Provided for non-commercial research and educational use...

Provided for non-commercial research and educational use only. Not for reproduction, distribution or commercial use.

This chapter was originally published in the book Progress in Brain Research, Vol. 204 published by Elsevier, and the attached copy is provided by Elsevier for the author's benefit and for the benefit of the author's institution, for non-commercial research and educational use including without limitation use in instruction at your institution, sending it to specific colleagues who know you, and providing a copy to your institution’s administrator.

All other uses, reproduction and distribution, including without limitation commercial reprints, selling or licensing copies or access, or posting on open internet sites, your personal or institution’s website or repository, are prohibited. For exceptions, permission may be sought for such use through Elsevier's permissions site at:

http://www.elsevier.com/locate/permissionusematerial

From: Marcos Nadal, The experience of art: Insights from neuroimaging. In Stanley Finger, Dahlia W. Zaidel, François Boller and

Julien Bogousslavsky, editors: Progress in Brain Research, Vol. 204, Amsterdam: The Netherlands, 2013, pp. 135-158.

ISBN: 978-0-444-63287-6 © Copyright 2013 Elsevier B.V.

Elsevier

Author's personal copy

CHAPTER

The experience of art:Insights from neuroimaging
7
Marcos Nadal1

Department of Basic Psychological Research and Research Methods,

University of Vienna, Wien, Austria1Corresponding author. Tel.:þ43-1-4277-47141; Fax: þ43-1-4277-47191,

e-mail address: [email protected]

AbstractThe experience of art is a complex one. It emerges from the interaction of multiple cognitive

and affective processes. Neuropsychological and neuroimaging studies are revealing the

broadly distributed network of brain regions upon which it relies. This network can be divided

into three functional components: (i) prefrontal, parietal, and temporal cortical regions support

evaluative judgment, attentional processing, and memory retrieval; (ii) the reward circuit, in-

cluding cortical, subcortical regions, and some of its regulators, is involved in the generation of

pleasurable feelings and emotions, and the valuation and anticipation of reward; and (iii) at-

tentional modulation of activity in low-, mid-, and high-level cortical sensory regions enhances

the perceptual processing of certain features, relations, locations, or objects. Understanding

how these regions act in concert to produce unique and moving art experiences and determin-

ing the impact of personal and cultural meaning and context on this network the biological

foundation of the experience of art–remain future challenges.

Keywordsbrain, art, aesthetics, neuroaesthetics, neuroimaging

1 THE EXPERIENCE OF ARTArt can move and affect us in powerful ways. It was art’s enthralling effects that

caused Saint Augustine to feel apprehensive about his experience of music. In his

Confessions, written between 397 and 400, he welcomed the virtuous power of sing-

ing to intensify religious faith and devotion. He felt compelled to confess, however,

that music also had a pernicious effect on him. Music’s charm sometimes completely

diverted his attention from the holy words:

Progress in Brain Research, Volume 204, ISSN 0079-6123, http://dx.doi.org/10.1016/B978-0-444-63287-6.00007-5

© 2013 Elsevier B.V. All rights reserved.135

http://dx.doi.org/10.1016/B978-0-444-63287-6.00007-5

136 CHAPTER 7 Insights from neuroimaging


Thus I fluctuate between peril of pleasure and approved wholesomeness; inclined

the rather (though not as pronouncing an irrevocable opinion) to approve of the

usage of singing in the church; that so by the delight of the ears the weaker minds

may rise to the feeling of devotion. Yet when it befalls me to be more moved with

the voice than the words sung, I confess to have sinned penally, and then had

rather not hear music (Augustine, 1909, p. 195).

The power to absorb people, and even to distract them for other matters, even im-

portant and sacred ones, was also recognized in other forms of art. Writing half a

millennia after Saint Augustine, Saint Bernard of Clairvaux complained that the

awe parishioners felt toward churches’ architecture, sculpture, and painting rivaled

with the awe they felt toward God.1 A late twelfth century chronicler described the

similar captivating influence that the paintings and stained glass windows in the

Bishop’s private Chapel at the Cathedral of Le Mans had on visitors:

The pictures painted in the chapel with unusual talent which have a more powerful

effect because they contain the shapes of living beings, and which absorb not only

the eyes, but also the mind of those who look at them, drew their gaze to such an

extent that they delighted in them, forgetting their personal affairs (transcribed by

Mabillon, 1682, pp. 367–368. Translated by Tatarkiewicz, 1970, pp. 172–173).

Although succinct, this last account identifies the fundamental elements of the ex-

perience of art. We engage with art perceptually, cognitively, and affectively. These

are recognized in today’s psychological and neuroscientific literature as the crucial

components of the art experience. They are the cornerstones of Leder et al.’s (2004)

and Chatterjee’s (2004b) well-known models. Leder et al. (2004) proposed a five-

stage psychological account of the cognitive processes involved in the experience

of art (Fig. 1). The first includes processes of perceptual analysis related to complex-

ity, symmetry, grouping, and organization. The second stage is concerned with the

analysis of the artwork’s familiarity, prototypicality, and meaning. It brings into play

the implicit and automatic integration of information with preexisting memory struc-

tures. Cognitive operations related to the recognition of style and content lead to ex-

plicit classifications in the third phase. This is followed by a stage in which specific

art- and self-related interpretations are developed. These stages influence, and are

influenced by, diverse affective processes. The model posits two different outcomes:

a cognitive state, emerging from earlier cognitive stages, and an affective state,

resulting from the continuous interactions among cognitive and affective mecha-

nisms. A crucial aspect of Leder and colleagues’ model (2004) is that the experience

takes place in a particular context whose significance is determined by a social dis-

course and that endows the experienced object with a particular meaning.

1“I say nothing of the immense height of churches, of their excessive length, superfluous breadth, lux-

urious fittings and exaggerated paintings, which, attracting the gaze of those in prayer, stand in the way

of their feelings” (Saint Bernard of Clairvaux, translated in Tatarkiewicz, 1970, p 190).

FIGURE 1

Leder and colleagues’ (2004) model of aesthetic experience of art.

Figure by Helmut Leder, with kind permission.


Figure 1



Chatterjee (2004b) proposed a compatible model, though formulated at a neuro-

functional level. He suggested that artworks’ simpler components, such as color and

form, are divided, extracted, and analyzed by early visual processes. In a subsequent

stage, intermediate visual processes cluster and segregate certain elements, forming

coherent representations. As regions of the artworks are processed in greater detail,

stored information becomes available, and objects are recognized and associated

with their meanings. The visual analysis and recognition elicits emotions associated

with the aesthetic experience. Emotion, perception, and memory provide the grounds

for the evaluation of the work. This model also includes feedback of information, via

attentional processes, from late visual levels and affective systems to early visual

processing stages.

The experience of art, thus, is a complex one. Although psychologists and neu-

roscientists have confirmed that the components highlighted in the description of vis-

itors’ responses to the Bishop’s Chapel at Le Mans constitute the foundations of the

experience of art, they have also shown how deceivingly simple this description is.

Our experience of art is the result of diverse and interdependent processes related to

perception, cognition (memory, attention, decision making), and affect. With the

advancement of knowledge about the human brain and its relation to cognitive

function, scientists have wondered about the neurobiological foundations of the ex-

perience of art. What brain mechanisms are responsible for producing this uniquely

human kind of experience? Can we explain Saint Augustine’s experience of music in

neuroscientific terms? Two main methods have been used to answer these sorts of

questions: analyzing the effects that brain lesions and neurodegenerative diseases

have on the appreciation of art and using noninvasive neuroimaging techniques to

measure the activity of brain regions while people engage with art.

2 BRAIN DAMAGE, NEURODEGENERATIVE DISEASE, AND ARTMost studies of the effects of neurological disorders on art have tended to focus on its

creation, rather than its appreciation (Alajouanine, 1948; Annoni et al., 2004; Bazner

and Hennerici, 2006; Bogousslavsky, 2005; Bonvincini, 1926; Budrys et al., 2007;

Chatterjee, 2004a; Colombo-Thuillard and Assal, 2007; Crutch et al., 2001; Drago

et al., 2009; Miller and Hou, 2004; Seeley et al., 2008; Zaidel, 2005; Zaimov et al.,

1969). To date, a few have examined the impact of neurological disorders on the ap-

preciation of art. Furthermore, given their anecdotal nature, conclusions can only be

tentative at this time.

Sellal et al. (2003) presented a case of an epilepsy patient who underwent left tem-

poral lobe resection, which only spared the hippocampus, the parahippocampal gyrus,

and the amygdala. During the first year after the surgery, the patient became aware that

heno longer enjoyed listening to rockmusicand thathenowpreferredCeltic orCorsican

polyphonic singing. His taste in the literature also shifted. He now preferred Kafkian-

inspired novels and not science fiction, as before. The authors reported that the patient

also began showing increased preference for realistic paintings, enjoying the small

1392 Brain damage, neurodegenerative disease, and art


details that previously went unnoticed to him. These changes in preference for art con-

trasted with his apparently unaffected preferences for food, fashion, and faces.

Griffiths et al. (2004) described the surprising case of a stroke patient who was

unable to experience emotion in response to music. The lesion affected mainly the

left insula but extended into the left frontal cortex and amygdala. He recovered

speech, which was also initially affected, after a year. However, he was still emotion-

ally unmoved by music a year and a half after the stroke, even though his perception

of diverse musical features was normal, and that he was able to take pleasure in other

activities. These observations led the authors to suggest that the perceptual and

emotional components of music processing rely on functionally and anatomically

distinct neural networks and that the insula is a crucial piece in the neural underpin-

nings of the emotional response to music.

Two studies of the effects of damage to the amygdala suggest that it plays a role

in the appreciation of art and design. Adolphs and Tranel (1999) reported substantial

differences in preferences for visual stimuli between two patients with bilateral

amygdala damage and a group of healthy controls. Both patients expressed

higher liking for three-dimensional geometrical shapes, landscapes, and color

arrangements than healthy control participants. This difference was especially strong

for the stimuli that controls liked least. Similar results were obtained from the

examination the musical processing of a patient with almost exclusive bilateral

damage to the amygdala (Gosselin et al., 2007). She was selectively incapable of

recognizing scary and sad music, but not happy music, despite being able to process

musical features correctly, even tempo and mode. Thus, it seems that the amygdala is

part of the mechanisms underlying aesthetic preference, especially in the experience

of disliking and in relation to negatively valenced stimuli.

Two strategies have been devised to overcome the limitations inherent to single-

patient accounts of the impact of neurological disorders on art appreciation: the anal-

ysis of preference consistency and the standardization of measurement instruments.

Halpern et al. (2008) and Halpern and O’Connor (2013) focused on the effects that

neurodegenerative disorders have on the consistency of preferences for art. Their re-

sults showed that artistically untrained patients suffering from Alzheimer’s disease

(Halpern et al., 2008) and frontotemporal dementia (Halpern and O’Connor, 2013)

were as consistent as healthy controls in their preference for visual artworks

presented at different times, even though Alzheimer’s patients were unable to

remember they had seen the stimuli before. Thus, despite Alzheimer’s disease

and frontotemporal dementia’s devastating effects on general cognitive function,

these disorders do not seem to prevent patients from experiencing art in a personally

meaningful and consistent way.

Does this mean that neurodegenerative diseases do not alter the experience of art?

Halpern’s studies (Halpern and O’Connor, 2013; Halpern et al., 2008) showed that

consistency of art experience is preserved in the face of Alzheimer’s disease and

frontotemporal dementia. They did not report, however, differences in preference

between patients and controls. Two reports of patients with frontotemporal dementia

suggest that this neurological disorder could have profound effects on the experience



of art. Geroldi et al. (2000) and Boeve and Geda (2001) describe how three patients

experienced marked changes in preference for music after the onset of frontotem-

poral dementia. The patients began compulsively listening to music they had not

particularly enjoyed previously, playing it for hours on end. Thus, although fronto-

temporal dementia patients seem to preserve the capacity for enjoying art, and their

preferences are consistent over time, this disorder can alter the kind of art these

patients enjoy.

The second strategy, developed by Chatterjee et al. (2010), involved the creation

of an instrument—the Assessment of Art Attributes—that allows testing the effects

of different neurological conditions on the perception of different aspects of visual

art. Specifically, this scale allows the measurement of six perceptual features, such as

color, balance, depth, and complexity, and six conceptual features, including abstrac-

tion, symbolism, or emotional expressiveness. In conjunction with voxel-lesion-

symptom-mapping, Bromberger et al. (2011) used the Assessment of Art Attributes

to show how specific brain lesions impair the appreciation of some of these art at-

tributes but not others. Their results indicated that patients with damage to different

regions within the right frontal, parietal, and lateral temporal cortices deviated sig-

nificantly from healthy participants when rating four of the six conceptual scales:

abstractness, symbolism, realism, and animacy. Brain damage, specifically to the

insula and the right temporal and frontal lobes, seemed to influence the appreciation

of only one of the formal attributes: depth.

Overall, the study of the impact of neurological disorders on the experience of art

argues against the existence of specialized brain mechanisms underlying the expe-

rience of art (Zaidel, 2005). The studies reviewed above suggest that any of the three

cornerstones of the experience can be affected, impairing the perception of an art-

work’s particular sensory features, its recognition or recollection, or the capacity

to emotionally engage with it. Most patients, however, are still able to recognize

and experience art in a meaningful and consistent way, even in the face of extensive

brain damage or disabling neurodegenerative diseases.

3 NEUROIMAGING STUDIES OF THE APPRECIATION OF ARTResearchers have used neuroimaging techniques to study healthy participants’ expe-

rience of art in controlled situations. This has allowed them to draw general conclu-

sions about the neural processes underlying this kind of experience. The emerging

picture suggests that at least three functionally distinct sets of brain regions underlie

the experience of art, roughly corresponding to its three main components: (i) pre-

frontal, parietal, and temporal cortical regions involved in evaluative judgment, at-

tentional processing, and memory retrieval; (ii) the reward circuit, including cortical

and subcortical regions, as well as some of the regulators of this circuit; and (iii) low-,

mid-, and high-level cortical sensory regions (Nadal and Pearce, 2011). We will

examine these three sets in turn.

FIGURE 2

Brain regions related to evaluative judgment, attention, and memory involved in the

experience of art.

1413 Neuroimaging studies of the appreciation of art


3.1 Activity of cortical areas involved in evaluative judgment,attention, and memory

Cela-Conde and colleagues (2004), Cupchik and colleagues (2009), Jacobsen and

Hofel (2003), Jacobsen et al. (2006), and Lengger et al. (2007) found evidence of

neural activity associated with evaluative judgment, decision making, and memory

while their participants were engaged with artworks (Fig. 2).

3.1.1 Dorsolateral prefrontal cortexCela-Conde and colleagues (2004) believe that increased left prefrontal activity ob-

served while their participants viewed photographs and paintings they regarded as

beautiful (Fig. 3) reflected the process of making a decision about their beauty.

Cupchik et al. (2009) also found greater activity in the left lateral prefrontal cortex

when participants were asked to approach the stimuli with an engaged aesthetic atti-

tude than when they were required to take an objective and detached approach. They

interpreted this as reflecting top-down control of perception. Lengger et al. (2007)

found that neural activity in the left frontal cortexwas lowerwhen participants viewed

modern artworks for which they had been given stylistic information than when they

had not. The authors believe that without the information, participants struggled to

categorize the stimuli and searched for related concepts in memory.

3.1.2 Anterior medial prefrontal cortexJacobsen and Hofel (2003) and Jacobsen et al. (2006) found that activity in the an-

terior medial cortex was greater when participants were asked to rate the beauty of

geometric patterns than when they were asked to rate their symmetry. Kirk (2008)

Figure 2

FIGURE 3

Results from Cela-Conde et al.’s (2004) experiment. Left panel shows activity while

participants viewed stimuli they regarded as beautiful. Right panel activity while participants

viewed stimuli they regarded as not beautiful.



found that activity in this region correlated with aesthetic appeal ratings of images

with unusual or unexpected figure-ground combinations. The authors believe that

this activity reflects the engagement of evaluative judgment processes.

Although all of these studies (Cela-Conde et al., 2004; Cupchik et al., 2009;

Jacobsen and Hofel, 2003; Jacobsen et al., 2006; Kirk, 2008) found that prefrontal

activity varied with evaluative judgments of paintings, photographs, and designs,

some located this activity in the dorsolateral prefrontal cortex and others in the an-

terior medial prefrontal cortex. Cela-Conde et al. (2011) provided a functional ex-

planation for this discrepancy. The frontomedial prefrontal cortex has been shown

to be involved in self-referential evaluative judgments (Northoff and Bermpohl,

2004; Zysset et al., 2002). Jacobsen and Hofel’s (2003) and Jacobsen et al.’s

(2006) use of abstract geometric patterns to elicit beauty judgments might have in-

creased the subjective elements of such judgments, where participants could only

base their decision on internally generated information. In contrast, Cela-Conde

et al.’s (2004) and Cupchik et al.’s (2009) use of paintings and photographs might

have encouraged participants to base their decisions about beauty on the richer ex-

ternal information provided by the stimuli, such as their style, explicit content, and

degree of artistry. This interpretation is in accordance with Christoff and Gabrieli’s

(2000) suggestion that while activity in the dorsolateral prefrontal cortex seems to be

primarily involved with information generated externally, activity in the frontome-

dian prefrontal cortex seems to reflect the engagement of processes related to the

evaluation and manipulation of internally generated information.

Owing to the techniques Cela-Conde et al. (2004) and Jacobsen and Hofel (2003)

used, their studies afford a degree of temporal precision the others do not. Both stud-

ies showed that the appreciation of art and design includes an early and brief

Figure 3



evaluation phase, grounded on internally or externally elaborated information, be-

tween 400 and 600 ms after stimuli onset. It seems unlikely that such evaluation ac-

tually represents the final decision elaborated by the participants. It is more plausible

that this early brain activity corresponds to the formation of an initial impression,

which can thereafter influence subsequent processes related to attention, perception,

response selection, and so on, and which could lead to decisions to continue or sus-

pend engagement with the artwork or design (Cela-Conde et al., 2011).

3.1.3 Ventrolateral prefrontal cortexJacobsen et al. (2006) found increased activity bilaterally in the inferior frontal gyrus

when participants were asked to rate the beauty of geometric designs as compared to

symmetry judgments, and Kirk (2008) found that activity in this region correlated

with aesthetic appeal ratings of images with unusual or unexpected figure-ground

combinations. The authors believed that this activity owes to the greater attentional

demands under these experimental conditions, which required transforming a non-

dichotomous judgment into a binary response (Jacobsen et al., 2006) or decoding

and processing complex visual stimuli (Kirk, 2008).

3.1.4 Temporal poleJacobsen et al. (2006) found that activity in the left temporal pole was greater when

participants gave beauty ratings to geometric designs than when they were asked to

rate their symmetry. They believe that this effect reflects the retrieval of information

frommemory to create a semantically and emotionally rich context as a reference for

valuing the visual stimuli. Kirk’s (2008) study revealed that activity in the left tem-

poral pole was greater when participants viewed photographs depicting abnormal

figure-ground pairings they found appealing than when they saw similar images they

found unappealing. In his view, this activity is related to participants’ use of prior

knowledge to organize the affectively salient combinations of figure-ground into

comprehensible scenes they could engage with (Kirk, 2008).

3.1.5 Posterior cingulate cortex and precuneusKirk (2008) found that activity in the posterior cingulate cortex (PCC) correlated

with aesthetic ratings of scenes portraying congruent figure-ground pairings. The au-

thor hypothesized that it reflected the role of semantic memory and familiarity in

participants’ responses to the stimuli. Jacobsen et al. (2006) found greater activity

in the PCC and precuneus when participants rated the beauty of geometric forms than

when they were asked to rate their symmetry. They believe that this activity reflects

processes of memory retrieval, either because the participants were taking into ac-

count patterns previously seen in the experiment or stored in memory from their ev-

eryday experience. Kirk et al. (2009a) found greater activity in the precuneus when

expert architects assessed the aesthetic appeal of buildings, an effect that was not

found for nonexperts. The authors believe that this activity reflects experts’ retrieval

of information stored in memory to create an appropriate context for rating.

FIGURE 4

Brain regions related to reward, affect, and emotion involved in the experience of art.



Taken together, these studies suggest that experiencing art, as well as design, en-

gages a network of cortical regions related to evaluative judgment, based on inter-

nally or externally elaborated information, the allocation of attentional resources,

and the retrieval of information from memory to contextualize the stimuli and judg-

ment. The weight of such processes in the experience of art depends on the kind of

task participants are required to perform, and the stimuli they were presented with.

3.2 The role of the reward circuit in the experience of artMany studies have shown that several regions that constitute the reward circuit have

a prominent role in the experience of art. These include cortical (anterior cingulate,

orbitofrontal, and ventromedial prefrontal) and subcortical (caudate nucleus, nucleus

accumbens) components, as well as some of the regulators of this circuit (amygdala,

thalamus, hippocampus). The regions most consistently reported in the neuroimag-

ing literature include the orbitofrontal cortex (OFC), the ventromedial prefrontal cor-

tex (VMPFC), the anterior cingulate cortex, insula, and nucleus accumbens (Fig. 4).

3.2.1 Orbitofrontal cortexActivation of the OFC has been observed while people rated the beauty of diverse

visual artworks (Ishizu and Zeki, 2011; Kawabata and Zeki, 2004), the aesthetic ap-

peal of photographs (Kirk, 2008; Kirk et al., 2009a,b; Lacey et al., 2011), or when

they listened to moving musical fragments (Blood and Zatorre, 2001; Blood et al.,

1999; Ishizu and Zeki, 2011). Based on numerous studies in other domains, this ac-

tivity is usually interpreted as reflecting the reward value of the presented artworks or

aesthetic stimuli. Activity in the medial OFC (mOFC) seems to be related in a

Figure 4



positive and linear manner to people’s liking or preference (Kawabata and Zeki, 2004;

Kirk, 2008; Kirk et al., 2009a,b). Activity in the lateral OFC (lOFC), however, seems

to be negatively correlated with appeal ratings (Kirk, 2008; Munar et al., 2012). This

functional dissociation is coherent with the lOFC’s role in the evaluation of punishers

and uncertainty, and with the involvement of the mOFC in the monitorization of

reinforcer reward value (Kringelbach and Rolls, 2004; O’Doherty et al., 2001).

During the experience of art, the OFC seems to be especially sensitive to mod-

ulation by environmental factors, such as semantic context, and by personal factors,

such as prior knowledge. For instance, Kirk et al. (2009a) reported that the belief that

visual stimuli were gallery artworks enhanced activity in the mOFC, which corre-

lated with increased aesthetic appeal scores awarded by participants (Fig. 5). Simi-

larly, Lacey et al. (2011) showed that activity in the mOFC was greater while

5

4

5

4

3

2

1

0Gallery label

% S

igna

l cha

nge

Computer label

3

2

1

0

FIGURE 5

Results from Kirk et al.’s (2009b) study. Upper panel: the figure shows activation in right

mOFC where the BOLD signal correlates with the first-order linear term for the contrast

(gallery–computer). The activation is overlaid on sagittal, coronal, and axial sections of the

canonical SPM structural image. Lower panel: Parameter estimates for voxels inmOFC for the

two conditions gallery (G) and computer (C), where the x-axis reflects the two stimulus

conditions and the y-axis shows BOLD signal changes. Error bars indicate 90% confidence

interval.

Copyright 2008 by Elsevier. Reprinted with permission.

Figure 5



participants viewed stimuli they regarded as artworks than while they saw stimuli

depicting similar content but which were not regarded as artworks.

3.2.2 Ventromedial prefrontal cortexThis region is functionally and anatomically related to the OFC. Kirk et al. (2009a)

found that labeling abstract images as gallery artworks increased activity in the fron-

topolar aspect of the VMPFC, as well as aesthetic appeal scores. Similarly, Harvey

et al. (2010) showed that when paintings were paired with the logos of companies

that were supposedly paying for participants’ involvement in the experiment, activity

in the VMPFC was greater than when the paintings were paired with other logos.

These increases were also accompanied by higher liking ratings for the paintings

when they were presented together with the sponsoring logo.

3.2.3 Anterior cingulate cortexAnother common finding is that anterior cingulate activity is higher when people

engage with artworks they like than when they do so with artworks they like less.

This effect has been observed with music, paintings, and architecture (Blood

et al., 1999; Brown et al., 2004; Cupchik et al., 2009; Kirk et al., 2009a;

Vartanian and Goel, 2004). Based on many prior findings, the role of the anterior

cingulate cortex in the experience of art has been related to participants’ monitoring

of their own affective state. People presumably use this information to guide their

evaluations and decisions regarding the object in question.

3.2.4 Insular cortexCupchik et al. (2009) found greater activity in the insula when participants engaged

subjectively with paintings, allowed themselves to experience the evoked mood and

feelings, and attended to their colors and composition, than when they viewed the

paintings in a detached manner and focused on their content. They attributed this

activation to the experience of emotion while participants viewing the paintings with

an aesthetic attitude. Brown et al. (2004) and Koelsch et al. (2006) also reported

greater insular activity while participants enjoyed musical fragments than while they

listened to fragments they did not enjoy. This suggests that the insula plays a funda-

mental role in the emotional response to music, confirming Griffiths and colleagues’

(2004) aforementioned neurological findings.

3.2.5 Nucelus accumbensFinally, many studies have identified activity in the subcortical components of the

reward circuit while people engage with art (Bar and Neta, 2007; Blood and

Zatorre, 2001; Blood et al., 1999; Brown et al., 2004; Cupchik et al., 2009;

Di Dio et al., 2007; Gosselin et al., 2007; Ishizu and Zeki, 2011; Kirk et al.,

2009a; Koelsch et al., 2006; Lacey et al., 2011; Mitterschiffthaler et al., 2007;

Salimpoor et al., 2011; Vartanian and Goel, 2004; Yue et al., 2007). This set of varied

brain regions is crucial for a variety of complex operations related to learning, reward

prediction and anticipation, emotions, and pleasure. The ventral striatum is one of the



key neurobiological elements contributing to the experience of art. Kirk et al.

(2009a) found that activity in the nucleus accumbens was greater while participants

viewed photographs of faces and buildings that were rated as highly appealing than

those rated as unappealing. This effect was independent of degree of expertise with

the architectural stimuli. The authors conjectured that this activity reflects the encod-

ing of the affective salience of the stimuli. Likewise, Lacey et al. (2011) showed that

activity in the ventral striatum was greater while participants viewed stimuli they

regarded as artworks than while they saw stimuli depicting similar content but which

were not regarded as artworks. Given that this brain region is involved in the gen-

eration of pleasurable experiences (Kringelbach and Berridge, 2009), the ventral

striatum might be responsible for the pleasurable aspect of our experiences with art.

Although their role has been examined separately, the nucleus accumbens,

the orbitofrontal, ventromedial prefrontal, anterior cingulate cortices, and insula

together constitute an extended network that processes different aspects of artworks

and contribute with the affective and emotional qualities of the experience of art.

Each of these regions interacts with the rest of the network. Moreover, they are

profusely intertwined with the set of brain regions related to evaluative judgment,

memory, and attention noted above, as well as with a different set of brain regions

involved in the analysis of perceptual features.

3.3 Enhancement of cortical sensory processesNeuroimaging studies have consistently identified increased activity in cortical

regions related to sensory processing while people enjoy art, possibly reflecting

an enhancement of perceptual analyses. Although this sort of brain activity is often

found unexpectedly, and its relevance generally overlooked, it is a common finding

in studies of painting, music, and dance (Fig. 6).

3.3.1 Occipital cortexVartanian and Goel (2004) asked their participants to rate their preference for a series

of abstract and representational artistic paintings. Results of the fMRI scans showed

that activity in occipital cortex, including bilateral fusiform gyri, correlated with

preference ratings for the visual stimuli. The more people preferred the paintings,

the greater the activity in the visual cortex. The authors suggest that this activity

could be related to the positive valence of preferred images or with the increased

visual attention they attracted. Kirk (2008) found that, independently of aesthetic

judgment, bilateral activity in the lateral occipital cortex was greater when partici-

pants rated the aesthetic appeal of images depicting congruent figure-ground pairings

than when rating images depicting unusual or unexpected pairings. These results are

congruent with Lacey et al.’s (2011), who showed that activity in several regions of

the occipital cortex was greater while participants viewed stimuli they regarded as

artworks than while they saw stimuli depicting similar content but which were

not regarded as artworks. Cupchik et al. (2009) aimed to dissociate the contribution

of bottom-up processes (hard-edge vs. soft-edge style) and top-down processes

FIGURE 6

Brain regions related to enhanced perceptual processing involved in the experience of art.



(objective and detached vs. subjective and engaged orientation) in aesthetic valua-

tion of artworks. Their results revealed an activation increase in bilateral occipital

gyri while participants viewed the artworks in relation to the baseline condition.

They believe that this reflected the recruitment of attentional resources for perceptual

analysis of the paintings’ visual details.

3.3.2 Parietal cortexCela-Conde and colleagues’ (2009) study revealed greater activity in bilateral angu-

lar gyrus when participants viewed paintings, photographs, and designs they

regarded as beautiful than when they viewed similar stimuli regarded as not beauti-

ful. The authors attributed this activity to the enhancement of spatial processing strat-

egies while viewing images considered by each participant as beautiful. Cupchik

et al.’s (2009) study mentioned in the preceding paragraph also revealed enhanced

activity in the superior parietal cortex while participants were aesthetically engaged

with soft-edge artworks. Based on this region’s involvement in spatial cognition and

visual imagery, the authors believe that such activity reflects participants’ endeavors

to elaborate coherent representations of the indeterminate forms.

3.3.3 Auditory cortexIn their fMRI study, Koelsch et al. (2006) asked 11 participants to listen to musical

excerpts and to rate their pleasantness or unpleasantness. The stimuli consisted of

instrumental fragments of joyful dance melodies and continually dissonant

Figure 6



counterparts. When compared to unpleasant stimuli, pleasant music was associated

with greater bilateral activity in Heschl’s gyri, location of the human primary audi-

tory cortex, and where fixed pitches are processed (Levitin and Tirovolas, 2009).

Koelsch et al. (2006) suggested that this effect owed to the positive affective valence

of pleasant fragments engaging top-down attentional mechanisms that increased ac-

tivity in the primary auditory cortex and thereby enhancing the perceptual analyses

of these fragments.

3.3.4 Representation of body and movementCalvo-Merino et al. (2008) asked participants with no dance expertise to rate how

much they liked a series of dance movements while their brain activity was scanned

with fMRI. Results revealed that liking scores were strongly correlated with activity

in the occipital cortex and premotor cortex. Both of these regions are involved in the

perception of bodies. In a subsequent study, Calvo-Merino et al. (2010) used transcra-

nial magnetic stimulation (TMS) to examine the contribution of the extrastriate body

area of the occipital cortex, involved in processing local body features, and the ventral

premotor cortex, involved in configural body processing, to the aesthetic valuation of

dance postures. Their results revealed that their lay participants’ aesthetic sensitivity

to dance postures was reduced to a greater extent when TMSwas applied to the extra-

striate body area than when it was to the ventral premotor cortex. The authors con-

clude from these results that early local perceptual processes in the extrastriate

body area make a significant contribution to aesthetic valuation of body stimuli.

3.3.5 Functional accountWhat role does the activity in these sensory regions play in the experience of art?

Biederman and Vessel (2006) argued that endomorphins and m-opioid receptors

are crucial mediators in this relation. Their hypothesis is based on Lewis et al.’s

(1981) observation that the density of m-opioid receptors on macaque cortical neu-

rons increases along visual, somatic, and auditory sensory processing hierarchies,

from primary sensory regions to association cortices. Biederman and Vessel

(2006) believe that such receptors, and their gradient distribution—especially the

great density in the parahippocampal cortex—represent the biological cornerstone

of pleasure derived from the acquisition of novel information: “If a stimulus contains

a great deal of interpretable information, it should lead to more neural activity in the

association areas and hence to a greater release of endomorphins and increased stim-

ulation of m-opioid receptors” (Biederman and Vessel, 2006, p. 251). This perceptual

pleasure is hypothesized to be independent of the reward circuit discussed earlier. In

an fMRI study, participants were asked to rate their preference for 200 different

scenes of landscapes, cityscapes, rooms, and so on (Yue et al., 2007). Confirming

their initial hypothesis, results showed stronger activity in the right parahippocampal

cortex while participants viewed highly preferred scenes than when they saw scenes

they did not prefer. In addition, activity in the right ventral striatum was also stronger

for stimuli rated as highly preferred. The authors suggest that activity in right



parahippocampal cortex, related to processing of perceptual pleasure, might engage

the ventral striatum, allowing a possible role for the conventional reward system.

Although Yue and colleagues’ (2007) study indeed showed increased activity in

the parahippocampal cortical “place area” while people viewed scenes they preferred

a lot, it does not represent a confirmation of the opioid substrate hypothesis, as the

authors themselves acknowledge. In fact, there is an alternative possibility that can-

not be discarded. Because opiates have effects on brain structures that are not directly

involved in reward (LeMerrer et al., 2009), m-opioid receptors along sensory proces-sing pathways are not necessarily related to the generation of pleasurable experi-

ences. They could function as modulators of common sensory and associative

operations (Koepp et al., 2009; Wise and Herkenham, 1982). In fact, Lewis et al.

(1981) did not relate the density gradient with pleasure. They believed that it played

an important role in selective attention, and in fact, Arnsten et al. (1983) confirmed

that opioids broaden the focus of attention in humans.

Is it possible that the enhanced sensory processes identified inVartanian andGoel’s

(2004), Calvo-Merino et al.’s (2008), Cela-Conde et al.’s (2009), Cupchik et al.’s

(2009), and Koelsch et al.’s (2006) studies owe to the effects of attention? Attention

canmodulate brain activity at almost every processing stage, from sensation to decision

making (Chun et al., 2011; Kanwisher and Wojciulik, 2000; Posner and DiGirolamo,

2000). Attention can influence perceptual processing along both the dorsal and ventral

visual pathways, and fundamentally in the fusiform and parahippocampal gyri

(Chelazzi and Corbetta, 2000). Attention modulates neural activity in sensory cortices

when selecting spatial locations, specific features, and even whole visual objects

(Kanwisher and Wojciulik, 2000). These forms of attention seem to operate through

similar principles: by increasing the sensitivity of neurons coding certain spatial loca-

tions (Hopfinger et al., 2000), features (Liu et al., 2007), or object recognition (Yantis

and Serences, 2003), thereby facilitating processing at attended locations, of attended

features, and of attended objects (Reynolds and Chelazzi, 2004).

However, the enhancement of sensory processing while engaged with art requires

an explanation that can cut across multiple sensory modalities. Vartanian and Goel’s

(2004), Cela-Conde et al.’s (2009), and Cupchik et al.’s (2009) participants viewed

paintings, Calvo-Merino et al.’s (2008) and Calvo-Merino et al.’s (2010) viewed dance

movements, and Koelsch et al.’s (2006) listened to musical fragments. Could the en-

hancement of perceptual processes during positive aesthetic experiences be a common

trait of the experience of painting, dance, and music? Ishizu and Zeki’s (2011) results

showing that appreciation of painting andmusic engages themOFC in concert with the

visual and auditory cortices, respectively, would suggest so. Indeed, in humans, atten-

tion modulates neural activity in the auditory cortex while attending to specific tone

sequences (Woldorff et al., 1993), in the somatosensory cortex while expecting tactile

stimulation at certain body locations (Johansen-Berg and Lloyd, 2000), in the gustatory

cortexwhile trying to detect tastes (Veldhuizen et al., 2007), and in the olfactory cortex

when sniffing for odorants (Zelano et al., 2005).

However, an increase in the activity of sensory cortical areas is also commonly

observed in response to emotionally significant pleasant and aversive sounds, voices,

printed words, images of faces, and complex scenes (Lang et al., 1998; Murphy et al.,



2003; Phan et al., 2002; Vuilleumier, 2005; Vuilleumier et al., 2004). Vuilleumier

(2005) believes that these effects are independent of the regular frontoparietal atten-

tional network and that they could result from feedback modulation from the amyg-

dala, which projects to all stages in the ventral visual processing pathway (Phan et al.,

2002; Vuilleumier et al., 2004). Thus, once the amygdala has identified relevant emo-

tional content in a stimulus, it enhances the activity of the cortical regions involved in

the representation of the relevant location, feature, or object (Compton, 2003).

Nevertheless, the fact that signals from the amygdala can enhance sensory proces-

sing in response to emotional stimuli does not preclude the possibility of concurrent

influence from frontoparietal attentional networks. In fact, under Vuilleumier’s

(2005) framework, signals from the amygdala to visual processing regions add

to—or under certain circumstances, even compete with—those imposed by the fron-

toparietal attentional network.When the emotional content of the stimuli is not strong

enough, attentional biasing signals may take precedence over signals from the amyg-

dala. In this scenario, it is possible that activity in cortical sensory regions that accom-

panied the experience of artworks in Vartanian and Goel’s (2004), Calvo-Merino

et al.’s (2008), Cela-Conde et al.’s (2009), Cupchik et al.’s (2009), and Koelsch

et al.’s (2006) studies reflects the confluence of signals from different sources related

to attentional and emotional mechanisms, mediated by a frontoparietal network and

the amygdala, respectively, aimed at biasing activity at different stages of sensory

processing. Both emotion and attention are closely related to prioritizing information

to be processed (Compton, 2003). The consequence of such enhanced activity would

be a deeper processing at earlier stages, resulting in an advantage over other stimuli or

features competing for attention (Murphy et al., 2003; Vuilleumier et al., 2004).

The effective connectivity study performed by Lacey et al. (2011), who asked

participants to view artistic and nonartistic stimuli matched for content, offers ten-

tative conclusions to this section. Their results revealed that activity in the ventral

striatum was driven by activity in the calcarine sulcus and the presupplementary mo-

tor area in the left hemisphere, and by activity in the hypothalamus, the posterior

frontal gyrus, and the lateral occipital complex in the right hemisphere. Conse-

quently, Lacey et al. (2011) suggested that the enhanced sensory processing observed

in the neuroimaging studies of aesthetic preference is not per se a correlate of pos-itive effect, as posited by Biederman and Vessel (2006), but a trigger for activity in

the ventral striatum, which in turn would generate the positive hedonic states asso-

ciated with liked stimuli, as observed in Yue et al.’s (2007) experiment.

As for the emotional or attentional mechanisms responsible for increased activity

in sensory cortices observed in the studies noted earlier (Calvo-Merino et al., 2008;

Cela-Conde et al., 2009; Cupchik et al., 2009; Koelsch et al., 2006; Vartanian and

Goel, 2004), Lacey et al.’s (2011) analysis of connectivity found no evidence that

amygdala drives activity in the visual regions. In fact, Lacey et al.’s (2011) connec-

tivity study shows that activity in the occipital gyri was driven predominantly by the

cingulate cortex and the posterior frontal gyrus, suggesting top-down influence from

attentional systems. In light of these results, it seems that the enhancement of per-

ceptual processes while experiencing art, at least in laboratory settings, owes primar-

ily to the effects of attention, rather than emotion.



4 CONCLUSIONS, LIMITATIONS, AND PROSPECTSOver the past decades, neuropsychological and neuroimaging studies have taught us

quite a lot about the biology of the art experience. We know that it is a complex ex-

perience. We might even conceive of it as a complex of experiences, of perceptual,

cognitive, and emotional experiences. There is no localized seat for art in the brain.

Rather, our experience of art emerges from the interaction among the nodes of a

broadly distributed network of cortical and subcortical brain regions. None of these

are specialized in responding to art alone, not even in the sense that one could think of

Broca and Wernicke’s regions as specialized for language processing. They all play

crucial roles in other domains of human experience, from perceiving small details in

the world or making small decisions to abstract reasoning or establishing social

relationships.

The distributed and unspecific quality of the neural underpinnings of the art ex-

perience might be the reason why it is resilient to neurological disorders. In spite of

the different effects that these disorders seem to have on the experience of art, pa-

tients continue to engage with art in personally meaningful ways, even though per-

ceptual, memorable, or affective qualities might escape them.

Neuroimaging studies suggest that the experience of art involves three functionally

discernible sets of brain activity, which roughly match the components highlighted in

the insightful description of visitors’ reactions to the Bishop’s chapel at Cathedral of

Le Mans, and current psychological models (Chatterjee, 2004b; Leder et al., 2004).

Appreciating art engages processes related to perception (attentional enhancement

of the analyses of certain features), cognition (evaluative judgment, attention, and re-

trieval of information frommemory), and affect (generation of pleasant feelings, emo-

tions, representation and anticipation of reward, and awareness of one’s own affective

state). These processes are performed in parallel, they are highly interrelated, and they

rely heavily on information feedback, making it impossible to describe anymeaningful

sequence of events. One cannot even say that an art experience begins with perception,

given the strong biasing influences that context, expectations, and prior knowledge

have even on very early perceptual processes (Churchland et al., 1994).

In light of the multilayered nature of the experience of art, it is fair to forgive Saint

Augustine for being distracted from the divine words. Listening to the music must

have given him pleasure. It might have soothed him, relaxed him, or aroused him,

but he was definitively moved by it. His attention might have been caught by some

parts of the melody, to which he listened intently. He might have recognized some of

the music, and thought that it sounded different, or similar to other tunes. It might

have brought back memories from people and places he used to know. Some of

the melodies might have been difficult to follow and required more effort. Concen-

trating on the words on top of all of this could not have been an easy task.

Despite how much science has reveled about the biology underlying art, to some

humanists, the studies reviewed earlier add little value to our understanding of the

experience of art (Tallis, 2008a,b). When the art experience is studied in the con-

trolled conditions of the laboratory, art loses crucial qualities that make it interesting

1534 Conclusions, limitations, and prospects


from the humanities perspective. To philosophers, art theorists, and art historians, art

is fundamentally a cultural construction subject to contextual variation, to historical

change, and to criticism; they value the way it flexibly adapts to different roles in

different cultures at different times (Davies, 2012). Humanists believe that consid-

ering artistic and aesthetic objects merely as physical elements—stimuli—strips

them of their essential historical, cultural, and intentional context and significance

(Margolis, 1980). Accordingly, their response to the scientific approach to art and

aesthetics commonly varies from mild skepticism to strong criticism, and to outright

hostility. Some humanists have even suggested that art is in principle not amenable to

scientific inquiry (Massey, 2009; Tallis, 2008a,b). Gopnik (2012), for instance,

believes that “The crucially artistic aspects of artworks are the kind of painfully com-

plex cultural and cognitive phenomena that are likely to escape experimental study,

at least for the foreseeable future” (Gopnik, 2012, p. 144). Others believe that

scientists simply miss the whole point of studying art and aesthetics (Currie,

2003). In a renowned paper, Dickie (1962) argued that scientific approaches to

aesthetics are completely irrelevant with regard to two of the main issues aesthetics

deals with: the logical problems of aesthetics, related to the meaning of aesthetic

notions and the veracity of descriptive and evaluative aesthetic assertions, and the

understanding of the aesthetic experience.

However, some of the questions philosophers ask are not well suited for psychol-

ogy or neuroscience (Fenner, 1992), such as what is art? or what is the definition of

art? Psychology and neuroscience should not be expected to address the same ques-

tions that philosophers and art theorists pose, or carry out their studies at the same

level of generality. It is unfair to judge the success of psychological or neuroscientific

approaches to art and aesthetics based on how they fair on philosophical or art-

theoretical issues. Maybe it is true that neuroscience will never be able to explain

why a given artwork is a good artwork. But maybe neuroscientists are more inter-

ested in explaining other issues, such as the biological mechanisms underlying peo-

ple’s enjoyment of some artworks and not others.

It is undeniable, however, that neuroscientists, and to a great extent psychologists

too, have overlooked the impact of context andmeaning on the experience of art. This

owes mainly to methodological constraints and to the strategy of starting small and

building up. Neuroimaging techniques require highly controlled environments. It is,

therefore, not easy to create contextual conditions that can bemanipulated experimen-

tally. Researchers have circumvented this problem by creating semantic contexts cre-

ated with verbal or visual cues, such as Kirk et al.’s (2009a) use of “gallery” and

“computer” labels to indicate that some stimuli had been taken from an art gallery

and others had been created using computer software, or Harvey and colleagues’

(2010) presentation of logos of companies that sponsored people’s participation in

the experiment or not. No doubt this is still a far cry from the idea of cultural and his-

torical context humanist scholars have in mind. It is, nonetheless, a first step in that

direction. The same could be said about psychological experiments showing that am-

biguity, knowledge, and information have crucial effects on the experience of art

(Belke et al., 2006, 2010; Leder et al., 2006; Russell, 2003; Temme, 1992).



In this chapter, I have outlined the basic role of several brain regions in the ex-

perience of music, painting, architecture, and dance. This analysis provides only a

static image of the biology of art appreciation. What we need now is to understand

the dynamic interactions among these components. Fortunately, some scientists are

already turning toward this issue (Lacey et al., 2011; Salimpoor et al., 2011).

However, only interdisciplinary collaboration among scientists—also between

scientists and philosophers, art theorists and historians—together with scientific in-

genuity, can lead to a true understanding of the biological foundations of the kind of

art experience that troubled Saint Augustine and marveled the Le Mans chronicler.

ReferencesAdolphs, R., Tranel, D., 1999. Preferences for visual stimuli following amygdala damage.

J. Cogn. Neurosci. 11, 610–616.

Alajouanine, T., 1948. Aphasia and artistic realization. Brain 71, 229–241.

Annoni, J., Devuyst, G., Carota, A., Bruggimann, L., Bogousslavsky, J., 2004. Changes in ar-

tistic style after minor posterior stroke. J. Neurol. Neurosurg. Psychiatry 76, 797–803.

Arnsten, A.F., Segal, D.S., Neville, H.J., Hillyard, S.A., Janowsky, D.S., Judd, L.L.,

Bloom, F.E., 1983. Naloxone increases electrophysiological measures of selective infor-

mation processing in humans. Nature 304, 725–727.

Augustine, A., 1909. The Confessions of St. Augustine. Translated by Edward B. Pusey. P. F.

Collier & Son Company, New York.

Bar, M., Neta, M., 2007. Visual elements of subjective preference modulate amygdala activa-

tion. Neuropsychologia 45, 2191–2200.

Bazner, H., Hennerici, M., 2006. Stroke in painters. In: Rose, F.C. (Ed.), The Neurobiology of

Painting. International Review of Neurobiology, vol. 74. Academic Press, San Diego,

pp. 165–191.

Belke, B., Leder, H., Augustin, D., 2006. Mastering style—effects of explicit style-related in-

formation, art knowledge and affective state on appreciation of abstract paintings. Psychol.

Sci. 48, 115–134.

Belke, B., Leder, H., Strobach, T., Carbon, C.C., 2010. Cognitive fluency: high-level proces-

sing dynamics in art appreciation. Psychol. Aesthet. Creativity Arts 4, 214–222.

Biederman, I., Vessel, E.A., 2006. Perceptual pleasure and the brain. Am. Sci. 95,

249–255.

Blood, A.J., Zatorre, R.J., 2001. Intensely pleasurable responses to music correlate with

activity in brain regions implicated in reward and emotion. Proc. Natl. Acad. Sci.

U.S.A. 98, 11818–11823.

Blood, A.J., Zatorre, R.J., Bermudez, P., Evans, A.C., 1999. Emotional responses to pleasant and

unpleasantmusiccorrelatewithactivity inparalimbicbrainregions.Nat.Neurosci.2,382–387.

Boeve, B.F., Geda, Y.E., 2001. Polka music and semantic dementia. Neurology 57, 1485.

Bogousslavsky, J., 2005. Artistic creativity, style and brain disorders. Eur. Neurol. 54, 103–111.

Bonvincini, G., 1926. Die Aphasie des Malers Vierge. Wien. Med. Wochenschr. 76, 88–91.

Bromberger, B., Sternschein, R., Widick, P., Smith, W., Chatterjee, A., 2011. The right hemi-

sphere in esthetic perception. Front. Hum. Neurosci. 5, 109. http://dx.doi.org/10.3389/

fnhum.2011.00109.

Brown, S., Martinez, M.J., Parsons, L.M., 2004. Passive music listening spontaneously en-

gages limbic and paralimbic systems. Neuroreport 15, 2033–2037.

http://refhub.elsevier.com/B978-0-444-63287-6.00007-5/rf0005






























http://dx.doi.org/10.3389/fnhum.2011.00109

http://dx.doi.org/10.3389/fnhum.2011.00109



155References


Budrys, V., Skullerud, K., Petroska, D., Lengveniene, J., Kaubrys, G., 2007. Dementia and art:

neuronal intermediate filament inclusion disease and dissolution of artistic creativity. Eur.

Neurol. 57, 137–144.

Calvo-Merino, B., Jola, C., Glaser, D.E., Haggard, P., 2008. Towards a sensorimotor aesthetics

of performing art. Conscious. Cogn. 17, 911–922.

Calvo-Merino, B., Urgesi, C., Orgs, G., Aglioti, S.M., Haggard, P., 2010.

Extrastriate body area underlies aesthetic evaluation of body stimuli. Exp. Brain Res.

204, 447–456.

Cela-Conde, C.J., Ayala, F.J., Munar, E., Maestu, F., Nadal, M., Capo, M.A., Marty, G., 2009.

Sex-related similarities and differences in the neural correlates of beauty. Proc. Natl. Acad.

Sci. U.S.A. 106, 3847–3852.

Cela-Conde, C.J., Agnati, L., Huston, J.P., Mora, F., Nadal, M., 2011. The neural foundations

of aesthetic appreciation. Prog. Neurobiol. 94, 39–48.

Cela-Conde, C.J., Marty, G., Maestu, F., Ortiz, T., Munar, E., Fernandez, A., Roca, M.,

Rossello, J., Quesney, F., 2004. Activation of the prefrontal cortex in the human visual

aesthetic perception. Proc. Natl. Acad. Sci. U.S.A. 101, 6321–6325.

Chatterjee, A., 2004a. The neuropsychology of visual artistic production. Neuropsychologia

42, 1568–1583.

Chatterjee, A., 2004b. Prospects for a cognitive neuroscience of visual aesthetics. Bull.

Psychol. Arts 4, 55–60.

Chatterjee, A., Widick, P., Sternschein, R., Smith II., W.B., Bromberger, B., 2010. The assess-

ment of art attributes. Empirical Stud. Arts 28, 207–222.

Chelazzi, L., Corbetta, M., 2000. Cortical mechanisms of visuospatial attention in the primate

brain. In: Gazzaniga, M.S. (Ed.), The New Cognitive Neurosciences. MIT Press,

Cambridge, MA, pp. 667–686.

Christoff, K., Gabrieli, J.D.E., 2000. The frontopolar cortex and human cognition: evidence for

a rostrocaudal hierarchical organization within the human prefrontal cortex. Psychobiol-

ogy 28, 168–186.

Chun, M.M., Golomb, J.D., Turk-Browne, N.B., 2011. A taxonomy of external and internal

attention. Annu. Rev. Psychol. 62, 73–101.

Churchland, P.S., Ramachandran, V.S., Sejnowski, T.J., 1994. A critique of pure vision. In:

Koch, C., Davis, J.L. (Eds.), Large-Scale Neuronal Theories of the Brain. The MIT Press,

Cambridge, MA, pp. 23–60.

Colombo-Thuillard, F., Assal, G., 2007. Persisting aphasia, cerebral dominance, and painting

in the famous artist Carl Fredrik Reutersward. In: Bogousslavsky, J., Hennerici, M.G.

(Eds.), Neurological Disorders in Famous Artists—Part 2. Frontiers of Neurology and

NeuroscienceKarger, Basel, pp. 169–183.

Compton, R., 2003. The interface between emotion and attention: a review of evidence from

psychology and neuroscience. Behav. Cogn. Neurosci. Rev. 2, 115–129.

Crutch, S.J., Isaacs, R., Rossor, M.N., 2001. Some workmen can blame their tools: artistic

change in an individual with Alzheimer’s disease. Lancet 357, 2129–2133.

Cupchik, G.C., Vartanian, O., Crawley, A., Mikulis, D.J., 2009. Viewing artworks: contribu-

tions of cognitive control and perceptual facilitation to aesthetic experience. Brain Cogn.

70, 84–91.

Currie, G., 2003. Aesthetics and cognitive science. In: Levinson, J. (Ed.), The Oxford Hand-

book of Aesthetics. Oxford University Press, Oxford, pp. 706–721.

Davies, S., 2012. The artful species. Oxford University Press, Oxford.

Di Dio, C., Macaluso, E., Rizzolatti, G., 2007. The golden beauty: Brain response to Classical

and Renaissance sculptures. Plos One 11, e1201.




















































Dickie, G., 1962. Is psychology relevant to aesthetics? Philos. Rev. 71, 285–302.

Drago, V., Foster, P.S., Skidmore, F.M., Heilman, K.M., 2009. Creativity in Parkinson’s

disease as a function of right versus left hemibody onset. J. Neurol. Sci. 276, 179–183.

Fenner, D.E., 1992. Modest aesthetic naturalism. J. Aesthet. Art Critic. 50, 283–289.

Geroldi, C., Metitieri, T., Binetti, G., Zanetti, O., Trabucchi, M., Frisoni, G.B., 2000. Pop

music and frontotemporal dementia. Neurology 55, 1935–1936.

Gopnik, B., 2012. Aesthetic science and artistic knowledge. In: Shimamura, A.P., Palmer, S.E.

(Eds.), Aesthetic Science. Connecting Minds, Brains and Experience. Oxford University

Press, New York, pp. 129–159.

Gosselin, N., Peretz, I., Johnsen, E., Adolphs, R., 2007. Amygdala damage impairs emotion

recognition from music. Neuropsychologia 45, 236–244.

Griffiths, T.D., Warren, J.D., Dean, J.L., Howard, D., 2004. “When the feeling’s gone”: a se-

lective loss of musical emotion. J. Neurol. Neurosurg. Psychiatry 75, 341–345.

Halpern, A.R., O’Connor, M.G., 2013. Stability of Art Preference in Frontotemporal Demen-

tia. Psychol. Aesthet. Creativity Arts 7, 95–99.

Halpern, A.R., Ly, J., Elkin-Frankston, S., O’Connor, M.G., 2008. “I know what I like”: sta-

bility of aesthetic preference in Alzheimer’s patients. Brain Cogn. 66, 65–72.

Harvey, A.H., Kirk, U., Denfield, G.H., Montague, P.R., 2010. Monetary favors and their in-

fluence on neural responses and revealed preference. J. Neurosci. 30, 9597–9602.

Hopfinger, J.B., Buonocore, M.H., Mangun, G.R., 2000. The neural mechanisms of top-down

attentional control. Nat. Neurosci. 3, 284–291.

Ishizu, T., Zeki, S., 2011. Toward a brain-based theory of beauty. PLoS One 6 (7), e21852.

http://dx.doi.org/10.1371/journal.pone.0021852.

Jacobsen, T., Hofel, L., 2003. Descriptive and evaluative judgment processes: behavioral and

electrophysiological indices of processing symmetry and aesthetics. Cogn. Affect. Behav.

Neurosci. 3, 289–299.

Jacobsen, T., Schubotz, R.I., Hofel, L., von Cramon, D.Y., 2006. Brain correlates of aesthetic

judgment of beauty. Neuroimage 29, 276–285.

Johansen-Berg, H., Lloyd, D.M., 2000. The physiology and psychology of selective attention

to touch. Front. Biosci. 5, 894–904.

Kanwisher, N., Wojciulik, E., 2000. Visual attention: insights from brain imaging. Nat. Rev.

Neurosci. 1, 91–100.

Kawabata, H., Zeki, S., 2004. Neural correlates of beauty. J. Neurophysiol. 91, 1699–1705.

Kirk, U., 2008. The neural basis of object-context relationships on aesthetic judgment. PLoS

One 3 (11), e3754. http://dx.doi.org/10.1371/journal.pone.0003754.

Kirk, U., Skov, M., Christensen, M.S., Nygaard, N., 2009a. Brain correlates of aesthetic ex-

pertise: a parametric fMRI study. Brain Cogn. 69, 306–315.

Kirk, U., Skov, M., Hulme, O., Christensen, M.S., Zeki, S., 2009b. Modulation of aesthetic

value by semantic context: an fMRI study. Neuroimage 44, 1125–1132.

Koelsch, S., Fritz, T., von Cramon, D.Y., Muller, K., Friederici, A.D., 2006. Investigating

emotion with music: an fMRI study. Hum. Brain Mapp. 27, 239–250.

Koepp, M.J., Hammers, A., Lawrence, A.D., Asselin, M.C., Grasby, P.M., Bench, C.J., 2009.

Evidence for endogenous opioid release in the amygdala during positive emotion. Neuro-

image 44, 252–256.

Kringelbach, M.L., Berridge, K.C., 2009. Towards a functional neuroanatomy of pleasure and

happiness. Trends Cogn. Sci. 13, 479–487.

Kringelbach, M.L., Rolls, E.T., 2004. The functional neuroanatomy of the human

orbitofrontal cortex: evidence from neuroimaging and neuropsychology. Prog. Neurobiol.

72, 341–372.






















http://dx.doi.org/10.1371/journal.pone.0021852


























157References


Lacey, S., Hagtvedt, H., Patrick, V.M., Anderson, A., Silla, R., Deshpande, G., Sathian, K.,

2011. Art for reward’s sake: visual art recruits the ventral striatum. Neuroimage 55,

420–433.

Lang, P.J., Bradley, M.M., Fitzsimmons, J.R., Cuthbert, B.N., Scott, J.D., Moulder, B.,

Nangia, V., 1998. Emotional arousal and activation of the visual cortex: an fMRI study.

Psychophysiology 35, 199–210.

Le Merrer, J., Becker, J.A.J., Befort, K., Kieffer, B.L., 2009. Reward processing by the opioid

system in the brain. Physiol. Rev. 89, 1379–1412.

Leder, H., Belke, B., Oeberst, A., Augustin, D., 2004. A model of aesthetic appreciation and

aesthetic judgments. Br. J. Psychol. 95, 489–508.

Leder, H., Carbon, C.C., Ripsas, A., 2006. Entitling art: influence of different types of title

information on understanding and appreciation of paintings. Acta Psychol. 121, 176–198.

Lengger, P.G., Fischmeister, F.P.S., Leder, H., Bauer, H., 2007. Functional neuroanatomy of

the perception of modern art: a DC-EEG study on the influence of stylistic information on

aesthetic experience. Brain Res. 1158, 93–102.

Levitin, D.J., Tirovolas, A.K., 2009. Current advances in the cognitive neuroscience of music.

Ann. N. Y. Acad. Sci. 1156, 211–231.

Lewis, M.E., Mishkin, M., Bragin, E., Brown, R.M., Pert, C.B., Pert, A., 1981. Opiate receptor

gradients in monkey cerebral cortex: correspondence with sensory processing hierarchies.

Science 221, 1166–1169.

Liu, T., Larsson, J., Carrasco, M., 2007. Feature-based attention modulates orientation-

selective responses in human visual cortex. Neuron 55, 313–323.

Mabillon, J., 1682. Veterum Analectum, vol. 3. Paris.

Margolis, J., 1980. Prospects for a science of aesthetic perception. In: Fisher, J. (Ed.), Perceiv-

ing Artworks. Temple University Press, Philadelphia, pp. 213–239.

Massey, I., 2009. The neural imagination. Aesthetic and neuroscientific approaches to the arts.

University of Texas Press, Austin.

Miller, B.L., Hou, C.E., 2004. Portraits of artists. Emergence of visual creativity in dementia.

Arch. Neurol. 61, 842–844.

Mitterschiffthaler, M.T., Fu, C.H.Y., Dalton, J.A., Andrew, C.M., Williams, S.C.R., 2007.

A Functional MRI Study of Happy and Sad Affective States Induced by Classical Music.

Hum. Brain Mapp. 28, 1150–1162.

Munar, E., Nadal, M., Rossello, J., Flexas, A., Moratti, S., Maestu, F., Cela-Conde, C.J., 2012.

Lateral orbitofrontal cortex involvement in initial negative aesthetic impression formation.

PLoS One 7 (6), e38152. doi:http://dx.doi.org/10.1371/journal.pone.0038152.

Murphy, F.C., Nimmo-Smith, I., Lawrence, A.D., 2003. Functional neuroanatomy of emo-

tions: a meta-analysis. Cogn. Affect. Behav. Neurosci. 3, 207–233.

Nadal, M., Pearce, M.T., 2011. The Copenhagen Neuroaesthetics conference: prospects and

pitfalls for an emerging field. Brain Cogn. 76, 172–183.

Northoff, G., Bermpohl, F., 2004. Cortical midline structures and the self. Trends Cogn. Sci. 8,

102–107.

O’Doherty, J.P., Kringelbach, M.L., Rolls, E.T., Hornak, J., Andrews, C., 2001. Abstract re-

ward and punishment representations in the human orbitofrontal cortex. Nat. Neurosci. 4,

95–102.

Phan, K.L., Wager, T., Taylor, S.F., Liberzon, I., 2002. Functional neuroanatomy of emotion:

a meta-analysis of emotion activation studies in PET and fMRI. Neuroimage 16, 331–348.

Posner, M.I., DiGirolamo, G.J., 2000. Attention in cognitive neuroscience: an overview. In:

Gazzaniga, M.S. (Ed.), The New Cognitive Neurosciences. MIT Press, Cambridge,

MA, pp. 623–631.

















































Reynolds, J.H., Chelazzi, L., 2004. Attentional modulation of visual processing. Annu. Rev.

Neurosci. 27, 611–647.

Russell, P.A., 2003. Effort after meaning and the hedonic value of paintings. Br. J. Psychol. 94,

99–110.

Salimpoor, V.N., Benovoy, M., Larcher, K., Dagher, A., Zatorre, R.J., 2011. Anatomically

distinct dopamine release during anticipation and experience of peak emotion to music.

Nat. Neurosci. 14, 257–262. http://dx.doi.org/10.1038/nn.2726.

Seeley, W.W., Matthews, B.R., Crawford, R.K., Gorno-Tempini, M.L., Foti, D.,

Mackenzie, I.R., Miller, B.L., 2008. Unraveling Bolero: progressive aphasia, transmodal

creativity and the right posterior neocortex. Brain 131, 39–49.

Sellal, F., Andriantseheno, M., Vercueil, L., Hirsch, E., Kahane, P., Pellat, J., 2003. Dramatic

changes in artistic preference after left temporal lobectomy. Epilepsy Behav. 4, 449–450.

Tallis, R., 2008a. The limitations of a neurological approach to art. Lancet 372, 19–20.

Tallis, R., 2008b. The neuroscience delusion. The Times Literary Supplement. April 9, 2008.

Tatarkiewicz, W., 1970. History of aesthetics. Medieval aesthetics, vol. II. Mouton, Warsaw.

Temme, J.E.V., 1992. Amount and kind of information in museums: its effects on visitors sat-

isfaction and appreciation of art. Visual Arts Res. 2, 28–36.

Vartanian, O., Goel, V., 2004. Neuroanatomical correlates of aesthetic preference for paint-

ings. Neuroreport 15, 893–897.

Veldhuizen, M.G., Bender, G., Constable, R.T., Small, D.M., 2007. Trying to detect taste in a

tasteless solution: modulation of early gustatory cortex by attention to taste. Chem. Senses

32, 569–581.

Vuilleumier, P., 2005. How brains beware: neural mechanisms of emotional attention. Trends

Cogn. Sci. 9, 585–594.

Vuilleumier, P., Armony, J., Dolan, R.J., 2004. Reciprocal links between emotion and atten-

tion. In: Frackowiak, R.S.J., Friston, K.J., Frith, C.D., Dolan, R.J., Price, C.J., Zeki, S.,

Ashburner, J., Penny, W. (Eds.), Human Brain Function. second ed. Academic Press,

San Diego, pp. 419–444.

Wise, S.P., Herkenham, M., 1982. Opiate receptor distribution in the cerebral cortex of the

rhesus monkey. Science 218, 381–389.

Woldorff, M.G., Gallen, C.C., Hampson, S.A., Hillyard, S.A., Pantev, C., Sobel, D.,

Bloom, F.E., 1993. Modulation of early sensory processing in human auditory cortex

during auditory selective attention. Proc. Natl. Acad. Sci. U.S.A. 90, 8722–8726.

Yantis, S., Serences, J.T., 2003. Cortical mechanisms of space-based and object-based

attentional control. Curr. Opin. Neurobiol. 13, 187–193.

Yue, X., Vessel, E.A., Biederman, I., 2007. The neural basis of scene preferences. Neuroreport

18, 525–529.

Zaidel, D.W., 2005. Neuropsychology of Art: Neurological, Cognitive, and Evolutionary

Perspectives. Psychology Press, Hove, England.

Zaimov, K., Kitov, D., Kolev, N., 1969. Aphasie chez un peintre. Encephale 58, 377–417.

Zelano, C., Bensafi, M., Porter, J., Mainland, J., Johnson, B., Bremner, E., Sobel, N., 2005.

Attentional modulation in human primary olfactory cortex. Nat. Neurosci. 8, 114–120.

Zysset, S., Huber, O., Ferstl, E., von Cramon, D.Y., 2002. The anterior frontomedian cortex

and evaluative judgment: an fMRI study. Neuroimage 15, 983–991.





http://dx.doi.org/10.1038/nn.2726






































Date post:	28-Feb-2019
Category:	Documents
Upload:	truonghuong
View:	215 times
Download:	0 times

Provided for non-commercial research and educational use...

Documents