Art reaches within: aesthetic experience, the self and the default mode network

Abstract

In a task of rating images of artworks in an fMRI scanner, regions in the medial prefrontal cortex that are known to be part of the default mode network (DMN) were positively activated on the highest-rated trials. This is surprising given the DMN's original characterization as the set of brain regions that show greater fMRI activity during rest periods than during performance of tasks requiring focus on external stimuli. But further research showed that DMN regions could be positively activated also in structured tasks, if those tasks involved self-referential thought or self-relevant information. How may our findings be understood in this context? Although our task had no explicit self-referential aspect and the stimuli had no a priori self-relevance to the observers, the experimental design we employed emphasized the personal aspects of aesthetic experience. Observers were told that we were interested in their individual tastes, and asked to base their ratings on how much each artwork “moved” them. Moreover, we used little-known artworks that covered a wide range of styles, which led to high individual variability: each artwork was rated highly by some observers and poorly by others. This means that rating-specific neural responses cannot be attributed to the features of any particular artworks, but rather to the aesthetic experience itself. The DMN activity therefore suggests that certain artworks, albeit unfamiliar, may be so well-matched to an individual's unique makeup that they obtain access to the neural substrates concerned with the self—access which other external stimuli normally do not get. This mediates a sense of being “moved,” or “touched from within.” This account is consistent with the modern notion that individuals' taste in art is linked with their sense of identity, and suggests that DMN activity may serve to signal “self-relevance” in a broader sense than has been thought so far.

Full text

FOCUSED REVIEW
published: 30 December 2013
doi: 10.3389/fnins.2013.00258
Art reaches within: aesthetic experience, the
self and the default mode network
Edward A. Vessel1*, G. Gabrielle Starr2and Nava Rubin3,4
1Center for Brain Imaging, New York University, New York, NY, USA
2Department of English, New York University, New York, NY, USA
3Center for Neural Science, New York University, New York, NY, USA
4ICREA and DTIC, Universitat Pompeu Fabra, Barcelona, Spain
Edited by:
Daniel S. Margulies, Max Planck
Institute for Human Cognitive and
Brain Sciences, Germany
Revi ewed by:
Luis M. Martinez, Spanish National
Research Council, Spain
Oshin Vartanian, University of
Toronto–Scarborough, Canada
Camilo J. Cela-Conde, Universidad de
las Islas Baleares, Spain
Dahlia Zaidel, University of
California, USA
Pablo Tinio, Queens College of the City
University of New York, USA
*Correspondence:
Edward A. Vessel, Ph.D. is a research
scientist at New York University’s Center
for Brain Imaging. Dr. Vessel’s research
combines brain imaging with behavioral
and computational approaches to study
how individuals are moved by, and get
pleasure from, visual experiences. He
received his PhD in Neuroscience from
the University of Southern California
in 2004.
ed.vessel@nyu.edu
In a task of rating images of artworks in an fMRI scanner, regions in the medial
prefrontal cortex that are known to be part of the default mode network (DMN) were
positively activated on the highest-rated trials. This is surprising given the DMN’s original
characterization as the set of brain regions that show greater fMRI activity during rest
periods than during performance of tasks requiring focus on external stimuli. But further
research showed that DMN regions could be positively activated also in structured
tasks, if those tasks involved self-referential thought or self-relevant information. How
may our findings be understood in this context? Although our task had no explicit
self-referential aspect and the stimuli had no aprioriself-relevance to the observers,
the experimental design we employed emphasized the personal aspects of aesthetic
experience. Observers were told that we were interested in their individual tastes, and
asked to base their ratings on how much each artwork “moved” them. Moreover, we used
little-known artworks that covered a wide range of styles, which led to high individual
variability: each artwork was rated highly by some observers and poorly by others.
This means that rating-specific neural responses cannot be attributed to the features of
any particular artworks, but rather to the aesthetic experience itself. The DMN activity
therefore suggests that certain artworks, albeit unfamiliar, may be so well-matched to an
individual’s unique makeup that they obtain access to the neural substrates concerned
with the self—access which other external stimuli normally do not get. This mediates
a sense of being “moved,” or “touched from within.” This account is consistent with
the modern notion that individuals’ taste in art is linked with their sense of identity, and
suggests that DMN activity may serve to signal “self-relevance” in a broader sense than
has been thought so far.
Keywords: neuroaesthetics, art, default mode network, self-relevance, medial prefrontal cortex (mPFC), fMRI,
visual, individual differences
INTRODUCTION
Theburgeoningfieldofneuroaesthetics attemptstoaddressthemysteriesofthehumanpreoc-
cupation with art by studying the underlying brain mechanisms. And, while understanding the
artistic creative process itself is certainly a formidable challenge, many of the open questions con-
cern the response to works of art by their viewers, listeners, and readers. What makes us so drawn
to certain artistic creations, so influenced and moved by them? In recent years, we have learned a
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Vessel et al. Art reaches within
KEY CONCEPT 1 | Neuroaesthetics
A multi-disciplinary field aimed at understanding the neural basis of aesthetic
experience and behavior. This includes interactions with art-objects as well
as aesthetic modes of interaction with non-art objects, such as faces, natural
objects, and scenes.
considerable amount from brain imaging studies about the neural
correlates of aesthetic experience and how they relate to sensory,
reward, and emotion neural processes (for reviews see Di Dio
and Gallese, 2009; Brown et al., 2011; Chatterjee, 2011; Nadal
and Pearce, 2011). One aspect that has so far received little inves-
tigation is that of individual differences: although it is widely
recognized that individuals can differ markedly in their aesthetic
response, previous research in neuroaesthetics tended to utilize
art pieces that were manipulated in a manner intended to have a
consistent effect on observers’ preferences or that were generally
highly regarded and often, widely known (e.g., the Mona Lisa). It
seems reasonable to expect that studying widely admired artwork
can help uncover the universal aspects of aesthetic experience. But
studying artworks that generate a diversity of responses can also
be valuable. Brain imaging can, in principle, be used to probe the
neural correlates of an experience in a manner dissociable from
the external stimuli that gave rise to this experience. In partic-
ular, it is possible to capitalize on the differences in individual’s
responses to artworks to search for commonalities in brain activ-
ity associated with the aesthetic experience itself, irrespective of
the stimulus properties of specific works of art that gave rise to it.
We have used this strategy in a recent study (Vessel et al., 2012)
and the results underscore its power and promise, by confirming
known results while at the same time revealing new and hitherto
unsuspected findings.
KEY CONCEPT 2 | Aesthetic experience
Aesthetics is a discipline concerned with the perception, appreciation, and
production of art. Aesthetic experiences, such as looking at paintings, lis-
tening to music or reading poems, are linked to the perception of external
objects, but not to any apparent functional use the objects might have.
Aesthetic experience involves more than preference, encompassing a vari-
ety of emotional responses ranging from beauty to awe, sublimity, and a
variety of other (often knowledge-based) emotions.
HIGHLY INDIVIDUALIZED RESPONSES TO VISUAL ART
As in much previous work in neuroaesthetics, we wished to com-
pare fMRI brain activity during observation of visual art that
elicited a high level of aesthetic appreciation with responses to
unappreciated artworks. But there was an important difference:
a primary goal of our study was to move away from the scenario
whereby different observers tend to respond similarly to the art
presented to them. (The rationale for this goal is explained below,
section Neural Correlates of Aesthetic Appreciation: Two Distinct
Activity Patterns). To achieve this, we collated a set of images of
two-dimensional visual artwork spanning a wide variety of peri-
ods, regions, styles and genres (fifteenth to twentieth century,
Western and Eastern works, including a range of representa-
tional and abstract genres). Importantly, although the images
were taken from museum collections, the artworks were not com-
monly reproduced and were therefore novel to our observers.
Moreover, the instructions to the participants emphasized that we
were interested in their own, individual response (rather than in
what may be the “normative” assessment of each artwork), and
that aesthetic experiences may come in a variety of forms: “The
paintings may cover the entire range from ‘beautiful’ to ‘strange’
or even ‘ugly.’ Respond on the basis of how much this image
moves you.” Each observer (N=16) was shown the same series
of 109 color artworks (in randomized order) while being scanned
using fMRI, and was asked to rate each artwork on a 4-point
scale according to these instructions. For a list of artworks and
other experimental details, see Vessel et al. (2012),Materials and
Methods and List of Artworks.
Analysis of the behavioral responses revealed that responses
were indeed highly individual: there was little agreement between
observers regarding how moving each painting was (0.13 average
correlation between the ratings of pairs of observers, computed
over the entire set of images; SD =0.17). This means that, on
average, each image was rated as highly moving by one sub-
set of observers and rated poorly by another subset of observers
(Figure 1). These results stand in contrast with the rather high
agreement obtained when observers make preference judgments
for real-world scenes [e.g., 0.46 between-observer correlation in
Vessel and Rubin (2010)] or attractiveness judgments for faces
[0.41 correlation between pairs of strangers in Bronstad and
Russell (2007);0.40inHoneköpp (2006)]. As we shall see below,
the low agreement between individuals in terms of their aesthetic
response is what allowed us to disentangle the external attributes
of specific stimuli from the internal (neural) states to which they
gave rise.
Another finding from the behavioral data that will play a
role in interpreting the brain imaging results is that, on average,
observers used the highest (“4”) rating significantly less than 25%
of the time (mean: 16.7%; SD =11.6%; 4 of 16 observers gave
more than 25% “4” responses). This is interesting given that there
was no special mention of the highest rating in our instructions,
and that in rating sensory/perceptual attributes (e.g., perceived
brightness) observers tend to distribute their responses across all
available options. That the observers in our experiments behaved
differently, and did not calibrate their responses so as to give a rat-
ing of “4” to roughly a quarter of the stimuli, suggests that they
reserved this response for images which met a certain internal
(and generally high) criterion.
NEURAL CORRELATES OF AESTHETIC APPRECIATION: TWO
DISTINCT ACTIVITY PATTERNS
The fMRI data were analyzed to compare responses during trials
in which the artworks were highly-rated with trials of low-rated
artwork. Contrasting brain activity between conditions that dif-
fer by the observers’ own responses, or performance, has been
used successfully in many domains of cognitive neuroscience
(e.g., studying neural correlates of memory encoding by contrast-
ing activity in subsequently-remembered and forgotten trials;
Brewer et al., 1998; Wagner et al., 1998). But in the context of
neuroaesthetics, extra care must be taken to dissociate neural
correlates of the aesthetic experience itself from other aspects of
brain activity elicited by the stimuli. As a simple example, sup-
pose observers are presented with a set of paintings comprised
mainly of portraits and landscapes, and suppose further that most
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Vessel et al. Art reaches within
FIGURE 1 | Aesthetic appreciation of visual art is highly individual.
(A) Two sample images from the set observers were shown. Images were
reproductions of museum artworks that are not commonly reproduced
(see Acknowledgments for image credits). Observers rated each image for
how much the artwork “moved” them on a scale of 1 (lowest) to 4
(highest). (B) Ratings of all 16 observers for the two images in (A). As was
typical for the artworks used in the experiment, observers differed widely
in their response to the pair of images. In particular, some observers rated
the top image (blue bars) to be highly moving, while others rated the
bottom image (red bars) to be highly moving. (For this bar plot, observers
were first sorted by their rating to the top image, then by their rating to
the bottom image).
of them happen to appreciate portraiture more than landscapes.
Face-selective brain regions would then likely show up in a con-
trast between highly-rated and low-rated trials, but is it warranted
to interpret their activity as pertaining to aesthetic experience?
In this case, the (conjured) agreement in aesthetic preference
is simple enough, and our knowledge of face-selectivity in the
brain sound enough, to easily discern that the activity can be
explained by other aspects of the stimuli (the types of objects
depicted). But in fact, such potential confounds are present when-
ever there is high agreement between observers about the art: the
highly-rated and low-rated trials in such cases correspond to dif-
ferent sets of (artwork) stimuli, which may well result in some
differential activation unrelated to the aesthetic experience they
produce. Conversely, high variability between different observers’
aesthetic judgments alleviates the potential confound: in the limit
of completely uncorrelated ratings, the highly-rated trials and
the low-rated trials contain identical sets of stimuli (each con-
tributed by a different observer to each set). This was therefore
our motivation in creating a stimulus set that generated highly
individualized responses: rating-specific neural responses would
then not be attributable to the features of any particular art-
works, thus allowing us to isolate neural correlates of the aesthetic
experience itself.
We performed several different analyses, using both statisti-
cal activation maps and regions of interest (ROIs) generated from
the same data set or from separate “localizer” runs. We first cre-
ated whole-brain activation maps by contrasting the group-level
brain response to the most moving trials (rated as “4”) with the
responses to the least moving trials (rated as “1”). This “4-vs.-1”
analysis revealed a network of regions distributed across poste-
rior, anterior, and subcortical structures (Figure 2A; note that, in
addition, extensive portions of visual sensory cortex were strongly
activated by all stimuli, but the magnitude of response did not dif-
fer by rating; Figure 2B). This is consistent with conclusions from
previous research using a variety of stimuli that multiple brain
regions are engaged during aesthetic appreciation (Aharon et al.,
2001; Blood and Zatorre, 2001; Cela-Conde et al., 2004; Kawabata
and Zeki, 2004; Vartanian and Goel, 2004; Jacobsen et al., 2006;
Koelsch et al., 2006; Di Dio et al., 2007; Kim et al., 2007; Yue et al.,
2007; Calvo-Merino et al., 2008; Fairhall and Ishai, 2008; Cupchik
et al., 2009; Ishizu and Zeki, 2011; Lacey et al., 2011; Salimpoor
et al., 2011; Jacobs et al., 2012; Kuhn and Gallinat, 2012). Note
that the large inter-observer variability in behavioral responses to
our stimulus set means that the common (group-level) activation
in the 4-vs.-1 contrast must reflect effects of the aesthetic experi-
ence itself, i.e., it could not be due to any attributes of particular
art stimuli that gave rise to this experience. This is because, at
the group level, the set of highly rated trials consisted mostly of
the same images as the poorly rated trials (recall that for every
image rated as high by one observer there was, on average, another
observer that rated it as low). This also means, however, that
our approach is more restrictive than that in some other studies,
which could give rise to differences in the activations observed.
We will not go here into details of comparing and contrasting the
loci of activation with those previously reported in the literature
(see Vessel et al., 2012). Instead, we focus below on those aspects
most relevant for a novel and intriguing finding: the activation by
highly moving stimuli of the default mode network (DMN).
The bar graphs surrounding the activation map in Figure 2A
show fMRI response magnitude as a function of observers’ ratings
for select ROIs, revealing that different ROIs exhibited distinct
response patterns. Moreover, ROIs could be grouped in two main
categories: for one set of ROIs, response magnitudes varied lin-
early with rating (right-side panels: lITS, lPHC, and lSTR). The
linear response pattern was observed in different variations in
terms of its relation to the baseline (“rest”) level: in occipi-
totemporal cortex, higher ratings were accompanied by linearly
changing BOLD signals that either increased well above a resting
baseline (lITS, and lPHC) or, in one case, decreased well below
it (rSTG, not shown). In subcortical regions, fMRI activity was
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Vessel et al. Art reaches within
FIGURE 2 | Distinct patterns of response to artworks as a function of
their ratings in a distributed network of brain regions. (A) Center panel:a
whole-brain analysis contrasting trials on which observers rated artworks as
highly moving (4) vs. trials where artworks were given the lowest rating (1),
showing a lateral (top) and ventral (middle) view of an inflated left
hemisphere, and a coronal section (bottom) through the striatum (data
thresholded at a False Discovery Rate of q<0.05 in volumetric space and
projected on a hemisphere of a single observer for visualization). Right-side
panels: a linear increase with rating was observed for the activation loci in
occipitotemporal cortex and some subcortical loci (shown here: left inferior
temporal sulcus, lITS; left parahippocampal cortex, lPHC; left striatum, lSTR;
see (Vessel et al., 2012) for additional ROIs and further detail). Left-side
panels: a nonlinear, “step”-like response pattern was observed in the anterior
activation loci; responses did not differ for images rated 1, 2, or 3, but were
significantly elevated for images rated 4 (shown here: left inferior frontal
gyrus par triangularis,lIFGt; left lateral orbitofrontal cortex, lLOFC).
(B) Extensive portions of early visual cortex were strongly activated by all
paintings, but the magnitude of fMRI response did not differ by rating.
suppressed below its resting level for low-rated stimuli and rose
progressively to above-rest for highly rated stimuli [lSTR, bottom
right panel; PRF, not shown; see Vessel et al. (2012) for ROIs not
shown here and further details]. Since the 4-vs.-1 contrast selects
for regions that responded differently to trials rated “4” compared
with trials rated “1,” the pattern of response for the intermediate
ratings of 2 or 3 in these regions is aprioriunknown. It is there-
fore noteworthy that responses in these ROIs followed a linear
trend so closely. Moreover, regions whose response patterns were
significantly non-linear all showed the same distinct pattern, as
follows.
A second category of regions revealed by the 4-vs.-1 contrast
were characterized by a distinct “step” pattern: fMRI responses
in those regions did not differ significantly for images rated 1,
2, or 3; only for the highest (4) rating was there a significant
difference in response magnitude, and it was marked and dra-
matic (Figure 2A, left-side panels; see Vessel et al., 2012 for other
examples; see also below, Figure 3). We performed several addi-
tional analyses in order to examine more closely the nature and
spatial distribution of these nonlinear “step” responses. A whole-
brain analysis contrasting the highest-rated trials with an average
of all other trials (4-vs.-321; Vessel et al., 2012)gaveusmore
power to detect regions that may not have reached the signifi-
cance threshold in the 4-vs.-1 contrast due to the lower number
of trials. A conjunction was subsequently computed to specifically
capture the regions that, while showing a differential response
to the highest-rated stimuli (“4”), showed no significant differ-
ences in responses within the lower ratings (1, 2, and 3). The
resulting statistical map contained large swaths of highly signif-
icant differences in several regions known to be part of the DMN,
and further examination indicated that the pattern of responses
in those regions consisted of a strong deactivation in trials rated
1, 2, or 3 (with no significant differences in magnitude), which
was greatly alleviated or even eliminated in the highest-rated tri-
als [“4”; see Vessel et al. (2012), Figure 6]. To better underscore
the commonalities and differences from what is currently known
about the DMN, below we represent our results in a different for-
mat than before, which is modeled after that used in the DMN
literature.
Figure 3A shows statistical activation maps contrasting the
task-induced fMRI responses with “Rest”—intervals interspersed
between the trials when only a blank screen was shown—overlaid
on the inflated surface of the left hemisphere. The maps were
generated separately for each of the four sets of trials correspond-
ing to the four possible ratings (from top to bottom: 1-vs.-Rest
to 4-vs.-Rest). Large regions in occipital cortex, as well as por-
tions of parietal and frontal cortex, showed activation above
rest for all four rating levels (warm colors, red-yellow). The
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FIGURE 3 | The default mode network (DMN) deactivation during task
performance is alleviated when viewing highly moving artworks.
(A) Lateral (left) and medial (right) views of an inflated cortical surface are
overlaid with statistical maps comparing fMRI responses during task (viewing
and rating of artworks) vs. “rest” periods. Maps were computed separately
for trials from each of the four possible ratings, 1 (top) to 4 (bottom). The warm
colors indicate greater fMRI response during task; the cool colors indicate
greater response during rest (“deactivation”; data were thresholded at a False
Discovery Rate of q<0.05 before projection onto one observer’s inflated
cortex). In trials rated 1, 2, or 3 (top three panels) there were deactivations in
medial prefrontal cortex (MPFC), posterior cingulate cortex (PCC), lateral
temporal cortex (LTC), temporoparietal junction (TPJ), and superior frontal
gyrus (SFG). The suppression was greatly reduced for the highest-rated trials
(4; bottom panel). (B) The spatial pattern of deactivation during the lower-rated
trials (1–3) closely resembles that of the default mode network [DMN; image
adapted with permission from Fox et al. (2005) Copyright 2005 National
Academy of Sciences, U.S.A.]. (C) Average fMRI response in the MPFC region
of interest (ROI) was markedly and uniformly below rest for trials rated 1, 2, or
3, but was not different from rest for the highest-rated trials (4). (D) fMRI
signal timecourse in the MPFC for the lower-rated trials (cyan) and the
highest-rated trials (magenta). Note that activity initially fell below its level
during rest also for the highest rated trials, yet it rapidly recovered and then
proceeded to increase above rest level. The fMRI response used for both C
and D was estimated from an ROI defined via a contrast of the response on
“4” trials vs. the other trials (4-vs.-321), conjoined with a map of regions
showing no difference in the low-rated trials. The timecourses for each rating
level were extracted by modeling the average timecourse from this ROI as a
set of four finite impulse response functions (Dale and Buckner, 1997).
cool colors (blue–green) denote regions that showed a reduced
fMRI signal during viewing and rating of the artworks, com-
pared to during rest. For the sets of trials rated 1, 2, or 3
(top three panels), extensive regions of reduced activity can be
seen; their anatomical loci and spatial distribution closely resem-
bles that observed in studies that contrasted activity during a
wide range of cognitive and perceptual tasks with periods of
rest (Shulman et al., 1997; Simpson et al., 2001), shown in
Figure 3B (adapted from Fox et al., 2005). Specifically, reduced
activity was observed in the medial prefrontal cortex (MPFC),
posterior cingulate cortex (PCC), precuneus (PCu), temporo-
parietal junction (TPJ), lateral temporal cortex (LTC) and supe-
rior frontal gyrus (SFG). Studies of blood flow and oxygen
utilization indicate that the baseline level of these regions—that
measured during rest—corresponds not to a lack of activity, but
rather to activity associated with an ongoing, organized “default
mode” of brain processing, which is suspended during perfor-
mance of tasks that require externally directed attention (Gusnard
and Raichle, 2001; Raichle et al., 2001). The reduced fMRI
response in regions of this default mode network (DMN) during
task performance is therefore widely referred to as deactivation
(although the mechanisms giving rise to it are not fully
understood).
KEY CONCEPT 3 | Default mode network
A network of brain regions typically found to be suppressed when observers
engage in externally oriented tasks, which includes the medial prefrontal
cortex (MPFC), posterior cingulate cortex (PCC), temporo-parietal junction
(TPJ), lateral temporal cortex (LTC), superior frontal gyrus (SFG) and the
hippocampus. Patterns of spatial correlation measured in the absence of
directed tasks (resting state fMRI) support this network structure and sug-
gest that the DMN is composed of midline hub regions (MPFC, PCC) and
two subsystems.
In contrast with the pattern observed for trials rated 1, 2, and
3, DMN regions showed markedly less deactivation during the
highest-rated trials (“4”; bottom panel in Figure 3A). Indeed, in
some portions of the DMN—most notably, in the MPFC—the
deactivation seems all but gone. ROI analysis confirmed that the
MPFC was strongly and uniformly deactivated for lower-rated
trials (1–3), but not at all during those trials when the artworks
were given the highest rating (4), resulting in a step-like response
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Vessel et al. Art reaches within
pattern [Figure 3C; for plots of several other DMN components,
see Vessel et al. (2012)].
THE DEFAULT MODE NETWORK AND SELF-REFERENTIAL
MENTAL PROCESSING
A defining characteristic of the DMN—indeed, how it was
discovered—is that it is suppressed when observers are engaged in
demanding tasks that require them to focus on external stimuli,
compared with its level of activity during passive viewing or peri-
ods of rest between the tasks (Shulman et al., 1997; Buckner et al.,
2008). The ubiquity of DMN deactivation during many different
cognitive tasks with a variety of stimuli and response demands,
along with studies of functional connectivity during rest, have
led to the view that the DMN represents a “task-negative”
network of brain regions that normally functions in an anti-
correlated manner from “task-positive” networks such as sensory-
semantic pathways and the dorsal attention network (Ingvar,
1979; Corbetta and Shulman, 2002; Fox et al., 2005; Buckner
and Carroll, 2007). The finding that, in our own task, the cor-
tical regions that overlap with previously identified components
of the DMN (MPFC, PCC, TPJ, LTC) showed significant deacti-
vation below their baseline (rest) level during a majority of the
trials, those rated 1–3 (Figure 3A, top three panels) is therefore
consistent with what is known about the DMN. From this same
perspective, the dramatic reduction of deactivation in the trials
rated “4” (Figure 3A, bottom panel) and its complete absence
in the MPFC (Figure 3C) therefore seems puzzling. But consid-
eration of additional findings about the DMN offers a potential
explanation.
Following its initial identification, further research showed
that the DMN regions can maintain their baseline activity not
only during periods of (waking) rest, but that they can escape
deactivation, or even become activated above baseline, also dur-
ing the performance of structured tasks. Ventral portions of the
MPFC are involved in affective decision making processes, includ-
ing (but not restricted to) encoding the subjective value of future
rewards and assessing the emotional salience of stimuli (Bechara
et al., 1999; Knutson et al., 2005; Kringelbach, 2005; Kable and
Glimcher, 2007; Schmitz and Johnson, 2007; Levy and Glimcher,
2011). The anterior and dorsal portions of MPFC are active in
tasks involving self-knowledge such as making judgments about
oneself as well as about close others (family and friends), self-
relevant moral decision-making (Reniers et al., 2012)andin
“theory of mind” tasks that require gauging others’ perspectives
(Zysset et al., 2002; Ochsner et al., 2004, 2005; Amodio and
Frith, 2006; Mitchell et al., 2006; Enzi et al., 2009; Andrews-
Hanna et al., 2010; Whitfield-Gabrieli et al., 2011). The PCC and
medial temporal lobe regions are active during tasks that involve
retrieving autobiographical memories as well as planning or sim-
ulating the future (Buckner and Carroll, 2007; Buckner et al.,
2008; Andrews-Hanna et al., 2010).
The DMN is thus emerging as a highly interconnected network
of brain regions that support self-referential mental processing
(Northoff et al., 2006). Such processing is, of course, ubiqui-
tous in everyday life and is undoubtedly important for normal
functioning. In experimental settings it can occur spontaneously
(e.g., as “mind wandering” during periods of rest) but it can also
be triggered in structured tasks, by external stimuli that cause
observers to draw on self-referential information (intentionally or
automatically), or to engage in inwardly focused attention. Could
this have been the case with the images that our observers rated
as “highly moving”? We propose that the answer is yes, as detailed
in the account provided below.
KEY CONCEPT 4 | DMN and self-referential mental processing
Structured tasks can activate the DMN if they require some self-referential
processing (e.g., introspection, autobiographical memory recall). Similarly, it
is presumed that the DMN is metabolically active during baseline non-task
periods (e.g., fixation or “rest” conditions) because observers engage in
such processes spontaneously.
INTENSE AESTHETIC EXPERIENCE: A (NON-PERSONAL)
EXTERNAL STIMULUS REACHES THE SELF
Taste in art is highly individual and can be hard to predict by
even the most well-informed bystander (e.g., Bell and Koren,
2007), yet it is strongly felt. Indeed, many individuals consider
their artistic taste to be an important part of their identity, their
sense of who they are. This is not limited to connoisseurs of
“high art”: from teenagers whose tumultuous struggles for self-
determination are conducted to the soundtrack of meticulously
compiled music collections, to adults of all ages who repeatedly
turntotheirfavoritegenresoffictionorfilmtoescapethetedium
of their daily lives, our taste in art is intertwined with the choices
we make about how to spend our time and with whom to spend
it, and as such it is part of who we are. How does this come about?
What gives certain artworks their mysterious “pull”? Our data
say nothing about this in terms of the attributes of the artwork
itself. (Whether this will remain a mystery forever or may yield to
future research is an interesting question that will not be discussed
here). But our results suggest that the strong effect of certain art-
works can be understood in terms of the physiological state they
generate and how this state is experienced, or interpreted, by the
observer.
We propose that certain artworks can “resonate” with an
individual’s sense of self in a manner that has well-defined physi-
ological correlates and consequences: the neural representations
of those external stimuli obtain access to the neural substrates
and processes concerned with the self—namely to regions of the
DMN. This access, which other external stimuli normally do not
obtain, allows the representation of the artwork to interact with
the neural processes related to the self, affect them, and possi-
bly even be incorporated into them (i.e., into the future, evolving
representation of self). This hypothesis gains considerable sup-
port from the way that the fMRI responses evolved over time in
the MPFC, the region most associated with evaluations of self-
relevance. As can be seen from the time course plots in Figure 3D,
immediately following stimulus presentation the fMRI signal in
the MPFC fell below baseline for all images, i.e., also for those
images that were (later) rated by the observer as highly mov-
ing (4). Thus, the initial predisposition of this DMN region was,
for all external stimuli, to deactivate. But in contrast with the
MPFC response to the artworks rated 1, 2, or 3, which was sup-
pressed during image presentation and remained below baseline
throughout the subsequent recovery (Figure 3D,cyanline),in
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Vessel et al. Art reaches within
the 4-rated trials activity started recovering soon after stimulus
presentation and then continued to rise above baseline (magenta
line). This is reminiscent of the MPFC recovery from deactivation
observed when a highly self-relevant stimulus such as one’s own
name is presented in a stream of self-irrelevant stimulation, as in
the “cocktail party effect” (e.g., Cherry, 1953; Bargh, 1982; Wood
and Cowan, 1995; Perrin et al., 2005). But why should a hitherto
unseen artwork, that has no aprioripersonal relevance for the
observer, have this effect of engaging the DMN system? Again, we
cannot say what attributes make specific artworks so exquisitely
attuned to an individual’s unique makeup. And yet this hypothesis
provides a coherent explanation of our data in that it is consistent
not only with what we know about the DMN, but also with what
we know about art.
Great art is, almost by definition, universal: the wide appeal it
commands comes from a connection with fundamental aspects of
human nature and human cognition (Kant, 1790/1987). Yet, at its
best, art in any of its forms—visual art, music, literature, etc.—
can feel strikingly personal. Intense aesthetic experience often
carries with it a sense of intimacy, “belonging,” and closeness with
the artwork. It may be hard to imagine that the experiences of
our observers, lying in an MRI scanner watching images of little-
known artworks selected by an experimenter who knew nothing
about them, reached the profound levels that give art its intense
power. And yet the data are compellingly in line with the phe-
nomenology of aesthetic experience: in the small subset of the
trials that observers rated as “highly moving,” DMN regions and
in particular the MPFC were released from deactivation and even
activated above baseline, a hallmark of self-relevant neural pro-
cessing. Perhaps the key to this was in our experimental design,
which relied on a stimulus set that maximized individual differ-
ences in behavioral response. As already mentioned, the original
motivation for this design was to measure neural correlates of
aesthetic experience in the absence of potential confounds with
effects of stimulus attributes. But the emphasis on a diversity of
artistic styles and topics may have, serendipitously, also increased
the chances that a few of the artworks resonate with each observer
in a particularly powerful way.
Note that the “resonance” between certain artworks and
observers’ sense of self that, we propose, occurs during intense
aesthetic experience, is different from explicitly self-referential
emotions such as pride, shame, guilt and embarrassment, as these
involve an appraisal of self-responsibility for an event (Silvia,
2012). It is also interesting to note in this context that intense aes-
thetic experience can sometimes be thrillingly bidirectional: not
only does the perceiver feel as if they understand the artwork,
but there is a sense that the artwork “understands” the perceiver,
expressing one’s own innermost thoughts, feelings, or values. The
latter sense points to the possibility that it is the artist, not the art-
work, who has understood something deep about the perceiver’s
experience; hence the intensely personal connection felt by many
people toward favorite artists who are, after all, strangers to them.
In some cases, this bidirectionality is accompanied by a perceived
or real congruence with the intentions of the artist (Jucker and
Barrett, 2011; Tinio, 2013). Thus, unlike in self-referential emo-
tions, in aesthetic experience the relation to others is not focused
on appraisal but on a sense of understanding, gained insight and
meaning. The extraction of meaning has been suggested previ-
ously as a primary factor of aesthetic experience (Martindale,
1984; Leder et al., 2004). But, while those authors suggest that
an appeal to self-related information is but one way in which
viewers extract meaning from artwork, the release of the DMN
from suppression on only the trials rated “4” suggests that, in fact,
self-relevance is an integral aspect of intensely moving aesthetic
experience.
What internal signal did the observers use to provide their
responses? It is tempting to think that they were able to detect
the unusual release from deactivation in the DMN when view-
ing artworks which they (later) rated “highly moving,” and that
they based their responses on this internal signal. Indeed, the
MPFC and PCC respond to self-relevant information even when
there is no explicit requirement to evaluate self-relevance, and
such information is in fact task-irrelevant (Moran et al., 2009;
Reniers et al., 2012). Perhaps observers conferred the highest
rating on those artworks that invoked in them a sense of self-
relevance, even though they were not instructed to do so, and
may well be unable to explicitly state this as their strategy. Yet
given the poor temporal information provided by fMRI, it is too
early to rule out the possibility that responses on the “4” trials
arose from posterior regions whose activity grew linearly with
rating or from other frontal regions that showed positive activa-
tion for only the “4” trials, and that the release from suppression
in the DMN for highly moving artworks occurred subsequent
to the evaluation. A recent MEG study of aesthetic appreciation
reported coherence between frontal midline, posterior and tem-
poral regions that was detectable 1 s after onset of images deemed
“beautiful” (1000–1500 ms analysis window) but not in an ear-
lier epoch (250–750 ms; Cela-Conde et al., 2013). This finding is
consistent with our proposal that the release of the DMN from
suppression for intensely moving artworks occurs subsequent to
an initial perceptual and semantic analysis, and early enough to
be a potential basis for response selection; however, it leaves open
the question of how, in time, explicit evaluation relates to these
dynamics.
A coactivation of the DMN and stimulus-driven sensory sys-
tem as we have observed for strongly moving aesthetic experi-
ences has so far not been reported in other contexts. Yet, if our
self identity is to be influenced by the world we inhabit, it may
be that similar moments should occur with greater frequency
than would be expected based on the current conceptualization
of the DMN as a network that is invariably suppressed during
mental activity which is directed at the external world. It may
be that our findings are just the “tip of the iceberg”—i.e., that
instances of resonance between external stimuli and internal, self-
related processing are more commonplace in daily life than what
has so far been captured in fMRI experiments in the labora-
tory. By that view, much of our existence may be well-served by
switching between periods of dominance of externally-directed
(“task-positive”) brain networks over the DMN and vice versa,
but those periods are punctuated by significant moments when
our brains detect a certain “harmony” between the external
world and our internal representation of the self—allowing the
two systems to co-activate, interact, influence and reshape each
other.
Frontiers in Neuroscience www.frontiersin.org December 2013 | Volume 7 | Article 258 |7

Vessel et al. Art reaches within
ACKNOWLEDGMENTS
This project was supported by an ADVANCE Research Challenge
Grant funded by the NSF ADVANCE-PAID award # HRD-
0820202 and by the Andrew W. Mellon Foundation (as a New
Directions Fellowship). Cloud Study, c. 1822. John Constable.
Oil on paper, 29.21 ×48.26 cm. The Frick Collection, Bequest of
Henrietta E.S. Lockwood in memory of her father and mother,
Ellery Sedgwick and Mabel Cabot Sedgewick, 2001.3.134. An
Ecclesiastic, c. 1874. Mariano José Maria Bernardo Fortuny y
Carbo. Oil on panel, 19 ×13 cm. The Walters Art Museum,
Bequest of William T. Walters. 37.150.
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Conflict of Interest Statement: The authors declare that the research was con-
ducted in the absence of any commercial or financial relationships that could be
construed as a potential conflict of interest.
Received: 10 April 2013; accepted: 09 December 2013; published online: 30 December
2013.
Citation: Vessel EA, Starr GG and Rubin N (2013) Art reaches within: aesthetic expe-
rience, the self and the default mode network. Front. Neurosci. 7:258. doi: 10.3389/
fnins.2013.00258
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