■ Neuroaesthetics is gaining momentum. At this early junc-
gence of experimental neuroaesthetics. I then suggest a few areas
ture, it is worth taking stock of where the field is and what lies
within neuroaesthetics that might be pursued profitably. Finally, I
ahead. Here, I review writings that fall under the rubric of neuro-
raise some challenges for the field. These challenges are not
aesthetics. These writings include discussions of the parallel or-
unique to neuroaesthetics. As neuroaesthetics comes of age, it
ganizational principles of the brain and the intent and practices of
might take advantage of the lessons learned from more mature
artists, the description of informative anecdotes, and the emer-
domains of inquiry within cognitive neuroscience. ■
within the last century, isolate and enhance different visual
What does neuroscience have to offer aesthetics? Neuro-
attributes. For example, Matisse emphasized color and
aesthetics, as a field, is gathering force (Skov & Vartanian,
Calder emphasized motion. Zeki suggests that artists en-
2009). As it grows, the field faces the challenge of being
deavor to uncover important distinctions in the visual
both true to its scientific roots and relevant to aesthetics.
world and discover visual modules that are segregated
The term aesthetics is used broadly to encompass the per-
functionally and anatomically within the brain.
ception, production, and response to art, as well as inter-
Parallelism claims point to the fact that artists are ex-
actions with objects and scenes that evoke an intense
perts of visual representations and part of their magic lies
feeling, often of pleasure. I focus on visual aesthetics,
in their creative expression of this expertise. For example,
although the principles also apply to music, dance, and lit-
Cavanagh (2005) also shows that images in paintings often
erature. The term neuroaesthetics is also used broadly as a
violate the physics of shadows, reflections, colors, and con-
domain that has something to do with properties of the
tours. Rather than follow physical properties of the world,
brain as it engages in aesthetics. I describe the kinds of
these painters reflect perceptual shortcuts used by our
writings that fall under the rubric of neuroaesthetics and
minds. Artists, in experimenting with forms of depiction,
examine what, in my view, is needed for the field to mature
discovered what psychologists and neuroscientists are
as a science, particularly as an experimental science. I then
now identifying as principles of perception. Livingstone
point to some questions worth pursuing in the near future
(2002) and Conway and Livingstone (2007) reveal how ar-
and conclude with challenges for the field.
tists make use of complex interactions between differentcomponents of vision in creating visual effects in theirpaintings. Livingstone suggests that the shimmering qual-
ity of water or the sunʼs glow on the horizon seen in someimpressionist paintings (e.g., the sun and surrounding
clouds in Monetʼs “Impression Sunrise”) is produced by
Writings on aesthetics by prominent neuroscientists high-
isoluminant objects distinguishable only by color. This
light parallels between properties of art and organizational
strategy plays on the distinction between the dorsal (where)
principles of the brain. Zeki (1999a, 1999b) should be
and ventral (what) processing distinction (Ungerleider &
credited for introducing neuroaesthetics into scientific dis-
Mishkin, 1982). The dorsal stream is sensitive to differ-
course. He exemplifies the parallelism approach and ar-
ences in luminance, motion, and spatial location, whereas
gues forcefully that no theory of aesthetics is complete
the ventral stream is sensitive to simple form and color.
without an understanding of its neural underpinnings.
Isoluminant forms are processed by the ventral stream
He suggested that the goals of the nervous system and
but are not fixed with respect to motion or spatial location,
of artists are similar. Both are driven to understand essen-
as the dorsal stream does not process this information.
tial visual attributes of the world. The nervous system de-
Thus, isoluminant forms are experienced as unstable or
composes visual information into such attributes as color,
shimmering. Conversely, because shape can be derived from
luminance, and motion. Similarly, many artists, particularly
luminance differences, she argues that artists can use con-trast to produce shapes, and leave color for expressive, rather
than descriptive, purposes. Livingstone highlights the way
2010 Massachusetts Institute of Technology
Journal of Cognitive Neuroscience 23:1, pp. 53–62
that combinatorial properties of visual attributes contrib-
problems with their language, attention, and ability to make
ute to our visual perception. Artists use these combina-
decisions. Despite these alterations in comportment and
torial properties to produce specific aesthetic effects.
cognition, Miller and Hou (2004) and Miller et al. (1998)
Ramachandran and Hirstein (1999) proposed a set of
discovered that some people with FTD develop a propen-
perceptual principles that might underlie aesthetic experi-
sity to produce art. They note that the art tends to be rea-
ences. They emphasize the “peak shift” phenomenon as of-
listic rather than abstract or symbolic. The art is most often
fering insight into the aesthetics of abstract art by relying on
visual and is highly detailed. The artists with FTD them-
Tinbergenʼs (1954) work on this phenomenon. Tinbergen
selves are intensely preoccupied with their art, suggesting
observed that sea gull chicks beg for food from their
that obsessive–compulsive traits acquired through their
mothers by pecking on a red spot near the tip of the
disease contributes to this artistic proclivity.
motherʼs beak. However, the chicks respond even more
Other cases of acquired obsessive–compulsive personal-
vigorously to a disembodied long thin stick with three red
ity traits have resulted in remarkable artistic output. Sacks
stripes near the end. Ramachandran and Hirstein propose
(1995b) described Franco Magnani, an Italian painter in San
that neural structures that evolved to respond to specific vi-
Francisco. Magnani painted hundreds of realistic scenes of
sual stimuli respond more vigorously (a shift in their peak
Pontito, an Italian town where he grew up. After a febrile
response) to underlying primitives of that form even when
illness, which was probably an encephalitis, he began to
the viewer is not aware of the primitive. Their hypothesis
paint obsessively. Pontito was the only subject of his art.
is that artists producing abstracts make explicit use of
Sacks speculated that Magnani had partial complex seizures
these visual primitives in evoking aesthetic responses in
and was, in part, demonstrating the obsessive personality
disorder sometimes associated with temporal lobe epi-
The parallelism approach to neuroaesthetics recognizes
lepsy ( Waxman & Geschwind, 1975). However, instead of
that the production and perception of art ought to con-
being hypergraphic verbally, as is more common among
form to principles of neural organization. Properties of art-
such people, he was hypergraphic visually. In a similar vein,
works and strategies used by artists have parallels in how
Lythgoe, Polak, Kalmus, de Haan, and Khean Chong (2005)
the nervous system apprehends and organizes its visual
reported the case of a builder with a subarachnoid hemor-
world. The question for brain–art parallelism is how to
rhage, who became an obsessive artist after he recovered
translate this starting point into programmatic research
from the initial injury. He began to draw hundreds of
with experiments testing falsifiable hypotheses.
sketches, mostly faces. He then moved to large-scale draw-ings, sometimes covering entire room, and confined his artto a few themes. The authors emphasize his perseverative
tendencies as critical to the emergence of his artistic skills.
A good example of informative anecdotes is observations
We also reported obsessive painting practices in an ar-
of the effects of neurological disease on the production of
tist with Parkinsonʼs disease following treatment with do-
art (Zaidel, 2005). The effect of brain damage on the capa-
pamine agonists (Chatterjee, Hamilton, & Amorapanth,
city to produce visual art stands in sharp contrast to its ef-
fects on many other human capacities. Diseases of the
A subset of autistic children produces striking visual
brain can impair our ability to speak or comprehend lan-
images (Sacks, 1995a). The most detailed description of
guage, to coordinate movements, to recognize objects, to
such a case was Nadia, as reported by Selfe (1977). Despite
apprehend emotions, and to make logical decisions.
severe developmental abnormalities, Nadia had remark-
Although diseases of the brain can certainly impair the abil-
able drawing skills. By the age of 3 she was drawing life-like
ity to produce art, in some instances, paradoxically, the art
horses. She drew intensively for a few moments at a time,
seems to improve. Elsewhere, I proposed that such para-
always copying images. She also focused on specific kinds
doxical improvements can be produced by a changing dis-
of images like horses, of which she drew hundreds of ex-
position to produce art, an enhanced visual vocabulary,
amples. Although Nadiaʼs abilities were striking, she is not
better descriptive accuracy, or enhanced expressivity
unique. Autistic children with striking drawing skills seem
(Chatterjee, 2006, 2009). Here, I outline the changes in
to focus on specific subjects and draw them repeatedly.
disposition and enhanced expressivity produced by these
These artists produce realistic images and tend to be
preoccupied by specific themes. Although the neural basisfor obsessive–compulsive disorders is not completely un-derstood, it is associated with a dysfunction of orbito-frontal
and medial-temporal cortices and fronto-striatal circuits
Neurological disorders that produce obsessive–compulsive
(Kwon et al., 2003; Ursu, Stenger, Shear, Jones, & Carter,
traits can also dispose people to produce art. Such a change
2003; Saxena et al., 1999). Notably, in the cases described,
in disposition to produce art is exemplified in a subset of
these regions could have been damaged and posterior
patients with fronto-temporal dementias (FTDs). FTDs
occipito-temporal cortices were presumably intact. The
can cause profound changes in personality. People with
preservation of posterior cortices ensures that the neural
FTD can be disorganized, socially disinhibited, and have
substrates for recognizing and representing faces, places,
and objects are preserved and are thus available to be the
changes in specific attributes of the art of any person with
subject of these artistsʼ obsessions.
brain damage. Charting such changes systematically will al-low us to identify the specificity of patterns of change thathappens to production of art following brain damage.
Among the most intriguing effects of brain damage on ar-tists is the class of phenomena in which the resulting art is
surprisingly appealing. Right hemisphere damage can pro-duce left spatial neglect in which patients are unaware
of the left side of space (Chatterjee, 2003). Artists with ne-
An experimental research program in visual neuro-
glect omit the left side of images that they draw or paint
aesthetics rests on two principles (Chatterjee, 2002, 2004a).
(Blanke, Ortigue, & Landis, 2003; Cantagallo & Sala,
First, visual aesthetics, like vision in general, has multiple
1998; Halligan & Marshall, 1997; Schnider, Regard, Benson,
components. Second, aesthetic experiences emerge from
& Landis, 1993; Marsh & Philwin, 1987; Jung, 1974). As they
a combination of responses to these different compo-
recover from their neglect, their use of line may still be
nents. The process by which humans visually recognize
impaired. Two examples show how changed spatial repre-
objects offers a framework from which to consider these
sentations can produce highly regarded art. Lovis Corinth,
components. Investigations can be focused on these com-
an important German artist, suffered a right hemisphere
ponents and on their properties in various combinations.
stroke in 1911. As he recovered, he resumed painting.
The nervous system processes visual information both
His self portraits and portraits of his wife showed clear
in hierarchical sequence and in parallel (Farah, 2000; Zeki,
changes in style, with details on the left sometimes left
1993; Van Essen, Feleman, DeYoe, Ollavaria, & Knierman,
out and textures on the left blended with the background.
1990). The sequential components of visual processing
Alfred Kuhn characterized his later work as shifting him
can be classified as early, intermediate, and late vision
into the circle of great artists (quoted in Gardner, 1975).
(Marr, 1982). Early vision extracts simple elements from
Heller (1994) reported the experience of Loring Hughes,
the visual environment, such as color, luminance, shape,
who after a right hemisphere stroke abandoned her pre-
motion, and location (Livingstone & Hubel, 1987, 1988).
morbid style of depictive accuracy. Instead, she turned
These simple elements are processed in different parts
to her own imagination and emotions. The artistic commu-
of the brain. Intermediate vision segregates some elements
nity responded well to her new images. The critic Eileen
and groups others together to form coherent regions in what
Watkins described her work as now delivering “an emo-
would otherwise be a chaotic and overwhelming sensory
tional wallop,” that was not present previously.
array (Ricci, Vaishnavi, & Chatterjee, 1999; Grossberg,
Changes produced by left hemisphere damage are ex-
Mingolla, & Ros, 1997; Vecera & Behrmann, 1997; Biederman
emplified in the Bulgarian painter, Zlatio Boiyadjiev, and
& Cooper, 1991). Late vision selects which of these coherent
the Californian artist, Katherine Sherwood. Boiyadjievʼs
regions to scrutinize and evokes memories from which ob-
premorbid artistic style was natural and pictorial and he
jects are recognized and meanings attached (Chatterjee,
tended to use earth tones in his paintings. Following the
onset of his aphasia, Boiyadjievʼs paintings have been con-
The hierarchical sequence of visual processing must be
sidered richer, more colorful, and containing more fluid
reflected in visual aesthetics (Chatterjee, 2004a; for related
and energetic lines (Brown, 1977; Zaimov, Kitov, & Kolev,
models that also incorporate broader contextual and cul-
1969). The imagery in his work became more inventive
tural factors, see Jacobsen, 2006; Leder, Belke, Oeberst,
and, at times, even bizarre and fantastical. Katherine
& Augustin, 2004)). Any work of art can be decomposed
Sherwood suffered a left hemisphere hemorrhagic stroke,
into its early, intermediate, and late vision components.
which also left her with an aphasia and right-sided weak-
Aesthetic perception can distinguish between form and
ness ( Waldman, 2000). Premorbidly, her images were de-
content (e.g., Woods, 1991; Russell & George, 1990), a dis-
scribed as “highly cerebral.” After her stroke, she felt that
tinction demonstrated experimentally (Ishai, Fairhall, &
she could not produce such images if she wanted. Her new
Pepperell, 2007). Similarly, scientists observe that form is
style has been described as “raw” and “intuitive,” with large
processed by early and intermediate vision, whereas con-
irregular circular movements. She says her left hand enjoys
tent is processed by later vision. Thus, the early vision fea-
an ease and a grace with the brush that her right hand
tures of an art object might be its color and its spatial
never had, and describes it as “unburdened.”
location. These elements would be grouped together to
These cases are but a few examples of the neuropsycho-
form larger units in intermediate vision. Grouping creates
logical effects of art (for comprehensive reviews, see
“unity in diversity,” a central notion of compositional
Chatterjee, 2004b, 2009; Zaidel, 2005). The next step is
to test the inferences made from these anecdotal observa-
Beyond perception, two other aspects of aesthetics
tions. To do so, we recently developed a tool, the Assessment
are important. The first is the emotional response to an
of Art Attributes (AAA) (Chatterjee, Widick, Sternschein,
aesthetic image; the second is how aesthetic judgments
Smith, & Bromberger, 2010). The AAA can quantify
are made. The anterior medial temporal lobe, medial
and orbito-frontal cortices, and subcortical structures
have not been modified greatly by experience. Some com-
mediate emotions in general, and reward systems in par-
ponents of beauty are undoubtedly shaped further by cul-
ticular (Berridge & Kringelbach, 2008; Breiter, Aharon,
tural factors (Cunningham, Barbee, & Philhower, 2002),
Kahneman, Dale, & Shizgal, 2001; OʼDoherty, Kringelbach,
but the universal components are likely to have distinct
Rolls, Hornack, & Andrews, 2001; Delgado, Nystrom, Fissell,
Noll, & Fiez, 2000; Elliott, Friston, & Dolan, 2000; Schultz,
Several studies report that attractive faces activate neural
Dayans, & Montague, 1997). Aesthetic judgments about
circuitry involved in reward systems, including orbito-
stimuli, as measured by preference ratings, are likely to en-
frontal cortex, the nucleus accumbens, the ventral stria-
gage widely distributed circuits, most importantly, dorso-
tum (Ishai, 2007; Kranz & Ishai, 2006; OʼDoherty et al.,
lateral frontal and medial frontal cortices. The general
2003; Aharon et al., 2001; Kampe, Frith, Dolan, & Frith,
point is that visual neuroaesthetics, like most complex bio-
2001), and the amygdala ( Winston, OʼDoherty, Kilner,
logical systems, is hierarchical and can be decomposed
Perrett, & Dolan, 2007). These regional activations are in-
into stable component subsystems (Simon, 1962). This
terpreted as reflecting emotional valences attached to at-
hierarchical organization is precisely what makes experi-
tractive faces (Senior, 2003). The particular emotional
mental approaches to aesthetics possible.
valences are those involved in the expectation of rewards
I have emphasized a cognitive neuroscience framework
and the satisfaction of appetites. The idea that attractive
for experimental neuroaesthetics. Another overarching
faces are rewarding stimuli, at least for men, is evident be-
framework to think about aesthetics comes from evolu-
haviorally. Heterosexual men discount higher future re-
tionary theorists. They make three kinds of arguments.
wards for smaller immediate rewards with attractive
First, beauty serves as a proxy for health and vigor in mate
female faces ( Wilson & Daly, 2004). Presumably, these pat-
selection. Second, beautiful objects are those that are com-
terns of neural activation reflect ways in which attractive
plex and yet are processed efficiently. And third, art mak-
faces influence mate selection (Ishai, 2007; Kranz & Ishai,
ing and appreciation serves an important ritualistic
function that enhances social cohesion. Space limitations
Perceptual features of faces, such as averageness, sym-
do not allow an adequate consideration of evolutionary
metry, the structure of cheekbones, the relative size of the
perspectives on beauty and art (see Brown & Dissanayake,
lower half of the face, and the width of the jaw, influence
2009; Cela-Conde et al., 2009; Dissanayake, 2008; Zaidel,
peopleʼs judgments of facial beauty (Penton-Voak et al.,
2005; Grammer, Fink, Moller, & Thornhill, 2003; Penton-
2001; Enquist & Arak, 1994; Grammer & Thornhill, 1994).
Voak et al., 2001; Etcoff, 1999; Rentschler, Jüttner, Unzicker,
Winston et al. (2007) found that left posterior occipito-
& Landis, 1999; Thornhill & Gangestad, 1999; Zahavi &
temporal activity was enhanced by facial attractiveness.
Zahavi, 1997; Symons, 1979 for relevant discussions). Ulti-
We conducted a study to examine the extent to which
mately, evolutionary and cognitive neuroscience approaches
facial attractiveness is apprehended automatically. Partici-
to aesthetics are likely to converge in informative ways.
pants judged the attractiveness or matched the identity ofpairs of faces. Attractiveness judgments evoked neural ac-tivity within a distributed network involving ventral visual
association cortices and parts of dorsal posterior parietal
Beauty is central to most peopleʼs concept of aesthetics
and prefrontal cortices (Chatterjee, Thomas, Smith, &
( Jacobsen, Buchta, Kohler, & Schroger, 2004). Of course,
Aguirre, 2009). We inferred that the parietal, medial, and
not all art is beautiful and artists do not always intend to
dorsolateral frontal activations represented neural corre-
produce beautiful things. However, beauty remains a cen-
lates of the attention and decision-making components
tral concept in discussions of aesthetic experiences. Un-
of this task. We also found positively correlated activity
derstanding the neural basis of the perception of and
within the insula and negatively correlated activations
response to beauty might give us insight into the percep-
within anterior and posterior cingulate cortex. We inferred
tion of and response to visual art. Facial beauty has re-
that these patterns represent the emotional responses to
ceived most attention in cognitive neuroscience.
attractiveness. Importantly, when subjects matched the
The response to facial beauty is likely to be deeply en-
identity of faces, attractiveness continued to evoke neural
coded in our biology. Cross-cultural judgments of facial
responses in ventral visual areas. This neural response was
beauty are quite consistent (Etcoff, 1999; Perrett, May, &
of a strength that was indistinguishable from the response
Yoshikawa, 1994; Jones & Hill, 1993). Adults and children
when participants considered beauty explicitly. We in-
within and across cultures agree in their judgments of fa-
ferred that this ventral occipital region responds to beauty
cial attractiveness (Langlois et al., 2000), suggesting that
universal principles of facial beauty exist. Infants look longer
Facial attractiveness has pervasive social effects beyond
at attractive faces within a week of being born, and the ef-
its specific role in mate selection (Palermo & Rhodes,
fects of facial attractiveness on infantsʼ gaze generalize
2007; Olson & Marshuetz, 2005). Attractive individuals
across race, sex and age by 6 months (Slater et al., 1998;
are considered intelligent, honest, pleasant, natural lead-
Langlois, Ritter, Roggman, & Vaughn, 1991). Thus, the dis-
ers (Ritts, Patterson, & Tubbs, 1992; Lerner, Lerner, Hess,
position to engage attractive faces is present in brains that
& Schwab, 1991; Kenealy, Frude, & Shaw, 1988), and are
viewed as having socially desirable traits, such as strength
beauty and complexity of the images evoked activity within
and sensitivity (Dion, Berscheid, & Walster, 1972). The cas-
orbito-frontal cortex. In a follow-up study using the same
cade of neural events that bias social decisions is likely to
stimuli (Hofel & Jacobsen, 2007), they found that beauty
be triggered by an early perceptual response to attractive-
generated a lateral positive evoked potential in a temporal
ness. We proposed that neural activity within ventral visual
cortices in automatic response to facial attractiveness
One might be disheartened that these studies inves-
serves as the initial trigger for this cascade (Chatterjee
tigating aesthetics, report inconsistent patterns of activa-
tion. Nadal, Munar, Capo, Rosselo, and Cela-Conde (2008)propose that these seemingly varied results of these studiesare compatible with the general model (Chatterjee, 2004a),
linking aesthetics to the neuroscience of visual and affective
A few studies have used art to examine the neural localiza-
processing as well as reward systems and decision-making.
tion of aesthetic processes. Although the goals in these stud-
Engaging visual properties of paintings increases activ-
ies are similar, their experimental approaches differ and
ity within ventral visual cortices ( Vartanian & Goel, 2004).
the results, at first glance, appear quite varied. Kawabata
Aesthetic judgments activate parts of dorsolateral prefrontal
and Zeki (2004) asked participants to rate abstract, still life,
and medial prefrontal cortices ( Jacobsen et al., 2005; Cela-
landscape, or portrait paintings as beautiful, neutral, or
Conde et al., 2004). In addition, emotional responses to
ugly. Not surprisingly, they found that the pattern of activ-
these stimuli activate orbito-frontal ( Jacobsen et al., 2005;
ity within ventral visual cortex varied depending on
Kawabata & Zeki, 2004) as well as anterior cingulate cortices
whether subjects were looking at portraits, landscapes,
(de Tommaso, Sardaro, & Livrea, 2008; Kawabata & Zeki,
or still lifes. In orbito-frontal (BA 11) cortex, they found
greater activity for beautiful than for ugly or neutral stimuli. In anterior cingulate (BA 32) and left parietal cortex
(BA39), they found greater activity for beautiful than forneutral stimuli. Only activity within orbito-frontal cortex in-
As neuroaesthetics moves forward, several domains could
creased with the beauty of all the painting types and the
be pursued profitably. Here, I suggest three: explorations
authors interpreted this activity as representing the neural
of the relationship of perception to aesthetic experiences,
underpinnings of the aesthetic emotional experience.
the nature of aesthetic judgment, and characterizing the
Vartanian and Goel (2004) used images of representa-
tional and abstract paintings in an fMRI study. They foundthat activity within the occipital gyri bilaterally and the left
anterior cingulate increased with preference ratings. They
also found that activity within the right caudate decreasedas preference ratings decreased. Representational paint-
As described above, visual art can be decomposed into dis-
ings evoked more activity within the occipital poles, the
tinct attributes such as color, line, texture, and form. Prom-
precuneus, and the posterior middle temporal gyrus than
ising questions for empirical research include a better
understanding of how these visual perceptual attributes
Cela-Conde et al. (2004) used magnetoencephalogra-
contribute to the aesthetic experience. Can we measure
phy to record event potentials when participants viewed
the contributions of these attributes? Some properties of
images of artworks and photographs. Participants judged
visual displays can be described with exquisite mathe-
whether or not the images were beautiful. Beautiful
matical precision (Graham & Field, 2007; Redies, 2007).
images evoked greater neural activity than not beautiful
These quantifiable parameters might also be used in neuro-
images over left dorsolateral prefrontal cortex with a la-
tency of 400–1000 msec. The authors infer that this region
How much of the aesthetic experience resides in a per-
is involved in making aesthetic judgments.
ceptual experience and how much resides in the emo-
Jacobsen, Schubotz, Hofel, and von Cramon (2005)
tional response to artwork? Paintings of landscapes are
used a different strategy to investigate the neural corre-
likely to activate the parahippocampus, still lifes lateral oc-
lates of beauty in an fMRI study. Rather than use actual art-
cipital cortex, and portraits the fusiform gyrus. Does
works as their stimuli, they used a set of geometric shapes
beauty modify these activations further? Perhaps these re-
designed in the laboratory. Participants judged whether
sponses simply reflect category-specific activations evoked
the images were beautiful or whether the images were
by perception itself and the aesthetic work is done within
symmetric. Participants found symmetric patterns more
reward systems. However, many feel that we perceive
beautiful than nonsymmetric ones. Aesthetic judgments,
beautiful objects more vividly than nonbeautiful objects.
more than symmetry judgments, activated medial frontal
Some studies show neural responses to beauty within ven-
cortex (BA 9/10), the precuneus, and ventral prefrontal
tral occipito-temporal cortex. Does ventral visual cortex
cortex (BA 44/47). The left intraparietal sulcus was con-
contain general “visual beauty detectors”? Because people
jointly active for symmetry and beauty judgments. Both
are inclined to look longer at beautiful things, are such
ventral visual activations a consequence of attention or is
not clear from the experiment, it demonstrates that the
there an independent aesthetic factor that modulates
same object, when viewed under different conditions,
neural activity? The relationship of attention and aesthetic
can evoke different neural responses.
perception remains to be sorted out.
Fairhall and Ishai (2008), Wiesmann and Ishai (2008),
Characterizing the Aesthetic “Reward”
and Yago and Ishai (2006) have used paintings as stimulito study object recognition and recall. In these studies, they
Beauty is a critically important aspect of how most peo-
find activations in limbic and prefrontal regions, suggesting
ple think of aesthetics ( Jacobsen et al., 2004). However,
that emotional and reward systems are activated automati-
aesthetics is not confined to beauty. Some artwork is spe-
cally even though participants are not making evaluations.
cifically designed to be provocative and disturbing. Ulti-
The apparent automaticity of our response to beauty or to
mately, a comprehensive program in neuroaesthetics
art is an area that invites further investigation.
would incorporate motivations in the creation of and the
One could also investigate the relationship of percep-
response to art that engage emotional systems beyond
tion to aesthetics in brain-damaged people. Some people
with brain damage probably do not perceive art in the
With respect to pleasure evoked by beauty or art, the
same way that non-brain-damaged individuals do, and
imaging studies reviewed here implicate orbito-frontal
their emotional responses to artwork may very well differ
cortex, the anterior and posterior cingulate, the ventral
from those of people without brain damage. Such neuro-
striatum including the nucleus accumbens, the caudate,
psychological investigations of aesthetic perception to date
and the amygdala as mediating the emotional response to
beauty or to artwork. Presumably, these structures differ intheir functions. We need a better sense of how the orches-tration of activity within these structures contributes to an
overall emotional aesthetic (Biederman & Vessel, 2006).
Recent cognitive neuroscience methods probe individual
Evolutionary arguments for the importance of beauty of-
differences. As these methods continue to develop, they
ten emphasize its importance in mate selection. Mate se-
could also be used to examine individual differences
lection is a utilitarian goal and the argument is that the
in aesthetic sensitivities. Aesthetic sensitivity has been re-
features that signal a desirable mate are the features we re-
ferred to as a “T-factor”, for taste (Eysenck & Hawker,
gard as beautiful. This utilitarian goal is at odds with an idea
1994; Eysenck, 1941). People can also develop taste with
proposed in the 18th century (Kant, 1790/1987) that the
training. Behavioral studies show differences in the way
aesthetic attitude is one of “disinterested interest.” On this
that art-experienced individuals and art-naïve individuals
view, aesthetic objects give pleasure without evoking addi-
engage with works of art (Locher, Stappers, & Overbeeke,
tional desires. Stated differently, what distinguishes the
1999; Hekkert & Van Wieringen, 1996). Understanding the
neural response to an aesthetic experience from other re-
neural basis for taste and the ways aesthetic judgment
warding experiences? Could neuroscience contribute to
might be modified with training would be of great interest.
an understanding of disinterested interest?
The studies conducted thus far suggest that parts of
Berridge and Kringelbach (2008) and Wyvell and Berridge
dorsolateral and medial prefrontal cortex are involved in
(2000) draw a distinction between “liking” and “wanting”.
making aesthetic judgments. These studies do not sort out
Liking seems to be mediated by the nucleus accumbens
whether these brain activations are specific to aesthetic
shell and the ventral pallidum mediated by opioid and
judgments or are part of neural systems that make judg-
GABAerigic neurotransmitter systems. By contrast, the
ments regardless of the domain under consideration. We
mesolimbic dopaminergic system, which includes the nu-
do not know if aesthetic judgments engage neural circuits
cleus accumbens core, might mediate wanting. Cortical
that are not engaged in other judgments.
structures, such as the cingulate and orbito-frontal cortex,
Another issue around aesthetic judgments is the institu-
may contribute further to conscious modulations of these
tional context in which art is usually viewed. For example,
liking and wanting experiences. This liking/wanting dis-
Leder et al. (2004) argued that the same object is ap-
tinction is made in a rodent model with experiments
prehended and evaluated differently when viewed “as art-
using sweet and bitter tasting stimuli. Whether the liking/
work.” Recently, Cupchik, Vartanian, Crawley, and Mikulis
wanting distinction generalizes to humans or to visual stim-
(2009) showed differences in brain activations when par-
uli remains to be seen. However, one might test the hy-
ticipants looked at art paintings in an “objective and
pothesis that a self-contained reward system exists and
detached” manner than in a “subjective and engaged”
forms the basis for aesthetic disinterested interest.
manner with an emphasis on experiencing the moodevoked by the paintings. They found greater activity in left
lateral prefrontal cortex in the latter condition, which theyregard as aesthetic, than when participants looked at paint-
These are early days in the neuroscience of visual aesthet-
ings in a detached manner. Although the cognitive mech-
ics. With a field so young, development in any direction
anism underlying this difference in activation patterns is
would be an advance. However, I suggest that practitioners
of neuroaesthetics might keep the following challenges in
world (Fairhall & Ishai, 2008; Wiesmann & Ishai, 2008;
mind: risks of reduction, distinguishing investigations prob-
Yago & Ishai, 2006). This line of work can be distinguished
ing the brain from those probing aesthetics, and adding
from those studies that use neuroscience to test hypoth-
value to our understanding of aesthetics using neuroscience.
eses about the nature of aesthetics.
Fechner (1860), a century and a half ago, made the dis-
tinction between an outer psychophysics and an inner
psychophysics. Outer psychophysics is the study of the re-lationship between psychology and the physical proper-
Experimental neuroaesthetics needs to conform to the con-
ties of stimuli. This kind of study has been the thrust of
straints of any experimental science. That is, experiments
empirical aesthetics ever since. Inner psychophysics is
need to be motivated by general frameworks and to test fal-
the study of the relationship between psychology and
sifiable hypotheses. Such experimental work would analyze
the physical (or physiological) properties of the brain.
specific components of the broader universe of
Fechner recognized that an inner psychophysics might
aesthetics to simplify the domain needs in a way that allows
be possible eventually. Neuroscience technologies such
experimental control. Cognitive neuroscience studies of lan-
as fMRI, ERPs, and transcranial magnetic stimulation now
guage, emotion, and decision-making are models of this ap-
provide the means of pursuing an inner psychophysics.
proach. Further, although qualitative analyses can certainly
The nature of the triangular relationships among psy-
provide important empirical information, quantification
chology, outer physics, and inner physics could be made
more easily provides ways to test hypotheses rigorously.
explicit. Conducting research that probes the relationship
The risk of decomposition and quantification is that re-
between outer and inner physics without direct recourse
duction attenuates the very thing we are most interested in
to psychology is possible. Here properties of objects, pos-
studying. Take the example of the aesthetic responses to
sibly aesthetic objects, would be/are used to probe the
beauty. Experimental aesthetics often addresses this issue
properties of the brain. In such experiments, finely charac-
by obtaining preference ratings from participants. One
terized stimuli are related to the spatial and temporal re-
might ask methodological questions about whether
sponse properties of neurons. Thus, one might find that
forced-choice approaches or Likert-scale ratings are a
the lateral occipital complex responds parametrically to
more stable measure of peopleʼs preferences. One might
some physical properties of objects, important informa-
ask whether judgments of interestingness are the same as
tion in its own right. The unanswered psychophysical
judgments of preference. Or one might explore the rela-
question would be, do lateral occipital complex neurons
tionship of complexity to either preference or to interest.
simply serve a classification function, distinguishing be-
These are legitimate and important questions to be pur-
tween objects and other visual stimuli like faces and
sued. However, the pursuit of such questions might easily
places, or do they also serve an evaluative function, being
obscure the basic question of how preference is related to
tuned to whether the configuration of objects are appeal-
aesthetic experience. Is preference a diluted version of
ing as in the rich tradition of still life paintings? To answer
the former? Or are deeply moving aesthetic experiences
this question, researchers would use the brain to probe
qualitatively different than those assessed in the laboratory
the psychology of aesthetics rather than using aesthetic
with preference ratings? What do neuroscientists make of
objects to probe properties of the brain.
notions such as “the sublime?” The sublime is an emo-
A danger in experiments designed to examine the rela-
tional experience mentioned frequently in aesthetics
tionship between inner and outer physics is that of making
(Kant, 1790/1987), but one that has, so far, had little trac-
inferences about the underlying psychology without ade-
tion in affective neuroscience. Reducing components of
quate investigation of the relevant behavior. This general
aesthetics to quantifiable measures risks inviting the pro-
problem is recognized in cognitive neuroscience as the re-
verbial problem of looking for the dropped coin under the
verse inference problem (Poldrack, 2006), where one uses
lamp because that is where things are visible, even if the
the location of neural activation to infer the underlying psy-
coin was dropped elsewhere. This problem is true for ex-
chological process. Such an inference is valid as a conclu-
perimental aesthetics in general, not just neuroaesthetics.
sion if this area is engaged in only one psychologicalprocess. Unfortunately, such one-to-one correspondencesbetween neural activation and psychological process are
Distinguishing Investigations Probing the Brain
rare in the brain. Findings of localized activations to specific
stimuli more often generate hypotheses about the mental
Art can be used to probe properties of the brain. Because
processes involved, rather than confirm these hypotheses.
brain systems devoted to aesthetics are complex and orga-nized hierarchically, processing art potentially provides a
unique window into the interactions of various subsys-
tems. For example, abstract paintings can be used as aprobe to investigate how the brain deals with indetermi-
This issue, in my view, is the most important challenge for
nate visual stimuli and tries to make sense of its visual
neuroaesthetics. If the goal is to understand aesthetics (as
opposed to understanding the brain), what does neuro-
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