
contributed to improving the manuscript.
Funding ’Open Access funding provided by the MIT Libraries’
Data Availability The data is available through the referenced articles.
Declarations
Conflict of interest In the past years, FS founded and received com-
pensation from BeSound SAS and Nested Minds LTD. Over the past 3
years, Dr. Pizzagalli has received consulting fees from Albright Stone-
bridge Group, Boehringer Ingelheim, Compass Pathways, Engrail
Therapeutics, Neumora Therapeutics (formerly BlackThorn Thera-
peutics), Neurocrine Biosciences, Neuroscience Software, Otsuka, Su-
novion, and Takeda; he has received honoraria from the Psychonomic
Society and American Psychological Association (for editorial work)
and from Alkermes; he has received research funding from the Brain
and Behavior Research Foundation, the Dana Foundation, Millennium
Pharmaceuticals, Wellcome Leap MCPsych, and NIMH; he has re-
ceived stock options from Compass Pathways, Engrail Therapeutics,
Neumora Therapeutics, and Neuroscience Software; he has a financial
interest in Neumora Therapeutics, which has licensed the copyright
to the probabilistic reward task through Harvard University. Dr. Piz-
zagalli’s interests were reviewed and are managed by McLean Hospital
and Partners HealthCare in accordance with their conflict of interest
policies. No funding from these entities was used to support the cur-
rent work, and all views expressed are solely those of the authors. All
other authors declare they have no competing interests. All other au-
thors have no conflicts of interest or relevant disclosures.
Open Access This article is licensed under a Creative Commons Attri-
bution 4.0 International License, which permits use, sharing, adapta-
tion, distribution and reproduction in any medium or format, as long
as you give appropriate credit to the original author(s) and the source,
provide a link to the Creative Commons licence, and indicate if changes
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copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.
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