Brain and Cognition 87 (2014) 52–56

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Brain and Cognition

journal homepage: www.elsevier.com/locate/b&c

Neural correlates of viewing paintings: Evidence from a quantitative

meta-analysis of functional magnetic resonance imaging data
Oshin Vartanian a,⇑, Martin Skov b,c
a
University of Toronto—Scarborough, Toronto, ON, Canada
b
Danish Research Centre for Magnetic Resonance, Copenhagen University Hospital Hvidovre, Copenhagen, Denmark
c
Decision Neuroscience Research Group, Copenhagen Business School, Copenhagen, Denmark

a r t i c l e i n f o a b s t r a c t

Article history: Many studies involving functional magnetic resonance imaging (fMRI) have exposed participants to
Accepted 8 March 2014 paintings under varying task demands. To isolate neural systems that are activated reliably across fMRI
Available online 4 April 2014 studies in response to viewing paintings regardless of variation in task demands, a quantitative meta-
analysis of fifteen experiments using the activation likelihood estimation (ALE) method was conducted.
Keywords: As predicted, viewing paintings was correlated with activation in a distributed system including the
Visual art occipital lobes, temporal lobe structures in the ventral stream involved in object (fusiform gyrus) and
Aesthetics
scene (parahippocampal gyrus) perception, and the anterior insula—a key structure in experience of emo-
Emotion
Default mode network
tion. In addition, we also observed activation in the posterior cingulate cortex bilaterally—part of the
Scene perception brain’s default network. These results suggest that viewing paintings engages not only systems involved
in visual representation and object recognition, but also structures underlying emotions and internalized
cognitions.
Ó 2014 Elsevier Inc. All rights reserved.

1. Introduction structures underlying visuospatial processing such as the parietal

Two theoretical models have proposed that aesthetic experi-
ence associated with exposure to works of art arises as a function
of the engagement of a distributed set of perceptual, cognitive, and
emotional processes (Chatterjee, 2003; Leder, Belke, Oeberst, &
Augustin, 2004). Uncovering the neural systems that underlie this
distributed functional architecture is one of the major goals of the
field of neuroaesthetics (Skov & Vartanian, 2009). Descriptive re-
views of studies to date have indicated that aesthetic experience
in response to viewing artworks is indeed a function of a distrib-
uted set of brain areas, each of which is hypothesized to underlie
a different component process modulated by task demands (see,
e.g., Cela-Conde et al., 2011; Nadal, Munar, Capó, Rosselló, &
Cela-Conde, 2008). For example, whereas explicit instruction to fo-
cus on subjective emotions while viewing artworks is more likely
to activate structures underlying the experience of visceral emo-
tion (e.g., anterior insula), explicit instruction to examine the ob-
jects that make up scenes in paintings is more likely to activate
⇑ Corresponding author. Address: Department of Psychology, University of
Toronto—Scarborough, 1265 Military Trail, Toronto, ON M1C 1A4, Canada.
E-mail address: oshinv1@mac.com (O. Vartanian).
lobes (Cupchik, Vartanian, Crawley, & Mikulis, 2009).
Recently, Brown, Gao, Tisdelle, Eickhoff, and Liotti (2011) con-
ducted a large-scale meta-analysis of neuroimaging studies of po-
sitive-valence aesthetic appraisal across sensory modalities. Their
aim was to highlight regions reliably activated during appraisal of
the valence of perceived objects in the visual, auditory, gustatory
or olfactory domains. They were motivated by their search for
the core processes underlying aesthetic evaluation. As a result,
although some studies that had used paintings as stimuli were also
included, the selected studies had necessarily used tasks involving
aesthetic evaluation, thereby excluding studies in which paintings
had been used to study sensory processing, decision making alone,
or passive viewing. In contrast, here we subject data from func-
tional magnetic resonance imaging (fMRI) studies in which partic-
ipants viewed paintings to a quantitative meta-analysis. Our aim
was different from Brown et al.’s in that we were motivated to re-
veal brain regions activated reliably as a function of exposure to
paintings regardless of variation in task demands (e.g., passive
viewing, active ratings, etc.). Given that paintings constitute a
key stimulus set across studies of neuroaesthetics, isolating the
neural structures that are activated in response to viewing them
will be a useful tool in teasing apart task-related and stimulus-re-
lated effects in future studies.

http://dx.doi.org/10.1016/j.bandc.2014.03.004
0278-2626/Ó 2014 Elsevier Inc. All rights reserved.
 O. Vartanian, M. Skov / Brain and Cognition 87 (2014) 52–56 53

Table 1
List of studies included in the meta-analysis.

Study N Peaks Analysis Task

Kawabata and Zeki (2004) 10 3 Contrast Aesthetic judgment
Vartanian and Goel (2004) 12 9 Parametric Aesthetic judgment
Fairhall and Ishai (2008) 12 12 Contrast Recognition
Kirk, Skov, Hulme, Christensen, and Zeki (2009) 14 7 Contrast Aesthetic judgment
Cupchik et al. (2009) 16 4 Contrast Active viewing
Wiesmann and Ishai (2010) 24 11 Contrast Recognition
Lebreton, Jorge, Michel, Thirion, and Pessiglione (2009) 20 26 Contrast Mixed judgment
Harvey et al. (2010) 87 6 Parametric Passive viewing
Lacey et al. (2011) 8 15 Contrast Active viewing
Kirk, Harvey, and Montague (2011) 40 5 Parametric Passive viewing
Huang, Bridge, Kemp, and Parker (2011) 14 6 Contrast Unrestricted
Ishizu and Zeki (2012a) 21 2 Parametric Aesthetic judgment
Vessel et al. (2012) 16 19 Contrast Active viewing
Ishizu and Zeki (2012b) 21 13 Contrast Aesthetic judgment
Silveira et al. (2012) 15 16 Contrast Affective judgment

Note. N = number of participants. Peaks = Foci of activation for selected contrast or parametric analysis. Aesthetic judgment = making a preference or beauty judgment,
passive viewing = viewing not coupled with instruction to form an attitude, active viewing = viewing coupled with instruction to form an attitude, mixed judgment = making
aesthetic and other judgments, recognition = memory task, unrestricted = subjects instructed to view each image as they pleased, affective judgment = judging the extent to
which one is affected by the painting.

We focused on the visual modality and paintings specifically for
two reasons. First, we were able to locate a sufficient number of
fMRI studies in this area to enable a meta-analysis. Second, both
models discussed above (Chatterjee, 2003; Leder et al., 2004) are
based primarily on vision. For this latter reason, we were able to
make predictions regarding the involvement of specific neural
structures across studies. First, we hypothesized that viewing
paintings would activate regions of the visual cortex involved in
processing of early, intermediate, and late visual features that
underlie painting perception, including color and form (Chatterjee,
2003; Greenlee & Tse, 2008; Wandell, Dumoulin, & Brewer, 2009).
Second, we hypothesized that structures involved in the percep-
tion of objects and spaces would also be activated, specifically
structures in the ventral stream tuned towards object recognition
(Grill-Spector & Sayres, 2008; Kanwisher & Yovel, 2009; Mishkin,
Ungerleider, & Macko, 1983; Ungerleider & Mishkin, 1982). Third,
it is almost universally assumed that a primary objective of art is
to evoke affective responses in the viewer, although whether the
brain’s emotion and reward systems would be activated across
studies with varying instructions remains an open question. Con-
veniently, the structures known to play a role in emotion and re-
ward are well established (Montague & Berns, 2002), including
the nucleus accumbens (Aharon et al., 2001), the ventral striatum
(Kampe, Frith, Dolan, & Frith, 2001), the orbitofrontal cortex
(O’Doherty et al., 2003; Winston, O’Doherty, Kilner, Perrett &
Dolan, 2006), and the insula (see Di Dio & Gallese, 2009). Therefore,
our third and exploratory hypothesis was whether viewing paint-
ings would activate the brains’ reward and/or emotion systems.
2. Material and methods
Studies were selected by conducting Boolean searches in Pub-
Med using the terms ‘‘painting’’, ‘‘art’’, ‘‘aesthetic’’, ‘‘beauty’’,
‘‘MRI’’, ‘‘brain’’, and ‘‘neuroimaging’’ in February 2014. This set of
papers was augmented by others in which participants viewed
paintings under non-aesthetic conditions. Extracted fMRI studies
were subsequently checked to ensure that (a) they involved view-
ing paintings,1 (b) they were comprised of neurologically healthy
and adult participants, (c) the analyses were whole brain rather than
1
For this meta-analysis we only selected studies that used paintings, resulting in
the exclusion of fMRI studies which had used sculptures or geometric patterns as
stimuli (e.g., Di Dio, Macaluso, & Rizzolatti, 2007; Jacobsen, Schubotz, Höfel, & von
Cramon, 2005).
exclusively region-of-interest (ROI), and (d) the complete list of acti-
vation peaks (i.e., foci) was published in the paper or made available
to us. This resulted in fifteen experiments, involving a total of 330
participants and 166 peaks of activation (Table 1).
2.1. Activation likelihood estimation
Our meta-analysis was conducted using the activation likeli-
hood estimation method (ALE) (Turkeltaub, Eden, Jones, & Zeffiro,
2002). ALE is a quantitative meta-analysis technique that high-
lights brain regions that are activated reliably across studies. Much
like traditional meta-analytic approaches, ALE’s advantages
include ‘‘seeing the ‘‘landscape’’ of a research domain, keeping sta-
tistical significance in perspective, minimizing wasted data,
becoming intimate with the data summarized, (and) asking
focused research questions’’ (Rosenthal & DiMatteo, 2001, p. 59).
In addition, the method has been shown to provide a reliable
means for conducting coordinate-based meta-analyses of func-
tional imaging data (Eickhoff, Bzdok, Laird, Kurth, & Fox, 2012).
We believe that meta-analyses and qualitative reviews are comple-
mentary, jointly providing windows into common and nuanced as-
pects of a domain, respectively.
ALE’s approach involves comparing activation likelihoods calcu-
lated from observed activation foci with a null distribution of ran-
domly generated activation likelihoods. It pools peak activation
coordinates across studies that have investigated an effect of inter-
est (Laird et al., 2005). For this meta-analysis all coordinates were
renormalized to Talairach space using the icbm2tal transformation
(Lancaster et al., 2007) implemented in the GingerALE 2.0 toolbox
(http://brainmap.org; Research Imaging Center of the University of
Texas Health Science Center, San Antonio, TX). The resulting coor-
dinates were used to generate ‘‘activation likelihoods’’ for each
voxel in the brain. For each focus, ALE computes each voxel as a
function of its distance from that focus using a three-dimensional
Gaussian probability density function centered at its coordinates.
This generates vectors of values for each voxel representing prob-
abilities of belonging to specific foci. These values are assumed to
be independent such that the existence of one focus does not give
information about whether another focus will occur. The vector
values are combined with the addition rule for log-probabilities,
yielding ALE statistics. Thus, the ALE statistic represents the prob-
ability of a certain voxel to belong to any of the included foci. Sig-
nificance tests are conducted by comparing the ALE statistic in
each voxel with a null distribution, generated via repeatedly
54 O. Vartanian, M. Skov / Brain and Cognition 87 (2014) 52–56

calculating ALE statistics from randomly placed activation foci.

This null distribution is then used to estimate the threshold result-
ing for a given false discovery rate (FDR). Finally, a cluster thresh-
old (minimum spatial extent of significant contiguous clusters) can
be applied. Here the FDR was set to p < .01, and the minimum clus-
ter size to 50 contiguous voxels. To ensure that the assumption of
independence was not violated, we selected only one contrast from
each study, comparing the main judgment or viewing condition
with a baseline (or rest) condition. In two cases the foci involved
a parametric analysis of brain activation co-varying with prefer-
ence ratings (Harvey, Kirk, Denfield, & Montague, 2010; Vartanian
& Goel, 2004). ALE can handle contrast and parametric foci within
the same analysis.

3. Results and discussion

As predicted, viewing paintings activated a distributed network

of structures in the brain (Table 2). Supporting our first hypothesis,
Fig. 1. Viewing paintings activated the middle occipital gyrus, the lingual gyrus, the
viewing paintings activated areas in the visual cortex including the inferior temporal cortex, the insula and the putamen.
lingual gyrus and the middle occipital gyrus, as well as the fusiform
gyrus (Table 2 and Fig. 1). These activations can be attributed to the
processing of various early, intermediate, and late visual features
of the stimuli embedded within paintings, including orientation,
shape, color, grouping, and categorization (see Chatterjee, 2003;
Greenlee & Tse, 2008; Wandell et al., 2009). Although not located
in the occipital lobes, the inferior temporal cortex (Table 2 and
Fig. 1) has a well-established role in visual representation of form
and color (Gross, 1992), and likely contributes to that process here.
Activation was also observed in the precuneus (Fig. 2), which we
attribute to visuospatial exploration of pictorial stimuli. Specifi-
cally, the precuneus has been activated in studies that facilitated
spatial exploration of paintings (see Cupchik et al., 2009; Fairhall
& Ishai, 2008).
Supporting our second hypothesis, we observed activation in
the fusiform gyrus and the parahippocampal gyrus (Table 2 and
Fig. 3). The fusiform gyrus is involved in object perception and rec-
ognition (Grill-Spector & Sayres, 2008), and its activation here
likely represents the detection of objects within paintings (e.g.,
faces) (Kanwisher & Yovel, 2009). The parahippocampal gyrus is in-
volved in the perception and recognition of places (Epstein & Fig. 2. Viewing paintings activated the precuneus.
Kanwisher, 1998), which explains its involvement while viewing
paintings rich in representations of scenes (e.g., landscapes). Our
analysis also activated the anterior temporal lobe (i.e., superior
temporal gyrus) (Fig. 4). Recent evidence suggests that this ante-
rior region of the temporal lobes is involved not just in semantic

Table 2
List of structures activated in the meta-analysis.

Structure BA x y z Cluster size (mm3)

Lingual gyrus 17 17 91 0 1776
Middle occipital gyrus 17 22 88 0 3928
19 33 79 12 592
19 33 75 15 336
Fusiform gyrus 19 27 59 10 376
19 28 57 10 216
37 40 45 18 304
Parahippocampal gyrus 37 31 41 14 2560
36 23 40 9 544
Inferior temporal gyrus 37 43 68 2 160
Superior temporal gyrus 38 41 17 15 216
Insula 13 39 18 5 80
Fig. 3. Viewing paintings activated the fusiform gyrus and the parahippocampal
13 34 17 4 392
gyrus.
Posterior cingulate cortex 30 8 52 11 520
30 12 51 10 176
Precuneus 7 27 62 45 120
memory—including our knowledge of objects—but also in higher-
Putamen – 19 7 0 72
order conceptual integration of information in relation to objects
Note. BA = Brodmann area. All coordinates are listed in Talairach space (see text). (e.g., how does a knife function?) (Bonner & Price, 2013; Patterson,
 O. Vartanian, M. Skov / Brain and Cognition 87 (2014) 52–56
Fig. 4. Viewing paintings activated the anterior temporal pole (superior temporal
gyrus) and the fusiform gyrus.
Nestor, & Rogers, 2007; Peelen & Caramazza, 2012). The activation
of this region indicates that the perception of paintings might trig-
ger higher-order semantic analysis of the represented objects be-
yond mere recognition.
With our third hypothesis we set out to explore whether view-
ing paintings would activate structures involved in emotion and/or
reward processing. Indeed, we observed activation in the anterior
insula bilaterally (Table 2 and Fig. 1). The anterior insula is known
to play a critical role in emotional processing (Craig, 2010), and is
part of the brain’s core affective system (Barrett, Mesquita,
Ochsner, & Gross, 2007). In addition, there was activation in the
putamen (Table 2 and Fig. 1). This structure in the basal ganglia
is reliably activated by the anticipation of rewards (Liu, Hairston,
Schrier, & Fan, 2011), and its activation here could signal the per-
ceived rewarding properties of paintings. The involvement of the
anterior insula and the putamen could be indicators of their contri-
bution to ‘‘continuous affective evaluation’’ (Leder et al., 2004),
potentially leading to a conscious evaluation during the later
stages of aesthetic information processing.
Somewhat unexpectedly, we also observed activation in the
posterior cingulate cortex bilaterally (Fig. 5). This region has
emerged as a key component of the brain’s ‘default network’
(Mason et al., 2007; see also Christoff, Gordon, Smallwood, Smith,
Fig. 5. Viewing paintings activated the posterior cingulate cortex and the fusiform
gyrus.
55
& Schooler, 2009), activated when we ‘‘maximize the utility of mo-
ments when we are not otherwise engaged by the external world’’
(Buckner, Andrews-Hanna, & Schacter, 2008, p. 1). This finding of-
fers an interesting angle to the study of paintings by highlighting a
process that many consider essential to deep appreciation of art-
works, namely a focus on inner emotions and thoughts (see Vessel,
Starr, & Gabrielle, 2012).
Finally, it is important to note that structures involved in visual
perception can also contribute to the computation of preferences.
For example, not only is the parahippocampal cortex involved in
scene perception, but its activity while viewing scenes is correlated
with pleasure (Biederman & Vessel, 2006; Yue, Vessel, &
Biederman, 2007). In fact, discovering the contributions of the
brain’s perceptual apparatus to preference formation holds one of
the great promises for the field of neuroaesthetics.
4. Conclusion
The aim of our study was to reveal the neural correlates of view-
ing paintings based on a quantitative meta-analysis of fMRI data.
Our results complement the ALE meta-analysis conducted by
Brown et al. (2011), geared toward uncovering the core processes
underlying aesthetic evaluation. This is because much like in
experimental aesthetics, many researchers in the field of neuroaes-
thetics have adopted paintings as their preferred stimuli for study-
ing aesthetic phenomena. As such, it is useful to know which brain
regions respond to a specific category of stimuli (i.e., paintings) in
addition to those that contribute reliably to positive-valence aes-
thetic appraisal. As researchers in the field move toward testing
progressively more fine-grained hypotheses, such meta-analytic
results should prove useful in interpreting findings and formulat-
ing predictions.
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