Spatial Vision, Vol. 21, No. 3–5, pp. 379–396 (2008)

Koninklijke Brill NV, Leiden, 2008.
Also available online - www.brill.nl/sv
Towards a framework for the study of the neural correlates
of aesthetic preference
MARCOS NADAL
1,∗
,ENRICMUNAR
1
, MIQUEL ÀNGEL CAPÓ
2
,
JAUME ROSSELLÓ
1
and CAMILO JOSÉ CELA-CONDE
2
1
Department of Psychology, Universitat de les Illes Balears, Crta Valldermossa s/n, km 7,5,
Palma de Mallorca 07122, Spain
2
Department of Philosophy, Universitat de les Illes Balears, Crta Valldermossa s/n, km 7,5,
Palma de Mallorca 07122, Spain
Received 21 March 2006; accepted 10 March 2007
Abstract—Aiming to provide a tentative framework for the study of the neural correlates of aesthetic
preference, we review three recent neuroimaging studies carried out with the purpose of locating
brain activity associated with decisions about the beauty of visual stimuli (Cela-Conde et al., 2004;
Kawabata and Zeki, 2004; Vartanian and Goel, 2004). We find that the results of the three studies
are not in line with previous neuropsychological data. Moreover, there are no coincidences among
their results. However, when they are mapped on to Chatterjee’s (2003) neuropsychological model of
aesthetic preference it becomes clear that neuroimaging data are not contradictory, but complementary,

possible means to work on a solid framework for future studies in this field. The
results of each of the reviewed neuroimaging studies will be discussed in reference
to previous literature, and they will be brought together with the aid of a recent
cognitive model of aesthetic preference.
Cognitive processes involved in aesthetic preference
Several views on aesthetic preference have been developed during the last century,
including Psychoanalysis, Gestalt theory and Empirical Aesthetics. Within the last
of these, aesthetic preference has been related with arousal (Berlyne, 1970, 1971),
prototypicality (Martindale, 1988; Martindale et al., 1988), and appraisals (Silvia,
2005), among other factors. Potentially, any of these perspectives could serve to
ground our interpretation of neuroimaging results. However, given that the present
work aims to provide a tentative relation between neural activity and cognitive
processes, we need a model that specifies the cognitive operations underlying
aesthetic preference, as well as their interactions, and that, at the same time, can
bridge the cognitive-neural levels.
Leder et al. (2004) have recently proposed a comprehensive model of visual aes-
thetic preference and perception, which includes five processing stages: perceptual
analysis of the visual stimulus, implicit memory integration, explicit classification,
cognitive mastering, and the emergence of a cognitive state, resulting from the pre-
vious stages, and an affective state, that results from the continuous interactions
between previous stages and affective systems in the brain. The cognitive state is
the source of aesthetic preference, while aesthetic emotion is grounded on the af-
fective state.
The different stages suggested by Leder and colleagues (2004) include several
operations, involving different variables known to affect aesthetic preference. How-
ever, their model is formulated at a general psychological level, which makes it
difficult to establish straightforward hypotheses about specific brain activity asso-
ciated with those operations. In contrast, Chatterjee’s (2003) framework for the
Neural correlates of aesthetic preference 381
neural correlates of aesthetic preference is directly grounded on visual neuroscience,

NEUROIMAGING STUDIES OF AESTHETIC PREFERENCE
There are currently four published neuroimaging studies concerned with the neu-
roanatomical correlates of aesthetic preference for visual stimuli. Three of them
have very similar objectives. Cela-Conde et al. (2004) aimed to locate “brain ar-
eas activated during the visual perception of aesthetic objects” (Cela-Conde et al.,
2004, p. 6321). Kawabata and Zeki (2004) wanted to verify whether “there are brain
areas that are consistently active across subjects when they perceive a painting as
being beautiful and, conversely, whether there are brain areas that are specifically
active when they view paintings that they consider to be ugly” (Kawabata and Zeki,
2004, p. 1699). Vartanian and Goel (2004) carried out their study to “determine
382 M. Nadal et al.
the neuroanatomical correlates of aesthetic preference for paintings” (Vartanian and
Goel, 2004, p. 893). Another similarity is that the three studies were designed to
contrast participants’ brain activity associated with positively and negatively rated
stimuli. Thus, while their brain activity was being recorded, participants created dif-
ferent stimuli conditions varying in beauty or preference as a result of their aesthetic
preference ratings.
A fourth study (Jacobsen et al., 2006) may appear to address the same question
as the aforementioned three. However, it differs in at least two very important
issues. First, whereas the other three studies were concerned with the differences in
brain activation when judging stimuli as beautiful or not beautiful, Jacobsen et al.
(2006) compared the whole process of aesthetic decision making with another kind
of decision, that of symmetry. Thus, the results of the studies by Kawabata and Zeki
(2004), Vartanian and Goel (2004) and Cela-Conde et al. (2004) refer specifically
to the neural correlates of judging stimuli as beautiful compared to those of judging
them as ugly, whereas Jacobsen et al. (2006) designed their experiment to identify
the neural correlates of judging the beauty of images compared to judging their
symmetry. Thus, results obtained by Jacobsen et al. (2006) refer to the neural
correlates of the judgment process itself. This difference is far from trivial; in fact
it has important consequences for the comparison of the results of this study with

many of the disadvantages of using artistic and decorative materials, and that the
use of simple visual patterns might engage different cognitive operations to those
that enable aesthetic appreciation in natural conditions. Furthermore, given that
symmetry is a very salient feature of the materials used by Jacobsen et al. (2006),
their results might be difficult to generalize to other stimuli whose symmetry is less
prominent. For these reasons, the remainder of the present work we will concentrate
on the studies by Kawabata and Zeki (2004), Vartanian and Goel (2004) and Cela-
Conde et al. (2004).
Summary of the neuroimaging results
Regarding the results of the three studies, we shall consider only the contrasts
performed between the conditions of positively and negatively valued stimuli.
Kawabata and Zeki (2004) and Vartanian and Goel (2004) obtained interesting
results when comparing brain activity before different categories of stimuli, such as
abstract vs representational, but such issues will not be commented on here, given
that this review is primarily concerned with the neural basis of general aesthetic
preference. The results of the three studies are illustrated in Fig. 1.
Kawabata and Zeki (2004) registered participants’ brain activity with fMRI
while rating the beauty of stimuli. They found that activity in the orbitofrontal
cortex was greater for stimuli classified as beautiful, while activity in the motor
cortex was greater for stimuli classified as ugly. Cela-Conde et al. (2004)
used magnetoencephalography (MEG) to record brain activity during the aesthetic
preference task. Their results showed that activity in the left dorsolateral prefrontal
cortex increased in late latencies (400–1000 ms) when participants judged stimuli as
beautiful, as compared to the non-beautiful condition. By means of fMRI Vartanian
and Goel (2004) found that the activity in the right caudate nucleus decreased as
preference ratings decreased, while activity in the left anterior cingulate gyrus and
bilateral occipital gyri, increased with preference ratings.
At least two issues merit comment. First, there seems to be a discontinuity
between these three neuroimaging studies and those carried out using lesion and
electroencephalographic methods. For instance, none of the neuroimaging studies

resolution of MEG, to test the suggested sequence of activity. But as it turned out,
the frontomedian activity Jacobsen and Höfel (2003) found associated with negative
ratings did not appear in any of the neuroimaging studies. In fact, frontal activity
detected by Cela-Conde et al. (2004) (dorsolateral), Kawabata and Zeki (2004)
(orbitofrontal) and Vartanian and Goel (2004) (anterior cingulate) was associated
with positive ratings. Additionally, in the study by Cela-Conde et al. (2004)
all brain activity correlating with aesthetic preference during the first second after
stimuli onset was limited to the left prefrontal dorsolateral cortex. Activity in other
areas, some of which were located in the right hemisphere, was also recorded, but
revealed no significant differences between the beautiful and ugly conditions. These
discrepancies may owe to the use of different kinds of stimuli. Jacobsen and Höfel
(2003) based their model on studies using patterns composed by simple geometric
forms, whereas Cela-Conde et al. (2004) used complex artworks and photographs.
The comparison of the results from neuroimaging studies on the assessment of
facial beauty (Aharon et al., 2001; O’Doherty et al., 2003; Senior, 2003) with
those from the three studies attempting to identify the neural correlates of aesthetic
preference reveals only one coinciding brain region. Kawabata and Zeki’s (2004)
study, the only one out of the three to include portraits, revealed significant activity
in the orbitofrontal cortex before stimuli classified as beautiful, just as was observed
with beautiful faces with high reward value.
Neural correlates of aesthetic preference 385
The second unexpected fact is the complete lack of coincidence among the results
of the three studies when comparing their results regarding the difference in brain
activity between the beautiful and ugly conditions (see Fig. 1). Although each of
the reviewed studies leaves room for improvement, we believe that their limitations
need not lead to an invalidation of their findings, at least until they are replicated or
experimentally disproved. On the other hand, none of these studies asserted that the
areas they had identified were the exclusive neural correlates aesthetic preference.
In fact they all acknowledged that these areas influence and are influenced by the
activity in other brain areas. It might be the case that the three studies captured

life, landscapes, abstract)) ×(original, classic, impressionist,
portraits) × filtered, altered) (=120) postimpressionist) +
(beautiful, neutral, 160 photos (=320)
ugly) (=192)
Procedure Pre-classification No No
386 M. Nadal et al.
The most obvious difference among the three studies is the use of a different neu-
roimaging technique (MEG) by Cela-Conde and colleagues (2004) with regards to
the other two studies (event-related fMRI ). The involvement of the detected brain
areas in aesthetic preference is inferred from different parameters. Whereas MEG
detects magnetic fields generated by excitatory and inhibitory postsynaptic poten-
tials in the dendrites of pyramidal neurons (Lounasmaa et al., 1996; Maestú et al.,
2005), fMRI offers an indirect measure of neural activity related with hemodynamic
and metabolic responses underlying neuronal events, probably reflecting input and
intracortical processes (Logothetis et al., 2001). Furthermore, both techniques have
different spatial and temporal resolutions, which require different exposure times
to the stimuli and interstimuli intervals (see Table 1). Verifying that indeed neu-
roimaging technique has an impact on the detected neural correlates of aesthetic
preference could be achieved by using a single protocol for MEG and fMRI, or by
the joint EEG and fMRI recording (Debener et al., 2006).
Another major difference among the three studies is the task that participants
were asked to perform. Kawabata and Zeki (2004) asked their participants to
rate the beauty of the stimuli on a 3-point scale (beautiful, neutral, ugly), whereas
Cela-Conde et al. (2004) used a dichotomous scale (beautiful, not beautiful). In
contrast, Vartanian and Goel’s (2004) participants were asked to rate their degree
of preference for the pictures on a 5-point scale. Leder and colleagues (2005)
suggested that preference ratings are associated with a strong affective or reward
component, and that the task of rating beauty might elicit a stronger cognitive
component. Hence, both tasks might have partially different neural correlates, as
suggested by studies of facial beauty (Aharon et al., 2001).

representational visual stimuli.
Finally, there are profound differences among the three procedures, which come
to reflect some of the divergences we have already pointed out. For instance, in
contrast to the studies by Vartanian and Goel (2004) and Cela-Conde et al. (2004),
Kawabata and Zeki (2004) included a stimuli pre-selection procedure. This might
have inadvertently elicited recognition processes in the task participants performed
while their brain activity was being recorded, as suggested by results from previous
studies (Cela-Conde et al., 2002; Nadal et al., 2006) that revealed an association
between mnemonic processes and aesthetic preference. The other procedural aspect
that is different among the studies is the preparation of stimuli. There are several
variables that have been shown to affect aesthetic preference of visual stimuli,
such as psychophysical variables (luminance, contrast, predominant wavelength),
complexity, novelty, prototypicality, and so on (e.g. Berlyne, 1970; Martindale
and Moore, 1988). Furthermore, it has been shown that some of these variables
have an impact on the neural correlates of visual processing (Daffner et al., 1998,
2000; Müller et al., 2003; Nicki and Gale, 1977; Sasaki et al., 2005). Whereas
Cela-Conde et al. (2004) homogenized the complexity, novelty, color spectrum and
luminance of the stimuli, in addition to their size and resolution, Vartanian and Goel
(2004) and Kawabata and Zeki (2004) did not report whether they carried out any
detailed procedure, beyond normalizing size and resolution, in order to control these
variables.
POSSIBLE RELATIONS BETWEEN BRAIN ACTIVITY AND COGNITIVE
PROCESSES INVOLVED IN AESTHETIC PREFERENCE
We have discussed the possible reasons for the lack of coincidence among the
results of three neuroimaging studies addressing the neural correlates of aesthetic
preference, concentrating on different procedural aspects. Given that aesthetic
preference is a process that involves multiple cognitive operations, which take
place in different brain areas and in different time frames, it is very possible that
the results of the three neuroimaging studies reviewed here, conditioned by their
respective experimental designs, refer to the neural correlates of only some of

Lane and colleagues (1999) found that emotional valence, arousal and attention
independently increased the activity in extrastriate cortex, suggesting that there is
a common influence that converges on early visual processing. They also noted
that the overlap of the activity patterns associated with emotion and attention could
be a reflection of their overlap at a behavioral level, in the sense that attentional
resources would automatically be recruited during emotional states. Maunsell
(2004) suggested that the brain might not even have different neuronal signals
related with attention and reward, broadly defined to include, in addition to primary
reinforcers, other factors that can motivate behavior, such as the preference for
places or stimuli. In this sense, the allocation of attention could be the representation
of the subject’s actual assessment of reward. Thus, occipital activity identified by
Vartanian and Goel (2004) might reflect enhanced processing of preferred stimuli,
related with attentional or affective processes.
Finally, the increase in cingulate activity could be a manifestation at an emotional
level of this attentional engagement of preferred stimuli. Bush and colleagues
Neural correlates of aesthetic preference 389
(2000) suggested that the anterior cingulate cortex is part of an attentional system
that regulates cognitive and emotional processing. They noted that the anterior
cingulate cortex can be functionally divided in two regions: a cognitive division
and an affective division. The latter encompasses the region identified by Vartanian
and Goel (2004), and has been related with the assessment of the relevance of
motivational and emotional information and the regulation of emotional responses
(Bush et al., 2000). The studies carried out by Lane and colleagues (1998) and
Hornak and colleagues (2003) support the notion that the anterior cingulate cortex
is involved in certain aspects related with the conscious awareness of emotions.
Turning to the results obtained by Kawabata and Zeki (2004), let us recall they
found that stimuli rated as beautiful were associated with an increased activity in
the orbitofrontal cortex, as Chatterjee (2003) had suggested, and low activity in
the motor cortex, while the inverse pattern was found for stimuli rated as ugly. It
seems quite clear that this study has essentially captured certain emotional processes

in the representation of the reward value of a stimulus, the motor cortex is involved
in keeping a degree of motivation proportional to the value of the reward (Roesch
and Olson, 2004). Thus, it could be that activity in the motor cortex registered
by Kawabata and Zeki (2004) reflects the motor readiness related with withdrawal
behavior elicited by ugly stimuli. However, at present it is not possible to decide
whether activity in this area is associated with the representation of the reward value
or this motor readiness.
The results of Cela-Conde and colleagues’ (2004) study showed that the activity
in the left dorsolateral prefrontal cortex (DLPFC) was greater when stimuli were
judged as beautiful than when they were judged as not beautiful. The involvement
of this area in aesthetic preference was anticipated by Chatterjee (2003). The
literature suggests that this area is involved in the process of decision-making based
on perceptual and/or affective information. Heekeren et al. (2004) showed that
activity in the left DLPFC increased with the easiness of a perceptual decision-
making task, and it decreased when the difficulty increased. Alternatively, it has
been shown that activity in the left DLPFC increases in association with positive
affect. Specifically, Davidson and Irwin (1999) suggested that the DLPFC might be
involved in the representation of goal states towards which elemental positive and
negative affective states are directed, relating specifically the left dorsolateral cortex
with the planning of approach goal-directed actions (Davidson, 2003). Krawczyk
(2002) provided an integrative view of the role of this area: “The left DLPFC
may play a privileged role in decision making that is better constrained, has fewer
options, and which may have preexisting reward characteristics that make for a more
confined set of rules for deciding” (Krawczyk, 2002, p. 661). Thus, the DLPFC
seems to participate in the conscious deliberation about different options, influenced
by emotional information from OFC and certain limbic areas. This is congruent
with Wallis and Miller’s (2003) hypothesis that information about rewards enters
the prefrontal cortex through the orbitofrontal cortex where it is passed on to the
prefrontal dorsolateral cortex, where it is used to control behavior.
A tentative framework for the study of the neural correlates of aesthetic preference

degree of ugliness of the images in the sample, including really unpleasant stimuli,
would elicit activity in lateral orbitofrontal cortex in an aesthetic preference task,
as observed by O’Doherty and colleagues (2001). The subcortical component of
reward value processing during aesthetic preference was registered by Vartanian
and Goel’s (2004) study, corresponding to activity in the caudate nucleus. Reward-
related activity in the motor cortex (Kawabata and Zeki, 2004) could either represent
the reward magnitude of ugly stimuli or the motor readiness elicited by them. The
subjective emotional experience associated with preferred stimuli was reflected in
Vartanian and Goel’s (2004) results. Activity in the anterior cingulate cortex seems
to be associated with the attentional regulation of emotional processes associated
with preferred stimuli.
Second, the enhancement of early visual processing (occipital cortex) has been
registered by Vartanian and Goel’s (2004) design. In principle, this effect could be
due to attentional mechanisms, emotional valence or arousal (Lane et al., 1999). It
is difficult to tell which of these factors is actually responsible for increased activity
in occipital areas before preferred stimuli, given that they all have similar influences
on these regions and because these factors have not been controlled in any of the
neuroimaging studies. However, as we have pointed out above, it is even possible
that in fact emotional and attentional influences on early visual processing owe to
the same neuronal and cognitive systems. This question can only be addressed
by further studies which manipulate or control emotional arousal and valence and
attention.
Finally, the decision-making process seems to have been highlighted by Cela-
Conde and colleagues’ (2004) design. However, at present it is not possible to
determine whether the activity they identified in the left dorsolateral prefrontal cor-
tex reflects decisions based on perceptual information, as Heekeren and colleagues’
392 M. Nadal et al.
(2004) results would suggest, or on information regarding reward value, as sug-
gested by Herrington and colleagues’ (2005) study. A third possibility is that the
decision associated with the activity in this brain region requires the integration of

of materials (artistic vs decorative; simple vs complex; novelty: first and second
exposures), and so on, must be systematically assessed. Additionally, the processes
included in Leder and colleagues’ (2004) model which are not accounted for by
that of Chatterjee (2003), such as the context in which the experience takes place,
pre-classification processes, the role of expertise and the influence of individual
differences must also be investigated and integrated with the results of current
studies. Another interesting issue that needs to be addressed is whether any of these
results can be extrapolated to judging the beauty of stimuli in the auditory modality.
Neural correlates of aesthetic preference 393
Acknowledgements
This work was supported by grant BSO2003-069094-C03 from the Dirección
General de Investigación. The authors would like to thank two anonymous
reviewers, as well as Oshin Vartanian and Martin Skov, for insightful comments
and helpful advice.
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Plate XII
M. Nadal et al.,Figure1.
M. Nadal et al.,Figure2.