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Progress in Neurobiology 94 (2011) 39–48

Contents lists available at ScienceDirect

Progress in Neurobiology
journal homepage: www.elsevier.com/locate/pneurobio

The neural foundations of aesthetic appreciation

Camilo J. Cela-Conde a,1, Luigi Agnati b,2, Joseph P. Huston c,3, Francisco Mora d,4, Marcos Nadal a,*
a
Human Evolution and Cognition (IFISC-CSIC), University of the Balearic Islands, Spain
b
Section of Physiology, Department of Biomedical Sciences, University of Modena and Reggio Emilia, Modena, Italy
c
Center for Behavioral Neurosciences, University of Düsseldorf, Düsseldorf, Germany
d
Department of Physiology, University Complutense of Madrid, Spain

A R T I C L E I N F O A B S T R A C T

Article history: The study of the cognitive and neural underpinnings of aesthetic appreciation by means of neuroimaging
Received 22 July 2010 techniques has yielded a wealth of fascinating information. Although the results of these studies have
Received in revised form 8 February 2011 been somewhat divergent, here we provide an integrative view of the early approaches, which identified
Accepted 11 March 2011
some of the core mechanisms involved in aesthetic preference. Then, a number of more specific issues
Available online xxx
under the perspective of recent work are addressed. Finally, we propose a framework to accommodate
these findings and we explore future prospects for the emerging field of neuroaesthetics.
Keywords:
ß 2011 Elsevier Ltd. All rights reserved.
Neuroaesthetics
Aesthetic appreciation
Neuroimaging
MEG
fMRI

Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
2. Neuroimaging techniques reach to the arts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
2.1. Psychological processes related with the visual appreciation of aesthetics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
2.2. Neural activity identified by the early neuroimaging studies. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
2.3. Models of cognitive processes underlying visual aesthetic preference . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
2.3.1. Temporal span . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
2.3.2. Decision and attention . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
2.3.3. Affective processes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
3. Outstanding issues raised in recent neuroimaging approaches to aesthetic appreciation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
3.1. Recognition and familiarity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
3.2. Bottom-up vs. top-down pathways . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
3.3. The influence of expertise . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
4. Summary and future prospects for neuroaesthetics. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47

Abbreviations: BA, brodmann area; DOC, dorsal occipital cortex; EEG, electroen- 1. Introduction
cephalography; FG, fusiform gyrus; fMRI, functional magnetic resonance imaging;
IOG, inferior occipital gyrus; IPS, intraparietal sulcus; MEG, magnetoencephalogra-
Objects created by visual artists, such as oil paintings,
phy; PFC, prefrontal cortex; TPJ, temporoparietal junction.
* Corresponding author. Tel.: +34 971 259899; fax: +34 971 173190. sculptures, etchings, drawings, watercolors and even glossies
E-mail addresses: cjcela@atlas.com.es (C.J. Cela-Conde), luigi.agnati@gmail.it, and decorative arts and crafts, constitute remarkable physical
luigi.agnati@unimore.it (L. Agnati), huston@uni-duesseldorf.de (J.P. Huston), expressions of the human capacity to produce and appreciate
francisco-mora@med.ucm.es (F. Mora), marcos.nadal@uib.es (M. Nadal). beauty. All human cultures have studied, discussed and engaged in
1
Tel.: +34 971 173029; fax: +34 971 173473.
2 these activities, though most often from a humanistic, subjective
Tel.: +39 0412207202.
3
Tel.: +49 211 8114296; fax: +49 211 8112024. and descriptive perspective. Neuroscientists and neuropsycholo-
4
Tel.: +34 91 394 1611; fax: +34 91 394 1628. gists have recently approached the traditionally philosophical field

0301-0082/$ – see front matter ß 2011 Elsevier Ltd. All rights reserved.
doi:10.1016/j.pneurobio.2011.03.003
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40 C.J. Cela-Conde et al. / Progress in Neurobiology 94 (2011) 39–48

of aesthetics aiming to characterize the neural and evolutionary The impact of degenerative diseases. Artists suffering from
foundations of our species’ capacity to appreciate beauty and art. Alzheimer’s disease seem to gradually lose the ability to represent
This approach, known as neuroaesthetics, has begun to provide the world with precision, but are able to make use of color and form
some insights into the neurobiological bases of aesthetic apprecia- in aesthetically appealing ways (Miller and Hou, 2004). Chatterjee
tion (Chatterjee, 2011; Dissanayake, 1992; Jacobsen, 2010; Miller, (2004a) notes that the production of these artists continues only as
2000, 2001; Nadal and Pearce, in press; Ramachandran and part of the general routines that the artists performed throughout
Hirstein, 1999; Skov and Vartanian, 2009; Zeki, 1999a,b). their lifetimes, and only if others around them provide the
Until recently, researchers could only empirically study the necessary setting to do so. Some patients with a specific form of
biological mechanisms underlying artistic and aesthetic apprecia- frontotemporal dementia, known as semantic dementia, which
tion and creation by examining single cases of patients suffering involves neural degeneration in the left anterior temporal lobe,
from brain lesions or neurodegenerative diseases. Despite their exhibit new interest in art. These patients approach their pictorial
great inherent interest, such accounts were often anecdotal and activity compulsively, in spontaneous spouts of creative produc-
difficult to understand in the absence of an adequate theoretical tion, often painting repetitive motifs, leading to progressive
framework. Chatterjee (2011) refers to them as ‘‘informative improvements to their painting. These depictions are usually
anecdotes’’ (Chatterjee, 2011, p. 54). Meaningful conclusions realistic, lacking in symbolism or abstract elements (Miller and
regarding the impact of different neurological conditions on Hou, 2004).
artistic and aesthetic activities have only emerged after the cases Other neurological conditions. Epilepsy and migraine have been
have been gathered and analyzed together by Bäzner and found to increase the variety of visual elements available to artists
Hennerici (2006), Bogousslavsky (2005), Chatterjee (2004a, who suffer from these conditions, as well as a source of inspiration
2006), Miller and Hou (2004) and Zaidel (2005, 2010). The main to them (Chatterjee, 2004a).
conclusions derived form these analyses merit examination before These observations reveal that neurological conditions can have
we turn to neuroimaging studies. diverse –and maybe even distinctive– effects on artistic produc-
General observations. Despite their proficient visuo-motor and tion. But what about their influence on the other side of the artistic
musical skills, artists are vulnerable to the same visual, motor, coin –appreciation? What can similar cases tell us about the
auditory and cognitive neuropsychological deficits that affect biological underpinnings of aesthetic appreciation? Given the very
other people. The difference, in Chatterjee’s (2004a) words, is that few studies that have been carried out on this issue, and their
artists manifest these deficits in strikingly eloquent ways. Most of anecdotal nature, we cannot offer a comprehensive response. At
the studied artists continued to be artistically motivated, produc- present we can only provide a tentative and provisional picture.
tive and expressive after the onset of their condition. Personal style The effect of strokes on artistic appreciation. Griffiths et al. (2004)
is usually preserved to a certain extent, probably due to the years describe a patient who suffered an infarct that left him unable to
devoted to practicing their skills (Zaidel, 2005). There is no direct experience emotion in response to music. The lesion mainly
relation between suffering from neuropsychological conditions affected the left insula, but extended into the left frontal cortex and
and improvised quality of art production. Chatterjee (2004a) notes amygdala. Speech, which was also initially affected, was recovered
several instances in which the effects of the condition on the after 12 months. However, even though his perception of diverse
artworks were aesthetically surprising and pleasing. musical features was normal, 18 months after the stroke the
The effect of strokes on artistic production. There is a noticeable patient was still emotionally unaffected by music, despite the fact
change in the work of most artists who have suffered a stroke. that during that time he was able to enjoy other activities. These
Many of them resume their creations, though only after observations led the authors to suggest that perceptual and
overcoming various forms of disability. Some of these artists emotional components of music processing might rest on
had to change from working with their dominant to their non- functionally and anatomically distinct neural networks, and that
dominant hand (Bäzner and Hennerici, 2006). Although lesions in the insula is a crucial piece in the neural underpinnings of the
either hemisphere seem to leave traces on the production of artists emotional response to music.
(Zaidel, 2005), spatial disorganization (perspective, third dimen- The impact of degenerative diseases. Halpern et al. (2008) found
sion, contours), neglect and distortion in facial representation are that artistically untrained patients with Alzheimer’s disease were
more obvious in the artworks of artists who suffered a right consistent in their preference for visual art on repeated presenta-
hemisphere stroke (Bäzner and Hennerici, 2006). tions, despite being unable to remember they had seen the art on
The impact of visual agnosia. The specific effect of visual previous occasions. It seems, thus, that the neurodegeneration that
agnosia on artists’ creations depends to a large extent on causes Alzheimer’s disease does not prevent these patients from
whether object recognition problems are related with their expressing aesthetic preference, and that their preferences are as
perceptual or conceptual features (Chatterjee, 2004a). In the first stable over time as those of healthy people. Two reports of patients
case, artists will often not render the overall form and with frontotemporal dementia describe how their preferences for
composition of the depicted objects, but will include some of music changed remarkably (Boeve and Geda, 2001; Geroldi et al.,
their conspicuous features. In the latter case, artists are still 2000). These three patients began compulsively listening to music
adept at drawing when copying, but seem completely incapable they had not previously enjoyed, playing it for hours on end.
when asked to draw from memory, and have to rely on their Other neurological conditions. Sellal et al. (2003) present a case of
knowledge of the world. an epilepsy patient who underwent left temporal lobe resection,
The impact of aphasia. Bäzner and Hennerici (2006) argue that which only spared the hippocampus, the parahippocampal gyrus,
there is little evidence indicating a significant impact of aphasia on and the amygdala. This case is interesting because the surgically
the creation of visual art, which suggests that verbal and visual removed brain region corresponds roughly to that which typically
production may be related with distinct output channels. degenerates in the form of frontotemporal dementia mentioned
However, Bogousslavsky (2005) and Chatterjee’s (2004a) revisions above. During the first year after surgery the patient became aware
reveal that whereas the production of some aphasic artists seems that he no longer enjoyed listening to rock music, and that he now
to be largely unaffected, other artists become more expressive, and preferred Celtic or Corsican polyphonic singing. His taste in
yet others begin producing works with different contents, literature also shifted, in this case from science fiction to Kafkian-
suggesting that this condition may be to broad as a window into inspired novels. The authors report that the patient also began
the biological bases of artistic and aesthetic production. showing increased preference for realistic paintings, enjoying the
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C.J. Cela-Conde et al. / Progress in Neurobiology 94 (2011) 39–48 41

small details that previously went unnoticed to him. These changes Thus, it seems that a focus on selected aspects of aesthetic
in aesthetic preference are in contrast with his unchanged appreciation cannot do justice to the whole panorama of what
preferences for food, fashion or faces. constitutes the appreciation of beauty by the mind/brain.
Damage to the amygdala. The crucial role of subcortical brain Explaining such appreciation requires at least a three-step
structures related with emotional processing in aesthetic appre- strategy:
ciation was revealed by two studies of the effects of damage to the
amygdala. Adolphs and Tranel (1999) report that there were (I) An account of the brain regions and neural networks that are
differences between the preferences for visual stimuli of two related with each phenomenon studied experimentally.
patients with bilateral amygdala damage and a group of healthy (II) An explanation of the diverse psychological aspects involved
controls. Both patients expressed higher liking for three-dimen- in the perceptual/appreciative tasks performed by partici-
sional geometrical shapes, landscapes and color arrangements pants.
than the control participants. This difference was especially clear (III) A general, integrative model of the cognitive tasks performed,
for the stimuli least liked by controls. Similar results were obtained that may eventually explain the qualitative aspects related
from the examination of musical preferences in a patient with with those cognitive tasks.
almost exclusive bilateral damage to the amygdala (Gosselin et al.,
2007). In this case, whereas the patient was able to correctly
process musical features, even tempo and mode, she showed Here, we will try to offer a preliminary approach to such a
selective impairment in the recognition of scary and sad music, but strategy. We will start with the four 2004–2006 initial studies,
not of happy music. Thus, it seems that the amygdala plays a role in providing a synthetic account of what they revealed about the
the affective processes underlying aesthetic preference, especially visual appreciation of beauty. Thereafter, we will refer to later
in the experience of disliking and in relation to negatively valenced experiments that have addressed certain outstanding matters
stimuli. related with aesthetic appreciation. Finally, we will try to assess
The advent and refinement of non-invasive neuroimaging how cognitive models of the appreciation of beauty fit this set of
techniques has allowed researchers to build upon these informa- results.
tive cases, and to address similar and new issues in healthy The focus of our analysis will be on the aesthetic experience,
subjects in controlled situations, and to correlate appreciation and insofar as it encompasses the concept of ‘‘beauty’’. This constitutes
enjoyment of music, painting, architecture or sculpture, among a narrow realm in modern aesthetics. Aesthetic qualities refer to
other forms of art, with the activity of several brain structures. In judgments derived from sensory contemplation of phenomena
the present review we first examine the psychological processes and, especially around the 18th century, the concept of beauty is
related to the visual appreciation of aesthetics and then bring intricately linked to theories of aesthetics. However, in many
together the results of early neuroimaging studies and provide an modern art forms, such as conceptual art, the prime role of beauty
integrative framework of the underlying neural and cognitive has diminished, and is even actively challenged. Thus, an ugly
foundations. We then show how the most recent neuroimaging object may still be a great work of art.
approaches have addressed certain specific issues and examine
how their results fit within this general framework. Finally, we 2.1. Psychological processes related with the visual appreciation of
point out some of the most relevant and challenging questions that aesthetics
arise when this framework is viewed from a broader perspective. In
this review we will only be concerned with the appreciation of Ever since Berlyne’s (1971, 1974) work, it has been known that
visual art, though many forms of human artistic production, such aesthetic appreciation rests on diverse psychological processes.
as music, literature, theatre, opera, and cinematography, share Pleasure, expectation, surprise, recognition and interest, to
important aspects with the visual arts. mention but a few of these, undoubtedly play a role in the
appreciation of beauty. Contextual features also seem to influence
spectators’ responses. Additionally, participants’ particularities,
2. Neuroimaging techniques reach to the arts including social, historical, cultural, biological, educational, and
personality variables, are also known to shape aesthetic experi-
In 2004, several neuroimaging studies began constructing the ences. Variables related with attention to the stimulus, the
basic picture of the neural correlates of visual aesthetic apprecia- motivation to attend and evaluate the stimulus, the emotional
tion (Cela-Conde et al., 2004; Kawabata and Zeki, 2004; Vartanian response to the stimulus and its rewarding properties contribute to
and Goel, 2004). Another paper on the same topic soon followed the prioritization and selection of all information, including visual
(Jacobsen et al., 2006). Due to the fact that these experiments were stimulation (Raymond and Narayanan, 2009).
designed independently, they employed different research strate- Researchers interested in the biological underpinnings of
gies and protocols. Thus, it is not surprising that the results aesthetic experiences are faced with the problem of dealing with
obtained were quite disparate (for an account about how these its complex and multifaceted nature. Can neuroimaging techni-
differences might have affected results, see Nadal et al., 2008). ques adequately deal with the interplay of such varied cognitive,
The relation between cognitive and neural processes involved affective, personal, social and cultural factors? The straightforward
in aesthetic appreciation is highly complex and intricate. The answer is that their strong methodological restrictions do not
activity of neural networks connecting diverse areas is still crucial allow the observation of brain activity reflecting the whole set of
for the simplest cognitive processes. Even leaving aside the more processes involved in aesthetic appreciation. As noted by
philosophical issue of ‘‘qualia’’, i.e., the intimate, personal Chatterjee (2011), researchers have responded to this challenge
experience of perception, the issue is formidably complex. The by studying the neural bases of only few specific aspects or
biological foundations of conscious experience seem to involve components of aesthetic appreciation at a time. This is done under
coordinated firing between groups of neurons belonging to several the tacit –as well as untested– assumption that a reasonably good
different, and often distant, areas. This notion was already understanding of the whole aesthetic experience can be gained
advocated by early scientific approaches to consciousness (Crick from the sum of its parts.
and Koch, 1990; Von der Malsburg, 1981; Cela-Conde and Marty, One favored option has been to study the isolated effects of
1997), and it is still defended today (Shulman et al., 2009). certain variables, such as expertise (Berkowitz and Ansari, 2010;
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42 C.J. Cela-Conde et al. / Progress in Neurobiology 94 (2011) 39–48

Kirk et al., 2009a; Müller et al., 2009) or context (Kirk et al., 2009b). The neuroimaging results could be summarized as follows:
But inevitably, neuroimaging methods impose serious restrictions
on the tasks that participants can be required to perform. (i) Kawabata and Zeki (2004) registered participants’ brain
Experimenters design such tasks guided by two constraints: the activity with fMRI (functional magnetic resonance imaging)
specific research hypothesis and the desire to capture meaningful while they rated the beauty of stimuli. They found that activity
aspects of their participants’ aesthetic experience. By far, the most was greater:
common task participants have been asked to perform in these – in the orbitofrontal cortex for stimuli classified as beautiful;
kinds of neuroimaging experiments is rating specific features of the – in the motor cortex for stimuli classified as ugly.
stimuli or certain aspect of their experience. This practical strategy (ii) By means of magnetoencephalography (MEG), Cela-Conde
necessarily involves losing a considerable amount of information et al. (2004) identified activity in the left dorsolateral
about many other non-measured aspects of the experience. So the prefrontal cortex. This activity was especially significant
choice of task is crucial. when participants judged stimuli as beautiful, as compared
A possible solution to this dilemma is provided by Eysenck’s to being non-beautiful, between 400 and 1000 ms after stimuli
(1940) general factor of aesthetic preference. The fact that this onset.
general factor was found to correlate with the ranks awarded to (iii) By means of fMRI, Vartanian and Goel (2004) found a double
odoriferous substances, several colors and a variety of polygonal correlation:
figures, led him to surmise that it was directly constrained by the – activity in the right caudate nucleus decreased as preference
properties of the nervous system. Crucially for the requirements of ratings for stimuli decreased;
neuroimaging studies, he believed that, with regards to visual – activity in the left anterior cingulate gyrus and bilateral
stimuli, this factor accounted for people’s appreciation of their occipital gyri, increased with preference ratings.
beauty (Eysenck, 1940). Several subsequent studies have found (iv) Although, as we have already pointed out, Jacobsen et al.’s
support for the notion that beauty ratings capture the aesthetic (2006) study differed from the previous three, they did
experience in a more meaningful way than ratings of, for instance, identify some differences when comparing brain activity
originality, interest, or pleasingness (Pickford, 1955; Looft and related with stimuli rated as beautiful and ugly, especially in
Baranowski, 1971; Seifert, 1992; Cupchik et al., 1992; Marty et al., the frontomedian and anterior cingulate cortex.
2003; Jacobsen et al., 2004).
This is not to say that rating the beauty of visual or auditory
2.3. Models of cognitive processes underlying visual aesthetic
stimuli, as is required from participants in many neuroimaging
preference
studies, rests on a simple single cognitive process. On the contrary,
any model of the cognitive processes involved in the appreciation
Chatterjee’s (2004b) theoretical model of the cognitive and
of beauty must also account for multiple factors, such as meaning
affective processes involved in visual aesthetic preference, based
and recognition, decision judgments, affective processes (pleasure,
on visual neuroscience, provides a means to organize the results
disgust), and reward. These factors multiply the complexity of
obtained in the 2004–2006 neuroimaging studies within a series of
visual aesthetic appreciation. In contrast, the experience of beauty
information-processing phases. The model posits the following
seems to be, introspectively speaking, something quite simple –
stages:
either we find an object to be more or less beautiful, or we find it
indifferent. Nonetheless, the complexity of the structure of
1. The elementary visual attributes of an artwork are processed
aesthetic appreciation and the relative simplicity of its manifesta-
like any other visual object’s (primary and secondary visual
tion do not collide head on. Since the timescale of the brain’s
brain areas).
functional activity is in the order of milliseconds, complex
2. Attentional processes redirect information processing to promi-
processes take place very quickly, leading to a qualitative, simple
nent visual properties, such as color, shape, and composition
subjective conclusion about the beauty of a stimulus. How to
(frontal–parietal networks).
understand such almost instantaneous complexity should be the
3. Attentional networks modulate processing within the ventral
objective of any model of aesthetic appreciation.
visual stream that leads to attributional networks – the ‘‘what
pathway’’, i.e., experience of the stimulus, attributes and
2.2. Neural activity identified by the early neuroimaging studies
contents, such as faces and landscapes (temporal lobe).
4. Feed-back/feed-forward processes linking attentional and
Four initial neuroimaging studies (Vartanian and Goel, 2004;
attributional circuits that enhance the experience of the visual
Kawabata and Zeki, 2004; Cela-Conde et al., 2004; Jacobsen et al.,
object.
2006) intended to identify the neural correlates of aesthetic
5. Emotional systems intervene in most cases (anterior medial
appreciation of visual stimuli. Three of them (Cela-Conde et al.,
temporal lobe, medial and orbitofrontal cortices, and subcortical
2004; Kawabata and Zeki, 2004; Vartanian and Goel, 2004)
structures) (Chatterjee, 2004b).
registered participants’ brain activity while they expressed their
preference for visual stimuli or rated their beauty. Their main aim
was to compare activity associated with stimuli appraised Jacobsen and Höfel’s (2003) proposal offers a complementary
positively and negatively by each individual participant. The view of the general cognitive processes involved in aesthetic
fourth study (Jacobsen et al., 2006) addressed a slightly different appreciation. On the grounds of several electroencephalographic
issue, asking participants to rate the beauty and symmetry of (EEG) experiments, they established a temporal sequence for
simple visual stimuli, namely, black and white geometric patterns processes in a two-stage model. During the first stage, which takes
created by the authors. This study compared neural activity while place at around 300 ms after stimulus onset, an initial impression
participants rated the beauty of images with activity while they is formed. This process is associated with anterior frontomedian
rated their symmetry. Thus, the results obtained concerning activity, mainly when participants consider stimuli to lack
beauty refer to the neural correlates of the judgment of beauty per aesthetic value. A deeper aesthetic evaluation, related with broad
se. It is important to note that all these researchers considered that right hemisphere activity, begins at close to 600 ms. A summary of
a subjective rating by the participants themselves was essential, Chatterjee (2004b) and Jacobsen and Höfel’s (2003) models of
given that it is likely that no general condition of ‘‘beauty’’ exists. aesthetic appreciation is given in Table 1.
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C.J. Cela-Conde et al. / Progress in Neurobiology 94 (2011) 39–48 43

In order to explore the relation between Chatterjee (2004b) and and Cela-Conde et al.’s (2004) reported activity refers to the
Jacobsen and Höfel’s (2003) models and the 2004–2006 neuroim- difference between ratings of ‘‘beautiful’’ and ‘‘not-beautiful’’.
aging results, we will address issues related with the temporal Hence, while there was more activity in the dorsolateral prefrontal
span of the cognitive processes, decision and attention, and cortex while Cela-Conde et al.’s (2004) participants viewed stimuli
emotional engagement, separately. they rated as beautiful than when they viewed the ones they rated
However, before we do so, we feel it is important to underscore as not-beautiful, there was more activity in the frontomedian
the fact that our main objective is to provide an integrative picture cortex while Jacobsen and Höfel’s (2003) participants performed
which can serve as the source for novel hypothesis. One of the main beauty judgments than when they performed symmetry judg-
challenges faced by the field of neuroaesthetics, as rightfully ments.
pointed out by Chatterjee (2011), is the reliance on the reverse Taking such methodological considerations into account allows
inference, that is, considering activity in specific brain regions to be us to understand the discrepancy between both studies. The
an indication of the involvement of given cognitive processes. The frontomedian prefrontal cortex has been shown to be involved in
impressive growth of our knowledge of the neural underpinnings self-referential evaluative judgments (Northoff and Bermpohl,
of aesthetic appreciation owes a great deal to neuroimaging 2004; Zysset et al., 2002), and it is to be expected that beauty
studies. In many cases, however, researchers have inferred that judgments, which are subject-based, would involve self-referential
participants were performing certain cognitive or affective processing to a greater degree than symmetry judgments, which
operations from the resulting brain activity. If such brain regions are object-based. Jacobsen and Höfel’s (2003) use of abstract
were known to be related with single specific cognitive processes geometric patterns to elicit beauty judgments might have
this would not be a problem. With very rare exceptions, though, increased the subjective elements of such judgments, where
this has not been the case – the same brain region may be involved participants could only base their decision on internally generated
in several diverse processes. But Poldrack (2006) does note that the information. In contrast, Cela-Conde et al.’s (2004) approach might
reverse inference can constitute the source for novel hypotheses have encouraged participants to base their decisions about beauty
that can be tested in subsequent studies. Hence, when integrating on the richer external information provided by the stimuli, such as
neuroimaging results with Chatterjee (2004b) and Jacobsen and their style, explicit content, and degree of artistry. This interpreta-
Höfel’s (2003) models in the following sections, we do so aiming to tion is in accordance with Christoff and Gabrieli’s (2000)
stimulate the formulation of hypotheses to be tested in future suggestion that while activity in the dorsolateral prefrontal cortex
studies, in line with Chatterjee’s (2011) caveat. seems to be primarily involved with information generated
externally, activity in the frontomedian prefrontal cortex seems
2.3.1. Temporal span to reflect the engagement of processes related with the evaluation
Only the study by Cela-Conde et al. (2004) was well suited, due and manipulation of internally generated information.
to the temporal resolution of MEG, to test the sequence of activities In sum, we suggest that Jacobsen and Höfel’s (2003) results
suggested by Chatterjee (2004b) and, especially, Jacobsen and reflect the greater involvement of self-referential processing of
Höfel (2003). It reported left prefrontal dorsolateral cortex activity internally generated information when rating the beauty of
in the range of 400–900 ms after stimulus onset. Thus, MEG studies geometric patterns than when rating their symmetry, and that
partially support the temporal sequence proposed by Chatterjee Cela-Conde et al.’s (2004) results reflect the greater involvement of
(2004b) and Jacobsen and Höfel (2003), but it is not fully consistent processing of externally provided information related with style,
with the localization of brain activity specified in Jacobsen and content, artistic status, and so on, while viewing images rated as
Höfel’s (2003) model. Specifically, whereas Jacobsen and Höfel’s beautiful. Together, however, both studies show that aesthetic
(2003) early activity was identified in the frontomedian cortex, experiences involve an early phase of evaluation, which can be
especially while participants viewed stimuli they rated as lacking mainly driven by internally or externally elaborated information. It
aesthetic appeal, Cela-Conde et al.’s (2004) early activity was seems implausible that such evaluation actually represents the
circumscribed to the dorsolateral prefrontal cortex, especially final decision arrived at by the participants. More likely, this early
while participants viewed stimuli they rated as beautiful. brain activity associated with evaluative processes represents a
We believe that this discrepancy can be accounted for by the fast impression formation which influences other processes
differences in the stimuli used in both studies and the kind of related with attention, perception, response selection, and so on,
statistical contrasts they explored. Whereas Cela-Conde et al. but which may be crucial for decisions to continue or interrupt
(2004) presented their participants with a broad array of artistic engagement with the stimulus.
and non-artistic abstract and representational visual stimuli,
Jacobsen and Höfel (2003) used a relatively homogeneous pool 2.3.2. Decision and attention
of abstract geometric patterns that varied in complexity and Decisions about the beauty of stimuli seem to be mediated by
symmetry. In addition, Jacobsen and Höfel’s (2003) results emerge the dorsolateral prefrontal cortex (Cela-Conde et al., 2004),
from contrasting beauty judgments with symmetry judgments, supporting Chatterjee’s (2004b) model. The prediction by Jacobsen

Table 1
Chatterjee’s (2004b) and Jacobsen and Höfel’s (2003) models of aesthetic appreciation. Comparison of the time course of brain activity underlying aesthetic appreciation
predicted by both models.

Temporal span Cognitive process Main regions involved

Chatterjee (2004b) Jacobsen and Höfel (2003)

Early latencies Visual processing Occipital

Intermediate latencies 300–400 ms Initial impression Anterior frontomedian
Undetermined latencies Attention given to stimuli shape and color features Frontal–parietal
Undetermined latencies Attributional experience of stimuli Temporal
Late latencies <600 ms Aesthetic evaluation Frontal–parietal Right hemisphere
Late latencies Enhancement of aesthetic experience Frontal–parietal–temporal
Undetermined latencies Emotional engagement Medial temporal
Deep aesthetic experience Medial and orbitofrontal
Subcortical
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and Höfel’s (2003) model related with negative aesthetic evalua- amygdala activity, but it entailed the comparison between their
tion is supported by the frontomedian prefrontal cortex activity portrait and non-portrait conditions, independently of their
identified by Jacobsen et al. (2006), who, importantly, used the aesthetic rating. This particular result is not surprising, given that
same kind of stimuli as Jacobsen and Höfel (2003). the amygdala is a crucial element in a distributed network of face
Regarding attention given to the stimuli as a function of their processing (Haxby et al., 2000), and its key role in analyzing and
symmetry, Jacobsen et al. (2006) reported that judgments of directing attention to visual social stimuli (Adolphs and Spezio,
beauty activated mainly the frontomedian cortex, but also the left 2006).
intraparietal sulcus of the symmetry network. The authors hold Winston et al.’s (2007) results, which showed that activity in
that aesthetic judgments of beauty rely on a network that partially the amygdala followed a non-linear response pattern in relation to
overlaps with another network that underlies evaluative judg- facial attractiveness suggest another possibility. Specifically,
ments based on social and moral cues. The attentional engagement activity was greater for highly attractive and highly unattractive
with visual stimuli during aesthetic experiences was also found by faces, compared to intermediately attractive faces. Thus, absence
Vartanian and Goel (2004). They reported activity in the occipital of amygdala activity reported in the neuroaesthetic literature may
cortex when participants rated their preference for visual stimuli, be due to the fact that neuroimaging studies have usually not
reflecting an enhanced processing of certain features of preferred included the extremely pleasant and unpleasant stimuli required
stimuli. These studies underscore the importance of attentional to detect amygdalar activity.
processes involved in aesthetic appreciation and their contribution
to deciding about the beauty of an object or one’s preference for it. 3. Outstanding issues raised in recent neuroimaging
approaches to aesthetic appreciation
2.3.3. Affective processes
Affective and emotional processes are an integral part of our After the 2004–2006 starting point, a number of articles
experience of artistic and aesthetic manifestations. It has been continued to explore distinct aspects of the neural underpinnings
shown, for instance, that the enjoyment of music involves activity of aesthetic appreciation. Hence, we will select certain aspects that
in brain regions responsible for pleasurable feelings (Blood and we consider particularly relevant due to their important relation to
Zatorre, 2001; Blood et al., 1999). A fact that became evident with the cognitive processes involved in aesthetic appreciation. These
the initial neuroimaging studies (Cela-Conde et al., 2004; Jacobsen aspects are: recognition and familiarity; bottom-up vs. top-down
et al., 2006; Kawabata and Zeki, 2004; Vartanian and Goel, 2004) is pathways; and the influence of expertise on the neural correlates of
that aesthetic appreciation of visual stimuli involves several aesthetic appreciation.
different affective processes.
Despite its temporal resolution advantages, the MEG technique, 3.1. Recognition and familiarity
used in Cela-Conde et al.’s (2004) study, is severely limited in its
access to signals from the orbital and ventromedial prefrontal Together with other collative variables, Berlyne (1971) consid-
cortex, anterior cingulate cortex, and all subcortical brain regions. ered familiarity to exert a strong influence on the psychobiological
Thus, that study was unable to offer any meaningful insights into mechanisms underlying aesthetic experiences. A number of
the affective side of aesthetic appreciation. However, the fMRI subsequent psychological studies have shown that people usually
studies performed by Jacobsen et al. (2006), Kawabata and Zeki prefer familiar stimuli, an effect currently explained under the
(2004) and Vartanian and Goel (2004) demonstrated that beauty umbrella of the processing fluency theory. Reber et al. (2004)
and preference ratings correlate with activity in several brain suggested that objects vary with regards to the fluency with which
regions related with positive affective states: temporal pole they are processed. Among the features that can facilitate
(Jacobsen et al., 2006), orbitofrontal cortex (Kawabata and Zeki, processing fluency they included goodness of form, symmetry,
2004), caudate nucleus, and anterior cingulate cortex (Vartanian clarity, figure–ground contrast, repeated exposure and familiarity.
and Goel, 2004). These results show that positive aesthetic Given that fluent processing is experienced as hedonically
experiences involve several different affective processes: the pleasurable, and that aesthetic experiences are strongly influenced
creation of a context based on past emotional experiences by affective states, it follows that positive aesthetic experiences
(Jacobsen et al., 2006), the assessment of the reward value of arise especially from fluent processing, such as that afforded by
the stimuli (Kawabata and Zeki, 2004), and the integration of prototypical exemplars of a category (Winkielman et al., 2006).
information from reward and cognitive processing cortical regions Fairhall and Ishai (2008) showed that the perception of familiar
to elaborate an appropriate response (Vartanian and Goel, 2004). content in works of art rests on several cognitive processes related
It is worth noting that none of the 2004–2006 neuroimaging with activity across a distributed cortical network. These processes
studies found significant activity in the amygdala associated with include object recognition, memory recall and mental imagery. In
aesthetic preference ratings. On the grounds of Adolphs and line with the processing fluency theory, Fairhall and Ishai (2008)
Tranel’s (1999) results, which suggested that the amygdala is showed that recognition was significantly influenced by the degree
involved in preference for visual stimuli that are generally of abstraction in artworks, strongly dependent on some of the
considered to be aversive, amygdala activation might have been features that Reber et al. (2004) and Winkielman et al. (2006)
expected in neuroimaging studies of aesthetic appreciation, showed to determine processing fluency, such as clarity or figure–
specifically when participants viewed stimuli considered as ugly. ground contrast. Specifically, participants rapidly recognized
The MEG neuroimaging technique used by Cela-Conde et al. familiar objects in representational pictures, but were slower to
(2004) does not allow the recording of activity in the amygdala, do so for abstract or almost-abstract paintings. The study showed
due to its spheroid structure. In addition, signals from medial that recognition of familiar objects in paintings relies on brain
temporal brain regions are notoriously difficult to register even regions related with the processing of object form, as Chatterjee’s
with fMRI unless special measures are adopted. Thus, one (2004b) model predicted, namely the inferior occipital gyrus (IOG)
possibility for the lack of significant activity associated with and fusiform gyrus (FG). Also, the dorsal occipital cortex (DOC),
aesthetic appreciation in the amygdala is that Kawabata and Zeki where configural relationships are processed, and the intraparietal
(2004), Vartanian and Goel (2004), and Jacobsen et al.’s (2006) sulcus (IPS), which is modulated by attentional demands, were
studies did not have adequate signal from this structure. It is true activated (Fairhall and Ishai, 2008). Vartanian and Goel’s (2004)
that one of Kawabata and Zeki’s (2004) contrasts did reveal results of enhanced activation in FG when representational
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C.J. Cela-Conde et al. / Progress in Neurobiology 94 (2011) 39–48 45

paintings were compared with filtered paintings, were also studies have set out to provide an explanation for them. The most
partially confirmed. that they have shown is that art-trained and naı̈ve participants
The recognition of meaningful, familiar content in art works, as perceptually explore artworks differently. Eye-tracking experi-
part of the representational paintings, not present in the abstract ments studying both kinds of participants’ exploration of paintings
ones, is mediated by activation in the temporoparietal junction have converged on the finding that increasing levels of expertise
(TPJ). Kawabata and Zeki (2004) reported that the left TPJ was are related with a deeper visual exploration of the overall
activated by ‘beautiful’ rather than ‘neutral’ art paintings. composition, rather then the constituent elements, which are
However, the aesthetic judgments of representational and the focus of art–naı̈ve experts. Experts spend significantly more
indeterminate paintings resulted in virtually identical ratings of time looking at background features, the composition, and the
affect (Ishai et al., 2007). Therefore, the stronger activation of TPJ, color contrasts, whereas untrained viewers spend more time
evoked by representational paintings, cannot be explained by the looking at individual figurative elements and exploring figures in
affective component of aesthetic preference. Crucially, Kawabata the centre and foreground (Nodine et al., 1993; Vogt and
and Zeki’s (2004) experimental protocol included a stimuli pre- Magnussen, 2007; Winston and Cupchik, 1992). These effects
selection procedure, which required participants to preview and may be stronger for abstract than representational paintings
rate all the stimuli they would be subsequently shown during the (Zangemeister et al., 1995). Such results are coherent with purely
fMRI scanning session. This might have inadvertently elicited behavioral studies such as Hekkert and van Wieringen’s (1996),
recognition processes in the task the participants performed in the which revealed that features such as color and degree of realism –
scanner. Prior studies have revealed that single exposures are whose variation alters the appearance of the elements but not the
enough for participants to recognize previously presented art- overall composition– had a stronger effect on the way naı̈ve
works and to bias aesthetic ratings (Cela-Conde et al., 2002; Nadal viewers appraise paintings.
et al., 2006). Even in the absence of explicit recognition, such pre- Neuroscientists are facing up to the challenge of relating these
scanning presentation might have enhanced processing fluency of differences in aesthetic appreciation and visual processing
the stimuli (Reber et al., 2004; Winkielman et al., 2006). strategies with differences between experts and nonexperts in
neural activity in specific brain regions. For instance, Calvo-Merino
3.2. Bottom-up vs. top-down pathways et al. (2005) asked expert ballet and capoeira dancers, as well as
non-dancers, to view short video clips of capoeira and ballet
Jacobsen et al. (2006) aimed to verify previous interpretations movements. Participants were instructed to rate how tiring they
of the amygdala as intervening in bottom-up processing (Wright thought each movement was –an irrelevant task– while their brain
et al., 2008) by means of an aesthetic appreciation task. As we have activity was registered using fMRI. Results showed that when
already commented, none of the 2004 neuroimaging experiments expert dancers viewed movements corresponding to their own
of aesthetic appreciation found activation of the amygdala. Jacobs style compared to the other style there was greater activity in
et al. (2006) were also unable to identify significant bottom-up premotor cortex, parietal areas and superior temporal sulcus,
amygdalar activity associated with the rated beauty of stimuli which are involved in biological motion perception. Such results
(Jacobs et al., 2006). However, they reported a top-down process were mirrored by Orgs et al.’s (2008) electroencephalographic and
related with aesthetic appreciation and argued that a top-down Cross et al.’s (2006) fMRI experiments.
enhancement of amygdalar activation seems to be required when Calvo-Merino et al. (2010a) subsequently attempted to further
expressing aesthetic preference for low-arousal stimuli. characterize the biological substrate of dance expertise. They asked
Cupchik et al.’ (2009) study was explicitly designed to 24 ballet dancers and 24 non-dancers to perform a visual
disentangle the top-down orienting of attention and bottom-up discrimination task between upright and inverted dance move-
perceptual facilitation involved in aesthetic appreciation. Regard- ments. The inverted presentation of stimuli which are usually
ing the bottom-up pathway, Cupchik et al.’ (2009) results revealed viewed upright is known to specifically disrupt processing and
that when subjects viewed softedge paintings after receiving subsequent use of relational features in configural analysis (Busey
instructions to engage aesthetically with the stimuli there was a and Vanderkolk, 2005; Diamond and Carey, 1986; Rossion and
greater activity in the left superior parietal cortex. The authors Curran, 2010). Experts were better at discriminating upright dance
attribute this activation to viewers’ attempts to resolve the movements than non-experts. However, no differences were found
indeterminate forms in softedge paintings to construct coherent for inverted dance movements. These results suggest that dance
images. They believe the top-down pathway to be related with the expertise enhances mainly configural –as opposed to featural–
activation of the left lateral prefrontal cortex. Specifically, the processing of dance movements. This result is particularly
authors attribute this activation to the top-down organization of interesting, given that there seem to be two specialized routes
perception when participants adopted an aesthetic orientation for processing human bodies, one of which processes bodies in a
towards the visual stimuli, which is in agreement with the general configural manner, involving areas of the dorsal visual system and
involvement of lateral prefrontal cortex in the control of cognition the premotor cortex, and another, which appears to be specialized
(Cupchik et al., 2009). This result fits with the activation during late in the processing of specific details of body posture, and is part of
latencies (>400 ms) of the left dorsolateral prefrontal, in Cela- the ventral visual processing stream, encompassing the extra-
Conde et al.’s (2004) study. striate body area (Urgesi et al., 2007).
Calvo-Merino et al.’s (2010b) study was designed to determine
3.3. The influence of expertise the relative contribution of these two body-processing streams to
aesthetic appreciation. In this study, transcranial magnetic
A number of psychological experiments have revealed signifi- stimulation was applied over the dorsal and ventral pathways
cant differences in aesthetic appreciation as a function of people’s while participants indicated the extent to which they liked several
experience and knowledge of art. This line of research stretches dance postures. Results showed that the procedure especially
from Barron and Welsh (1952) and Munsinger and Kessen’s (1964) interfered with aesthetic appreciation when the stimulation was
pioneering work, to Winston and Cupchik (1992) and Hekkert and produced over the dorsal pathway, that is, the one underlying
van Wieringen’s (1996) more recent studies. Although such configural processing (Urgesi et al., 2007). These results reveal that
differences in aesthetic appreciation between groups with and general configural aspects of dance play a greater role in aesthetic
without art experience have been described many times, few appreciation than specific features of body postures. Although this
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46 C.J. Cela-Conde et al. / Progress in Neurobiology 94 (2011) 39–48

particular study did not include expert dancers, it follows from this as we noted at the beginning, there is no straightforward link from
line of researchers that expertise in dance enhances the neural a particular kind of neural network activation to the subjective
mechanisms underlying configural body processing, the same ones experience of considering a given stimulus to be ‘‘beautiful’’ –
that seem to be crucial for aesthetic appreciation of dance. leaving aside, moreover, that there is no clear definition of
Kirk et al. (2009a) designed a study to characterize the ‘‘beauty’’. Instead of a linear scheme of stimulus–activation–
biological underpinnings of architecture expertise. The authors appreciation, akin to a bottom-up perceptive ‘‘module’’ postulated
wished to determine whether architecture experts and nonexperts by Fodor’s (1983) architecture of mind, we would like to propose a
showed any differences in brain activity related with processes of different view, initially grounded on Chatterjee’s (2004b) model.
perception, memory or reward while viewing buildings. Architects Table 2 shows how Chatterjee’s (2004b) model can be related to
and non-architects were asked to indicate, on a 5-point scale, how neuroimaging results on aesthetic appreciation.
appealing they found a series of photographs of buildings, stimuli The interactions among cognitive and affective processes
for which only the group of architects were experts, and faces, predicted by the model have not been adequately tested yet,
stimuli for which both groups evidently had equal expertise. due to the lack of sufficient temporal resolution of the fMRI
The results showed that, independently of appeal, there were technique. The notion of connectivity has largely been considered
differences in activity between architects and non-architects in the by anatomists, neurologists and psychologists to be at the core of
hippocampus and precuneus while viewing the buildings. In explanations of consciousness (Crick and Koch, 1990). Since the
addition, there was significant activity related with appeal ratings times of von der Malsburg (von der Malsburg and Schneider, 1986)
in three distinct brain regions. First, activity in the medial consciousness has been associated with the presumed synchroni-
orbitofrontal cortex correlated positively with the level of appeal zation of neuronal network activity in the establishment of
for the groups of expert and nonexperts while viewing both kinds neuronal ensembles (Young and Eggermont, 2009). Thus, any
of stimuli, faces and buildings. However, the increase in activity in connectivity identified between brain regions known to be
this brain region was stronger for the architects viewing images of relevant for specific mental processes, or their altered functioning,
buildings. Second, anterior cingulate activity was also significantly is of great interest for the delineation of the mind’s architecture.
associated with appeal ratings, and it was similar for both groups of Determination of the correlation between fMRI time series is
participants while viewing faces. This was not so, however, when grounded on the promise of discovering ‘‘functional connectivity’’
participants viewed images of buildings. Here activity increased between brain sites. This is a common procedure, with its pros and
with appeal for experts and decreased for nonexperts. Third, cons. However, a general, methodological question can be posed,
activity in the nucleus accumbens correlated positively with namely, whether the temporal resolution of fMRI records is
appeal for both groups of participants and both stimuli kinds (Kirk adequate to gauge neural synchronization. Given that fMRI
et al., 2009a). averages neural activity in the order of seconds, this time interval
Overall, Calvo-Merino et al. (2010b) and Kirk et al.’s (2009a) seems too long to deduce the presence of functional connections
study reveal how expertise in visual aesthetic activities, such as responsible for the initial evaluation of visual stimulation. It seems
dance and architecture, can lead to differences in perceptual, that other neuroimaging techniques, such as MEG or EEG, which
representational and affective processes involved in aesthetic have a much higher temporal resolution, might be combined with
experience. Experts seem to rely more than nonexperts on fMRI in the experimental studies in order to obtain a higher
configural processing and their stored knowledge of facts accuracy regarding the temporal span of neural activation.
pertaining to their particular domain of expertise while viewing However, in addition to these and other methodological issues
visual stimuli depicting objects from such domain. They also (Nadal and Pearce, in press), the emerging field of neuroaesthetics
attribute greater reward value and engage self-monitoring also faces challenges of a conceptual nature. Probably the most
strategies to a greater extent that nonexperts. These differences controversial unresolved matter is whether the neurosciences
seem to be in contrast with the common activity observed in brain should be limited to exploring the biological foundations of aesthetic
structures involved in core pleasure sensations, such as the nucleus experience, or whether this approach can also make a meaningful
accumbens. If anything, these studies have clearly shown that contribution to our understanding of artistic activities. Four
expertise has a complex and multifaceted impact on the neural arguments have been put forward to support the contention that
correlates of aesthetic appreciation, and that this line of research neuroaesthetics cannot account for artistic behavior: (i) its strict
constitutes fertile ground for future inquiry. focus on beauty, preference, liking, and so on; (ii) its search for
general principles and neglect of particular aspects; (iii) its disregard
4. Summary and future prospects for neuroaesthetics for contextual features; (iv) its attempts to reduce art production
and appreciation to general neurobiological mechanisms.
Each of the studies mentioned above focuses on particular Regarding the first of these four arguments, Brown and
aspects of the neural correlates of aesthetic appreciation. However, Dissanayake (2009) have explicitly stated that neuroaesthetics

Table 2
The relation between Chatterjee’s (2004b) model and the neuroimaging results on aesthetic appreciation. It shows how Chatterjee’s (2004b) neurocognitive model of
aesthetic appreciation provides an integrative framework to interpret the overall pattern of results obtained by neuroimaging results.
Attentional processes Fronto-parietal networks Fairhall and Ishai (2008)
Attributional networks Ventral visual stream Fairhall and Ishai (2008)
Vartanian and Goel (2004)
Experience of the aesthetic quality Feed-back/feed-forward links Cela-Conde et al. (2004)
Jacobsen et al. (2006)
Cupchik et al. (2009)
Emotional reward enhancement of aesthetic experience Anterior medial temporal lobe Kawabata and Zeki (2004)
Medial and orbitofrontal cortices Cela-Conde et al. (2004)
Subcortical structures Vartanian and Goel (2004)
Jacobsen et al. (2006)
Jacobsen et al. (2006)
Kirk et al. (2009a,b)
Kirk et al. (2009a,b)
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cannot in principle deal with the arts, on account of its emphasis on ful contributions to understanding the human propensity to
aesthetic phenomena: ‘‘Aesthetic emotions are unquestionably an produce and appreciate art? We believe that there is no certain a
integral part of the arts, but they are neither necessary nor priori response to this question. There is no reason why studies
sufficient to characterize them. Thus, a narrow focus on aesthetic cannot in principle address some of the aforementioned issues. In
responses is ultimately a distraction from the larger picture of fact, we believe that such criticisms may constitute the source for
what the arts are about’’ (Brown and Dissanayake, 2009, p. 54). interesting avenues to be explored in future studies. Researchers
Indeed, many artistic manifestations around the world and could, for instance, decide to study the neural correlates of
throughout history have been completely unrelated to beauty. aesthetic appreciation under different contextual conditions,
Besides portraying beauty, art has been –and continues to be– whether different kinds contextual information influences the
created to intimidate, to express sorrow, to extol a society’s neural processes underlying aesthetic appreciation for different
grandeur, to make us reflect on humanity’s and our own existence, kinds of artworks in different ways, and so on. It is up to
among many other functions. In fact, contemporary western art neuroscientists to come up with ingenious designs for studies that
has seen many instances of works that are conceived as a reaction can begin overcoming some of the limitations pointed out above
against beauty as an artistic concept. As noted by Danto (1997), and, as a consequence, producing findings which are relevant to
‘‘the connection between art and aesthetics is a matter of historical other approaches to aesthetic appreciation, such as art theory or
contingency, and not part of the essence of art’’ (Danto, 1997, p. philosophy.
25). Moreover, artworks rarely serve a single purpose. They have
different meanings for different people, in different context, and at Acknowledgements
different times. Brown and Dissanayake (2009) firmly believe that
neuroaesthetics cannot grasp the emotional, formal, cultural, and
Camilo José Cela-Conde and Marcos Nadal were supported by
intentional plurality inherent to the production and perception
the research grant HUM2007-64086/FISO awarded by the Spanish
of art.
Ministerio de Educación y Ciencia.
The second issue refers to the different emphasis placed by the
traditional approach to the arts and neuroaesthetics on the general
and the particular. Neuroaesthetics, in its approach to art, seeks References
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