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Essay Neuroaesthetics and the Trouble with Beauty Bevil R. Conway1,2*, Alexander Rehding3 1Program in Neuroscience, Wellesley College, Wellesley, Massachusetts, United States of America, 2Department of Neurobiology, Harvard Medical School, Boston, Massachusetts, United States of America, 3Department of Music, Harvard University, Cambridge, Massachusetts, United States of America The famous nineteenth-century psycho- physicist Gustav Fechner was also a poet and art critic. Armed with the tools of science, Fechner sought to reconcile his various interests. He would doubtless be interested by technological developments in neuroscience that have revealed the operations of neurons at cellular resolution and have enabled us to peer almost unnoticed into each other’s working brains. But can these tools advance our understanding of aesthetics beyond Fech- ner’s insights ? The nascent field of neuroaesthetics claims it can. Here we consider what questions this new field is poised to answer. We underscore the importance of distinguishing between beauty, art, and perception—terms often conflated by ‘‘aesthetics’’—and identify adjacent fields of neuroscience such as sensation, perception, attention, reward, learning, memory, emotions, and decision making, where discoveries will likely be informative. Aesthetics and Neuroscience Aesthetics has a complex history. The term derives from the Greek ‘‘perception’’ and was coined by Alexander Baumgarten in 1750 as the study of sensory knowledge. But following Immanuel Kant’s Critique of Judgment in 1790 , aesthetics began focusing on the concept of beauty, in nature and in art. During the nineteenth century, the term became largely synony- mous with the philosophy of art. These three connotations—perception, beauty, art—point in different directions but are often conflated in neuroaesthetics. Kant is a preferred philosopher among neuroaestheticians, no doubt because of his towering stature in the history of Western thought. He pursued a universal- ist approach to beauty, an appealing concept for neuroscientists because it suggests a discrete neural basis. But Kant’s concept of beauty has been severely criticized in light of the prevailing plural- ism of artistic styles. To make matters more complicated, there is no consensus on the nature of beauty. Kant’s under- standing of beauty was predicated on an attitude of ‘‘disinterested contemplation’’ , whereas Friedrich Nietzsche roundly dismissed this notion and underlined the impact of sensual attraction . For the poet John Keats, beauty equaled truth , while Stendhal, the French novelist, char- acterized beauty as the ‘‘promise of happiness’’ . More recently, Elaine Scarry described beauty as an urge to repeat . While each of these theories is respected, not one is universally accepted. Partly this diversity of opinions is connect- ed to the different functions that beauty holds within various philosophical systems, being sometimes viewed in connection with epistemology or with ethics. One goal of neuroaesthetics is to get to the bottom of the problem of artistic beauty. How can this be accomplished? Experiences of beauty are often deeply moving, and their importance to the human condition invites a neuroscientific explanation. But while deep emotional reactions are often associated with beauty, being moved does not always indicate an instance of beauty. Consider hearing about a disaster, celebrating a sports victory, or smelling a long-forgotten scent. These experiences are better described as ‘‘sympathy,’’ ‘‘elation,’’ and ‘‘memory,’’ rather than experiences of beauty. If neuroaesthetics is to be concerned specif- ically with beauty, it must draw distinc- tions between mechanisms for such dispa- rate reactions. Since many experiences of beauty are related to art, neuroaestheti- cians have focused their attention on the analysis of artworks. For example, Rama- chanran , Zeki , and Kandel  have presented case studies focusing on classical Indian art, American and Euro- pean modernists, and the Viennese Seces- sionists. Explicitly or implicitly, these studies aim to extract rules that would lead to a practical definition of beauty, connecting features of objects and neural activity. Zeki, for instance, argues that the power of Alexander Calder’s sculptures derives from the black-and-white moving parts, potent activators of the brain’s motion-processing center. It may be no coincidence that the art these three authors hold up relates to the culture in which they were each raised. One potential danger in aesthetic projects is to universalize one’s subjective convic- tions and assume that an experience of beauty is common to all. Projecting from individual subjective experience is decep- tive, for there is ample evidence that notions of beauty vary between cultures and are mutable even within a culture— just think of fast-changing trends in fashion. Moreover, the equation (art=beauty) rests on shaky ground. Throughout history, artists have created deeply moving art- work that is emphatically not beautiful; Goya’s Saturn Devouring One of His Sons (Figure 1) provides a famous historical example. Large swaths of twentieth- century art have greatly expanded—or entirely disavowed—notions of beauty. Such distinctions may seem picky, but interdisciplinary work such as neuroaes- thetics relies on shared principles, and requires heightened attention to concep- tual clarity. Essays articulate a specific perspective on a topic of broad interest to scientists. Citation: Conway BR, Rehding A (2013) Neuroaesthetics and the Trouble with Beauty. PLoS Biol 11(3): e1001504. doi:10.1371/journal.pbio.1001504 Published March 19, 2013 Copyright: 2013 Conway, Rehding. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Funding: This work was supported by the National Science Foundation (Grant 0918064). The NSF had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Competing Interests: The authors have declared that no competing interests exist. * E-mail: firstname.lastname@example.org PLOS Biology | www.plosbiology.org 1 March 2013 | Volume 11 | Issue 3 | e1001504 Neuroscience has provided a heuristic outlining how sensory signals are pro- cessed by the nervous system to yield behavior [10,11]. Signals from sensory epithelia such as the retina or basilar membrane are processed in the cerebral cortex by a series of areas that compute descriptions of the world: what or where objects are. These brain areas send signals to other brain structures that are respon- sible for evaluating options against expect- ed rewards—attaching significance to the sensory descriptions—and ultimately for making decisions, guided by learning, memory, and emotions. Below we argue that a successful neuroaesthetics will include the study of each of these stages of processing as they relate to handling, encoding, and generating aesthetic expe- riences, rather than an attempt to derive a single universal neural underpinning of what constitutes beauty. First Steps in Neuroaesthetics: Sensation, Perception, and Art One approach commonly included under the umbrella of neuroaesthetics involves examining art objects in muse- ums. Here the complication of establishing ‘‘beauty’’ is obviated by treating artworks as products of a massive empirical exper- iment. By analogy with evolutionary theory, the assumption is that the tiny number of works that survive the selective pressures exerted by collectors, cultural institutions, and fads are enriched for the strength of their effects on the nervous system. Using this approach, studies have uncovered various artistic strategies re- flecting fundamental operations of the neural mechanisms for sensation and perception [7,8,12–14]. For example, de- pictions of shadows in paintings often do not correspond to the light sources that cause them . Such unnoticed devia- tions from veracity reveal important ad- aptations of the brain to ecological pres- sures during evolution and development— in the case of shadows, the relationship of objects to light sources is in flux and therefore not a stable feature. Similarly, analysis of portraits has been insightful, showing that the outer contour of a face is more important for face recognition than the precise configuration of features . And paintings by Paul Cezanne, Henri Matisse, and Claude Monet show how these artists capitalized upon the neural mechanisms of color . This line of research is often described as the neuro- science of art, rather than neuroaesthetics, since it does not test for beauty . The approach may reveal the perceptually Figure 1. Goya y Lucientes, Francisco de, S a t u r n d e v o u r i n g o n e o f h i s s o n s (1821–1823). Mural transferred to canvas. 143.5 cm681.4 cm. Museo del Prado, Madrid. doi:10.1371/journal.pbio.1001504.g001 PLOS Biology | www.plosbiology.org 2 March 2013 | Volume 11 | Issue 3 | e1001504 relevant properties of visual stimuli— contributing to aesthetics as Baumgarten defined it—but these properties are nei- ther necessary nor sufficient features of beautiful objects. An Alexander Calder sculpture may consist of optimal stimuli for the brain’s motion center, but this aspect of the work does not make it beautiful. The art simply provides a fascinating demonstration of the compu- tations of the brain’s motion-perception circuits, and the genius of the artists for discovering them. It is an open question whether an analysis of artworks, no matter how celebrated, will yield universal principles of beauty. Compositional principles such as the golden ratio are intriguing possible universals, and captured the attention of Fechner, but despite mathematical appeal, the golden rectangle is not the favorite rectangle shape of most people . One possible almost-universal may be the appeal of certain female facial features (symmetry, high cheekbones, large eyes) and a 0.7 waist-to-hip ratio  or high body mass index . Explanations for these preferences depend on a correlation between the attributes and reproductive fitness. Yet celebrated representations of female beauty across history can deviate considerably from the 0.7 rule, and ratio preferences vary across cultures [21,22]. Depictions of reproductive fitness can be sexually appealing and contribute to aesthetic appeal, but such depictions are, again, neither necessary nor sufficient for beauty. Another possible universal con- cerns the intriguing discovery that painters typically center one eye along the hori- zontal axis of a picture , taken to indicate ‘‘hidden principles…operating in our aesthetic judgments.’’ But the trend towards eye-centering has declined dra- matically during and after avant-garde movements such as those led by Picasso . Whether this decline is attributable to the relative decline of beauty as a driving force in artistic creation or indi- cates a cultural shift in aesthetic prefer- ences is unclear. Using celebrated works as empirical data to understand beauty might be a worthwhile gambit, but we doubt that conclusions can be extended across peo- ples, times, and cultures. The only univer- sal feature of beauty besides our capacity to experience it appears to be its mutabil- ity, itself perhaps a topic for neuroscience. A Beauty Center? Fechner was well aware of the pitfalls of philosophical aesthetics and aimed to reformulate the field ‘‘from the ground up.’’ His appreciation of the inherently subjective nature of beauty led him to start with feelings of pleasure and displeasure elicited by art, since these constituted for him the bottom line beyond which further analysis was impossible. Contemporary neuroscience has gone much further. A recent study claims that ‘‘all works that appear beautiful to a subject have a single brain-based characteristic, which is that they have as a correlate of experiencing them a change in strength of [fMRI] activity within the mOFC [medial orbito- frontal cortex]’’ . Leaving aside meth- odological challenges [25,26], is such a correlation meaningful to understanding aesthetics? Subjectivist studies such as these over- come the difficulty of defining beauty by asking the participants to first rate visual objects or sounds [24,27]. Brain activity of each subject is then assessed to their own set of ‘‘beautiful’’ versus ‘‘ugly’’ stimuli. Four experimental-design chal- lenges surface. First, the options are necessarily restricted, and might not in- clude a truly beautiful choice—the study design tests preferences, not beauty. Second, different subjects likely interpret the instructions in radically different ways. Third, the use of different stimulus sets in different subjects makes it difficult to control for differences in low-level stimu- lus features, which likely drive different patterns of neural activity. And fourth, the experiment requires that a given object retain a fixed preferred status, and one that is not modulated by context, which we know is unlikely. As Fechner showed, mere exposure changes judg- ments of preference in favor of the familiar option. Brandishing fMRI does not circumvent these problems. More- over, fMRI has cripplingly low spatial and temporal resolution, and the relationship between the measured signal and under- lying neural activity is indirect. In addi- tion, fMRI experiments often only report regions that show differential activation between pairs of conditions (e.g., response to beautiful greater than response to ugly); such an analysis is misleading in situations in which all brain regions show significant but slightly different levels of activity for the different conditions, as is likely the case in considerations of beauty. Brain imaging provides a blurry, although seductively glossy, view of brain function. And by finessing a definition of beauty, these sorts of studies sidestep what is at the heart of our interest in beauty: the connection between physical stimuli, spe- cifically those crafted by human hands, and our response. Nonetheless, a discovery that every person’s experience of beauty (however vaguely defined) correlates with activity within a specific brain region would be surprising, since it would seem more likely that a complex reaction (beautiful!) would hinge not on the absolute level of activity within a single brain center but rather on the pattern of activity across many distrib- uted brain regions—specifically those re- sponsible for perception, reward, decision making, and emotion. Indeed, a broader reading of the literature reveals that the mOFC is not uniquely associated with experiences of beauty and may be neither necessary nor sufficient for these experi- ences. The mOFC appears to be part of a large network of brain regions that sub- serves all value judgments. For example, elevated activity within the mOFC is reported in studies of neuroeconomics in which subjects are asked to assign value to a selection of choices and are never asked to consider the beauty of the choices [11,28–30]. The mOFC has also been implicated in impulse control and self- regulation , in changing decision thresholds that influence whether infor- mation should be expressed in an evalu- ation , in attentional processes that underlie emotion-congruent judgment , and in moral decision making . Ascribing responses of the mOFC to experiences of beauty is premature; many experiences depend on these processes without being beautiful [27,35–38]. If the mOFC plays a critical role in mediating beauty, one might expect that strokes of the region would impair expe- riences of beauty. Strokes of the mOFC are rare, but the limited evidence suggests they affect self-related systems such as self- evaluation [39,40] and do not impact a person’s ability to experience beauty. Alternatively, strokes in other brain re- gions can, paradoxically, enhance creativ- ity, providing support for the notion that the expression of beauty depends on a broad, distributed network. Frontotempo- ral dementia can produce an acquired obsessiveness that is often linked to enhanced art production, usually of ex- tremely detailed works . In addition, strokes of the left hemisphere, which often cause aphasia, can produce hyperexpres- siveness . What Questions Can Neuroaesthetics Answer? Inspired by the power of polling, in 1994 a pair of artists, Komar and Melamid, set out to determine ‘‘USA’s most wanted painting.’’ The painting was formulated on PLOS Biology | www.plosbiology.org 3 March 2013 | Volume 11 | Issue 3 | e1001504 the basis of a thousand people’s responses to questions of their favorite color, favorite setting, and favorite subjects. The resulting painting is absurd, showing that a compo- sition with everything that people find beautiful does not make a beautiful paint- ing. Rational reductionist approaches to the neural basis for beauty run a similar risk of pushing the round block of beauty into the square hole of science and may well distill out the very thing one wants to understand. There is a popular conception of beauty as a fixed attribute of objects, a notion that much of current neuroaes- thetics depends upon. But there is a distinction between abstract notions of beauty and our experience of it—consider a specific example in which you have experienced beauty. Beauty is an analog, not binary, condition that varies in complex ways with exposure, context, attention, and rest—as do most perceptual responses. In trying to crack the subjective beauty nut with scientific, objective information, we also run the risk of fueling a normative, possibly dangerous campaign through which science is required to valorize our experience. Should we deny someone’s experience of beauty if the mOFC is not activated? Obviously not. But the question underscores the danger of reverse infer- ence, a technique used in brain-imaging studies which posits that activation of a brain region indicates the presence of a stimulus . Reverse inference is almost always invalid because single brain struc- tures almost never regulate single specific experiences. Insofar as beauty is a product of the brain, correlations between brain activity and experiences of beauty must exist. At what spatial scale, and within what brain regions, do we find these correlations? What functions do the brain regions implicated serve in other behaviors? What signals during development and experi- ence are responsible for wiring up these circuits? And perhaps most critically, how does the activity of these circuits integrate across modalities and time to bring about the dynamic, elusive quality of beauty? To address these questions, the field is thirsty for carefully conducted experiments that distinguish responses to beauty from those involved in more general value-based decision tasks such as self-evaluation or selecting a juice for lunch. But any such experiments are caught on the same stubborn thorn—the lack of a cogent, universally accepted definition of beauty. One should not always demand a precise definition to make headway, but it might turn out that the philosophers’ disagree- ment is symptomatic: maybe there is no universal concept beyond the human capacity to experience beauty. Our cau- tion about neuroscience’s focus on beauty differs from the skepticism that attended scientific study of other subjective phe- nomena such as illusory contours (or even consciousness); in the case of illusory contours, the subjective experience to a given physical stimulus is universal. So, what is neuroaesthetics supposed to study? Experiences of beauty typically require attention and are accompanied by feelings of pleasure [11,27,44]. In the same way that basic studies at the interface of sensory neuroscience and art have been productive—not in addressing why art objects are beautiful but in uncovering the strategies that artists use to generate artwork—basic investigations of the mech- anisms of attention, decision making, reward, and emotion [11,28,29,45–47] could inform neuroaesthetics. The field will benefit from developing models relat- ing observations from the humanities to the careful neuroscience that has uncov- ered computations at cellular resolution within the value-judging structures of the monkey brain. These structures, not coincidentally, are analogous to those identified in fMRI studies of beauty in humans. Some neurons within these structures encode the value of the choices on offer, while others encode the value of the selected choice. Moreover, the neurons adapt on different timescales, displaying ‘‘menu-invariant’’ firing at short timescales and adaptable behavior on longer time- scales. This adaptation may account for our ability to make choices across vastly different scales, for example from a restaurant menu in one instance and from houses offered for sale in the next instance . It seems entirely reasonable—even likely—that these neurons are also impli- cated in the thorny task of deciding what is beautiful. Reformulated in this way, neu- roaesthetics is decoupled from beauty and can exploit advances across a range of empirical neuroscience, from sensory en- coding to decision making and reward. There may well be a ‘‘beauty instinct’’ implemented by dedicated neural machin- ery capable of producing a diversity of beauty reactions, much as there is lan- guage circuitry that can support a multi- tude of languages (and other operations). A need to experience beauty may be universal, but the manifestation of what constitutes beauty certainly is not. On the one hand, a neuroaesthetics that extrapo- lates from an analysis of a few great works, or one that generalizes from a single specific instance of beauty, runs the risk of missing the mark. On the other, a neuroaesthetics comprising entirely sub- jectivist accounts may lose sight of what is specific to encounters with art. Neuroaes- thetics has a great deal to offer the scientific community and general public. Its progress in uncovering a beauty instinct, if it exists, may be accelerated if the field were to abandon a pursuit of beauty per se and focus instead on uncovering the relevant mechanisms of decision making and reward and the basis for subjective preferences, much as Fech- ner counseled. This would mark a return to a pursuit of the mechanisms underlying sensory knowledge: the original concep- tion of aesthetics. Acknowledgments We thank Caroline Jones and David Hilbert for useful discussions and also thank the Wellesley College Neuroscience Program. References 1. Fechner GT (1872) Vorschule der a¨sthetik (2 volumes). Leipzig: Breitkopf und Ha¨rtel. 2. Kant I (2001 ) Critique of power of judgment. Guyer P, translator. Cambridge: Cam- bridge University Press. 3. Nietzsche F (1989 ) On the genealogy of morals. Kaufmann WA, translator. New York: Vintage Books. 4. Keats J (2010) Ode on a Grecian urn and other poems. Whitefish (Montana): Kessinger Publish- ing, LLC. 5. Stendhal [pseud. 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Yu DW, Shepard GH Jr (1998) Is beauty in the eye of the beholder? Nature 396: 321–322. 22. Marlowe F, Wetsman A (2001) Preferred waist-to- hip ratio and ecology. Pers Individ Dif 30: 481– 489. 23. Tyler CW (1998) Painters centre one eye in portraits. Nature 392: 877. 24. Ishizu T, Zeki S (2011) Toward a brain-based theory of beauty. PLoS ONE 6: e21852. doi:10.1371/journal.pone.0021852 25. Vul E, Pashler H (2012) Voodoo and circularity errors. Neuroimage 62: 945–948. 26. Kriegeskorte N, Simmons WK, Bellgowan PS, Baker CI (2009) Circular analysis in systems neuroscience: the dangers of double dipping. Nat Neurosci 12: 535–540. 27. Blood AJ, Zatorre RJ (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. 28. Kable JW, Glimcher PW (2009) The neurobiol- ogy of decision: consensus and controversy. Neuron 63: 733–745. 29. Padoa-Schioppa C, Cai X (2011) The orbitofron- tal cortex and the computation of subjective value: consolidated concepts and new perspec- tives. Ann N Y Acad Sci 1239: 130–137. 30. Plassmann H, O’Doherty J, Rangel A (2007) Orbitofrontal cortex encodes willingness to pay in everyday economic transactions. J Neurosci 27: 9984–9988. 31. Mehta PH, Beer J (2009) Neural mechanisms of the testosterone-aggression relation: the role of orbitofrontal cortex. J Cogn Neurosci 22: 2357– 2368. 32. Hughes BL, Beer JS (2012) Medial orbitofrontal cortex is associated with shifting decision thresh- olds in self-serving cognition. Neuroimage 61: 889–898. 33. Bhanji JP, Beer JS (2011) Unpacking the neural associations of emotion and judgment in emotion- congruent judgment. Soc Cogn Affect Neurosci 7: 348–356. 34. Tsukiura T, Cabeza R (2010) Shared brain activity for aesthetic and moral judgments: implications for the Beauty-is-Good stereotype. Soc Cogn Affect Neurosci 6: 138–148. 35. Cloutier J, Heatherton TF, Whalen PJ, Kelley WM (2008) Are attractive people rewarding? Sex differences in the neural substrates of facial attractiveness. J Cogn Neurosci 20: 941–951. 36. Lacey S, Hagtvedt H, Patrick VM, Anderson A, Stilla R, et al. (2010) Art for reward’s sake: visual art recruits the ventral striatum. Neuroimage 55: 420–433. 37. O’Doherty J, Winston J, Critchley H, Perrett D, Burt DM, et al. (2003) Beauty in a smile: the role of medial orbitofrontal cortex in facial attractive- ness. Neuropsychologia 41: 147–155. 38. Tsukiura T, Cabeza R (2010) Remembering beauty: roles of orbitofrontal and hippocampal regions in successful memory encoding of attrac- tive faces. Neuroimage 54: 653–660. 39. Feinberg TE, Venneri A, Simone AM, Fan Y, Northoff G (2009) The neuroanatomy of asoma- tognosia and somatoparaphrenia. J Neurol Neu- rosurg Psychiatry 81: 276–281. 40. Beer JS, Lombardo MV, Bhanji JP (2009) Roles of medial prefrontal cortex and orbitofrontal cortex in self-evaluation. J Cogn Neurosci 22: 2108–2119. 41. Miller BL, Cummings J, Mishkin F, Boone K, Prince F, et al. (1998) Emergence of artistic talent in frontotemporal dementia. Neurology 51: 978– 982. 42. Chatterjee A (2004) The neuropsychology of visual artistic production. Neuropsychologia 42: 1568–1583. 43. Poldrack RA (2006) Can cognitive processes be inferred from neuroimaging data? Trends Cogn Sci 10: 59–63. 44. Vartanian O, Goel V (2004) Neuroanatomical correlates of aesthetic preference for paintings. Neuroreport 15: 893–897. 45. Schultz W (2011) Potential vulnerabilities of neuronal reward, risk, and decision mechanisms to addictive drugs. Neuron 69: 603–617. 46. Schultz W, O’Neill M, Tobler PN, Kobayashi S (2011) Neuronal signals for reward risk in frontal cortex. Ann N Y Acad Sci 1239: 109–117. 47. Shenhav A, Greene JD (2010) Moral judgments recruit domain-general valuation mechanisms to integrate representations of probability and mag- nitude. Neuron 67: 667–677. 48. Padoa-Schioppa C (2009) Range-adapting repre- sentation of economic value in the orbitofrontal cortex. J Neurosci 29: 14004–14014. PLOS Biology | www.plosbiology.org 5 March 2013 | Volume 11 | Issue 3 | e1001504 GH Jr (1998) Is beauty in the eye of the beholder? Nature 396: 321–322. 22. Marlowe F, Wetsman A (2001) Preferred waist-to- hip ratio and ecology. Pers Individ Dif 30: 481– 489. 23. Tyler CW (1998) Painters centre one eye in portraits. Nature 392: 877. 24. Ishizu T, Zeki S (2011) Toward a brain-based theory of beauty. PLoS ONE 6: e21852. doi:10.1371/journal.pone.0021852 25. Vul E, Pashler H (2012) Voodoo and circularity errors. Neuroimage 62: 945–948. 26. Kriegeskorte N, Simmons WK, Bellgowan PS, Baker CI (2009) Circular analysis in systems neuroscience: the dangers of double dipping. Nat Neurosci 12: 535–540. 27. Blood AJ, Zatorre RJ (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. 28. Kable JW, Glimcher PW (2009) The neurobiol- ogy of decision: consensus and controversy. Neuron 63: 733–745. 29. Padoa-Schioppa C, Cai X (2011) The orbitofron- tal cortex and the computation of subjective value: consolidated concepts and new perspec- tives. Ann N Y Acad Sci 1239: 130–137. 30. Plassmann H, O’Doherty J, Rangel A (2007) Orbitofrontal cortex encodes willingness to pay in everyday economic transactions. J Neurosci 27: 9984–9988. 31. Mehta PH, Beer J (2009) Neural mechanisms of the testosterone-aggression relation: the role of orbitofrontal cortex. J Cogn Neurosci 22: 2357– 2368. 32. Hughes BL, Beer JS (2012) Medial orbitofrontal cortex is associated with shifting decision thresh- olds in self-serving cognition. Neuroimage 61: 889–898. 33. Bhanji JP, Beer JS (2011) Unpacking the neural associations of emotion and judgment in emotion- congruent judgment. Soc Cogn Affect Neurosci 7: 348–356. 34. Tsukiura T, Cabeza R (2010) Shared brain activity for aesthetic and moral judgments: implications for the Beauty-is-Good stereotype. Soc Cogn Affect Neurosci 6: 138–148. 35. Cloutier J, Heatherton TF, Whalen PJ, Kelley WM (2008) Are attractive people rewarding? Sex differences in the neural substrates of facial attractiveness. J Cogn Neurosci 20: 941–951. 36. Lacey S, Hagtvedt H, Patrick VM, Anderson A, Stilla R, et al. (2010) Art for reward’s sake: visual art recruits the ventral striatum. Neuroimage 55: 420–433. 37. O’Doherty J, Winston J, Critchley H, Perrett D, Burt DM, et al. (2003) Beauty in a smile: the role of medial orbitofrontal cortex in facial attractive- ness. Neuropsychologia 41: 147–155. 38. Tsukiura T, Cabeza R (2010) Remembering beauty: roles of orbitofrontal and hippocampal regions in successful memory encoding of attrac- tive faces. Neuroimage 54: 653–660. 39. Feinberg TE, Venneri A, Simone AM, Fan Y, Northoff G (2009) The neuroanatomy of asoma- tognosia and somatoparaphrenia. J Neurol Neu- rosurg Psychiatry 81: 276–281. 40. Beer JS, Lombardo MV, Bhanji JP (2009) Roles of medial prefrontal cortex and orbitofrontal cortex in self-evaluation. J Cogn Neurosci 22: 2108–2119. 41. Miller BL, Cummings J, Mishkin F, Boone K, Prince F, et al. (1998) Emergence of artistic talent in frontotemporal dementia. Neurology 51: 978– 982. 42. Chatterjee A (2004) The neuropsychology of visual artistic production. Neuropsychologia 42: 1568–1583. 43. Poldrack RA (2006) Can cognitive processes be inferred from neuroimaging data? Trends Cogn Sci 10: 59–63. 44. Vartanian O, Goel V (2004) Neuroanatomical correlates of aesthetic preference for paintings. Neuroreport 15: 893–897. 45. Schultz W (2011) Potential vulnera