Culture and Brain (2022) 10:1–26

https://doi.org/10.1007/s40167-021-00102-z

REVIEW ARTICLE

Neural representations of visual aesthetic experience (VAE):

a meta‑analysis

Xiyu Feng1 · Jing Gan1 · Xiaoqi Huang1 · Siyang Luo1 

Accepted: 15 May 2021 / Published online: 6 July 2021

© The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature 2021

Abstract
The present study intended to investigate the generic nature of visual aesthetic expe-
rience. Researchers have not agreed upon what constitutes visual aesthetic experi-
ence, and the present study proposed that visual aesthetic experience is comprised
of at least two components: enhanced visual processing and positive emotional and
reward experience. We applied a general activation likelihood estimation meta-anal-
ysis to 42 functional magnetic resonance imaging experiments described in 37 pub-
lished studies. The general activation likelihood estimation revealed activation in the
left orbitofrontal cortices and bilateral anterior cingulate cortex, which was thought
to be related to emotional and reward processes, and activation in the right fusiform
gyrus. In addition, a conjunction analysis of passive viewing tasks and tasks with
explicit instructions showed activation in the anterior cingulate cortex/orbitofrontal
cortex, and contrast analysis revealed stronger activation in the anterior cingulate
cortex/orbitofrontal cortex during the passive viewing task without explicit instruc-
tions to make aesthetic evaluations, suggesting that stronger emotional experiences
occur under such conditions. A conjunction analysis of groups with different cul-
tural backgrounds showed activation in the ventral anterior cingulate cortex/orbit-
ofrontal cortex, suggesting that there are universal cultural components of visual
aesthetic experience. Together, our findings complement the existing literature by
including all kinds of visual stimuli that could induce an aesthetic experience in the
viewer and contributes to our understanding of aesthetics by showing that it involves
enhanced visual sensation and positive emotional and reward experience.

Keywords  Functional magnetic resonance imaging · Visual aesthetic experience ·

Meta-analysis · Activation likelihood estimation

Xiyu Feng, Jing Gan, and Xiaoqi Huang have contributed equally to this work.

* Siyang Luo
[email protected]; [email protected]
1
Department of Psychology, Guangdong Key Laboratory of Social Cognitive Neuroscience
and Mental Health, Guangdong Provincial Key Laboratory of Brain Function and Disease, Sun
Yat-Sen University, Guangzhou 510006, China

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Introduction

What is visual aesthetics? Researchers have studied aesthetic experience from many
perspectives using various research methods, and we intend to use meta-analysis to
investigate visual aesthetics in the most generic sense. The premise of this research
is that aesthetics can be studied scientifically (Palmer et al., 2013).

Existing theories of visual aesthetic experience

Palmer et al. (2013) proposed three possible viewpoints for defining aesthetics. The
present study refers to one of the three definitions: aesthetic experience is a set of
psychological processes that corresponds to some stable neural activation pattern.
Therefore, it is necessary to identify the exact psychological processes involved.
Several theories of aesthetic processing have been proposed. Brielmann and Pelli
(2018) categorized existing models and research as either stimulus-focused, i.e.,
aiming to identify a set of object properties in the visual domain that contribute to
aesthetic pleasure, or response-focused, i.e., investigating the mechanisms (includ-
ing neural processes) that underlie aesthetic judgments. Previous attempts have not
identified a stable set of object features, although some features (curvature, symme-
try) seem to have universal appeal. Influential response-focused theories are listed
below.
In terms of neural activation, Chatterjee et  al. (2016) identified related neural
mechanisms and proposed that aesthetic experiences are mental states arising from
the interaction of emotion-valuation, sensory-motor, and meaning-knowledge neu-
ral systems; the interactions of these systems during aesthetic experiences resem-
ble those that occur during nonaesthetic engagements with objects. They further
explained that aesthetic experiences can occur without explicit aesthetic judgments,
that the three systems can interact during the early stages rather than exerting their
effects sequentially, and that the systems may not contribute equally. Skov and Nadal
(2019) proposed that the assumption that aesthetic experiences are psychologically
or neurobiologically distinctive from other “nonaesthetic” experiences cannot be
supported by empirical evidence and that aesthetic experience pertains to an inter-
action between sensory processing and neural activity in the reward circuit that is
subject to the influence of contextual factors such as relevance, physiological needs
and behavioral goals.
Researchers have also examined the behaviors of art viewing in detail. The Vienna
Integrated Model of top-down and bottom-up processes in Art Perception (VIMAP)
(Pelowski et al., 2017) was constructed to provide a detailed explanation of the ways
people respond to art. It integrates bottom-up, artwork-derived processes, which
involve how visual art is perceived and integrated with prior knowledge, and top-
down mechanisms, which explain how viewers behave or adapt according to their
processing experience, such as the extent of self-relevance and schema congruence.
By examining empirical evidence, Nodine et  al. (Nodine et  al., 2008) introduced
an information-processing stage model, according to which aesthetic experiences

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Neural representations of visual aesthetic experience (VAE):… 3

with visual art occurs in two phases: viewers first generate a global first impres-
sion of the art with quick glances, and if they find it sufficiently interesting, they
then proceed to scrutinize the art in more detail and develop appreciation. Likewise,
Graf and Landwehr (2015) developed the Pleasure-Interest Model of Aesthetic Lik-
ing (PIA Model), which views aesthetic preferences as the result of two hierarchical
processes: the first is a stimulus-driven process from which aesthetic preferences are
derived. If processing motivation is sufficient, perceiver-driven processing follows
and gives rise to fluency-based aesthetic evaluations.

The present study

Skov and Nadal proposed (2020a) that three sets of problems are addressed in a sci-
entific investigation of aesthetics: factors that determine the individual outcomes
of an aesthetic appreciation event, the mechanism by which sensory information
activates processes in the reward circuit, and the computational mechanisms that
make up the reward circuit. With the intention of laying groundwork for the study
of dynamic interaction, in this research, we performed a meta-analysis to identify
brain regions that contribute to visual aesthetic experience (VAE). It may be futile
to argue in favor of one definition of VAE over another without considering context.
Here, we set boundary conditions for what constitutes the VAE of interest and its
characteristics in the most generic sense while providing principles for screening the
literature for analysis.

Subjectivity of VAE

First, VAE is subjective. This humanistic characteristic is supported by the failure of

stimulus-focused research to identify a common set of objects that are able to elicit
an aesthetic experience in all groups. Therefore, only studies with rating paradigms
were selected. Rating paradigms were preferred over forced-choice paradigms for
their accurate reflection of subjective judgments.

The VAE of interest is positive valence

Second, the VAE of interest is positive valence. Although various complex emotions
can be evoked while viewing art (Silvia, 2012), based on the assumption that emo-
tions can be described in multidimensional space with one dimension of valence, we
decided to limit our investigation to positive valence to clearly identify related brain
regions. In addition, Skov and Nadal showed evidence that aesthetic emotions are
not distinctly different from emotions observed during other forms of sensory valu-
ation in terms of their psychological components or neurobiological underpinnings
(Skov & Nadal, 2020b), refuting Menninghaus’s theory (Menninghaus et al., 2019)
that suggested otherwise.
Hence, in the screening procedure, we carefully viewed and selected the terms
used to ensure that participants were instructed to reflect on the psychological

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 4 X. Feng et al.

processes involved in the positive appraisal of targeted objects. One unconfirmed

assumption was that individuals describe the same set of feelings and internal pro-
cesses using the same set of words, but if the contrasting of positive appraisals with
negative appraisals occurs within the individual, problems with ambiguous connec-
tions between the signifier and the signified can be avoided.
Positive affect (PA) states (Burgdorf & Panksepp, 2006) are related to regions
such as the ventral striatum system (VS), orbital frontal cortex (OFC) and amygdala,
all of which are part of the cortical-basal ganglia circuit at the heart of the reward
system (Haber & Knutson, 2010), along with the anterior cingulate cortex, ventral
pallidum and midbrain dopamine neurons. Researchers have identified shared and
varied representational styles in these regions. Specifically, the orbital frontal cortex
and ACC have been found to be activated in relation to various PA states (Burgdorf
& Panksepp, 2006) representing information about reward outcomes and contextual
aspects of rewards. In particular, the dACC is involved in monitoring potential con-
flict situations. The ventral striatum also has a complicated representation of reward-
related information; it responds to PA (Burgdorf & Panksepp, 2006), anticipation of
reward, reward outcomes, reward prediction errors and tracking uncertainty (prob-
ability), delay or effort, and different regions exhibit characteristic responses. While
some researchers have found that positive emotions tend to reduce amygdala activa-
tion (Burgdorf & Panksepp, 2006), others have suggested that this activation may
not reflect the value of a stimulus. The ventral pallidum was not reviewed due to
the methodological limitations of separating it from the VS. Lastly, it has been sug-
gested that midbrain activation increases during reward anticipation (Haber & Knut-
son, 2010).
Based on known representational styles within the PA- and reward-related sys-
tem and more direct evidence concerning VAE (Brown et al., 2011; Jacobsen et al.,
2006; Lacey et al., 2011; Vartanian & Skov, 2014; Vessel et al., 2012), we focused
on and expected to observe the involvement of the orbital frontal cortex (Brown
et al., 2011; Jacobsen et al., 2006; Vartanian & Skov, 2014), anterior cingulate cor-
tex, and ventral striatum (Lacey et al., 2011; Vessel et al., 2012) (Hypothesis 1). We
did not involve the anticipation process because no cue was shown prior to targeted
stimuli. We left the amygdala out of the discussion because the main analysis con-
centrated on how positive experiences differ from neutral or negative experiences
(instead of contrariwise).

Spontaneity of VAE

Third, VAE is a spontaneous process (instead of an intentional task), i.e., it occurs

independent of instructions associated with an aesthetic task. Hofel (2007) and
Schacht (2008) contrasted ERP components observed under conditions with or
without explicit instructions to contemplate or rate beauty-related attributes and
suggested that while symmetry analysis occurred spontaneously, aesthetic apprecia-
tion appeared to require intention. Nevertheless, related evidence seems insufficient
due to concurrent changes in other processes, and the interpretation contradicted

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Neural representations of visual aesthetic experience (VAE):… 5

findings that the aesthetic value of objects can influence viewers’ attention (Rolke
et al., 2019), memory (Rolke et al., 2019), affective response (Makin et al., 2012)
and neural activities (Calvo-Merino et al., 2008; Lacey et al., 2011) even when no
instruction are provided.
All of the collected studies included viewing of objects (shown on screen) and
performing rating tasks for them, which can be categorized with respect to the tem-
poral relationships of the two procedures: we used the term passive viewing tasks to
describe studies in which the participants performed only the viewing procedures
in the scanner (as we included only fMRI studies) and performed the rating tasks
at a later time, and we used the term explicit response tasks to describe studies in
which the participants performed the two procedures simultaneously in the scan-
ner. In light of this, we expected to find that the same regions were involved in pas-
sive viewing tasks and explicit response tasks, except for those that were specifically
involved in the completion of questions (Hypothesis 2).

Target objects for VAE

Fourth, VAE can be elicited by objects in general (rather than exclusively by art
objects or sets of events). Brown et al. (2011) proposed the idea of naturalizing aes-
thetic processing because it can be equated with object-appraisal processes.
Note that aesthetic experience is often used to describe engagement with art
objects. However, we did not focus on such processes (Graf & Landwehr, 2015;
Nodine et  al., 2008; Pelowski et  al., 2017) because concepts and ideas regard-
ing what constitutes art were formed fairly recently, are vaguely defined and are
hotly debated in philosophy. Skov and Nadal (2020a) reviewed the creation of art
and indicated that art objects are not different from other “nonartistic” objects in
nature, other than that they belong to a culturally meaningful category. Furthermore,
researchers found that the brain areas involved in aesthetic responses to artwork
overlap with those involved in the appraisal of objects of evolutionary importance
(Brown et  al., 2011). No existing evidence suggests that a unique set of computa-
tional principles has developed specifically for processing art (Agnati et al., 2007).
Instead of considering how the brain represents and experiences art (Skov & Nadal,
2020a), we were interested in aesthetics as the study of how and why sensory stimuli
acquire hedonic value. Consequently, during the screening procedure, we included
not only art objects but also a broader range of targets, including food, faces, bodies,
scenery and symbols. This is congruent with Chatterjee (2016) and Skov and Nad-
al’s views (2019) that aesthetic experiences are not distinctive from “nonaesthetic”
experiences, psychologically or neurobiologically.

Visual processing of VAE

Once a potential visual stimulus is effectively processed, it can elicit a feeling of

beauty, regardless of the duration of stimulus exposure (Brielmann et  al., 2017).
Is a beautiful stimulus processed differently from a mundane stimulus? Some

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researchers have suggested that this might be the case. Previous research found that
VAE involves the ventral visual stream (Boccia et  al., 2016; Vessel et  al., 2012).
Vartanian and Goel (2004) found an enhancement of early visual processing upon
viewing beautiful scenes. One plausible explanation is that aesthetic objects elicit
greater attention, consequently leading to enhanced early visual processing, which is
characterized by enhanced activation of the ventral visual cortices. This is supported
by single-cell recording studies in monkeys and by functional brain imaging and
event-related potential studies showing that selective attention can modulate neu-
ral processing in the visual cortex (Gazzaley & Nobre, 2012; Reynolds & Chelazzi,
2004; Watanabe et al., 2011). Accordingly, we examined whether the involvement
of the ventral visual stream changes with aesthetic value, without making strong
assumptions.

Contextual factors in VAE

Fifth, as VAE has developed as an evolutionarily shared experience (Agnati et al.,

2007) and is not a modern product, anyone should be capable of experiencing the
process. Hence, contextual factors such as targeted objects, cultural background and
expertise in certain fields should affect only when or where people experience aes-
thetics rather than how they experience it.
Some theories have taken into account how various factors can influence art
experiences. Perceptual fluency theory (Reber et al., 2004) proposed that aesthetic
experience is a function of the perceiver’s processing dynamics, namely, the more
fluently the perceiver can process an object, the more positive his or her aesthetic
response is. Shimamura (2013) further proposed that the aesthetic response to art
objects is the result of an interaction between the intention behind the art and the
viewer’s sensory, knowledge-based and emotional responses. Bullot and Reber
(2013) introduced a psycho-historical framework for the science of art apprecia-
tion that combined the psychological and historical approaches to a theory of art
and emphasized the effect of art-historical information on art appreciation. Redies
(2015) proposed a unifying model that combined the formalist aspect (based on the
intrinsic beauty of artwork) and contextual aspects (accounting for individual and
culturally dependent aspects) of aesthetics, and the predictive coding framework
(Kesner, 2014) implies that the mindset for generating expectations and predictions
about visual arts is constituted by culture-related skills and knowledge.
What effects do culture have on VAE? Researchers suggest that aesthetic experi-
ences such as aesthetic chills appear to be universal (McCrae, 2007), supporting the
fifth assumption. People across cultures share a taste for certain objects (Che et al.,
2018) because aesthetic preference can emerge from basic perceptual and valuation
processes that are common to all humans, such as those for higher-contrast paint-
ings (van Dongen & Zijlmans, 2017), mid-range fractal patterns (Street et al., 2016),
divine proportions (Pittard et al., 2007), and landscape paintings over portraits (Bao
et al., 2016); however, cultures can have different opinions regarding other features,
such as noses (Broer et al., 2012), color (Taylor et al., 2013), and skin color (Chen

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Neural representations of visual aesthetic experience (VAE):… 7

et  al., 2019, 2020), along with a preference for paintings from one’s own culture
over those from other cultures (Yang et al., 2019).
In testing the fifth assumption, we conducted an independent analysis of studies that
recruited participants from the same cultural background and compared culture-spe-
cific results, with the expectation that we would find that shared regions were activated
for different cultural groups (Hypothesis 3).

VAE and similar processes

We defined the VAE of interest by setting the aforementioned boundary conditions (or
assumptions). If VAE is defined as a natural entity separable from other similar, albeit
qualitatively different, psychological processes, such as moral judgment (Tsukiura &
Cabeza, 2011), ecstatic drug experience (Sumnall et  al., 2006), and self-referential
thought (Vessel et al., 2013), then the underlying recruitment of neural activities may
be different. In testing this idea, we performed a “decoding” analysis (Yarkoni et al.,
2011) to examine the similarity of activation patterns between the VAE of interest and
other processes in the existing repository. Briefly, this analysis utilized a large gener-
ated database of mappings of neural and cognitive states and estimated the likelihood
that new activation maps (in this case, the activation map of the VAE of interest) were
associated with specific psychological terms, thereby “decoding” psychological con-
structs based on patterns of brain activity. This decoding process was carried out via a
web interface (http://​neuro​synth.​org).
By definition, we anticipated similarity between the patterns for the VAE of interest
and processes such as object viewing, reward processing or PA, and we expected to
find similarity between VAE and the aforementioned processes (Sumnall et al., 2006;
Tsukiura & Cabeza, 2011; Vessel et al., 2013) that were previously explored.
Many researchers have pointed out the problem of heterogeneity of results in the
field of neuroaesthetics. Nadal et  al. (2012) tentatively explained that differences are
caused by methodological variables such as the paradigms used, the composition of
materials used, and the composition of participant groups. Therefore, we performed an
exploratory analysis to examine how methodological variables affected the results.
Testable hypotheses are listed below for clarity: First, we hypothesized that VAE
would involve stronger activation of positive affect and reward-related system, includ-
ing the orbital frontal cortex, anterior cingulate cortex and ventral striatum. Second,
we hypothesized that, due to the spontaneity of VAE, same regions would be activated
in passive viewing tasks and explicit response tasks, except for those that were spe-
cifically involved in the completion of questions. Third, we hypothesized that shared
regions would be activated for different cultural groups during VAE, whose activation
patterns should be independent of contextual factors once occurred.

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 8 X. Feng et al.

Method

Paper selection

A systematic method was adopted for the literature review. The search
was carried out using PubMed and Web of Science. This meta-analy-
sis included studies that involved the aesthetic evaluation of visual stim-
uli. Relevant articles were identified using the following search terms:
(“Beauty” < OR > “aesthetic” < OR > “aesthetics” < OR > “esthetic” < OR > “neu-
roaesthetic” < OR > “neuroaesthetics” < OR > “neuroesthetic” < OR > “aes-
thetic judgment” OR “pleasure” < OR > “pleasant” < OR > “pleasant-
ness” < OR > “valence”) < AND > “visual” < AND > (“fMRI” < OR > “functional
magnetic resonance imaging” < OR > “functional MRI”) < AND > (“rat-
ing” < OR > “judgment” < OR > “preference” < OR > “evalua-
tion” < OR > “appraisal” < OR > “liking”). The search yielded a total of 370
studies. The reference lists of these studies and relevant meta-analyses were also
scrutinized. A total of 394 papers published from 2000 to 2019 were found.
The inclusion criteria for the current investigation were as follows:

1. Inclusion of whole-brain analyses performed using functional magnetic resonance

imaging (fMRI).
2. Provision of coordinates of activation foci in either the Montreal Neurological
Institute (MNI) or Talairach reference space.
3. Subjects were healthy individuals.
4. All neuroimaging studies had to include a visuoperceptual control condition to
exclude all activation that was not directly related to the VAE.
5. The tasks involved aesthetic evaluations of the positive valence of the presented
stimuli, including ratings of pleasantness, attractiveness, and liking.
6. There could be no pharmacological manipulation.
7. Only group studies were included (e.g., no case studies, meta-analyses, or
reviews).

Each article was carefully read to ensure that it fulfilled the criteria. A total of
37 papers including 42 experiments and 380 foci from 779 participants remained
(Supplementary Table  1). Regarding the aforementioned methodological vari-
ables (types of stimuli and types of tasks), we included 13 experiments involv-
ing faces, 5 experiments involving paintings, 3 experiments involving bodies, 23
experiments involving the rating of attractiveness, 6 experiments involving the
rating of liking, 7 experiments involving pleasantness, 36 experiments involv-
ing explicit responses and 6 experiments involving passive viewing. We also
classified the papers in terms of their participants’ cultural backgrounds, which
resulted in 6 experiments involving people from East Asia (e.g., Japan, China)
and 35 experiments involving individuals from Western countries (e.g., America,
Canada) (Fig. 1).

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Neural representations of visual aesthetic experience (VAE):… 9

370 Records identified through

24 additional records identified
Web of Science & PubMed
from related meta-analyses
database search

356 studies excluded

38 studies without inclusion of whole-brain analysis using fMRI
394 Records identified 5 studies without provision of coordinates of activation foci in Montreal
Neurological Institute or Talairach reference space
39 studies not using healthy individuals
110 studies without inclusion of visuo-perceptual control condition
154 studies not related to ratings of pleasantness, attractiveness, and liking
38 studies included 1 studies using pharmacological manipulation
9 studies without group study design

Meta-analysis for rating of Meta-analysis for rating of Meta-analysis for

body /face/ painting pleasantness /attractiveness/ liking explicit response/passive viewing
(56 / 109 / 21 foci) ( 75 / 196 / 78 foci) ( 349 / 31 foci)

Fig. 1  Flowchart of the literature search and selection process

Activation likelihood estimation (ALE) meta‑analysis

A series of meta-analyses was performed using the revised ALE method in Gin-
gerALE 3.0.2 software (http://​brain​map.​org/​ale) with MNI coordinates (Eickhoff
et  al., 2009, 2012; Turkeltaub et  al., 2015). This method offered a random-effects
approach for assessing clustering between experiments to identify consistent brain
regions with specific activation. When coordinates were reported in Talairach
spaces, a conversion tool within GingerALE was used to transform the coordinates
to correspond to the MNI space.
A general ALE meta-analysis was performed on 380 foci from all selected exper-
iments to identify consistent activation during aesthetic experience. Experiments
that could be clearly classified were also used to perform separate ALE analyses of
different types of visual stimuli (faces, bodies, paintings) and types of tasks (rating
of attractiveness, rating of liking, rating of pleasantness, explicit response, passive
viewing) to test the effect of methodological variables. It is important to clarify that
the term “explicit response” was used in reference to tasks that included explicit
instructions that asked the participants to conduct evaluations while they were in
the scanner, while “passive viewing” referred to tasks in which the participants pas-
sively viewed images that were presented in the scanner and performed their evalu-
ations while they were outside the scanner. We followed the methodological guide-
lines in  the GingerALE User Manual (https://​brain​map.​org/​ale/​manual.​pdf). These
meta-analyses were given a voxel threshold of p < 0.001 and a cluster-level thresh-
old of p < 0.05 with 5000 permutations to find convergence with a minimum cluster
size automatically determined by GingerALE. The cluster-level threshold of p was
corrected with Family‐wise error (FWE) method. FWE thresholds are more con-
servative so p < 0.05 is recommended (Eickhoff et al., 2009, 2012; Turkeltaub et al.,
2015).
Following the single-dataset ALE analyses, conjunction and contrast anal-
yses were conducted to identify both common and unique areas within the

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methodological variables. The conjunction analysis was performed for task types
(explicit response ∩ passive viewing), and contrast images between the two types
were created by subtraction (explicit response—passive viewing, passive viewing—
explicit response). Clusters in these analyses were given a threshold of p < 0.05 with
5000 permutations and a minimum cluster size of 50  ­mm3 (Robert et  al., 2018).
The generated displays were plotted using the image processing software Mango 4.1
(Lancaster et al., 2012).

Decoding activation maps

To distinguish the content of the mental processes of VAE during different tasks,
we conducted an exploratory analysis using an online tool, the Neurosynth Image
Decoder (Yarkoni et  al., 2011). After we uploaded the activation map resulting
from our ALE analyses, we used this tool to compute the spatial correlations r
of the uploaded map and the concept-based meta-analysis maps in Neurosynth’s
database. The r values reflected the correlation between the two maps across all
voxels. Based on the r values, the tool provided a ranked list of concepts that
activate regions similar to those activated in the uploaded map. The correlation of
each concept indicated the probability that the concept was used in a study based
on the activation presented in the uploaded map.
We applied the Neurosynth Image Decoder to activation maps of general VAE,
VAE in explicit response tasks, VAE in passive viewing tasks and VAE for each
stimulus type (body, face, painting). The decoding results (the lists of concepts
related to our activation maps) enabled us to reference the possible targets pro-
cessed during VAE.

Results

General ALE meta‑analysis

The general ALE meta-analysis of VAE revealed 2 clusters of activation (Fig. 2;

Table  1). The areas encompassed a bilateral range of the ACC, with one peak
located in the left ACC/OFC (x/y/z − 6 44 − 4). The OFC is a vast and heteroge-
neous region that can be broadly divided into three main sections based on ana-
tomical and cytoarchitectonic considerations: a posterior OFC region, an ante-
rior OFC region and a ventromedial prefrontal cortex (vmPFC) region (Haber &
Knutson, 2010; Sescousse et al., 2013). The OFC and ACC are components of the
reward and emotional systems (Beckmann et al., 2009; Haber & Knutson, 2010),
whose activation was suggested by Hypothesis 2. Consistent with Hypothesis 1,
VAE also enhanced activation of the ventral visual cortices, as represented by the
other cluster we found with a peak in the fusiform gyrus (50 − 70 − 8).

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 Table 1  Regions showing consistent activations across general studies of VAE
Region (Hem) Volume ­(mm3) BA ALE Z x y z
value

Anterior cingulate/orbitofrontal 5416 32 0.036 6.355 −6 44 −4

cortex (L)
Anterior cingulate (R) 24 0.027 5.128 8 34 − 10
Fusiform gyrus (R) 1320 19 0.022 4.431 50 − 70 -8

BA = Brodmann areas; *Brodmann areas were not applicable; Hem = hemisphere; Reference Space = MNI
Neural representations of visual aesthetic experience (VAE):…
11

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 12 X. Feng et al.

Separate ALE meta‑analyses for each stimulus type

We performed 3 single ALE meta-analyses to test for the specific brain areas that
were engaged for each stimulus type (faces, paintings, bodies). The results are pre-
sented in Fig. 2 and Table 2.
Face conditions were associated with activation of the left ACC (− 4 36 2), OFC
(− 18 46 − 16) and superior frontal gyrus (− 20 54 − 10). For the painting condition,
we observed activation of the left ACC/OFC (− 2 48 − 6) and the bilateral ACC (2
36 − 10, − 2 38 − 10). Activation of the right inferior frontal gyrus at 50 8 28 was
shown in the body condition.

Separate ALE meta‑analyses for each task type

Single ALE meta-analyses were also performed for the different task types to iden-
tify the related brain regions. First, we performed single meta-analyses for the pas-
sive viewing tasks and the explicit response tasks, which differed only in terms of
whether the participants were given explicit instructions to conduct evaluations
while in the scanner. For the passive viewing tasks, we observed activation of the
left ACC/OFC at -2 48 -6. The ALE analysis revealed activation of the bilateral
ACC, left OFC and right fusiform gyrus for explicit response tasks (8 34 − 10, − 4
42 − 4, 50 − 70 − 8).
Additionally, we conducted 3 single meta-analyses based on the rating tasks (rat-
ing of attractiveness, rating of pleasantness, rating of liking). The rating of pleasant-
ness revealed activation of the left precentral gyrus (− 50 4 28) and ACC/OFC (− 6
46 0), as well as the ACC (0 40 -6), fusiform gyrus (48 − 68 − 8) and middle frontal
gyrus (− 28 − 4 50) in the right hemisphere. In addition, we observed activation of
the bilateral ACC (6 34 − 10, − 4 36 0), left ACC/OFC (− 8 42 − 6) and medial pre-
frontal cortex (− 2 52 4) during the rating of attractiveness, while the ALE analysis
did not provide any significant cluster for the rating of liking. The results are pre-
sented in Fig. 2 and Table 2.

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Neural representations of visual aesthetic experience (VAE):… 13

Fig. 2  Results of the general meta-analysis and separate meta-analyses for methodological variables. Sig-
nificant clusters are displayed on sagittal, coronal and axial slices

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 14 X. Feng et al.

Table 2  Regions showing consistent activations across VAEs of (A) each stimulus type and (B) each task
type
Condition Region (Hem) Vol- BA ALE x y z
ume value
­(mm3)

A. Stimulus type
 Faces Anterior cingulate 912 24 0.015 − 4 36 2
(L)
Superior frontal 672 10 0.017 − 20 54 − 10
gyrus (L)
Orbitofrontal 32 0.012 − 18 46 − 16
cortex (L)
 Paintings Anterior cingulate/ 1488 32 0.015 − 2 48 − 6
orbitofrontal
cortex (L)
Anterior cingulate 24 0.008 − 2 38 − 10
(L)
 Bodies Inferior frontal 1027 9 0.021 50 8 28
gyrus (R)
B. Task type
 Rating of attractiveness Anterior cingulate 3424 24 0.023 6 34 − 10
(R)
Anterior cingulate/ * 0.017 − 8 42 − 6
orbitofrontal
cortex (L)
Medial prefrontal 10 0.013 − 2 52 4
cortex (L)
 Rating of pleasantness Anterior cingulate 1040 * 0.017 0 40 − 6
(R)
Anterior cingulate/ 32 0.015 − 6 46 0
orbitofrontal
cortex (L)
Precentral gyrus 632 6 0.017 − 50 4 28
(L)
Fusiform gyrus 560 19 0.014 48 − 68 − 8
(R)
Middle frontal 472 6 0.018 − 28 − 4 50
gyrus (R)
 Rating of liking No cluster found
 Passive viewing Anterior cingulate/ 648 32 0.015 − 2 48 − 6
orbitofrontal
cortex (L)
 Explicit response Anterior cingulate/ 4968 24 0.032 − 4 42 − 4
orbitofrontal
cortex (L)
Anterior cingulate 24 0.027 8 34 − 10
(R)
Fusiform gyrus 856 19 0.021 50 − 70 − 8
(R)

BA = Brodmann areas; *Brodmann areas were not applicable; Hem 

= hemisphere; Reference
Space = MNI

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Neural representations of visual aesthetic experience (VAE):… 15

Comparisons between explicit response and passive viewing

To further investigate methodological variables, we performed conjunction and

contrast analyses of VAE involving 2 types of tasks—explicit response and pas-
sive viewing—to identify their common areas and unique features. The conjunc-
tion analysis revealed consistent activation in 1 cluster encompassing the left ACC/
OFC at − 4 48 − 6 (Fig. 3a, Table 3). No significant cluster was found in the explicit
response—passive viewing contrast analysis, whereas 1 activated cluster that
included the left ACC/OFC (− 1 50 − 8) was found in the inverse contrast analysis
(Fig. 3b, Table 3).

Fig. 3  a Results of conjunction analysis (passive viewing ∩ explicit response). b Results of contrast anal-
ysis (passive viewing—explicit response). Significant clusters are displayed on sagittal, coronal and axial
slices

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 16

13
Table 3  Conjunction and contrast analysis for passive viewing and explicit response
Condition Region (Hem) Volume ­(mm3) BA Z x y z

Passive viewing ∩ explicit response Anterior cingulate/orbitofrontal cortex (L) 272 * \  − 4 48  − 6
Passive viewing—explicit response Anterior cingulate/orbitofrontal cortex (L) 408 32 2.075  − 1 50  − 8
Explicit response—passive viewing No cluster found

BA = Brodmann areas; * = Brodmann areas were not applicable; Hem = hemisphere; Reference Space = MNI
X. Feng et al.
Neural representations of visual aesthetic experience (VAE):… 17

VAE in Western and Eastern cultural contexts

Human cognition, motivation, emotion, and behavior are shaped by cultural val-
ues and norms. Considerable interobserver variability among different cultures and
social groups has been found in previous aesthetic preference experiments (Palmer
et al., 2013). Since the aforementioned results showed that aesthetic processing had
many components involving evolutionary, anatomical, cognitive, personality and
emotional factors, VAE is obviously culture-related. Cultural contexts might pro-
vide explanations of why beauty may be attributed to some things but not to others
(Jacobsen, 2006). Following this idea, we divided the selected papers into 2 cat-
egories, i.e., Western and East Asian, and then performed conjunction and contrast
analyses to compare VAE in the 2 culture groups (Fig. 4).
The conjunction analysis revealed the 3 clusters presented in Fig.  4 (b), which
included the ventral anterior cingulate (vACC)/OFC (8 36 − 10, − 6 36 2) and left
superior frontal gyrus (− 20 16 54); consistent activation identified cross-cultural
commonality in the emotional and higher cognitive processing (e.g., working mem-
ory) of VAE.

Fig. 4  a Results of separate meta-analyses for East Asian and Western cultures. b Results of conjunction
analysis between the two culture groups

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 18 X. Feng et al.

Decoding the contents of VAE

We decoded the activation maps resulting from the above meta-analyses of general
VAE, VAE in the explicit response task, VAE in the passive viewing task and VAE
for each stimulus type (body, face, painting) to identify constructs related to these
types of VAE.
The correlated constructs are displayed in Fig. 5. For general VAE, related con-
structs were value (with a correlation of 0.249), reward (0.227), money (0.172),
viewing (0.172), valence (0.151), incentive (0.141), motivational (0.132), self-
referential (0.125), face recognition (0.091), engagement (0.088) and recognition
(0.054). The decoding results support the view that VAE comprises a complex mix
of images, feelings and thoughts. Explicit response VAEs exhibited a similar pattern
to that of general VAE, whereas passive viewing VAEs showed lower correlations
with value (0.064), reward (0.058), money (0.043), valence (0.047), and incentive
(0.007) and higher correlations with face recognition (0.124), engagement (0.124),
and recognition (0.11).
Value and valence were common constructs in the VAE of all 3 stimuli types.
Specifically, related constructs for the VAE of bodies comprised attention (0.247),
action observation (0.206), monetary (0.2), incentive (0.191), motivational (0.182),
working memory (0.164) and dopaminergic (0.132). The VAE of faces was related
to incentive, monetary, motivational and dopaminergic, with lower correlations than
VAE for bodies; the VAE of faces also showed similarities with maps for gambling
(0.116), pain (0.078) and probable inverse of action observation (− 0.032). The
VAE of paintings had a group of related constructs that was distinct from the afore-
mentioned VAE types and included including self-referential (0.182), engagement
(0.159), social (0.082) and mentalizing (0.061).

Fig. 5  Meta-analytic decoding results. a Decoding of the general ALE meta-analysis results; b Decod-
ing of the ALE meta-analysis results for each task type; c Decoding of the ALE meta-analysis results
for each stimulus type. Related concepts and their spatial correlation with activation maps for VAE are
displayed. For example, given the specific voxel location of the activation of general VAE, (a), there was
a 0.23 probability that the concept “reward” was used in a study when this activation was present

13
Neural representations of visual aesthetic experience (VAE):… 19

Discussion

Our first aim was to find converging evidence to identify the brain regions that are
most consistently involved in VAEs by performing an ALE meta-analysis across all
studies. Consistent with previous findings suggesting that the ACC and OFC may
play important roles in the process of aesthetic appreciation (Yeh et al., 2015), we
found that VAEs activated areas in the ACC and OFC. Our results are consistent
with a review of the neurobiology of beauty, which revealed that the visual experi-
ence of beauty was correlated with activity in the ACC and OFC (Yeh et al., 2015;
Zeki et al., 2014). Regions in the ACC and OFC have been implicated in aesthetic
emotional and reward processing (Cupchik et al., 2009; Di Dio et al., 2007; Ishizu &
Zeki, 2011). Specifically, an important guiding principle related to ACC functioning
is that emotional and cognitive information is processed separately. The two major
subdivisions of the ACC have distinct functions. Evidence suggests that our find-
ing of ACC activation can be regarded as a rostral-ventral affective division (Bush
et  al., 2000). Furthermore, activation of the rostral ACC was found to be related
to increases in the level of expected reward (Marsh et  al., 2007). Additionally, as
we speculated that VAEs involve enhanced activation of the ventral visual cortices,
we indeed observed activation of the fusiform gyrus, which is a component of the
ventral visual stream (Ptito et  al., 2012). The fusiform gyrus is involved in object
perception and recognition (Grill-Spector & Sayres, 2008), and its activation likely
represents the detection of objects across different aesthetic types of stimuli.
However, in contrast to Hypothesis 1, we did not observe any activation of
the VS. One probable explanations is that VS has been found to be recruited in
multiple forms of positive affective states and has a complicated representational
style of reward information (Burgdorf & Panksepp, 2006), with psychostimulants
such as dopamine acting as contributing factors. For example, while increaseing
neural activation is found in anticipation of reward in the VS, the actual receipt
of a monetary reward is found in concurrence with decreased activities (Knutson
et  al., 2001). Although there is some evidence suggesting VS showed stronger
activation during art-images viewing (Lacey et al., 2011), with a wider range of
visual stimuli included in this study, it is possible that its recruitment cannot be
manifested via the current methods (further investigations could apply more fine-
grained methods such as multivariate pattern analysis).
Additionally, consistent with our Hypothesis 2, which stated that the passive
viewing tasks and the explicit response tasks would show similar patterns of acti-
vation, activation of the ACC/OFC could be observed both in the passive view-
ing tasks and the explicit response tasks. This finding suggests that VAEs occur
spontaneously and automatically. Our brains do indeed conduct some evalua-
tions of stimuli during tasks, even when explicit decisions are absent (Montague
et al., 2014). Surprisingly, the fusiform gyrus did not activate during this stage;
however, future research could explore whether this finding can be attributed to
reduced attention under the passive viewing condition compared to the explicit
response condition.

13
 20 X. Feng et al.

In further analysis, the passive viewing—explicit response contrast analysis

revealed 1 cluster that included the left ACC and OFC, while no significant clusters
were identified in the explicit response -passive viewing contrast analysis. The func-
tions of the attention system can be classified into the following two distinct types:
bottom-up attention, which refers to attentional guidance driven by external factors
to salient stimuli due to their innate properties, and top-down attention, which refers
to the internal guidance of attention based on current goals and prior knowledge
(Katsuki & Constantinidis, 2014). Based on this definition, aesthetic experiences
include a bottom-up process, and the rating of aesthetic experiences is related to
a top-down process. Focused task demands can lead to attention fatigue when par-
ticipants process information in a top-down way (Crossan & Salmoni, 2019). We
speculate that the top-down process may cause cognitive resource depletion and that
participants can be more involved in aesthetic processing under a passive viewing
condition.
Previous research has illustrated that people with different cultural backgrounds
may share some universal aesthetic experiences (McCrae, 2007); thus, we sought to
gain further insight into the universal components of aesthetics through a conjunc-
tion analysis of different culture groups (a Western group and East Asian group).
The conjunction analysis revealed activation of the vACC/OFC. The vACC has
been robustly implicated in reward processing during social evaluation (Rigney
et al., 2018). Activation of the vACC/OFC may suggest that aesthetic emotional and
reward processing is a culturally universal component shared by people from East
Asian and Western countries. We also expect further evidence from future research
to support this implication.
In addition to the general meta-analysis, we classified the included studies based
on the stimulus types (paintings, body and face) and task types (rating of pleasant-
ness/liking/attractiveness) and performed separate meta-analyses for exploratory
purposes. In terms of the stimulus types, body stimuli were notably associated with
one cluster in the right inferior frontal gyrus (IFG). Previous works have lent consid-
erable weight to the hypothesis that the right IFG is of great significance in response
inhibition. For example, patients with frontal lobe lesions, including lesions of the
right IFG, exhibit impairment in inhibitory control tasks (Floden & Stuss, 2006).
The existence of a centrally located inhibitory mechanism that can suppress irrel-
evant responses has been suggested (Hampshire et al., 2010). Therefore, it may be
inferred that the right IFG was activated because nude or partially draped figures
were included under the body conditions, which required adult participants to inhibit
their sexual desire or feelings to focus on their task. In addition to involving the
ACC and OFC, face stimuli were associated with activation of the left superior fron-
tal gyrus (SFG), which has been shown to contribute to higher cognitive functions,
particularly working memory (Boisgueheneuc, 2006). We assumed that this finding
occurred partly because when the participants evaluated the attractiveness of face
stimuli, they compared the current face stimuli with familiar faces in their working
memory and provided their evaluation. Regarding the task types, we did not observe
any activation in the rating of the liking condition. This finding might be due to the
low statistical power of the test, since we included only 6 studies with this condi-
tion, a smaller number than was available for the other two task types. In most cases,

13
Neural representations of visual aesthetic experience (VAE):… 21

activation of the ACC and OFC could be observed when we grouped the studies
differently according to the methodological variables, except for the body stimulus
condition and the rating of liking condition. In contrast, we did not observe activated
clusters in the fusiform gyrus except for the rating of pleasantness condition. Future
research should delve deeper into the mechanisms underlying these incongruent
results and provide further explanations for our results.
Finally, in the present study, we used the Neurosynth database to meta-analyti-
cally decode the functional role of activation maps of general VAEs, VAEs during
explicit response tasks and passive viewing tasks, and VAEs of each stimulus type
(body, face, painting) separately. The results showed that constructs related to the
activation regions in general VAE and VAEs during explicit response tasks showed
almost the same patterns. As we anticipated, the process of VAE was similar to the
process of object viewing, feelings of reward and PA, including incentives, motiva-
tional feelings and a sense of value, to a certain extent. Previous research concerning
the aesthetic experience of works of art suggested that stimuli for aesthetic experi-
ence can cause observers to draw on self-referential information, and an appeal to
self-related information provides a way to extract meaning from the stimuli (Vessel
et al., 2013). This may explain why activation regions in general VAEs and VAEs
during explicit response tasks are related to the self-referential construct. In addi-
tion to the relation of the activation regions to PA, as we hypothesized, the decod-
ing results of the 3 stimulus types (bodies, faces and paintings) indicated that their
related constructs showed some specificity. VAEs of bodies showed higher corre-
lations with attention, action observation, and working memory than VAEs of the
other two stimuli, which may indicate that people pay more attention to body stim-
uli and that the VAE of bodies demands more cognitive resources than the VAE of
other two stimulus types. The VAE of paintings exhibited higher correlations with
self-referential constructs because people may rely on personal information to make
sense of painting stimuli, whereas they do not need to extract meaning from body
and face stimuli.

Limitations and future directions

In this study, we performed several separate meta-analyses to detect any nuances

related to methodology variables, but at the cost of reduced statistical power—only
a small number of studies were included in each separate ALE meta-analysis. This
loss of statistical power may explain why certain regions (e.g., the fusiform gyrus)
appeared in the general meta-analysis but were not consistently shown across the
separate meta-analyses. Future research could consider task forms other than rating
tasks to include more studies, although this approach may involve the risk of inter-
ruption by some irrelevant concepts. Moreover, neuroaesthetics is a growing field
of research of interest among researchers (Nadal et al., 2012). Future meta-analyses
may be able to draw more reliable conclusions once more recently published studies
are included.
Another limitation that should be noted is that cognitive dispositions are crucial
for aesthetic experience (Marković, 2012). Based on existing models of aesthetics,

13
 22 X. Feng et al.

we proposed (in the definition) that VAEs involve the integration of visual input,
knowledge and experience, followed by a positive-valence judgment or decision. We
did not aim to test this hypothesis in the current meta-analysis, and it is unlikely
that the cognitive component of VAEs could be validated using the paradigm in this
study because the neural activation pattern related to the cognitive component (if
this is the case) may not differ across various contents of thoughts/feelings. Thus, if
we were to compare the evaluation of an aesthetic experience condition to the evalu-
ation of neutral or unattractive experience condition, the result would not include a
condition corresponding to cognitive component because this component would be
cancelled out by the shared pattern of the two conditions. In future studies, it may be
possible to compare tasks involving the evaluation of aesthetics to tasks that do not
involve this cognitive process (e.g., tasks that require the evaluation of symmetry) to
determine whether the cognitive process is a constituent of VAEs.

Conclusion

The aim of our study was to reveal the neural correlates of visual-related stimuli
based on a quantitative meta-analysis of fMRI data. In this meta-analysis, we spe-
cifically limited the aesthetic evaluation task types to prevent irrelevant concepts
from confounding the validity of the results. In addition, to complement the pre-
vious ALE meta-analyses focusing on specific stimulus types (Boccia et al., 2016;
Brown et al., 2011; Bzdok et al., 2011; Vartanian & Skov, 2014), this meta-analysis
attempted to include all types of visual stimuli to improve the generalizability of our
results, which showed consistent activation in the ACC, OFC and fusiform gyrus.
Moreover, the current meta-analysis is the first to compare a passive viewing condi-
tion and an explicit response condition to identify similar patterns of activation; the
results showed that aesthetic evaluations are automatic processes and indicated that
stronger activation in the ACC/OFC can be observed under passive viewing con-
ditions, likely reflecting that a stronger emotional experience occurred under pas-
sive viewing conditions. In addition, the conjunction analysis of different cultural
groups (a Western group and an East Asian group) provided some insight into the
universal cultural components of VAEs. The activation of the vACC/OFC may sug-
gest that aesthetic emotional and reward processing is a culturally universal com-
ponent shared by participants with East Asian and Western cultural backgrounds.
Finally, the decoding results indicated that the VAE process showed a certain degree
of similarity to the processes involved in object viewing, feelings of reward and PA,
including incentives, motivational feelings and a sense of value.

Supplementary Information  The online version contains supplementary material available at https://​doi.​
org/​10.​1007/​s40167-​021-​00102-z.

13
Neural representations of visual aesthetic experience (VAE):… 23

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