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John Cacioppo, Penny Visser, Cynthia Pickett. Social Neuroscience

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edited by John T. Cacioppo, Penny S. Visser and Cynthia L. PickettSocial people about thinking
Social
Social Neuroscience Series John T. Cacioppo and Gary C. Series Editorial Board Ralph Adolphs, C. Sue Carter, Richard J. Davidson, Martha K.McClintock, Bruce S. McEwen, Michael J. Meaney, Oaniel L. Schacter, Esther M.Sternberg, Steve S. Suomi, and Shelley E. Tayloredited by John T. Cacioppo, Gary G. Berntson, Ralph Adolphs, C. Sue Carter, Richard J.Davidson, Martha K. McClintock, Bruce S. McEwen, Michael J. Meaney, Daniel L. Esther M. Sternberg, Steve S. Suomi, and Shelley E. edited by John T. Cacioppo and Gary G. Berntsonedited by John T. Cacioppo, Penny S. Visser, and Cynthia L.
Social People Thinking about Thinking edited by John T. Cacioppo, Penny S. Visser, and Cynthia L PA Bradford BookThe MIT PressCambridge, London, E
2006 Massachusetts Institute of All rights reserved. No part of this book may be reproduced in any form by any electronic ofmechanical means (including photocopying, recording, or information storage and retrieval)without permission in writing from the MIT Press books may be purchased at special quantity discounts for business or sales use. For information, please email special_sales@mitpress.mit.edu or write to Special Sales ment, The MIT Press, 55 Hayward Street, Cambridge, MA This book was set in Stone Sans and Stone Serif by SNP Best-set Typesetter Ltd., Hong Kong, andwas printed and bound in the United States of Library of Congress n-Publication Social neuroscience : people thinking about thinking people / edited by John T. Cacioppo,Penny S. Visser, and Cynthia L. p. cm.(Social neuroscience "A Bradford Includes bibliographical references and ISBN-13 978-0-262-03335-0 (hbk. : alk. ISBN 0-262-03335-6 (hbk.: alk. paper)1. Cognitive neuroscience. 2. Neuropsychology. 3. Social psychology. I. Cacioppo, John T. II.Visser, Penny S. III. Pickett, Cynthia L. IV. QP360.5.S636 200610 98765432
Foreword viiMahzarin R. Preface xi1 Reasoning about Brains 1Gary G. Bemtson2 Neurological Substrates of Emotional and Social Intelligence: Evidence Patients with Focal Brain Lesions 13Antoine Bechara and Reuven 3 Neural Substrates of Self-Awareness 41Debra A. Cusnard4 Thinking about Others: The Neural Substrates of Social Cognition 63Jason P. Mitchell, Malia F. Mason, C. Neil Macrae, and Mahzarin R. 5 Four Brain Regions for One Theory of Mind? 83Rebecca 6 Theory of Mind and the Evolution of Social Intelligence 103Valerie E. Stone7 Investigating Cortical Mechanisms of Language Processing in Social Context 131Howard C. Nusbaum and Steven L. Small8 Orbitofrontal Cortex and Social Regulation 153Jennifer S. 9 A Pain by any other Name (Rejection, Exclusion, Ostracism) still Hurts the Same:The Role of Dorsal Anterior Cingulate Cortex in Social and Physical Pain 167Matthew D. Lieberman and Naomi I.
10 The Social Neuroscience of Stereotyping and Prejudice: Using Event-RelatedBrain Potentials to Study Social Perception 189Tiffany A. Ito, Geoffrey R. Urland, Eve Willadsen-Jensen, and Joshua 11 Race and Emotion: Insights from a Social Neuroscience Perspective 209Nalini Ambady, Joan Y. Chiao, Pearl Chiu, and Patricia 12 Animal Models of Human Attitudes: Integration across Behavioral, and Social Neuroscience 229Elizabeth A. Phelps and Mahzarin R. 13 Characterizing the Functional Architecture of Affect Regulation: Answers and Outstanding Questions 245Kevin N. Ochsner14 What Is Special about Social Cognition? 269Ralph 15 Social Neuroscience: A Perspective 287Marcus E. Contributors 297Index 299
Foreword: Sciences Newest Brain Child, Social NeuroscienceOne can do little about the timing of ones birth. So it is a matter of unearned goodfortune to be present when the evolution of ones science happens to be in a momentthe sidelines but a long way away, the rumblings and grumblings of the shaping ofelder and younger, and the uncharted territory ahead.spot suitable for a rest. John T. Cacioppo is such a person, and he predates the development was remarkable in that he was self-taught and drew from every strandthat linked behavior, brain, and social world. John is able to stack level-of-analysisupon level-of-analysis, shunning none in favor of another. He is both a superb reductionist and a committed integrationist. To him nothing is more satisfactory thanto see it all line up. Johns passion for social neuroscience, his prescience regarding its inevitability, and his pulling it all toward the center so that nothing topples off the path of discovery are among the reasons that many of us are able to be
sometimes to draw in others and sometimes to be complained about by passersby. Westart of this century. Younger entrants to the eld are Stakanovites who are also out-they are bound by a desire to understand social animals and to do so by observingEach of us performs countless numbers of such mental acts every day, and social neu-knowledge and social cognition broadly, self-awareness, emotion regulation, and atti-tudes, beliefs, and memory involving social groups. In this volume, Cacioppo, PennyS. Visser, and Cynthia L. Pickett have paid special attention to gathering diverseher research happened to be human faces. Now, already, stronger credentials wouldbe necessary. The fundamental questions that draw social neuroscientists togetherperception and have no interest whatsoever in social cognition. Instead, social neuro-studied by measuring mental and brain function. If a brain imaging study, for instance,looks somewhat infantile compared with traditional behavioral ones, theres a reason.breakthroughs that will allow faster advances than are currently visible. Everybody
Amidst the hullabaloo, a few matters deserve to be mentioned, even if they are no-of work that become relevant have no boundary; many elds become of interestgoing to be to be a lumper. This eld will be kind to those who are naturals, or readyleverage and even create new domains of study. However, unlike the past, going deepcannot be a solitary journey. Collaborations may look more like baseball teams, andmay even bring a sense of unease, because every single piece of knowledge is not goingto be equally accessible to every expert on the team. Psychologists, neuroscientists,and physicists will have to develop appropriate levels of faith in the others expertiseas well as posing challenges to anothers familiar assumptions.Third, developing respect for disciplines not ones own should not be underesti-The brain, as an object of study, belongs to anybody who has one. It would be silly
still ll popular books (and journal articles to a lesser extent) will die away.wiping off the cobwebs of twentieth-century simplicities about human nature and
half of the twentieth century was the computer, a solitary device with massive infor-mation-processing capacities. At the dawn of the twenty-rst century, this metaphorseems dated. Computers today are massively interconnected devices with capacitiesthat extend far beyond the resident hardware and software of a solitary computer. Itless broadband interconnectivity to humans for millennia. Just as computers haveware of a single computer, the human brain has come to be recognized as havingthrough, but that extend far beyond, a solitary brain.either. Human infants will not survive to contribute to the gene pool unless theyability to think, to develop and use tools, and to work together. It may be that the, however, derives not simply from an individuals repro-ductive success but more critically from the success of ones children to reproduce.Hunter-gatherers who, in times of famine or danger, chose not to return to share theirfood or protection may have survived to hunt and reproduce again, but their genesspring who survived to contribute to the human gene pool. In short, humans have
NeuropsychologiaJournal of Personality and Social Psy-Biological PsychiatryJournal of Cogni-tive NeuroscienceNeuroimageAnnual Review of NeuroscienceCurrent Opinion in NeurobiologyTrends inmation processing; more specically, on mechanisms underlying people thinkingabout thinking people. Recent work in evolutionary biology and in social neurosciencewithin a given disciplinary perspective, when among the strengths of social neuro-science is its multidisciplinary approach. This book therefore draws heavily on thedisciplines and levels of analysis. Accordingly, the volume draws on research usingthis literature. The goal, however, was to provide an illustrative rather than exhaus-tive treatment of the topic of people thinking about thinking people. We hope this
Work on this book was made possible by funding from the National Science Foun-the social, cognitive, and neurosciences for more than a quarter-century, and we areindebted for its support. We also thank Tom Stone, the cognitive science, linguistics,philosophy, and psychology editor at MIT Press, for his efforts and backing; and toFinally, we thank Gary Berntson, Marc Raichle, Gn Semin, Wendi Gardner, and John
brain circuits and mechanisms underlying social reasoning, and the extent to whichthese circuits and mechanisms are specialized and/or specic for social reasoning.include but are not limited to person perception, social categorization and reasoning,tion. The special (brain) mechanisms debate has a long and enduring history, includ-ing the possibility of specialized perceptual and cognitive mechanisms for socialother chimpanzee vocalizations that were matched for intensity, duration, and fre-quency composition. Moreover, infant orangutans did not display this response to thetory reaction to some simple physical property of the stimulus per se. These ndingstionally endowed through evolutionary history, and may play an important role insocial perception and evaluative judgment. The example seems tting in the currentcontext, as Premack and Woodruff (1978) rst introduced the construct theory ofAnother example of the special nature of social perception comes from studies of1Reasoning about BrainsGary G. Berntson
of the underlying mechanisms of this stress-disease link. Unfortunately, standard lab-oratory stressors such as restraint and shock are ineffective in triggering HSV1 reacti-vation in mice. Research in our laboratory, however, discovered that reactivation couldresponse comparable with that of restraint stress. Rather, it appeared to have some-ing. The question remains, however, as to whether such selective effects imply specialmechanisms. This is a topic to which we will return shortly.Moreover, they are not properties of neuronal circuits or even the outputs of these circuits. Functions are at least one step removed from outputs, as they represent the specic neuronal circuits. It is in this sense that we use the construct of functionalThe history of brain localization is a colorful one, framed in the extreme by theclassic contest of the nineteenth century between the ultra-localizationist and phre-ist view, and Friedrich Goltz stridently challenging that position (see Phillips et al.,1984; Young, 1968). The controversy extended well into the twentieth century, withGary G. Berntson
in the contemporary literature, but developments and insights during the latter halfof the twentieth century led to partial resolution of this issue that, as is often the case,connected circuits () could underlie memory anddegree of functional specicity, exist within sensory and motor systems. On the otheroperations mediated by distributed, interacting cortical-subcortical circuits. This is notto say that these circuits are nonspecic or equipotential, just that they do not havea punctuate cortical representation (Sarter, Berntson, & Cacioppo, 1996; Uttal, 2003).Moreover, these distributed circuits may anatomically many circuits.category errortures, circuits, patterns of activity etc.). Because psychological constructs, for the The absence of such isomorphism limits logical interpretation (Cacioppo & Tassinary,behaviors such as eating, drinking, grooming, defensive responses, and predatory-likebehavior. Distribution elds for eliciting such diverse behaviors often overlapped,however, and some electrode loci could evoke several behaviors depending on theneuroanatomical specicity in motivational neurosubstrates (Valenstein, 1975), a
like responses, predatory-like stalking attack, or both. These responses are behaviorallyDespite this overlap, however, the distributions were doubly dissociated in thedigitated circuits may be represented within the central area of overlap. A stochasticGary G. Berntsonelectrode points yielding defensive threat and predatory attacklike responses in a hypothalamic-relevant area which is expanded to the right. Observed probabilities of threat [ved probabilities of threat [p(Attackobs)] are illustrated, together with the probability of both threat and attackresponses elicited by the same electrode [p(ThreatAttackobs)]. The latter compares closely withthe expected probability [p(ThreatAttackexp)] based on the joint occurrence of independentevents. Results suggest an overlapping distribution of specic differentiated behavioral systems.Results further show that stimulation of a single locus can induce more than one response andthat stimulation of divergent points within a distribution eld can induce the same behavior.
circuits were involved there should be a statistical independence among the behav-response, given the other, should be greater than the overall probability of thatAs illustrated in gure 1.1, results support statistical independence of these circuits.0.25). Within the overlap area, the observed probability of loci yieldingfrom imaging studies (for more complete discussions see Cacioppo, Tassinary, &distributed circuitsdifferent neural activations can yield the same behavioralcurrently. To state the converse, to the extent to which functional systems overlap,the same pattern of brain activation across contexts and occasions (within the limits of res-olution) does not necessarily imply the same behavioral process. tive and social neuroscience often begins with an observation of the following sort:
(Cacioppo et al., 2000). We return to this specialcase below.Finally, another inherent complication arises in the analysis of brain imaging results.isolate a more limited aspect of brain circuits involved in the social event. The veryTo some extent the substractive approach is necessitated by the difculty of dealingcephalic structures outside classic sensory and motor systems are implicated in socialprocesses. Moreover, there are subcortical systems that regulate all areas of the corti-cal mantle and thus affect social processes indirectly. An example is the basal fore-brain cortical cholinergic system, which supplies cholinergic innervation to all corticaltive operations (Berntson, Sarter, & Cacioppo, 2003; Sarter, Bruno, & Givens, 2003).This system is an especially important component of circuits underlying socialAlzheimers disease that is attributable in part to degeneration of the basal forebrainGary G. Berntson
So How Do We Proceed?neurology of social cognition and social behavior. Contributing to this challenge isfor attention, sensory and perceptual processing, motivation, memory access and asso-such as theory of mind and social regulation have complex mappings onto multipleneurobehavioral substrates. Moreover, category errors can arise when social constructslevel of organization. An integrative, interdisciplinary approach across levels of organ-are known to interact. Multilevel research will be necessary to form bridges amongdisciplines, calibrate and rene constructs to minimize category errors, and ultimately,achieve a truly interdisciplinary social neuroscience. A need remains for research with a more restricted focus, of course, and it is not necessary that all researchersBasic principles of multilevel analysis, which frame issues and organize research1.The principle of multiple determinismcorollary of proximitylevel analysis may be more straightforward for more proximal levels or organization.
2.The principle of reciprocal determinismone way. Consequently, although the multilevel perspective entails the ability to relate3.Finally, the principle of nonadditive determinismentrenched in the psychological literature. It became apparent quite early, however,nic relation to behavior. Duffy (1962) wrote, There appears to be some degree of gen-the psychological literature, partly because of its utility, despite overwhelming evi-but rather, that many ascending anatomical systems subserve arousal- or activation-like functions (Sarter, Bruno, & Berntson, 2003). These parallel systems are anatomi-target selection (Sarter, Bruno, & Berntson, 2003). In contrast, dopaminergic projec-Gary G. Berntson
distinct aspects of behavior. These ndings offer an excellent opportunity for multi-level research to clarify the nature and dimensions of arousal processes in a neuro-Overviewin which multilevel research efforts provide reciprocal benets to both biological andwith other interdisciplinary approaches. These include studies of the neuropsycho-netic brain stimulation, and even animal research in which more invasive brainTo return to the earlier question of whether social processes reect special brainmechanisms, it is simply too early to tell. We do not know enough about either socialments and conceptual relations be couched in the language of space. Rather, they willhave to incorporate fundamental underlying processes that subserve social psycho-Journal of Comparative and Physiological Psy-
Berntson, G. G. & Boysen, S. T. (1989). Specicity of the cardiac response to conspecic vocal-Pan troglodytesBehavioral NeuroscienceBerntson, G. G., Sarter, M., & Cacioppo, J. T. (2003). Ascending visceral regulation of corticalEuropean Journal of NeuroscienceCacioppo, J. T. & Berntson, G. G. (1992). Social psychological contributions to the decade of theCacioppo, J. T., Berntson, G. G., Sheridan, J. F., & McClintock, M. K. (2000). Multi-level integra-Cacioppo, J. T. & Tassinary, L. G. (1990). Inferring psychological signicance from physiologicalCacioppo, J. T., Tassinary, L. G., & Berntson, G. G. (2000). Psychophysiological science. In J. T.Cacioppo, L. G. Tassinary, & G. G. Berntson (Eds.), Duffy, E. (1962). Activation and Behavior.New York: Wiley.The Organization of Behavior: A Neuropsychological Theory.New York: Wiley.History of PsychologyMagoun, H. W. (1963). The Waking Brain.Padgett, D. A., Sheridan, J. F., Dorne, J., Berntson, G. G., Candelora, J., & Glaser, R. (1998). SocialProceedings of the NationalPhillips, C. G., Zeki, S., & Barlow, H. B. (1984). Localization of function in the cerebral cortex.Premack, D. & Woodruff, G. (1978). Does the chimpanzee have a theory of mind? Quan, N., Avitsur, R., Stark, J. L., He, L., Shah, M., Caligiuri, M., et al. (2001). Social stress increasesJournal of NeuroimmunologySarter, M., Berntson, G. G., & Cacioppo, J. T. (1996). Brain imaging and cognitive neuroscience:Towards strong inference in attributing function to structure. AmericanPsychologistGary G. Berntson
Sarter, M., Bruno, J. P., & Berntson, G. G. (2003). Reticular activating system. In L. Nadel (Ed.),, Vol. 3 (pp. 963967). London: Nature Publishing Group.Sarter, M., Bruno, J. P., & Givens, B. (2003). Attentional functions of cortical cholinergic inputs:What does it mean for learning and memory? Neurobiology of Learning and MemoryUttal, W. R. (2003). The New PhrenologyThe Limits of Localizing Cognitive Processes in the Brain.Valenstein, E. S. (1975). Brain stimulation and behavior control. Young, R. M. (1968). The functions of the brain: Gall to Ferrier (18081886),
Thorndike (1920) as the ability to perceive ones own and others internal states,social intelligence overlap with neural systems subserving the processing of emotionsto neural structures that subserve emotions and feelings, but not those that subserve2Neurological Substrates of Emotional and Social Intelligence:
ications range from changes in internal milieu and viscera that may not be percep-tible to an external observer (endocrine release, heart rate, smooth muscle contraction)to changes in the musculoskeletal system that may be obvious to an external observerResponses aimed at the brain lead to (1) central nervous system release of certainmodication of the state of somatosensory maps such as those of the insular cortex,and (3) modication of the transmission of signals from the body to somatosensoryregions. The ensemble of signals as mapped in somatosensory regions of the brainitself provide the essential ingredients for what is ultimately perceived as a feeling, aphenomenon perceptible to the individual in whom the signals are enacted. Thus emotions are what an outside observer can see or, at least, can measure. Feelings arewhat the individual senses or experiences subjectively.specic brain center. Rather, it is a function of a neural circuitry that involves severalbrain structures interconnected through several neural pathways. Subsequent work byPaul McLean provided further elaboration on the original Papez circuit by adding theDamasios description of neural systems that subserve emotions and feelings (1999,system as described originally. The functional anatomy of this system includes the tionally competent object is basically the object of ones emotion. It can be present
or recalled from memory. In neural terms, images related to the emotional object arerepresented in one or more of the brains sensory processing systems. Regardless ofavailable to a number of emotion-triggering sites elsewhere in the brain. Two of theseemotional object is recalled from memory. However, the general function is the same;To create an emotional state, the activity in triggering sites must be propagated tochanges in the bodys response to an emotional object. Signals from body states arelevel of visceral sensory nuclei in the brainstem. Representations of these body signalsalso form at the level of the insular cortex and lateral somatosensory cortices (SII androceptive signals to conscious perception (Damasio, 1999; Craig, 2002). There is someas subserving the mechanisms of emotions and conscious feelings. The following isfrequent and less intense than positive emotions (Damasio, 1999). Many laboratory
16Antoine Bechara and Reuven Bar-On
brainstem/ hypothalamus cortex cortex cortex the viscera THE WORLD THE BRAIN visceral motor visceral responses
brains sensory processing systems. This information, which can be derived from the environ-ment or recalled from memory, is made available to the amygdala and orbitofrontal cortex, whichemotion comprises endocrine and somatomotor responses as well. Visceral sensations reach
Substrates of Emotional and Social Intelligence17
resonance data and obtained with Brainvox. Circles highlight the region showing bilateral amyg-
(Tranel and Hyman, 1990; Adolphs et al., 1995).More specically, laboratory experiments reveal that the amygdala plays a criticalin two different ways: (1) in an impulsive or automatic way, which we referred to asprimary induction, and (2) in a reective or thoughtful way that we referred to as sec-ondary induction (Bechara et al., 2003).Primary inducers are stimuli or entities that are innate or learned to be pleasurableor aversive. Once they are present in the immediate environment, they automatically,quickly, and obligatorily elicit an emotional response. An example of primary induc-snake. Semantic information such as winning or losing a large sum of money, whichinstantly, automatically, and obligatorily elicits an emotional response, is anotherSecondary inducers are generated by recall of a personal or hypothetical emotionalevent (thoughts and memories about the primary inducer) that, when they arebrought to working memory, slowly and gradually begin to elicit an emotionalresponse. Examples of secondary inducers include the emotional response elicited bythe memory of encountering or being bitten by a snake, the memory or imaginationof gaining or losing a large sum of money, and recall or imagination of the death ofnecessary for triggering emotional states from primary inducers. It couples the featuresof primary inducers, which can be processed subliminally by way of the thalamus(LeDoux, 1996; Morris et al., 1999) or explicitly by early sensory and high-order-emotional state (feeling) associated with the inducer. This emotional state is evokedtor structures such as the ventral striatum, periacquenductal gray, and other brainstemtent stimuli such as snakes (Kluver & Bucy, 1939; Zola-Morgan et al., 1991; Aggleton,
1992), suggesting that the object of fear can no longer evoke a state of fear. AnimalOscar, or a lottery ticket can instantly, automatically, involuntarily, and obligatorilyelicit an emotional response. Emotional reactions to gains and losses of money, forexample, are learned responses because we were not born with them; however,through development and learning, they become automatic. We do not know howthis transfer occurs. However, we have presented evidence that patients with bilateralesting that humans tend automatically, involuntarily, and obligatorily to show a pleas-Oshercon, 2001).et al., 1997). However, the amygdala appears to respond only when stimuli are actu-the KluverBucy syndrome described in monkeys (Kluver & Bucy, 1939; Zola-Morgan
humans is much milder.Lesions of the Insular-Somatosensory Cortexsomatosensory, and adjacent cortices), especially on the right side, which demon-placidity. It is most commonly associated with right hemisphere lesions (as opposedto left), particularly inferior parietal cortices that include primary and secondary
in the right parietal region that includes the insula and somatosensory cortices (SII, SI). The left
somatosensory cortices and insula. The most frequent causes of this condition arestroke, glial tumors, and head injury.The classic example of anosognosia is paralysis on the left side of the body, withrst few days after onset. Within a few days or a week, these patients will begin tothey minimize the implications of the impairment. They might say, I am righthanded, I dont care if my left hand moves or not. In the chronic epoch (3 monthsabilities; however, defects in appreciation of acquired cognitive limitations may persistMany laboratory experiments identied problems processing emotional informationfailure to observe social convention. One example is Supreme Court Justice WilliamO. Douglas, who is described in Damasios Descartes ErrorMore specically, once emotional states from primary inducers are induced, signalssomatosensory cortices (insular-SII, SI cortices, cingulated cortices). After an emotionalstate has been triggered by a primary inducer and experienced at least once, a patterna specic primary inducer will then operate as a secondary inducer. Secondary induc-cic primary inducer and generate a weaker activation of the emotional state than ifit were triggered by an actual primary inducer. For example, imagining the loss of alarge sum of money (secondary inducer) reactivates the pattern of the emotional statebelonging to an actual experience of money loss (primary inducer). However, the emo-
Reception of body signals at the level of visceral sensory nuclei in the brainstemsignals at the level of the insular-somatosensory cortices and posterior cingulated cor-tices is perceived as a feeling. Evidence from functional neuroimaging studies suggestsplays a role in the generation of feelings from autobiographical memory. Support forthe idea that the insula and SII and SI cortices are necessary for feeling to occur is sup-ported by clinical observations in subjects with focal brain lesions (Berthier et al., 1988;Damasion, 1994, 1999). We also have preliminary evidence showing that when aprimary inducer, such as an aversive loud sound, induces a somatic response in normalobservations).lated, thus encompassing Brodmanns areas (BA) 25; lower 24 and 32; medial aspectsfunctioning. Previously well-adapted individuals become unable to observe social con-
in impulse control, disinhibition, and antisocial behavior. For instance, they may utterwithout regard to rules of decency.With more extensive damage, the patient may suffer a condition known as akineticmentary motor area. The condition is a combiation of mutism and akinesia. Lesionsbral artery territories, and in some cases, from rupture of aneurysms of the anteriorcommunication artery or anterior cerebral artery. It may also result from parasagittalbilateral lesions appears to differ only in relation to course of recovery: with unilat-
Figure 2.4
(although generally hypophonic). With extensive prompting, the patient may repeatProvided that the amygdala and insular-somatosensory cortices were normal duringdevelopment, emotional states associated with secondary inducers develop normally.Generating emotional states from secondary inducers depends on cortical circuitry incritical substrate in the neural system necessary for triggering emotional states fromsecondary inducers. It serves as a convergence-divergence zone, in which neuronensembles can couple (1) a certain category of event based on memory records in high-words, the VM cortex couples knowledge of secondary inducer events to emotionalSeveral studies support this notion. We conducted investigations using paradigmsfor evoking emotional states, namely, from the internal generation of images relatedto emotional situations (emotional imagery). We predicted that patients with VMsadness, fear, and anger. After a brief description of each memory was obtained, theand family disputes. However, they had difculty reexperiencing the actual emotionsof feeling the emotions. This was especially marked in the case of sadness (Tranel et
They showed activations or deactivations in the VM region during recall and imageryences, consistent with observations made in monkeys with orbital lesions (Butter etal., 1963, 1968; Butter & Snyder, 1972). The lesions produced a clear reduction inwas involved in the lesion. This suggestion is preliminary and we are still investigat-conditioning studies in animals showing that the VM cortex is not necessary forby a primary aversive unconditioned stimulus such as electric shock (Davis, 1992a, b;neural circuitry of emotions and feelings (gure 2.1) also experience disturbances inemotions and emotion regulation. One example is pathologic laughing and crying.This condition is also referred to as forced, or spasmodic, laughing and crying, or the syndrome of pseudobulbar palsy. It may occur after bilateral interruption of
nerves (motor V, VII, ambiguus, hypoglossal). Thus it appears that disinhibition or lossThe disorder is observed after strokes, amyotrophic lateral sclerosis, or multiple scle-in muscles supplied by corresponding cranial nerve nuclei. Most important is thatthese patients have inappropriate outbursts of laughter and crying: with the slightestthe opposite happens: the mere mention of the patients family or the sight of thedoctor provokes a spasm of uncontrollable crying. Of interest, most often patientsing: they cry, but they do not feel sad; they laugh, but they do not feel happy.condition is linked to the cerebellum (Parvizi et al., 2001). Specically, the conditionter or crying to the cognitive and situational context of a potential stimulus, becomeing in inadequate expression of the proper emotion (Parvizi et al., 2001). In any case,the corticobulbar and cerebellar views are not contradictory, except that they differ ina primary disturbance of feelings.duction of controlled laboratory studies designed to examine the effects of such brain-stem lesions on emotion, emotion regulation, and social behavior.ing emotions and observing social convention. In striking contrast, those with exten-sive damage in areas that spare the neural circuitry that is critical for emotions andfeelings, may express a variety of cognitive problems related to memory, language,perception, and so on. However, very seldom do they express improper social behav-
that subserve emotions and feelings, but not neural structures that subserve cold cog-right parietal cortex that involved the insular and surrounding somatosensory cor-in the parieto-occipital cortex on the left side. We note that these patients had damageAll patients were drawn from the Division of Cognitive Neurosciences Patient Registry at the University of Iowa. Those in the experimental group had stable focalsurgical removal of a meningioma. In the right insular-somatosensory region, damagepatients) or UrbachWeithe disease (1 patient). In the control group, the lesions weregroups were matched with respect to gender, age, level of education and handednessWe used three measures in examining these patients: the Bar-On EQ-i for assessingemotional intelligence; semistructured interviews for assessing social functioning; andstandard neuropsychological tests for assessing cognitive intelligence, perception,memory, and executive functioning.The Bar-On Emotional Quotient inventory (EQ-i) is a self-report measure of emotion-ally and socially intelligent behavior that provides an estimate of ones underlying
employs a 5-point response format ranging from very seldom or not true of me (1) tovery often true of me or true of me (5). The subjects responses render a total EQ scoredence, and self-actualization); (2) interpersonal EQ (empathy, social responsibility, andcontrol); (4) adaptability EQ (reality testing, exibility, and problem solving); and (5)an exaggerated positive or negative manner, respectively. This is an important psy-patients self-awareness of their acquired decits is limited (anosognosia).Raw scoresTable 2.1
Demographic Data on Participating Patients
Demographic dataControl groupExperimental groupZp level (2-tailed)Number1112Men47Women75Age (yrs)47.143.50.460.644Age range (yrs)24742163Education (yrs)14.613.70.850.398Right1012Left10
The Mann-Whitney U test was applied to compare average age and years of education between
social status were evaluated with a series of semistructured interviews and rating scales(Tranel et al., 2002). Briey, this evaluation entailed a comprehensive assessment ofwas unaware of the objectives and design of the study. For each social domain evalu-Table 2.2
EQ-i Subscales
EQ-i subscalesEI competencies measuredSelf-regardTo perceive, understand, and accept oneself accurately.Emotional self-awarenessTo be aware of and understand ones emotions.AssertivenessTo express ones emotion and oneself effectively andconstructively.IndependenceTo be self-reliant and free of emotional dependency on others.Self-actualizationTo strive to achieve personal goals and actualize ones potential.EmpathyTo be aware of and understand how others feel.Social responsibilityTo identify with ones social group and cooperate with others.Interpersonal relationshipTo establish mutually satisfying relationships and relate well toStress toleranceTo manage emotions effectively and constructively.Impulse controlTo control emotions effectively and constructively.Reality testingTo validate ones feeling and thinking objectively with reality.FlexibilityTo adapt and adjust ones feelings and thinking to newProblem solvingTo solve problems of a personal and interpersonal natureeffectively.OptimismTo be positive and look at the brighter side of life.
HappinessTo feel content with oneself, others, and life in general.
Cognitive Intelligence, Perception, Memory, and Executive FunctioningCognitive intelligence (IQ) was measured with the Wechsler Adult Intelligence Scale(WAIS-III). Perception was measured with the Benton Facial Discrimination Test andthe Benton Judgment of Line Orientation Test. Memory was measured with the ReyAuditory Verbal Learning Test (RAVLT), the Benton Visual Retention TestrevisedWisconsin Card-Sorting Test (WCST), the Trail-Making Test (TMT), and ControlledOral Word Association Test (COWA). Possible existence of psychopathology wasassessed with the Minnesota Multiphasic Personality Inventory (MMPI). More detailedThe results of these measures were intriguing. Tests of cognitive intelligence, perception, memory, and executive functioning revealed that patients in the The very small samples in both groups do not allow closer examination of differ-insular-somatosensory cortices, and amygdala). However, in light of the proposedearlier, one can speculate on how each subgroup might differ in terms of scores onspecic competencies of each subscale of the EQ-i if the samples were larger.
Table 2.3
Neuropsychological Test Scores for the Control Group (n
Neuropsychological testsControl groupExperimental groupZp level (2-tailed)WAIS-IIIFull IQ97.7105.31.170.241Verbal IQ99.2107.91.320.186Performance IQ95.7102.81.420.155Benton faces44.443.90.500.620Benton lines25.424.60.810.420MemoryWAIS-IIIDigit span11.010.80.160.869BVRTCorrect7.47.40.120.908Errors3.84.00.580.565RAVLTTrials 1 to 510.112.01.530.12730-minute recall8.19.30.460.648Recognition28.928.80.910.361Complex gure (Rey-O)Copy32.230.90.550.585Delay20.719.50.220.827Trail-making test A34.839.60.060.957Trail-making test B86.879.90.330.745Perseverative errors10.217.70.440.662Categories5.85.10.710.476COWA37.040.40.530.596
See text for abbreviations of these tests.
Table 2.4
MMPI Test Scores for the Control Group (n-11) and Experimental Group (n
Personality testsControl groupExperimental groupZp level (2-tailed)1 (Hs)57.357.60.340.7322 (D)53.063.01.260.2093 (Hy)64.357.60.690.4934 (Pd)58.761.91.030.3045 (Mf)52.755.00.570.5686 (Pa)50.357.10.920.3597 (Pt)49.361.31.380.1688 (Sc)52.063.31.480.1399 (Ma)54.353.90.230.8180 (Si)49.752.90.340.731
Hs, hypochondriasis; D, depression; Hy, conversion hysteria; Pd, psychopathic deviate; Mf, Table 2.5
(n 12)
Postmorbid changes inControl groupExperimental groupZp level (2-tailed)Employment01.1702.703.110.002Social functioning01.0002.302.840.005Interpersonal01.0002.402.870.004Social status01.0001.903.400.001Total social change04.1709.303.000.003
The MannWhitney U test was applied to compare scores for signicant differences.
Specically, given that the self-regard and emotional self-awareness competencies aremostly concerned with perception and awareness of ones emotions, we would antic-ipate that patients with right insular-somatosensory cortex damage (those withpendence, and self-actualization competencies involve expression of ones emotionsFinally, all the competencies measured by the interpersonal, stress-management,adaptability, and general mood subscales involve emotional and social competenciesTable 2.6
EQ-i scales ControlExperimentalZp level
(emotional intelligence)groupgroup(2-tailed)Total EQ101.1082.33.330.001100.0081.83.230.001Self-regard099.1083.82.400.016Emotional self-awareness100.9090.11.480.139Assertiveness103.6082.63.210.001Independence097.7087.31.580.115Self-actualization099.4086.82.250.024099.6091.61.360.175Empathy098.6089.81.240.216Social responsibility101.5095.31.140.254Interpersonal relationship098.8092.80.830.406Stress management EQ104.8089.12.620.009Stress tolerance100.1083.22.560.011Impulse control108.3096.92.130.033100.0086.32.280.023Reality testing099.8091.01.080.280Flexibility100.3086.82.380.017Problem solving100.6088.32.160.031099.9083.33.270.001Optimism099.0083.53.020.003Happiness100.9085.82.710.007
Emotional intelligence was assessed by the EQ-i, and the MannWhitney U test was applied to
It is important to note that the specic type of brain injury sustained by subjectsHowever, the EQ-i proved to be successful in detecting abnormal levels of emotionalquate construct validity when used with individuals who otherwise may be unawareeven lower without the EQ-is correction factor that automatically adjusts scale scoresreason or another.Only patients with injury to the neural circuitry subserving emotions and feelingsabsence of psychopathology. The experimental group had many disturbances in socialtional and social intelligence. Yet, no differences were observed between experimen-tal and control groups with respect to IQ, executive functioning, perception, memory,or signs of psychopathology. Similarly, the groups did not differ with respect to demo-(EQ) exist in spite of the fact that the two groups were well matched on demographicand EQ in the clinical sample. Together, these results suggest that complex cognitiveprocesses that subserve social competence, which appears to constitute a distinct form
Rather, these processes depend on known brain mechanisms related to expression ofcan have an ill effect on ones ability to cope effectively with daily demands.nitive intelligence. This was also conrmed by David Van Rooy (personal communi-cation, April 2003), who suggests that no more than 5 percent of the variance ofgence may not be strongly related, reinforcing the notion that they may be separatethis chapter and those reviewed by Van Rooy and colleagues (Van Rooy & Viswes-varan, in press; Van Rooy et al., submitted), but neurological evidence also suggestssuccinctly, the ventromedial prefrontal cortex appears to be governing basic aspectsfrontal cortex of nonclinical subjects who were engaged in various cognitive exercisesexercise personal judgment in decision making (Bar-On et al., 2003). Specically, on
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The capacity for self-awareness, or reexivity, as it was dubbed by William Jamesself to take itself as the object of its own view. Thus, it inheres aspects of both processrepresent ones own states (motoric, mental, emotional, perceptual, visceral), as onesa variety of disciplines. Yet, until recently, very little could be said about its neuralinformation in this regard. Historically, however, it has tended to deal with the sub-personal domain, addressing phenomena such as sensory and motor systems, andfunctional domains and processes such as language, memory, and attention, usingvarious information-processing models. Until recently, little emphasis was placed onimaging methodology, which has traditionally been grounded in the transformationrecently, however, certain technical advances in imaging methodology (stronger3Neural Substrates of Self-Awareness
however, not only illustrate evolutionary considerations for the development of anfor consciousness more generally, for example, facilitates the planning and rehearsalof intended behavior (Edelman & Tononi, 2000), with salient operations thus beingplanning and rehearsal depend on the ability to make perceptual discriminations onOf signicance, investigators from diverse research traditions independently con-crimination and evaluation in the service of guiding and optimizing certain behaviors.through a challenging physical environment. Various modern philosophers grapplingsized relationships between self-awareness and the subjects status as a discriminatingof his intentional action. Work in human developmental psychology also underscoresdevelopmental stages that parallel increasing competences in perceptual discrimina-In the social psychological literature, in particular (Carver, 2003), some debate hascontent, has on subsequent behavior and experience. It is generally viewed, however,
ciated with an evaluation of the agents behavior, whether that be relative to a per-sonal standard or one that the agent perceives as socially derived.Obviously, such self-focused attention and scrutiny do not always dominate the owof subjective experience, as they clearly do not have to. Much of our behavior, par-ticularly automatic or well-learned behavior, is in fact more effectively executedwithout our direct conscious supervision and evaluation of our ongoing performance.However, given our nature as highly developed social creatures with a signicantsocial learning, circumstances in which we are induced to become more self-aware areclearly ubiquitous. Notably, these circumstances include not only novel ones in whichwe must effect behaviors relatively naively, but also those in which we perceive our-selves as an object of social attention. It is in such circumstances, as well as in theanticipation of such circumstances, that self-awareness is likely to come to the foreinvolving reentrant connection of new perceptual categorizations with previouslyestablished categorizations in memory systems. Stabilization in an act of awarenessa process occurring along with the continual updating of these memory systems. This processing lag, however slight, which involves a bootstrapping of memory and continuing perception, thus inheres an element of assessment. In the case of highly evolved and socially rened capacity for self-awareness. Notably, this work illus-awareness. In particular, it provides data suggesting neural mechanisms that poten-Neural Substrates of Self-Awareness
Functional Imaging and Self-AwarenessGreenberg in Woolsey et al., 1996; Ueki, Linn, & Hossmann, 1988). The change in thevascular response to the neural activity, is a temporally delayed and dispersed reec-tion of underlying neural activity, beginning about two seconds after the latters onsetHistorically, initial interest in and interpretation of functional brain imaging studiesfocused on correlates of imaging signal increases(activations). However, researchersare now increasingly aware of the fact that signal decreases(deactivations) also occur.their origin and signicance (Tootell et al., 1998; Smith, Singh, & Greenlee, 2000;sively between or within perceptual systems, such that decreases in activity may appearin auditory or somatosensory cortices when engaging in a task involving visual per-Ghatan et al., 1998), or within portions of the somatosensory or visual systems whilethem (Drevets et al., 1995; Shmuel et al. 2002, 2001; Tootell, Tsao, & Vanduffel, 2003).
the processing of information expected to carry behavioral signicance by lteringout unattended sensory input.In addition to such signal decreases within sensory cortices, however, a special setof areas in association cortices have been observed to exhibit decreases in activity incontext of this discussion, and its signicance is detailed below. What are importantto note here, however, are physiological characteristics of imaging signal decreases.In particular, such signal decreases are not at all likely to be directly attributable tolocal inhibitory processes. This is because the latter are known to require energy andappear as likely to be associated with signal increases as excitatory processes are the neurovascular coupling involved in inhibitory processes are less well understood.Also, and most important, recent neurophysiological observations (Gold & Lauritzen,actual decreases in neuronal activity, particularly in local eld potentials, which reectNeural Substrates of Self-Awareness
Transverse images of positron emission tomographic data averaged across nine cognitive tasks (n
In the average adult human, the brain represents about 2 percent of total body weight,but accounts for about 20 percent of oxygen consumed and, thus, calories (energy)consumed by the body (Clark & Sokoloff, 1999). In relation to this very high rate ofterms often less than 5 percent of the ongoing metabolic activity of the brain in thatof the brains metabolic requirements, but evidence suggests that it may inhere func-tionally signicant signaling processes.Several lines of evidence strongly suggest this.spectroscopy (Sibson et al., 1998; Shulman, Hyder, & Rothman, 2001; Hyder,Rothman, & Shulman, 2002) that indicate that up to 75 percent to 80 percent of thetaneous electrical activity that does not bear an obvious relationship to specic sensoryor motor tasks (Arieli et al., 1996; McCormick, 1999; Tsodyks et al., 1999; Sanchez-Vives & McCormick, 2000; Shu, Hasenstaub, & McCormick, 2003), and is thought notsimply to represent noise (see Ferster, 1996; McCormick, 1999). This spontaneousin the physiologic reservoir of baseline activity, where oxidative metabolism is theprimary source of energy, a very efcient strategy has developed to manage largethe situation for activations, where glycolysis assumes an important role as a sourcecic, rapidly varying, and time-limited information processing required for immedi-One view suggests that it may serve a preparatory or facilitatory processing roleprocess is a necessary and expensive component of brain function but it is not asso-ciated with information processing with any inherent functionality. However, others
have proposed that information processing may, in fact, be a property of such a state(Tononi & Edelman, 1998; Shu et al., 2003), and here functional brain imaging withfrom the observation of the previously mentioned imaging signal decreases and, inparticular, those that occur in a specic set of association cortices during the per-Two observations using novel approaches to the study of the resting state (awake,regions that are among those regularly exhibiting the commonly observed decreasesNeural Substrates of Self-Awareness
of directed visual attention) varied inversely with alpha power (812Hz), whereas apower in a beta band (1723Hz), an EEG band thought to be associated with cogni-Self-AwarenessData that support the view that circumstances exist in which this set of brain
Figure 3.2Map of resting-state neural connectivity for the posterior cingulate cortex (PCC) (blue arrow indi-cates the PCC peak used for the analysis). Labels A H designate the signicant clusters in orderof descending tscore (Greicius et al., 2003). Note correspondence of this map with the patternof the common decreases (gure 3.1). See plate 3 for color version.
have been investigated by varying levels of anesthesia and, in one study, by means ofregional decreases in blood ow were observed that, in the cortex, were most promi-recovery from a vegetative state, which is dened as impaired awareness in the settingpatients recovery of consciousness largely consisted of medial parietal and lateral pari-controls, which returned to normal after patients recovery (Laureys et al., 2000). Inregions. Such studies usually involve the presentation of visual or auditory stimuliNeural Substrates of Self-Awareness
serving some process of explicit self-focus or self-awareness (and, in the case of otherones own mental state, with or without an affective component, or attributing Such episodic memory processes as well as explicit spatial perspective taking haverior cingulate, precuneus, and retrosplenial cortex (Maguire, 2001; Vogeley & Fink,oneself in large-scale space (Maguire, 2001). In a summary of work considering avariety of empirical data, including functional imaging data (Vogeley & Fink, 2003),A long history of neuropsychological lesion data indicates that this right lateral pari-etal region plays an important role in attending to aspects of ones personal space(Behrmann, 1999; Farrell & Robertson, 2000) as well as ones extensions (e.g., by meansof tool use) into that space (Ackroyd et al., 2002; Maravita, Spence, & Driver, 2003).2003; Farrer & Frith, 2002; Jackson & Decety, 2004).region (Craik et al., 1999; Kelley et al., 2002; Kjaer, Nowak, & Lou, 2002; Mitchell,
callosum or more dorsally. By contrast, to date, little evidence suggests that portionswith a low-level control condition such as visual xation). Rather, these ventral areason the part of the subject (Bush, Luu, & Posner, 2000; Nagai et al., 2004; Simpson et al., 2001a, b), so that the degree to which a behavior or task may be perceived ascommon set of decreases. This region, the insula, has been observed to be engaged incrosshair. With this maneuver, typical activations were seen in the visual cortex ashowever, there was evidence of several of the common decreases, in this case involv-also observed in the insula. It is thus possible that such decreases in the insula wereobserved in this study and are not observed more routinely in other studies showingcondition is not included as it was in our study.Neural Substrates of Self-Awareness
does not point to a unitary common system supporting the subjective experience ofa unied self (Gillihan & Farah, 2005). The position taken here, however, is that thepresence of such a system is suggested if ones conceptual view takes into accountwithin the purview of attention. When such information is salient, some of these areasated. Notably, the magnitudes of some of these regions decreases in activity have beenobserved to vary with such things as task difculty (McKiernan et al., 2003) and thesubjects emotional state (Simpson et al., 2001b).emotions, percepts, or other visceral experiences, in a particular behavior context. Theeffects of such factors as perceived task difculty and engendered emotional response,also suggest possible sources of differences in process and reactivity that individualsWe are, in fact, beginning to employ such a model in investigating effects of manip-ulations of objective self-awareness, as suggested by Duval and Wicklund (1972) andIn a pilot study, normal undergraduates underwent fMRI scanning while perform-consisted of informing the subjects that they were being observed and exposing themwas assessed by subjects self-report (verbal thought listing) (Cacioppo, Von Hippel, &
appearance and/or behavior. Subjects self-reports included content such as wonder-consciousness and were affected in their performance, however. Also, of particularinterest were the preliminary imaging ndings in those expressing the most self-Neural Substrates of Self-Awareness
camera on to that with camera off), one observes increased activity in portions of medial pre-et al., 1997) and social pain (Eisenberger, Lieberman, & Williams, 2003), as well as in the leftdorsolateral prefrontal cortex (typically associated with aspects of working memory) and left
recruit information-processing resources in the service of attention to more proximaldetails of ones performance). When personally relevant phenomena (interoceptivesense-data, ones orientation in space, ones mental states, episodic memories, personalhowever, such targeting typically results in a dissociation of the activity in these brainregions from each other, and either little or no deactivation or actual activation ofunclear. I also contend that many of these data, particularly those regarding thestate, underscore the dynamic character of self-awarenessawareness of ourselves (or,rather, a representation of some aspect of our so-called selves)often only eetinglyin the cognitive foreground, depending on the attentional resources that we are moti-vated to devote to aspects of our selves in any behavioral circumstance.Social psychologists have noted that, under certain circumstances, we may beencouraged or cued to devote more attentional resources to aspects of our selves current behavior and desired goal states. Clearly, however, competition between moti-formance may, at other times, be adjudicated smoothly and yield the ready
of self-awareness in any such circumstances, remain to be elucidated. Such details andadaptive challenges of many demanding personal and social circumstances.1.For purposes of clarity, it is useful to dene more precisely what we mean by a physiologicalFundamental to this notion of a physiological baseline is the observation that, in normal adultthe ratio of oxygen consumption to oxygen delivery. This ratio (sumption between gray matter and white matter, as well as among gray matter areas.In contrast, activations are characterized by a local fall in the average resting OEF, which resultswith eyes closed. It is thus constituted by an absence of activation in terms of an average OEF.equilibrium exists between local metabolic requirements necessary to sustain a continuing levelof neural activity and blood ow in that region. We have suggested that this equilibrium stateConsequently, areas with a reduction in this equilibrium OEF are regarded as activated (neuralNeural Substrates of Self-Awareness
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harnassed arbitrary symbols into languages capable of expressing an innite numberior, namely, social behavior. Although obvious examples of sociability are to be foundGibson, 1990; Ristau, 1991). Moreover, many of these abilities, such as recognizing(even when their beliefs conict with ones own), do not appear to have ready homo-logues among other animals (Gallup, 1985; Povinelli, Parks, & Novak, 1991; Tomasellohas prompted some observers to suggest that social cognition represents the primaryfocus of evolutionary change in humans (Kamil, 2004; Tomasello, 1999).Recently, researchers in the neurosciences have turned their attention to under-4Thinking about Others: The Neural Substrates of Social CognitionJason P. Mitchell, Malia F. Mason, C. Neil Macrae, and Mahzarin R. Banaji
2001; Blakemore, Winston, & Frith, 2004; Ostrom, 1984). As Blakemore et al. (2004)asked, are the general cognitive processes involved in perception, language, memory...sufcient to explain social competence or, over and above thesesupported by observations that the cognitive challenges posed by social interactionthat are not generally required in the rest of everyday life. For example, appropriateings, and beliefs underlying that individuals behavior (Baron-Cohen, 1995), whilestable dispositional characteristics (personalities) that inuence their actions. Finally,order to act in a socially appropriate manner as well as to manipulate the othersAt a minimum, these informal observations of the special problems posed by sociala distinct set of mental processes. How, then, does one proceed to gather evidence insupport of this possibility? Within several other elds of psychology, neuroimagingand neuropsychological research have proved critical to the resolution of similar the-oretical debates. In particular, these techniques have contributed signicantly tothe relation between implicit and explicit memory (whether implicit memory shouldbe considered a degraded form of explicit memory or the product of an entirely dif-ferent memory system) persisted even after years of accumulated behavioral dataobservations of patients with neuropsychological disorders (Gabrieli et al., 1995), andneuroimaging (see Schacter & Buckner, 1998) showed that the basis of implicitmemory was both neuroanatomically and functionally distinct from explicit memory.J. P. Mitchell and colleagues
In a different controversy, neuroimaging data were crucial in showing that visualimagery relies on the very same neural mechanisms as actual visual perceptionaccount that imagery relies on nonsensory, propositional knowledge (Pylyshyn, 1981,nition relies on its own set of mental processes or instead piggybacks on other, moregeneral processes of memory, inference, executive planning, and so on. By demon-science research has generally supported a view of social cognition as distinct fromcognition is the medial prefrontal cortex (PFC), although research also suggests thatof stimulus. Whereas peoples behaviors are understood through consideration of their underlying motivations and feelings (intellectual curiosity, teenage angst),objects such as jackhammers, avocados, and Jeepsindeed everything other thanpeopleare governed by external forces. Because appropriate social behavior is pred-Until recently, studies of how the brain represents semantic knowledge focused onwhat unexpected, nding from this research is that the brain appears to organize sometypes of semantic knowledge in a category-specic manner (Warrington, 1975). Thatis, knowledge about various kinds of objects (animals, tools) appears to be subservedorganizing principles underlying such category specicity, several inuential theoriessuggested that the brains semantic representations of a class of object center around
instance, because most tools are dened by their function and not some arbitrary phys-ical property, such as color, motor regionssuch as left premotor cortexare involvedbe represented by brain regions involved in the visual perception of animate objectsHaxby, & Martin, 1999). Moreover, recent work suggests that regions of the motorthrow,or (Hauk, Johnsrude, & Pulvermuller, 2004).Is semantic knowledge about other people also represented in such a category-of having mental states, do discrete brain regions subserve knowledge about peopleas mental agents? Alternately, could our understanding about the characteristics ofother people simply rely on the same brain regions known to subserve semantic pro-cessing more generally (perceptual and functional representations)?of other people in a discrete manner. In an initial study, Mitchell, Heatherton, and(denoted by common American forenames, e.g., John, Mary), fruits (banana, grape),previously implicated in object-knowledge tasks, specically, left-lateralized infer-sumably, a concomitantly different set of cognitive processes, than did similar deci-J. P. Mitchell and colleagues
While suggesting that distinct neural representations may subserve knowledge aboutthese representations. First, do observed functional differences between person anddescriptors. Second, do these observed differences extend to knowledge about the psy-To examine directly the extent to which brain regions observed in our earlier worktargetstheir potential psychological states (e.g., curious, frightened, angry) or theirunobservable physical parts (e.g., lung, heart, liver). An equal number of words could
systems that subserve semantic knowledge, object judgments engaged an extensive region of leftin qualitatively distinct regions of the prefrontal cortex, specically, medial PFC (blue-greenscale). T-maps from comparisons of object and person judgments are overlaid on coronal slices
not serve as potential descriptors (e.g., onic, metallic) and parts (nozzle, rudder). Critically, the words that could be used to describe people were pretested to be equallyresponse to each of these four types of trials, we observed that activity in the medialresponding to parts trials). However, the medial PFC did appear to generalize towith mental imagery (occipital and parahippocampal gyri), identical judgments aboutpeople yielded activity in the medial PFC. Together, these data suggest that activity insome brain regions, such as the medial PFC, specically subserves social-cognitive rep-Arguably the most important social-cognitive challenge is understanding the forcesthat govern other peoples behavior. Unlike inanimate objects, the behavior of peoplecan often be attributed to unobservable mental states. According to Dennett (1987),perceivers understand other people and predict how they will act by adopting theA fair proportion of neuroimaging and neuropsychological research on social cog-nition has focused on understanding brain mechanisms that subserve the capability J. P. Mitchell and colleagues
which perceivers must infer the mental states of others. For example, in some of theas well as stories that instead required understanding physical causality. Similarly, of the characters in them. Finally, more recent work has had participants playing interactive games that require second-guessing ones opponent, such as the childrensgame rock, paper, scissors (Gallagher et al., 2002), and compared activations whensubjects thought they were playing against a human opponent versus a computer.mind of another person. In each of these studies, greater activity was observed ininformation about a persons stable, idiosyncratic characteristics. For better or worse,
form theories of their stable qualities. Decades of research in social psychology suggestimportant additional inuences on behavior, such as situational constraints (Gilbertpersons behavior, as well as our tendency to explain others behaviors in terms oflate 1970s, researchers began examining mechanisms underlying perceivers abilitiesWorth, 1989; Hamilton, Katz, & Leirer, 1980; Hastie & Kumar, 1979; Srull & Wyer,1989; Wyer, Bodenhausen, & Srull, 1984). Participants were typically given informa-asked to memorize the information. Subsequently, participants were asked to retrieveall the information that had been presented. Researchers observed a number of intrigu-ing dissociations in memory performance between the social-cognitive (impressionparticipants memory was typically better after impression formation than after inten-patternsof memory performance differed across the twoMoreover, impression formation often led to increased memory for information thatwas inconsistent with participants expectations of the targets personality (Hastie &Kumar, 1979; Srull, 1981). For example, if a target was rst described as honest, infor-the participant was trying to form an impression of the person (Hartwick, 1979; Hastie& Kumar, 1979).In making sense of such differences, researchers generally suggested that social-cognitive processing prompts deeper, more elaborative encoding of the sort that generally supports episodic memory (Craik & Lockhart, 1972), such as generation ofJ. P. Mitchell and colleagues
item associations (Hastie & Kumar, 1979; Srull, 1981). However, a second possibilityon these issues by distinguishing between the circumstances under which (1) pro-Neuroimaging research suggest that impression formation does indeed prompt dis-tinct kinds of cognitive processing (Mitchell, Macrae, & Banaji, 2004). In one study,the information, the cognitive processes engaged during memorization may vary fromunderconstrained for event-related neuroimaging.) Subsequently, participants per-formed an associative memory task during which they were asked to match statementsmation and sequencing; specically, greater activation for impressionformationalongpression formation was very different from that during sequencing, brain differencesbetween the tasks are somewhat unsurprising. However, the design of this experimentof subsequent memory. Specically, encoding trials were retroactively conditionalizedformation, sequencing) as well as subsequent memory success; that is, whether an
hits than for subsequent misses. Of importance, no signicant difference was observedate between sequencinghitsandsequencingmissespart of the sequencing task, subsequent memory success was correlated only withence was observed between impression hits and impression misses in this regionwith subsequent memory for sequencing, but not for impression formation. ByJ. P. Mitchell and colleagues
Dorsomedial PFC
-0.4 -0.2 Impression Sequencing ImpressionSequencing
subsequent memory success were observed as a function of the orienting task performed duringwas observed between hits and misses for sequencing trials (gray solid and striped bars) in thisfor subsequent hits than misses, and no difference was observed in this region for impression
memory differences after social and nonsocial tasks result from the operation of theseHowever, just as for research on the neural systems that subserve semantic knowl-object. To address this possibility, a follow-up study examined the functional neu-of statements that ostensibly described an associated target. However, only half thesequencing for person trials only), a very similar region ofdorsomedial PFC was observed, providing a direct replication of earlier ndings.However, of critical interest in this experiment was whether or not the medial PFCof the targets. To address this question, the neural response to forming an impressionspecically subserves social-cognitive processing, results revealed a weak response inthe response observed for the sequencing task. In other words, although the dorso-necessary for appropriate social behavior, the ultimate output of a system for socialcognition must be the direct guidance of such behavior. As Fiske (1992) pointed out,anothers mind), but is unable to act appropriately based on that information, will
ior, including the selection and initiation of appropriate actions (is it permissible to(is it a good idea to snort derisively at the search committee members uninformedHowever, relative to the amount of extant work on the intrapersonal aspects of socialcognition, little neuroscience research has examined the demands of social behavior. This state of affairs is due at least in part to the strict contraints ofbehavior has tended to come from research in patients with neuropsychological dis-selective failure on theory of mind tasks (including second-order false belief and fauxto the attention of neurologists because of marked changes in social behavior, includ-ing personality changes, lack of empathy, and inappropriate social interactions, noto hold a job, lost his life savings in a series of bad business ventures, and was divorcedand divorced a second time.and medial PFC, Elliot had unimpaired intelligence, language, and working memory.J. P. Mitchell and colleagues
The distinctiveness of social behavior is also underscored by research on the autis-of the disorder, one characteristic of autism is a profound disengagement from theinterest, a disorder known as Williams syndrome appears in many ways to present thebut are profoundly impaired in nonsocial domains (Tager-Flusberg, Boshart, & Baron-In another interesting line of research, patients with frontal variant frontotemporaldementia (FTD) were examined on a series of mental tasks of varying degrees of dif-culty (Gregory et al., 2002). FTD is a progressive disorder that results in degenerationpatients with lesions to medial PFC, including personality changes, lack of empathy,and socially inappropriate behavior. Meta-analytic procedures over large numbers ofneural degeneration is most severe in a circumscribed region of PFC highly similar tomedial regions observed in neuroimaging studies of social cognition. Gregory et al.compared performance of such patients with that of individuals with Alzheimersability to mentalize about the minds of other people. Future research is necessary toexamine whether separate systems exist for such guidance of social behavior, or
mental states and behavior of other people? One inuential theory suggests thatknowledge of ones own mind can be used successfully to help infer the mind oftheory, proposes that one valuable source of information about the thoughts, feelings,in anothers mind can certainly be gleaned from a variety of sources, (emotionalA good deal of overlap appears to exist between brain regions that subserve thinkingabout other people and those that subserve thinking about oneself. Specically, in addi-ponent of tasks that require participants to assess ones own qualitities or currentfeelings (Johnson et al., 2002; Kelley et al., 2002; Macrae et al., 2004; Schmitz, examined the neural basis of the self-relevance effect in memory, whereby participantstypically demonstrate enhanced episodic memory for information that has beenrelated to oneself (Rogers, Kuiper, & Kirker, 1977). Participants incidentally encoded adescribed their own personality, described the personality of current president GeorgeW. Bush, or appeared in uppercase or lowercase letters. Results indicated that self-judg-Later research revealed that activity in this region of medial PFC also cor-relates with subsequent memory for items that were encoded in a self-referential man-Together, these observations that medial PFC appears to subserve thinking aboutmay rely on a common set of cognitive processes, these disparate lines of research con-J. P. Mitchell and colleagues
should engage in simulation only when ones task is to infer the current mental statesmental state attribution (e.g., when looking for a friends familiar face in a crowdedbar), one need not simulate the minds of others based on ones own. Second, simu-to the person in question. If a person thinks very differently (perhaps because of cul-or predicting his behavior can be achieved through consideration of ones own.participants were asked to mentalize about the targets internal states by judging howcomponent, namely, indicating how symmetrical (left to right) each face appeared.performed during the initial presentation (mentalize,nonmentalize) as well as howsimilar each participant felt he or she was to each target (similar, dissimilar). As pre-dicted on the basis of extant research on social cognition, the contrast of mentalizingfor those for whom the mentalizing task was performed. Specically, for faces in theobserved for faces in the nonmentalizing task. These results are consistent with the perceiver to make simulation an appropiate basis for understanding that persons mind.sessed by any other class of stimuli. To make sense of anothers behavior we must accurately identify these mental states and consider how, in combination with stable
damage. Indeed, the fundamental importance of human sociability both to everydaylife as well as to cultural achievements of the species prompted some observers tosuggest that social cognition may have been one of the primary engines of humanevolution. For example, Tomasello (1999) argued convincingly that what sets neuroimaging and neuropsychological methods, researchers have begun to providetriangulating support for the notion that separate systems do indeed subserve socialand nonsocial cognition. The promise of future research in this domain is to begin to1.Of course, the ability to use neuroimaging and patients with neuropsychological disorders tothese assumptions is compelled philosophically or empirically, nor has serious challenge been2.A somewhat counterintuitive nding from this study was that the medial PFC was not engagedby judging the personality characteristics another person (George W. Bush), despite consistentobservations that this region accompanies similar social-cognitive tasks. One possibility is thatvines and can be pressed into wine. Recent research suggested that the region of medial PFCTrends in Cognitive ScienceJ. P. Mitchell and colleagues
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Gallagher, H. L., Happ, F., Brunswick, N., Fletcher, P. C., Frith, U., & Frith, C. D. (2000). Readingthe mind in cartoons and stories: An fMRI study of theory of mind in verbal and nonverbalNeuropsychologiaGallagher, H. L., Jack, A. I., Roepstorff, A., & Frith, C. D. (2002). Imaging the intentional stanceNeuroimageGallup, G. G., Jr. (1985). Do minds exist in species other than our own? Neuroscience and Biobe-Gilbert, D. T. & Malone, P. S. (1995). The correspondence bias. Goel, V., Grafman, J., Sadato, N., & Hallett, M. (1995). Modeling other minds. NeuroreportGregory, C., Lough, S., Stone, V., Erzinclioglu, S., Martin, L., Baron-Cohen, S., et al. (2002). Theoryof mind in patients with frontal variant frontotemporal dementia and Alzheimers disease: Proceedings ofHamilton, D. L., Driscoll, D. M., & Worth, L. T. (1989). Cognitive organization of impressions:Journal of Personality and Social PsychologyHamilton, D. L., Katz, L. B., & Leirer, V. O. (1980). Cognitive representation of personality impres-Journal of Personality and Social Hartwick, J. (1979). Memory for trait information: A signal detection analysis. Journal of Experi-Hastie, R. & Kumar, P. A. (1979). Person memory: Personality traits as organizing principles inmemory for behaviors. Journal of Personality and Social PsychologyHauk, O., Johnsrude, I., & Pulvermuller, F. (2004). Somatotopic representation of action wordsNeuronJohnson, S. C., Baxter, L. C., Wilder, L. S., Pipe, J. G., Heiserman, J. E., & Prigatano, G. P. (2002).Trends in Cognitive ScienceKelley, W. M., Macrae, C. N., Wyland, C. L., Caglar, S., Inati, S., & Heatherton, T. F. (2002). Findingthe self? An event-related fMRI study. Journal of Cognitive NeuroscienceImage and Brain: The Resolution of the Mental Imagery DebateJ. P. Mitchell and colleagues
Macrae, C. N., Moran, J. M., Heatherton, T. F., Baneld, J. F., & Kelley, W. M. (2004). Medial pre-frontal activity predicts memory for self. Cerebral CortexMartin, A. (2001). Functional neuroimaging of semantic memory. In R. Cabeza & A. KingstoneHandbook of Functional Neuroimaging of CognitionMason, M. F., Baneld, J. F., & Macrae, C. N. (2004). Thinking about actions: The neural sub-Cerebral CortexMitchell, J. P., Banaji, M. R., & Macrae, C. N. (in press-a). General and specic contributions ofNeuroImageMitchell, J. P., Banaji, M. R., & Macrae, C. N. (in press-b). The link between social cognition andJournal of Cognitive NeuroscienceMitchell, J. P., Heatherton, T. F., & Macrae, C. N. (2002). Distinct neural systems subserve personProceedings of the National Academy of SciencesMitchell, J. P., Macrae, C. N., & Banaji, M. R. (2004). Encoding specic effects of social cognitionon the neural correlates of subsequent memory. Journal of NeuroscienceMitchell, J. P., Macrae, C. N., & Banaji. M. R. (in press). Forming impressions of people versusNeuroImageMontague, P. R., Berns, G. S., Cohen, J. D., McClure, S. M., Pagnoni, G., Dhamala, M., et al.NeuroImageOstrom, T. M. (1984). The sovereignty of social cognition. In R. S. Wyer & T. K. Srull (Eds.), Parker, S. T. & Gibson, K. R. (1990). Journal of Comparative PsychologyPylyshyn, Z. (1981). The imagery debate: Analogue media versus tacit knowledge. Trends inRogers, T. B., Kuiper, N. A., & Kirker, W. S. (1977). Self-reference and the encoding of personalJournal of Personality and Social Psychology
Salmon, E., Garraux, G., Delbeuck, X., Collette, F., Kalbe, E., Zuendorf, G., et al. (2003). Pre-roimageSchacter, D. L. & Buckner, R. L. (1998). Priming and the brain. NeuronSchmitz, T. W., Kawahara-Baccus, T. N., & Johnson, S. C. (2004). Metacognitive evaluation, self-NeuroimageSrull, T. K. (1981). Person memory: Some tests of associative storage and retrieval models. Journalof Experimental Psychology: Human Learning and MemorySrull, T. K. & Wyer, R. S. (1989). Person memory and judgment. Tager-Flusberg, H., Boshart, J., & Baron-Cohen, S. (1998). Reading the windows to the soul: Evi-dence of domain-specic sparing in Williams syndrome. Journal of Cognitive NeuroscienceTomasello, M. (1999). . Cambridge: Harvard UniversityTomasello, M. & Call, J. (1997). Warrington, E. K. (1975). The selective impairment of semantic memory. Quarterly Journal ofWyer, R. S., Bodenhausen, G. V., & Srull, T. K. (1984). The cognitive representation of personsand groups and its effect on recall and recognition memory. Journal of Experimental Social Psy-Zysset, S., Huber, O., Ferstl, E., & von Cramon, D. Y. (2002). The anterior frontomedian cortexand evaluative judgment: An fMRI study. NeuroimageJ. P. Mitchell and colleagues
In developmental psychology, the paradigmatic task for assessing a childs ability toreason about the mental states of others (theory of mind) is the false belief task(Wimmer & Perner, 1983; for reviews of this literature see Flavell, 1999; Wellman &Lagattuta, 2000; Wellman, Cross, & Watson, 2001). In the standard version of this task(the object transfer problem), the child is told a story in which a characters beliefthe characters knowledge. In the original version, Maxi puts his chocolate in the greenof the characters belief (Where does Maxi characters action (Where will Maxi green these questions, even ones that do not specically query a belief content, require thechild to pay attention to Maxis belief, and not the actual location of the chocolate(Dennett 1978; Premack & Woodruff, 1978). Dozens of versions of the false beliefdren and among task versions (Wellman, Cross, & Watson, 2001), in general childrenbelief problems as denitive theory of mind tasks (Fletcher et al., 1995; Goel et al.,1995; Gallagher et al., 2000; Brunet et al., 2000; Vogeley et al., 2001; Saxe & Kanwisher, 2003; Ruby & Decety, 2003; Grezes, Frith, & Passingham, 2004). Thesewhen subjects are required to reason about someones false belief, including the medial 5Four Brain Regions for One Theory of Mind?
junction extending into the posterior superior temporal sulcus (left: LTPJ, right: RTPJ;be a specialized neural substrate for theory of mind? In other words, is reasoning aboutother minds a necessary condition for the activity of each of these regions? Is it a suf-to the subjects reasoning about other people? That is, the coactivation of at least fourdients of theory of mind. Can we identify and distinguish those ingredients?Certainly the RTPJ and the MPFC, and to a slightly lesser extent the LTPJ and PC,are good candidates for the neural substrate of theory of mind. However, questionsabout the distinct contributions of these regions remain open. Contrary to sometational ingredients contributed by the RTPJ, LTPJ, MPFC, and PC are not yet known.read false belief problems could be a specialized neural substrate for a theory of mind.To illustrate how neuroimaging data can help to answer this question, a relatively in-depth characterization of the functional prole of the RTPJ is in order. It establishes
Temporoparietaljunction (TPJ)bilateralMedial prefrontal
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Figure 5.1
mind appears to be both a necessary and a sufcient condition for a robust BOLDHowever, a few caveats are in order to preclude some common confusions. From1.That the RTPJ is the sole (or most important) brain region responsible for reason-2.That a functional region of interest is a single thing, and/or a functionally homo-3.That theory of mind is a simple, homogeneous process.region of the brain is solely, or even primarily, responsible for carrying out inferencesing to problems, executing responses, and so on. Furthermore, the LTPJ, MPFC, andPC share many of the features of the RTPJs response prole. Data from the RTPJ areastonishingly clean and reliable, and so serve as a useful illustration.(hundreds of millions of them) are functionally homogeneous. Undeniably, there Finally, theory of mind itself is not a simple computation, and must not be identi-and neuroimaging data, see Saxe, Carey, and Kanwisher (2004) and Frith and FrithFour Brain Regions for One Theory of Mind?
subjects reasoned about the false belief of a character in a story (Fletcher et al., 1995;Gallagher et al., 2000; Vogeley et al., 2001), a cartoon (Gallagher et al., 2000; Brunetet al., 2000), an imaginary ill-informed protagonist such as Christopher Columbus(Goel et al., 1995) or a medical lay person (Ruby & Decety, 2003), relative to when noGallagher et al., 2000; Vogeley et al., 2001), judgments about the true function of anobject (Goel et al., 1995) or medical condition (Ruby & Decety, 2003).Based on these preliminary hints, what might be the role of the RTPJ? One possi-bility is that the RTPJ is genuinely engaged in the core responsibility of a theory ofmind: achieving a representation of other peoples mental states. However, there are1.Making causal inferences, specically, inferences about invisible causal processes;2.Maintaining two competing representations of reality (actual state of affairs andreality represented in the characters head) and/or inhibiting prepotent responses3.Representing the presence of a person or intentional action in the story, prerequi-sites for the operation of a theory of mind, but not its fundamental core.Could the RTPJ response observed in previous studies reect a domain-general processments of physical causality, such as breaking a laser beam to set off an alarm (Fletcheret al., 1995; Gallagher et al., 2000). However, whereas all of the stories in previoussome control stories did so. We therefore devised a new version of false belief storiesphysical (Saxe & Kanwisher, 2003).We had subjects read theory of mind stories that described a characters action
jects read theory of mind stories compared with MI stories. In fact, the response tothe RTPJs role in causal inference, if any, is truly specic to reasoning about mentalone of these representations in order to facilitate a response based on the other. AnyTo test this hypothesis, we used a control condition that would force subjects torepresent literal pictures of the world, like photographs and maps (Saxe & Kanwisher,reality. Adults, like children (Zaitchik, 1990), found false photograph problems slightlyFour Brain Regions for One Theory of Mind?
88Rebecca Saxe
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The percentage signal change with respect to passive xation in four brain regions when sub-jects read false belief and false photograph stories. Time (seconds) on the x axis (Saxe, unpub-
Four Brain Regions for One Theory of Mind?
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PC1.20.80.400.40.81.2
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(c) Posterior cingulate (PC). (d) Left temporoparietal junction (LTPJ).
yet denied to the RTPJ responsibility for theory of mind proper; that is, for attribut-ing mental states. Rather, they suggest that the RTPJ reects a precursor for genuinetheory of mind: identifying and representing people, and/or peoples physical behav-of another person (i.e., a human faceHaxby, Hoffman, & Gobbini, 2000, 2002),movements of a person (body motionVaina et al., 2001; Grossman & Blake, 2002;animate motionCastelli et al., 2000; Allison, Puce, & McCarthy, 2000), the inten-cartoon or story (but no false belief attribution) than to scrambled sentence or picturestanding other people. However, our results suggest a different resolution.presence of a person in the stimulus to the RTPJ response (Saxe & Kanwisher, 2003).To conrm that the RTPJ is recruited specically during the attribution of mentalstates, we created new stories in which a character was introduced immediately, butinformation about that characters mental state was delayed (Saxe & Wexler, in press).The rst sentences described the characters background (nancial, geographical). Sixseconds later, sentences describing what the character wanted or believed were addedbelief stories), and when the background of a character was described in the story but
Four Brain Regions for One Theory of Mind?
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Response in the first 6 secondsImmediateDelayed
dition, stories began with a characters mental state. In the delay condition, the rst section ofthe story (corresponding to time points 48) provided information about a characters social oravailable only in the second section of the story (time points 1014). These data were collectedfrom 12 nave subjects in a 3.0T scanner at Massachusetts General Hospital in the CharlestownNavy Yard (Saxe & Wexler, in press).(Saxe & Wexler, in press).
higher when mental state information was available (average percentage signal changetest). Clearly, the response of the RTPJ reects not just the onset of the story or the detected presence of a person, but the beginning of mental stateThese data rule out the possibility that the RTPJ is involved in detecting, perceiv-believed, and not an action (Saxe & Wexler, in press). However, the response of thePerceiving another persons intentional action is therefore not necessary to inducea robust response in the RTPJ. Other data suggest that it is not sufcient either. If thetional action should produce an even larger response. On the contrary, when subjectslower than while subjects read false belief stories (Saxe, Perrett, & Kanwisher, unpub-bution of mental statesthe core of a theory of mindand not merely in a precur-sor process. Neither perception of a person nor of an intentional action is sufcientto elicit a robust response in the RTPJ. Mental state attribution is necessary.Kanwisher, unpublished data). These results highlight the importance of regions ofThere is one more criterion for a candidate neural substrate of theory of mind: its rolemust not be limited to the attribution of false beliefs. Observers attribute false beliefs
is particularly dependent on the subjects ability to juggle simultaneous competingA neural substrate for theory of mind should be recruited whenever the subject hasbeliefs and reasonable desires is robust (Saxe & Wexler, in press).Therefore, the RTPJ met our initial criteria for a neural substrate of theory of mind.boring posterior superior temporal sulcus as a precursor of genuine theory of mind.Finally, its response generalizes beyond the attribution of false beliefs.So far, we have focused exclusively on the RTPJ, but our introduction named fourthe LTPJ, PC, and MPFC. Why are all four of these brain regions activated together?and memory, especially forhas the advantage of a much broader scope than any single study, canvassing theFour Brain Regions for One Theory of Mind?
However, meta-analyses have at least two important limitations. The rst is were signicant in one brain region and not signicant in another. But a differenceof signicances is not necessarily a signicant difference. To conclude that two brainregions make signicantly different contributions to a task, it is necessary to comparethe two response proles directly. The second drawback is that a meta-analysis Therefore, to evaluate what, if anything, is each brain regions distinct contributionto theory of mind, the ideal is to compare directly (1) the response of each regionent regions to one another. Differences among these four brain regions emerge undersuch scrutiny. The overall pattern so far, however, is one of striking functional simi-larity. Distinguishing the contributions of each region to reasoning about other mindswill be a critical problem for future research.tive to scrambled sentence controls. To illustrate the difculty of identifying the sibility, the authors own assumption, is that subjects did not reason about mentalaction control stories, but the MPFC was not. However, this conclusion is incompat-
the observed dissociation between the RTPJ and MPFC is desirable.native very plausible. In one of the human action stories, for example, a burglar inthe door. But once inside he accidentally disturbs a small animal that runs out thebank door, breaking the beam and setting off the alarm. To me, this story stronglyHowever, what are we to make of the fact that the MPFC did not respond to the stories?inhibitory processing (Decety & Sommerville, 2003). That is, the MPFC, unlike thereality, one of which is false, independent of whether the story involved any mentalstates. Unfortunately, this conclusion was undermined by further investigation.subjects attribute true beliefs and reasonable desires to a character. In our data, neitherLTPJ and PC as well; gure 5.5). Second, PSC relative to xation in the MPFC forinanimate objects (Saxe & Kanwisher, 2003). The response during the rst six secondsFour Brain Regions for One Theory of Mind?
during the same interval of stories that allowed immediate mental state attributiontest; see gure 5.4; Saxe & Wexler, in press).core of theory of mind, representing and reasoning about mental states. Moreover, wehuman action stories to those in the original study, we found an intermediate responsein both the RTPJ and MPFC. We speculated that the relatively unconstrained human
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Figure 5.5
MPFC is consistent with reports of the MPFCs generally high resting metabolic rate(Raichle et al., 2001). However, the functional signicance of this difference is notclear. As described, the latency and magnitude of selectivity in the MPFC is overallremarkably similar to those observed in the RTPJ, and so the difference seems to lieinitial criteria for a region genuinely involved in theory of mind. Of course, these fewfor instance, or that the RTPJ is only involved in precursors of theory of mind.The question for the future remains: within the domain of theory of mind, what isone hint is available. Two studies had subjects play a simple game, such as rock,paper, scissors, in two conditions: in some trials, subjects believed that they werewere playing against a human (although unfortunately, neither region of interestgestive, but their interpretation is unclear. Selective MPFC activation could reectarousal and emotion associated with playing a human opponent, or choosing onesown actions in a social interaction, or predicting a human opponents reply. Greenother hand, might be more involved in inferring the content of another personsbeliefs and desires (which are simple and unchanging in a game such as rock, paper,scissors). These are just post hoc speculations however; more work is necessary.PC and LTPJFour Brain Regions for One Theory of Mind?
false photograph or mechanical inference stories (Saxe & Kanwisher, 2003). Neitherregions response was higher to pictures of people than pictures of objects, or to verbaldescriptions of people relative to verbal descriptions of objects (Saxe & Kanwisher,be sufcient and (mostly) necessary for involvement of the LTPJ and PC, just like theHowever, data from our most recent experiment do contain one small hint aboutthe distinctive contribution of these two regions: responses of the PC and LTPJ are notperfectly selective for the availability of mental state information (Saxe & Wexler, inmental state information was not yet available. That is, the LTPJ and the PC both diddissociation means. The LTPJ and the PC may be activated in anticipation of needingto reason about the characters mental states, in forming an impression of the per-character from long-term memory. Greene and Haidts (2002) proposal that the PC isthese hypotheses directly.Developmental psychology has so far proved to be a very rich vein for neuroimagingminers interested in the neural substrate of theory of mind. The canonical false beliefControl conditions adopted from developmental psychology, such as false photographpsychology, however, social neuroscience may have to move on. Important questionsfor the future will not be whether given brain regions are involved in theory of mindingredients of theory of mind indexed by activation in the RTPJ, MPFC, PC, and LTPJ
Nancy Kanwisher. Also, thanks to Anna Wexler, Lindsey Powell, Laura Schulz, YuhongJiang, Johannes Haushoffer, Tania Lombrozo, and Jonah Steinberg for comments onAllison, T., Puce, A., & McCarthy, G. (2000). Social perception from visual cues: Role of the STSTrends in Cognitive ScienceBloom, P. & German, T. P. (2000). Two reasons to abandon the false belief task as a test of theoryBrunet, E., Sarfati, Y., Hardy-Bayle, M. C., & Decety, J. (2000). A PET investigation of the attri-NeuroimageCastelli, F., Happe, F., Frith, U., & Frith, C. (2000). Movement and mind: A functional imagingstudy of perception and interpretation of complex intentional movement patterns. NeuroimageDecety, J., Chaminade, T., Grezes, J., & Meltzoff, A. N. (2002). A PET exploration of the neuralNeuroimageDecety, J. & Sommerville, J. A. (2003). Shared representations between self and other: A socialcognitive neuroscience view. Trends in Cognitive SciencesKinds of Minds: Towards an Understanding of Consciousness. New York: Flavell, J. H. (1999). Cognitive development: Childrens knowledge about the mind. Fletcher, P. C., Happe, F., Frith, U., Baker, S. C., Dolan, R. J., Frakowiak, R. S., et al. (1995). Otherminds in the brain: A functional imaging study of theory of mind in story comprehension.Transactions of the Royal Society of LondonGallagher, H., Jack, A., Roepstorff, A., & Frith, C. (2002). Imaging the intentional stance in aNeuroimageGallagher, H. L. & Frith, C. D. (2003). Functional imaging of theory of mind. Trends in Cogni-Four Brain Regions for One Theory of Mind?
Gallagher, H. L., Happe, F., Brunswick, N., Fletcher, P. C., Frith, U., & Frith, C. D. (2000). Readingthe mind in cartoons and stories: An fMRI study of theory of mind in verbal and nonverbalNeuropsychologiaGoel, V., Grafman, J., Sadato, N., & Hallett, M. (1995). Modeling other minds. NeuroreportTrends in Cognitiveoneself and others: An fMRI study. NeuroimageGrossman, E. D. & Blake, R. (2002). Brain areas active during visual perception of biologicalNeuronHaxby, J. V., Hoffman, E. A., & Gobbini, M. I. (2000). The distributed human neural system forface perception. Trends in Cognitive SciencesHaxby, J. V., Hoffman, E. A., & Gobbini, M. I. (2002). Human neural systems for face recogni-Biological PsychiatryHooker, C. I., Paller, K. A., Gitelman, D. R., Parrish. T. B., Mesulam, M. M., & Reber, P. J. (2003).Cognitive Brain ResearchMcCabe, K., Houser, D., Ryan, L., Smith, V., & Trouard, T. (2001). A functional imaging study ofProceidings of the National Academy of Sciences ofPelphrey, K. A., Mitchell, T. V., McKeown, M. J., Goldstein, J., Allison, T., & McCarthy, G. (2003a).Brain activity evoked by the perception of human walking: Controlling for meaningful coher-Journal of NeurosciencePelphrey, K. A., Singerman, J. D., Allison, T., & McCarthy, G. (2003b). Brain activation evokedby perception of gaze shifts: The inuence of context. NeuropsychologiaPremack, D. & Woodruff, G. (1978). Does the chimpanzee have a theory of mind? Raichle, M. E., MacLeod, A. M., Snyder, A. Z., et al. (2001). A default mode of brain function.Proceidings National Academy Sciences of the United StatesRuby, P. & Decety, J. (2003). Effect of subjective perspective taking during simulation of action:A PET investigation of agency. Nature NeuroscienceSaxe, R., Carey, S., & Kanwisher, N. (2004). Understanding other minds: Linking developmentalSaxe, R. & Kanwisher, N. (2003). People thinking about thinking people: fMRI investigations oftheory of mind. Neuroimage
Saxe, R. & Wexler, A. (in press). Making sense of another mind: The role of the right temporo-NeuropsychologiaSaxe, R., Xiao, D.-K., Kovacs, G., Perrett, D. I., & Kanwisher, N. (2004). A region of right poste-rior superior temporal sulcus responds to observed intentional actions. NeuropsychologiaVaina, L. M., Solomon, J., Chowdhury, S., Sinha, P., & Belliveau, J. W. (2001). Functional neu-roanatomy of biological motion perception in humans. Proceidings of the National Academy of Sci-Vogeley, K., Bussfeld, P., Newen, A., Herrmann, S., Happe, F., et al. (2001). Mind reading: Neuralmechanisms of theory of mind and self-perspective. NeuroimageWellman, H. M., Cross, D., & Watson, J. (2001). Meta-analysis of theory-of-mind development:Wellman, H. M. & Lagattuta, K. H. (2000). Developing understandings of mind. In S. Baron-Cohen, H. Tager-Flusberg, & D. J. Cohen (Eds.), New York: Oxford University Press.Wimmer, H. & Perner, J. (1983). Beliefs about beliefs: Representation and constraining functionof wrong beliefs in young childrens understanding of deception. Zacks, J. M., Braver, T. S., Sheridan, M. A., Donaldson, D. I., Snyder, A. Z., Ollinger, J. M., et al.(2001). Human brain activity time-locked to perceptual event boundaries. Nature NeuroscienceZaitchik, D. (1990). When representations conict with reality: The preschoolers problem withFour Brain Regions for One Theory of Mind?
I would like to place social cognition, and thus the social brain, in an evolutionarywith other primates (Yoder et al., 1996; Foley, 1997). Many of our social behaviors areall of these abilities affect our social behavior and cognition. These abilities, however,evolutionary biology and primatology have revealed surprising cognitive capacities in7 mya (Foley, 1997; Gibbons, 2002; gure 6.1). African and Asian monkeys (commonMcGrew, 2001). These discoveries have forced psychologists and neuroscientists toSocial behavior can be dened as any interaction with members of ones ownspecies. Social cognition, then, is the information-processing architecture that enablesus to engage in social behavior. Social neuroscience is the study of how the brainimplements the information-processing architecture for sociality. A key question forsocial neuroscience is to what extent brain systems subserving social behavior are6Theory of Mind and the Evolution of Social IntelligenceValerie E. Stone
that subserve multiple areas of cognition. Research in patients with neurologicalprocesses, but on some very general and powerful cognitive abilities that are uniqueValerie E. Stone
Old World54261814
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and humans. Time is measured in mya (millions of years ago), except where marked kya analyses and was compiled from Yoder et al. (1996), Foley (1997), Gibbons (2000), and Wildman
memory, to name a few (Suddendorf, 1999; Corballis, 2003). The remarkable changesgeneral cognitive capacities, particularly those subserved by the frontal lobes. One(1997) made such a case for the general perceptual computations that subserve facecognitive system will perform the function it was selected for most efciently, andthat it will have design features specic to that function (Tooby & Cosmides, 1992).However, executive function and recursion clearly fail that test, as they are equallyuseful in memory, language, social cognition, and tool manufacture, and have noMuch of our social behavior, however, is similar to that of other primates and othermammals. It is parsimonious to assume that many of the brain systems that subservedescent are inherited from a common social mammal ancestor, and will have commontions. In comparative biology, this is known as homology, to indicate that two speciesAlthough some domain-specic brain systems for sociality may have been preservedfor tens of millions of years, social neuroscientists should be wary of assuming toomuch social specicity in social cognition. With each domain of social cognitionattachment, hierarchy negotiation, cooperation, in-groupout-group categorizationTheory of Mind and the Evolution of Social Intelligence
Social Cognition in Monkeys, Apes, and Humans: Components of Theory of MindAs an example of how to approach a topic in social neuroscience in this way, I wouldlike to review research on our ability to understand other peoples mental states, a cog-nitive capacity known as theory of mind (ToM). Humans make inferences about anddesires, goals, intentions, attention, knowledge, and beliefs. Thus ToM encompasses a(Baron-Cohen, 1995; Wellman & Liu, 2004). By breaking ToM down into components,First, however, we must clarify some issues of terminology, as different scholars and different elds sometimes use the term theory of mind differently. The develop-mental and primate literature on theory of mind makes distinctions among severaledge, belief, perception. Sometimes ToM is used to refer specically to the ability torepresent the contents of ones own and others mental states, something that youngerchildren cannot do. Theory of mind is often seen as equivalent to metarepresentation,y of mind is often seen as equivalent to metarepresentation,or She saw that [the lion had escaped from its cage] (Leslie, 1987; Perner, 1991;infer require metarepresentational inferences. For example, inferring another personsemotional state does not require representing someone elses representations, but onlyhis or her external appearance; She looks angry. For this reason, Leslie and Frith(1990) proposed that inferring emotional states should not be considered ToM. Infer-ring others intentions, goals, and desires is another gray area for theory of mind terminology, as such inferences also do not necessarily require metarepresentation.chology, ToM is used broadly to mean inferring a variety of mental states, not limitedDevelopmental psychology can reveal the building blocks and components of ToM,Valerie E. Stone
tives have each component (Saxe, Carey, & Kanwisher, 2004; Suddendorf, 1999). Thein each component. Many building blocks of ToM are present in our ape relatives (Suddendorf, 1999; Suddendorf & Whiten, 2001; Hare, Call, & Tomasello, 2001;Below, I discuss developmental studies on how humans develop ToM and relevantIn reviewing the developmental literature, I remain agnostic on theories of how ToMdevelops, whether through simulation, theory building, or modular maturation.Rather, I focus on when certain abilities emerge and which abilities might sharecommon processes. From such a review, it is clear that we have gaps in research onthe neural basis of ToM. For example, although we have ample research on systemsunderlying our ability to detect eye gaze direction, there is almost no research on whatresearch on theory of mind in the brain does not include control conditions withequivalent task demands (I include some of my own research in this), and thus leavesopen the question of whether certain areas are involved specically in ToM. As a cor-rective to this, I suggest taking a careful look at components of ToM to see how futureresearch might take account of these issues.Building Blocks of ToM: Inferring Goals and IntentionsInfants from very early on begin to distinguish actions that are intentional, and todiscern an actors goal. Between ve and nine months of age they can differentiateaccidental from intentional behavior (Woodward, 1999), and by fteen months theyPan troglodytestional actions (Call & Tomasello, 1998). Monkeys do not seem to make this distinc-tion. Assuming homology, this would put the date for this ability in a commonagent was looking at the point reached for, with looking at indicated by eye Theory of Mind and the Evolution of Social Intelligence
limb movement direction. Such integration is a necessary part of the cognitive archi-After apes branched off from monkeys, further elaborations of this architecture mayBuilding Blocks of ToM: Joint AttentionBetween the rst and second years, children treat others gaze direction as a source ofinformation, indicative of that persons focus of attention. In joint attention, emerg-erately into another persons line of view, or that he or she is using protodeclarativethe object (Baron-Cohen, 1995; Franco & Butterworth, 1996).see (Mossler, Marvin, & Greenberg, 1976; Liben, 1978; Flavell et al., 1981), and callto a parent on the telephone, Mommy, look at my new Mickey Mouse watch!). Thus,tents of others mental states. The childs representation may only integrate informa-tion about whether or not an adult is paying attention (binary yes-no), the roughing adults attention (holding things up, pointing). Nevertheless, children at this agebecome more active in trying to affect others attention.in gaze monitoring and learned to use protodeclarative pointing. In a later study, theChimpanzees and orangutans have clearly been observed to engage in gaze monitor-another animal or human experimenter can see an object clearly, and situations Valerie E. Stone
in which the others gaze is occluded (Hare et al., 2000). Chimpanzees seem to usethe empirical standard for joint attention is pointing with a nger, there may be aNo patient or neuroimaging research to date has focused specically on joint atten-tion. It is an important stage in ToM development, however, and thus this is a Building Blocks of ToM: Pretend Playing involves decoupling the pretend reality (this is my baby) from perceptual realitystrongly posited that pretense involves representing ones own and others mental[the doll is a baby]. However, children at this age still fail perspective-taking tasksand make systematic errors about what others can and cannot see (Mossler, Marvin,convincing case that they can represent the contents of a playmates mental states.stand the representational nature of pretend play, the more parsimonious alterna-tive hypothesis is that they treat pretense as a special kind of action (Wellman & Lagattuta, 2000). Neuroscience research into the brain substrates for pretense mightcurrently being done, such as the supplementary motor area. If pretense does involverepresenting the contents of others mental states, the same areas active for ToMTheory of Mind and the Evolution of Social Intelligence
ToM and Implicit Mentalistic Understanding: Acting Based on Others Mental Statesprivate, cannot be observed directly, and can change or not change independent ofreality. I refer to this understanding as mentalism, to denote understanding of theproperties of mental things. Wellman (1990) discussed this as belief-desire psychol-ogy. Children at about age three also begin to demonstrate implicit knowledge of thecontents of others mental states, although not explicit knowledge. This aspect of ToM,as she likes, and seem to understand that peoples attitudes and emotions towardvarious objects can be used to predict what they will do (Wellman & Lagattuta, 2000;Wellman, Cross, & Watson, 2001). Thus, understanding desire may bootstrap off of anunderstand that different peoples desires are distinct, that, for example, they dontbetween eighteen and twenty-four months (Wellman & Woolley, 1990; Repacholi &Gopnik, 1997; Wellman & Liu, 2004). Likes, wants, and desires are private mental statesAdult human theory of mind involves understanding belief before they can talk about it or understand it explicitly.To test whether children or primates know about someone elses knowledge state,one has to distinguish their representation of someone elses mental state from theirrepresentation of the state of reality. If one probes what a subject thinks someone elseValerie E. Stone
be the key test of ToM. But what does it mean to understand false belief? Does someoneunderstand it if he or she can act based on someone elses false belief, but cannot talkTwo basic kinds of false belief tasks have been used with children, location-changestory (and shown pictures to go with the story, or the story is acted out with toybut it is rare that three-year-olds can pass it (Wellman et al., 2001).However, three-year-olds can pass an implicit version of the task. Perner anddren had to act quickly, without time for deliberation. Children age 36 months wereability to track the other persons belief state, true or false. These same children failedchildren generally cannot pass false belief tasks (Pratt & Bryant, 1990). Children werebox but looking away, and then asked which girl knew what was in the box. This taskdoes not require explicitly reporting the contents of the girls mental state, and thusthat knowledge can change independent of reality, and that such changes are linkedto perception, seems to emerge around age three years.Pan troglodytesstanding of knowledge and ignorance. In chimpanzee society, if two animals see theTheory of Mind and the Evolution of Social Intelligence
of reality. The mentalism that emerges with the understanding of desire is thus furtherextended into implicit understanding of belief (Wellman & Lagattuta, 2000). Evenwith evidence for implicit belief understanding, however, we still have no water-tight1976; Liben, 1978; Flavell et al., 1981). To represent others mental contents explic-itly, another cognitive ability must emerge rst.Theory of Mind Proper: Metarepresentationy of Mind Proper: Metarepresentationment about belief can be true whether or not the proposition that the belief repre-sents is true. Understanding this representational nature of knowledge and beliefmeans understanding the way that epistemic mental states refer to propositions aboutthe world. Mentalism does not sufce for understanding representation. Rather, a newstep in ToM development must occur, metarepresentation, the ability explicitly to represent representations as representations (Perner, 1991; Leslie, 1994; Baron-Cohen,Valerie E. Stone
mistaken. Passing an explicit false belief task is certain evidence of theory of mindHowever, the converse is not true. Many other cognitive abilities also contribute toformance depends. In particular, solving such tasks depends on executive control,what the other persons mental state is (Carlson & Moses, 2001; Flynn, OMalley, &Wood, 2004; Carlson, Moses, & Claxton, 2004). In fact, children can pass false beliefinhibitory control is required, for example, by making the current state of reality lesssalient (Wellman & Lagattuta, 2000). False belief tasks also depend on workingmemory and sequencing, as the subject has to keep in mind all the elements of thestory as it unfolds in order, and how those elements are changing with respect to eachdecits in inhibiting a prepotent response or in working memory could easily fail anitive ability, embedding/recursion. Explicitly to represent X represents , embedding/recursion. Explicitly to represent X represents tion] requires the ability to embed one proposition in another. If metarepresentationis simply one type of recursion rather than a separate ability, difculties with recur-self-representation, creativity, episodic memory and future planning (a.k.a. mentaltime travel), metamemory, and counterfactual reasoning (Shimamura, Janowsky, &Squire, 1990; Knight & Grabowecky, 1995; Suddendorf, 1999; Suddendorf & Fletcher-Flinn, 1999; De Villiers, 2000; Shimamura, 2000). Thus, recursion and metarepresen-with mentalism to produce what we call explicit theory of mind. Indeed, evidencesociated from metarepresentation and counterfactual reasoning (Saxe & Kanwisher,cognitive abilities besides ToM, it would not be socially specic.Theory of Mind and the Evolution of Social Intelligence
representation (De Villiers, 2000). De Villiers proposes that the ability to form embed-vides the representational structure necessary for explicitly representing belief andknowledge. Sentences such as Agent says that X, however, are about observablestates. Thus, the metarepresentational ability that is necessary to use and understandship between mental states and reality. In development, the ability to use and under-clauses, precedes the ability to pass false belief tests (De Villiers & Pyers, 2002; Smith,Apperly, & White, 2003). Although not a strict test of cause and effect, this suggeststhat a general metarepresentational capacity could be necessary before children canThe idea that explicit ToM is dependent on the metarepresentational competencenecessary for such complex grammatical structures is consistent with results on thethey lack explicit metarepresentation (Call & Tomasello, 1999). They also do not showrefer to things have never been observed to use complex syntax at all, much less anyof either episodic memory or future planning, also abilities that depend on metarep-resentation (Suddendorf & Busby, 2003). Thus, metarepresentation and recursion seemmental statesresults in having an explicit ToM in humans. Below, I discuss neuro-science research on ToM, and interpret the ndings in terms of implicit mental stateNeuroscience Research on ToM: Metarepresentation ToMSocial neuroscience has been studying ToM for less than a decade, and thus researchin this area is still very much in its infancy. We are only now beginning to learn fromhad to work out over the past thirty years. Much ToM research in neuroscience hasnot been done with proper controls for working memory, inhibitory demands of tasks,Valerie E. Stone
of research in this area claims variously that ToM might be processed in superior et al., 1998, 2003; Gallagher et al., 2000, 2002; Fine, Lumsden & Blair, 2001; Happ,Mahli, & Checkley, 2001; Stuss, Gallup, & Alexander, 2001; Gregory et al., 2002; of the complexity of ToM and the many cognitive abilities that may contribute to suc-cessful performance of ToM tasks, not to mention ToM developments after age four,I believe it is not surprising that the brain basis of ToM has not been narrowed downToM in different studies could be that these areas may be subserving different aspectsof ToM.Given the review of ToMs components, I believe the following four questions haveto be answered before we will have a clear answer about ToM in the brain.1.Do patients fail ToM tasks because of non-ToM task demands? Do patients showdecits on a ToM task but not a control task that has the same executive function (EF)2.Are ToM and EF independent? Do patients decits in ToM correlate with decitson EF measures tapping into relevant areas of EF: inhibitory control and workingmemory? Can some patients perform highly on relevant EF measures while beingimpaired in ToM? Are there patients with EF decits who can perform well on ToM3.Does inferring belief require different brain systems as inferring other mental states4.Is metarepresentation/recursion separable from ToM? Can some patients performpoorly on ToM measures while still being able to perform well on other tasks requir-Below, I discuss how social neuroscience has or has not provided answers to theseTheory of Mind and the Evolution of Social Intelligence
ent kinds of tasks. I will focus primarily on patient research, as neuroimaging researchKanwisher, 2004), and as only patient research can answer questions about whetheran area is crucial for a particular ability.Do patients fail ToM tasks because of non-ToM task demands? AreToM and EF independent? Science consists of nding evidence that is consistent withrological damage is impaired on a ToM task, the obvious alternative hypothesis is thatthe patient failed because of task demands that have nothing to do with ToM, suchas inhibitory control or working memory. The correct way to test ToM in patients isOne way to control for task demands is to vary non-ToM task demands to see if thismakes a difference in ToM performance. When patients who had lesions in left dor-solateral frontal cortex (DFC) had to hold the elements of the story in workingmemory (which is the standard false belief task format), they often failed the falsebelief task (66% correct). However, they performed almost at ceiling on these sametasks when we removed the working memory load (98% correct), showing that theirToM metarepresentational capacities were intact (Stone et al., 1998). Clearly, patientscan fail a false belief task because of non-ToM task demands.ures and ToM performance should be reported, as these can be informative. Patientsspective taking and deception, but these tasks had strong working memory andinhibitory demands, as they had to track a sequence of actions involving hiding anToM component. Because these patients were impaired on some EF measures as well,it is difcult to interpret their decits as truly reecting decits in ToM, particularlysince no direct correlation between ToM performance and EF measures was reported.Patients with frontotemporal dementia (OFC damage) were impaired on making ToMwould infer what someone else wanted from that persons eye gaze direction (WhichValerie E. Stone
way because of impulsivity and lack of inhibitory control rather than because of failurein ToM per se. These patients were also impaired on EF measures, although again, nodirect correlations between EF and ToM were reported.Researchers working with patients with frontal damage can learn from the examplebelief task in patients with TPJ lesions. In the control tasks, memory and inhibitoryToM rather than non-ToM demands.One can also control for non-ToM task demands by using different kinds of ToMtasks. Happ et al. (2001), Stone et al. (1998), and Gregory et al. (2002) got aroundsome of the demands in ToM tasks by assessing ToM in neurological patients withoutfaux pas (Happs Strange Stories Task, and Stone and Baron-Cohens Faux PasRecognition Task). Decits on ToM questions were found on these tasks in a patienttask were seen in patients with damage to OFC (Stone et al., 1998; Gregory et al. 2002),However, even in using ToM tasks that are not false belief tasks, it is still importantto look at the relationship to EF. Happ et al. (2001) report that the patient with medialfrontal damage had severe EF decits, particularly in inhibition and working memorytasks, so there remains some question as to why this patient failed ToM tasks. Gener-ures, but a relationship was seen between perseverative errors on the Wisconsin andFaux Pas task performance (Gregory et al., 2002). This correlation may have beencult to conclude that ToM decits are independent of EF decits.There are examples, however, of patients who have impaired ToM without impairedEF. One patient from Gregory et al. (2002) showed a striking dissociation between hisToM performance, which was poor, and his EF performance, which was close to ceiling(Lough, Gregory, & Hodges, 2002). Stone et al. (2002) reported a patient from theimpaired on a variety of ToM tasks and had intact executive function. The patient wasTheory of Mind and the Evolution of Social Intelligence
person, which could possibly tap into ToM, but performed normally on a control taskparticularly inhibition tasks, but was severely impaired on a variety of ToM tests,including false belief tasks (Fine, Lumsden, & Blair, 2001). Thus, his poor performanceand unimpaired on EF (Stone et al., 2003). In looking for ToM decits independentof EF decits, patient research points to the amygdala, OFC, and TPJ as possible keymental states (desire or intention), or are the same brain areas involved? Many ToMresearchers in neuroscience have used tasks that measure multiple types of inferences,(Strange Stories Task, and Faux Pas Recognition Task, Saxe & Kanwisher, 2003).Attributing intentions is an early building block of ToM, something that great apesand very young children can do. Attributing desires taps into mentalism, but does notrequire metarepresentation. Thus, it is important to try to tease apart whether all ofWe can answer these questions if we use more ne-grained methods. Researchersdesire, and intention separately. Patients with OFC damage were impaired on theGregory et al., 2002) were reected in statements such as, Well, he meant to put accidentally or intentionally. Only the most severely affected patients with fron-with false belief tasks (Gregory et al., 2002).Valerie E. Stone
tational ToM inference comes from results with the same OFC-damaged patients whowere tested in Stone et al. (1998) on the Soap Opera Task, which measures ability tomental states requiring no metarepresentation (zero order) were all about characterssuch as, Tim fancies Maria, or The children like Easter candy.thought that Sue believed that Mary thought that X. Control statements werematched for grammatical complexity with ToM questions, and had equal levels ofmade no more errors on ToM than on non-ToM statements. Thus, with a more dif-cult task tapping metarepresentation in both ToM and non-ToM linguistic statements,in any metarepresentational aspects of ToM.The picture of OFCs role in ToM is complicated by neuroimaging results concern-although it is difcult to obtain a good signal from OFC in fMRI. However, a recent2004).Since the task looked at expectations, rather than belief statements withor medial frontal cortex, but is consistent with a generator in left OFC. However, itTheory of Mind and the Evolution of Social Intelligence
also be important to see the results for true and false belief items separately, to ruleabout false belief. It may take research with a technology such as magnetoen-verbal and nonverbal ToM tasks, to clarify the meaning of these studies. I believe themost parsimonious interpretation of all results on ToM and OFC may be that OFC isUnfortunately, it has been difcult to separate belief and desire in both patient andWith both the medial frontal and the amygdala patients, it is possible that their under-In future research in patients, I believe it is important to report results separately forrately. There are also a range of mental states related to desire (adoration, disgust), andto these areas being involved in mentalism more generally, rather than just in belief.Is metarepresentation/recursion separable from ToM? Developmental andevolutionary considerations point to mentalism; that is, an understanding of theValerie E. Stone
neuroimaging research. No published patient research has addressed the question ofwhether areas involved in ToM might be involved in other non-ToM tasks that requireity of ToM and metarepresentation in fMRI. The false photograph task requires anfrontal pole was signicantly greater during belief tasks. However, the researchers didrepresentation. Clearly, TPJ and medial portions of frontal pole and temporal pole seem to be involved in mentalism, and possibly, this ability can be separated With respect to TPJ in particular, further research distinguishing it from areasinvolved in recursion would be helpful. Some regions very close to the parts of TPJthat were reported for ToM (Saxe & Kanwisher, 2003) have also been found to be activeet al., 2001). These areas do not completely overlap with the TPJ areas for ToM, butboth ToM and non-ToM control questions. Using this test with patients with medialdissociations between recursion and mentalism in ToM inferences. A study directlycomparing the processing of embedded sentences and ToM in fMRI would solidify Neuroimaging research with implicit belief tasks would also be important. The rststudy ever to test ToM with fMRI used a task that required only implicit inferencesTheory of Mind and the Evolution of Social Intelligence
ToM, often including instructions to think about characters motivations or mentalstates (Liu et al., 2004; Saxe et al., 2004). As ToM inferences in everyday life are madeon the y, implicitly, tasks requiring deliberate inferences may not tap into theseprocesses in the same way. Implicit belief attribution tasks, perhaps styled after thisrst one used, or after Perner and Garnhams (2001) implicit false belief task (withoutresearch in this area, as they might help identify areas that are involved in mentalismThe Maturation of ToM Research in NeuroscienceDecades of research on ToM in developmental psychology and primatology have givenus a detailed picture of its precursors and components. Developmental research intowhy children have difculty with false belief tasks, in particular, provides insight intohow people can seem to be impaired on certain ToM tasks because of limitations innon-ToM cognitive abilities. Neuroscience research on ToM is just beginning to takethese methodological lessons into account. We are also just starting to make distinc-tions between ToM and other related abilities such as recursion.I believe the mentalism evident in childrens and primatess understanding of desiredevelopment and evolution. Although further research will solidify this conclusion,in ToM tasks because they subserve mentalism, rather than metarepresentation andToM, the mentalism required to understand that belief states and desires can changeindependent of reality, and form the neural basis of implicit mental state under-with other great apes. To the extent that brain regions are involved in specicallyMetarepresentation, recursion, and executive control are not at all limited to ToM.They enable language, complex tool manufacture, future planning, episodic memory,Valerie E. Stone
Overall, areas involved in the social aspects of ToM seem well positioned anatomi-research using both patient data and neuroimaging, testing a variety of both ToM andnon-ToM tasks that require executive control, metarepresentation, and recursion, canclarify these issues. For now, we can understand the processing of ToM in the brainscience research on ToM matures, and takes these complexities into account, we willhave a clearer picture of how brain areas involved in ToM might have evolved andhow different components of ToM interact with each other.1.In an unexpected contents task, the subject is shown a container that is clearly labeled as ifit contains one kind of thing; for example, a candy box clearly indicates that it contains candy.there. Control questions usually ask about what was true originally, what the subject thoughtoriginally, and what is true right now.2.Some would describe desire statements as involving rst-order intentionality (Dennett, 1987).When I use zero-order, rst-order, second-order, and third-order here, I am referring to the levelMindblindness: An Essay on Autism and Theory of Mind.Theory of Mind and the Evolution of Social Intelligence
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personal interaction. Consequently it provides the foundation for society, culture, andchurch) and almost all of our daily interactions with other individuals depend on language. To researchers interested in understanding language processing, this is, ofcourse, all very basic and assumed. Yet, little of the research focused on mechanismsthat mediate language processing takes this into account explicitly. Such researchtional architecture of the language system.Language Is Treated as Cold and Not Cognitivetive psychology from each other. However, interactions between social psychology andyears. For example, research in social cognition is informed by work on mechanisms ofattention and automaticity (Bargh & Chartrand, 1999; Wegner & Bargh, 1998) and cat-egorization (Hugenberg & Bodenhausen, 2004; Wittenbrink, Hilton, & Gist, 1998).However, basic research on language processing has been affected only to a small degreeby work in social psychology, despite the entirely social nature of language use. As brainarising in the context of purely cognitive functions (Siegal & Varley, 2002).For over half a century, the scientic study of language was disproportionately inu-enced by the perspective of linguistics (Chomsky, 1957). Linguistic investigation 7Investigating Cortical Mechanisms of Language Processing in
represents just one way to study language, and psychology, sociology, computerscience, anthropology, neurobiology, and philosophy all play important roles inrange of language research, particularly in psychology and neurobiology. The rstguage areas of the brain can be understood apart from other complementary regionsbe several equally explanatory theoretical accounts of syntax. He was convinced thatition) to distinguish among equally explanatory syntactic theories. Needless to say,theory of syntax. But one could draw the inference that the entire enterprise of sep-be misleading, taking scientic inquiry down a garden path.one being simple and linear and the other complex and nonlinear. However, this sep-aration into systems based on apparently different modes of behavior led researchers
tionarily expert, mandatory, and automatic syntactic processor has given rise to a greatdeal of research and controversy (Appelbaum, 1998; Gareld, 1987; Sperber, 2001). Assuch interaction whereas opponents support it; however, this debate has had severaland without consideration for either the speakers or the listeners intentions or goals.other words, it is assumed that the impact or possible importance of a messagestion. On the other hand, this is a theoretical foundation that shifts research away frombasic evolutionary forces that actually shaped the biological development of linguis-Dewey (1896) pointed out that although researchers can analytically decompose ability or isolability leads to research questions and methods that would not otherwiseroscience research (Barrie, Freeman, & Lenhart, 1996; Freeman, 2003; Freeman &
explicitly distinguished from phrenology (Fodor, 1983), Gazzaniga (1985) proposed amore specic cortical modularity. Whereas Fodors (and most linguists and psychol-for specic cognitive functions. Indeed, on the face of it, some examples seem clear.blindsight in which dorsal-stream visual information is preserved compared with casesin which ventral-stream information is preserved (Weiskrantz, 1986). In these cases,derive from basic sensory systems such as vision. When it comes to language func-tion, however, which is often treated as the prototype for cortical modularity The landmark ninetieth-century work of Broca (1861) and Wernicke (1874) shapedtion between anatomical locations of brain injury and disruption of particular lan-anterior or posterior cortical regions, and research questions emerging from this divi-versus working memory decits for Brocas area). However, the clear divisions betweenBrocas aphasia as expressive aphasia and Wernickes aphasia as receptive aphasia areWith increasing use of neuroimaging measures, methods of lesion analysis and psy-
performance (Chomsky, 1965; de Saussure, 1959). Over the past fty years we havecourse, using this approach. However, this approach may be limited when it comesfrom its grounding in behavior. By shifting to studying language use rather than lin-guistic competence, however, we may gain, rather than lose, in our ability to under-approach is that it considers how language processing, in service of specic goals anduses, interacts with a broad set of neural circuits that are involved in more generalactivity, working memory, and attention. A tendency in neuroimaging research hasbeen to try to isolate language processing from these other processes using a varietyof analytic and design methods. However, it is important to remember that languagenitive, sensorimotor, and affective operations in addition to linguistic ones. To studyand affective mechanisms, so that research on the biology of language processing
A common feature of both lesion analysis and much psychological research is theby a collection of functionally independent units (Fodor, 1983). Work in dynamicsystems theory (Freeman & Barrie, 1994) suggests an alternative approach: rather thanof the older holographic view of the brain as a mass of equipotential tissue (Lashley,1950). We do not assume that all parts of the brain participate equally in all behav-and functionally separate process. Rather, we postulate that the neural circuits thatinteractive, and operate differently depending on their dynamic patterns of activity.mental context, producing different modes of processing in different circumstances,leading to specic patterns of behavior. In some cases, the apparent specializations ofcomponential boxological models of behavior (Neisser, 1976). For example, lan-(Clark, Carpenter, & Just, 1973). This decomposition provided the basis for importantThese information-processing models assumed, however, that each processing stageThis approach to cognitive research has continued through recent times. Just asbrain as modular, consisting of functionally specialized and independent locations(e.g., Shallice, 1988). In the study of language, the frontal operculum (Broca, 1861)
and posterior superior temporal region (Wernicke, 1874) played special roles in thislocalizational view, representing sites for language production (early view) or syntaxtively. In part, these componential views are rooted in other studies of biological spe-motor and sensory functions, replicating the notion of structure-function relation-ships found elsewhere in biology.systems to putative functional components. In psychological research, this compo-from and complementary to the change produced by damage to a different Ultimately, this conceptual framework is the basis for many studies in functionalonance imaging (fMRI). In research on language and the brain, some studies focusedon validating certain models derived from information-processing psychology, which
Typical tasks used to study language in the brain include, at different levels of lan-lexical decision (lexical level), or grammaticality judgment (sentence level). To carrythe phonological patterns of the words, thereby exercising phonological processing.linguistic property of an utterance exercises the same kind of processing (same mech-A study conducted in our laboratory illustrated this concern and the nature of thetion tasks (one phonological, one auditory), both calling for stimulus comparison andplanned motor behavior, we intended to isolate those neural processing componentsthat mediate phonological segmentation. We concluded that it is the process of seg-ulatory recoding, that appears to involve left...inferior and middle frontal [gyri]actuallysegment the speech stream into phonemes before recognizing the phonemes,the basic units of speech perceptual analysis, or are syllables or diphones or onset-rime structures the basic unit of perception? Although these are standard assumptionsin much speech research, and may reect consistency in information conveyed inspeech (Studdert-Kennedy, 1981), this does not necessarily license a neural reality for
the specic nature of the task comparison, as we later learned. A follow-up study, usinglogical discriminations during real conversations (thus making discrimination a veryHowever, this leaves us with a very real question: which result is more indicative ofreal phonological perception, involvement or noninvolvement of the frontal lobe? Ifone nonspeech-control task emphasizes working memory and the motor system morethan another, this will moderate the appearance of neural activity in the frontal regionby an a priori theoretical assumption, which may be of questionable validity. This par-ticular issue is broader than the study of language in the brain. Tasks that involvejudgments of any kindself-relevance, affective valenceinvolve working memory,
tioning. To understand language use, rather than competence, however, it is importantaffective, and motor systems. With this research goal, it is likely that assumptionsTo understand the difference between a behavioral psychology experiment and itsogy studiesresponse time and accuracy. Behavioral measures, such as response timeand accuracy, typically give us a relatively univariate view of language processing, pro-in this network over time (Nyberg & McIntosh, 2001). Every subprocess can manifestprocesses of interest. However, in a neuroimaging study, manipulated targetsubprocesses and ancillary subprocesses are all manifest distributed across the depend-ent measure. We call this difference between behavioral and neurophysiological meas-Brain imaging offers the opportunity to observe all the components operating in parallel, overlapping, and distributed in time. However, unlike response timeior in a very different way. It is important to note that in neuroimaging, dependentmeasures are themselves directly linked to system components of interestanatomy.
Variation in one dependent measure is no longer a reection of the entire chain ofA corollary issue then is that the decompositional or subtractive approach totions can obscure activity due to psychological processing under investigation. We callcessing of interest in these studies. However, due to the dependent measure problem,This has meant that for the results of imaging studies to be interpretable, it necessaryity to be subtracted off using appropriately matched control conditions. Yet thisassumption seems questionablewe know that complex motor circuits interact withworking memory (Cohen et al., 1997; Smith et al., 1998) are also closely identied with(Grafton, Hazeltine, & Ivry, 1998; Morecraft & van Hoesen, 1998; Picard & Strick, 1996).
study, perhaps to avoid their investigation, this approach affords a natural opportu-neurology. In fact, the Chicago school of psychology emphasized the study of cogni-naturalistic phenomena (Dewey, 1896; James, 1904). Furthermore, Brunswik (1955)argued that psychological research should contrast conditions that display the fullrange of natural variation observed in behavior. Whereas true ecologically valid lan-in the context of telephone conversation. Although most theories of speech percep-as the talker, can change the perception of speech. For example, for Caucasians, seeing
same speech. However, it is important to begin to understand the mechanism bywhich visual information can change auditory speech perception.Visual information showing movements of the mouth and lips during talkingmay report hearing an emergent phoneme (e.g., /t/). Visual and auditory informationjointly specify a percept that is present in neither modality but combines informationciency as measured by memory capacity (Goldin-Meadow et al., 2001), suggesting anaction among sources of information but not how this interaction takes place.However, neuroimaging reveals more directly the neural systems that are active whendifferent sources of information are available to the comprehender. Subjects werewhile seeing the storyteller (audiovisual), or just seeing the storyteller (visual). Wecondition than in either other condition (Skipper, Nusbaum, & Small, 2002, 2005).Moreover, the presence of the visuomotor information changed the laterality of thebetween face information and acoustic speech in more traditional speech perceptionstories in this study, and not perform an additional judgment task. If we had designed
working memory used during judgment could have masked Brocas area activityobserved during comprehension. However, the limitation of this approach is thatstories and some of what they remember, these measures are not sufciently sensitivewithout changing either. We can think of this as a kind of Heisenberg uncertaintyprinciple in cognitive neuroimaging research.of speech perception. Visual information from mouth movements is combined withauditory information to determine a new interpretation of an utterance. This by itselfwould not suggest much of a paradigmatic change to language research and wouldguage processing more broadly. Rather than think about language as a signal (multi-cant act that was part of the basic force shaping the evolution of the brain. By thisconstrual, the listeners goal may not be to interpret the linguistic message but to inter-would suggest that communicative behavior, broadly construed, should be affected bya conversational partners behavior, even beyond the simple process of interpretation.verges on the speech of the other, in terms of speaking patterns. This vocal accom-modation or indexical mimicry is increased between members of the same social groupone interlocutor is trying to persuade the other of something (Giles & Coupland,1991). Moreover, this behavioral convergence in a conversation is not restricted to
ior. And this can serve to link interlocutors socially, increasing the sense of interper-sonal afliation (Lakin & Chartrand, 2003) as well as shifting attention from ones selfClearly, in face-to-face communication the motor system plays an important role inlanguage. Whereas this motor mimicry may be mediated in part by the observation-social consequences of such interpersonal interaction. Such motor mimicry changesinvolved in the motor system, attention, working memory, and affect.Roland (1993) proposed that frontal cortex can generate sensory expectations thatcortex in just this way, specically because of the relationship between dorsolateralmemory systems. His theory is that affective goals and evaluations are important inshifting attention for perception. This process of shifting attention involves frontaland prefrontal systems changing the sensitivity of posterior sensory processes, pre-phonetic perception. The race of a talkers face (shown as a static image) can changethe perception of the persons speech (Rubin, 1992) and the gender of a talkers facewithout dynamic real-time perceptual input about a talkers mouth movements, expec-shaped by expectations even when they do not involve information about a talkers
We measured cortical activity of listeners presented with sine wave speech before andmotor network involving inferior frontal gyrus and superior parietal cortex (Wymbs,Nusbaum, & Small, 2004). Similarly, changing listeners expectations that the sameadvantage suggesting a change in cortical processing (Luks, Nusbaum, & Levy, 1998).Changing a listeners expectations about speech fundamentally changes the patternof cortical processing that mediates perception of speech.Summary and ConclusionEvery utterance is inherently ambiguous. Comprehension of an utterance depends onunderstanding the speakers intended meaning. Given the inherently nondeterminis-use more information than is present in the utterance itself, such as race, gender, andsocial status of the speaker, to interpret the message. However, none of these cate-gories is itself assessed deterministically. Listeners must use a variety of weak con-ential process (Nusbaum & Schwab, 1986; Small & Rieger, 1982). Listeners may formsuggests that comprehension will depend on working memory (for holding commu-and attention shifts, affective systems (for evaluating sources of information and forsetting goals), perceptual processes, and motor processes. This view of language pro-By shifting the focus of research questions from meta-linguistic tasks to under-standing language use in service of social interaction, brain imaging allows us inves-tigate the breadth of neural mechanisms that interact during real language behavior.
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tations and the expectations of others (social regulation). Self-perception processes and perceptive-processes of other people have been theorized to be central to the regulation of social behavior. However, studying the role of self-perception and person-perception processes in the regulation of behavior has been challenging. Individuals are socialized from a very early age to regulate their behavior. Just asralistic studies. However, the fact still remains that variance in social regulation isneeded to make hypotheses about the underlying role of self- and person-perceptionperson-perception processes in social regulation. Neuropsychological populationsregulatory dysfunction and healthy control individuals. Functional neuroimagingused in social-personality psychology, neuroscience methods may provide additional8Orbitofrontal Cortex and Social Regulation
Social Regulation: A Brief Overviewpresence of food is to approach it. However, people do not simply react to their envi-dards, in turn, inuence their behavior (Baumeister & Heatherton, 1996; Carver &Scheier, 1990; Higgins, 1987). Therefore, descriptions of human social behavior mustism contribute to perception of stimuli and to selection of behavioral responses. Forexample, even a hungry person may not immediately reach for the last cookie attheir weight or appearing gluttonous. In this case the persons response was mediatedothers. In summary, social regulation occurs when people exert control over theirself-regulatory decits. If so, comparisons between these people and healthy controlswill be useful for understanding whether self- and person-perception processes are nec-essary for social-regulation. The complexity of social-regulatory processes suggests thatassociated with cognitive or emotional control are a good place to begin. Generally,cortical sites. More specically, the orbitofrontal cortex (gure 8.1) is richly connected
anterior cingulate, and somatosensory areas I and II (Adolphs, 1999; Brothers, 1996).tively associated with poorly regulated social behavior.was a railroad worker who accidently shot a tamping iron through his head. Harlow,the physician who treated him, noted that after his injury, Gage could walk, talk andremember things just ne, but he did have problems with his social behavior. Forto the extent that persons of delicacy, especially women, found it impossible to1986, p. 99). Similarly, more recent descriptions of patients with orbitofrontal lesions
Orbitofrontal cortex indicated with a white circle on a sagittal brain slice.
damage does impair social regulation (table 8.1; Beer, 2002; Beer et al., 2003; to regulate their behavior, in many cases, behaving with strangers in ways that weremore appropriate for interactions with close others. In one study, patients withlikely to include unnecessary personal information when answering questions (Beer orbitofrontal and dorsolateral prefrontal lesions and healthy controls (Beer, 2002).priate when speaking to a stranger. Finally, in the third study, orbitofrontal damage wasTable 8.1
Examples of Social Disinhibition Associated with Orbitofrontal Damage
Social impairmentStudyProfane, irreverentHarlow, 1848, cited in MacMillan, 1986Impaired problem-solving for Saver & Damasio, 1991Riding on a hospital gurneyBlair & Cipolotti, 2000Practicing karate in the waiting room,Rolls et al., 1994Unrestrained, disinhibited actionsKretschmer, 1956, cited in Stuss & Benson, 1992Puerile, jocular attitude, sexuallyBlumer & Benson, 1975disinhibited behavior, inappropriateDigression from conversation topicKaczmarek, 1984Teasing inappropriately, excessive Beer, 2002
Excessive self-disclosureBeer et al., 2003
Together these results suggest that orbitofrontal cortex is critically involved in ensur-The Role of Self-Perception and Person Perception in Self-RegulationTwo possible mechanisms by which orbitofrontal cortex may support appropriatesocial functioning are self- and person-perception processes. Both processes are given
0100%
Figure 8.2
Carver & Scheier, 1990; Gray & McNaughton, 2000; Higgins, 1982). Specically, indi-doing and determine if it is affecting other people in the intended manner.Self-Perception Processes and Social RegulationThe fundamental role of self-perception processes in social regulation is exempliedby consensus on the importance of monitoring ones behavior to ensure appropriate-and/or expectations of others (Carver & Scheier, 1990; Gray & McNaughton, 2000;research on self-focused attention. When people are placed in conditions of self-because the frequency of monitoring is increased (Carver & Scheier, 1990; Gibbons,focusing peoples attention on themselves. Experimental inductions of self-focusedof themselves (Carver & Scheier, 1990; Gibbons, 1990; Robins & John, 1997).Another suggested role between self-perceptive processes and social regulationcomes from research on the accuracy of self-perception. Whereas the importance ofself-perception for social regulation is somewhat agreed upon, the motivation under-lying self-perceptions most benecial for social regulation has been debated. Someand that such ination promotes adaptive social behaviors (Taylor & Brown, 1988),perceptions (Heatherton & Ambady, 1993; Robins & Beer, 2001).Self-Perception Processes and Orbitofrontal CortexA host of research supports the role of orbitofrontal cortex, as well as other frontallobe areas, in self-perception processes identied as necessary for social regulation(monitoring, insight accuracy). Clinical characterizations and empirical research ofcately involved in self-monitoring (Beer, 2002; Lhermitte, 1986; Luria & Homskaya,
and healthy controls (Beer, 2002). Participants took part in a task that required them to regulate how much personal information they revealed. Transcripts of intimacy of self-disclosure. A measure of accurate self-perception was computed Not all self-processes are associated with the orbitofrontal cortex. Imaging research
political gure and syllabic structure. Similarly, fMRI studies consistently show thatright BA 9 and 10 has increased activity when observing ones own face versus thatPerson-Perception Processes and Social Regulationperceptiveness (Goffman, 1967). To ensure smooth social interactions, people must beskilled at perceptiveness, that is, inferring interpretations others place on ones acts(Goffman, 1967). Presumably, perceptiveness includes interpreting both evaluationsregulation comes from research on self-disclosure and liking. Self-disclosure is gov-& Miller, 1994; Cozby, 1973). The reciprocal element makes it necessary to properlyunderstand the perspective of a conversation partner. Understanding mental and includes making correct attributions for a partners intentions and motivations and isconsidered essential for relationship satisfaction (Harvey & Omarzu, 1997).Person-Perception Processes and Orbitofrontal CortexBoth lesion and imaging researches suggest that prefrontal cortex, includinglarly the right side, is critically involved in theory of mind on a visual perspective task (Stuss, Gallup, & Alexander, 2001), another group found that patients with
theory of mind tasks (Stone, Baron-Cohen, & Knight, 1998). However, in this study,sity of a coherent description to ensure that the audience understood the speaker. Itare implicated in making inferences about anothers mental states. In two imagingstudy, participants had to decide whether a person living in the fteenth centurytion was assessed in relation to story comprehension that required participants tomake either a mental or physical inference about a characters actions. When the twoShamay-Tsoory et al., 2003). Similarly, they were impaired at inferring emotional statesfrom pictures of various emotional facial expressions (Hornak, Rolls, & Wade, 1996),and shame (Beer et al., 2003). However, orbitofrontal patients had no trouble infer-ring the feelings of story characters that had been on the receiving end of a social faux
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Nevertheless, we suggest a certain type of reduction. We propose that social paincally, we suggest that some of the basic neural mechanisms that support the experi-spearean title). Evidence from both animal and human research literatures suggest thatdistressing component of both forms of pain. Moreover, once a connection is estab-Finally, we reexamine the function of dACC versus rostral anterior cingulate (rACC)9A Pain by any other Name (Rejection, Exclusion, Ostracism) still
is involved in affective processes. However, this view does not ultimately hold up todifferent dichotomy of function can account for the previous view, such that bothnectionist constraint-satisfaction network. Alternatively, rACC can best be understoodWe should say from the outset that although we focus on the dACC as a regionquently, this region is ripe for consideration. This should not be taken to mean thatis not; right ventral prefrontal cortex is another. Although we do not focus on otherregions of the pain matrix (Peyron, Laurent, & Garcia-Larrea, 2000), such as the peri-acqueductal gray, insula, and somatosensory cortex, the reader should not be surprisedthese lines (MacDonald & Leary, in press; Panksepp, 1998).my leg hurts. Similarly, we describe social pain with phrases such as, she broke myreference to physical pain terminology. In fact, English speakers have no other way todescribe the feelings associated with social pain (MacDonald & Leary, in press).MacDonald and Leary (in press) recently examined whether this linguistic overlap
countries, to provide typical ways of describing social pain. In each country, socialpain descriptions relied on physical pain words. This evidence, at the very least, sug-humans around the world. It does not in itself, however, say much about whetherspread so widely.An Evolutionary Story for the Linket al., 2002). Finally, the social attachment system may have piggybacked onto ordependence on their caregivers. Whereas lack of food, water, shelter, and defenseneeds is entirely contingent on the continuing relationship with a caregiver. Forin infancy, because meeting them is critical to meeting all other needs.could serve as a solid foundation for the creation of this attachment system. Whereas
that attention and other biological resources can be mobilized to prevent greater injuryand promote survival, the social pain system produces social pain in response to socialinjuries so that attention and other biological resources can be mobilized to preventthese injuries and promote survival. It is of interest that mammals are also the rstWe have chosen to focus on the role of the dACC in both social and physical paintors in the brain (Vogt, Wiley, & Jensen, 1995), and thus may have been one of theprimary sites of action in Panksepps work on the social and physical pain-alleviatingical pain processes in humans. Finally, a number of studies with nonhuman mammalssuggest that the ACC, and perhaps the dACC specically, is involved in the experi-dACC, rACC, somatosensory cortex, insula, periacqueductal gray, and right ventralsensory, distressing, and regulatory components of pain. Somatosensory cortex andinsula are primarily, although not exclusively, linked with the sensory aspects of region of ones body that is in pain and other sensory features including intensity.Periacqueductal gray, rACC, and right ventral prefrontal cortex are more frequentlyFinally, dACC is generally associated with the subjectively distressing component ofSensory intensity and subjective distress associated with pain are often highly cor-with sensory intensity likened to the radios volume and subjective distress likened to
versus outdoors at a barbeque or ones sensitivity to or tolerance of loud noises. Thus,tions or across different people, the same degree of sensory intensity might produceThe consequences of damaging neural structures associated with the sensory andchronic pain problems sometimes underwent cingulotomies, a procedure that involvesnicant pain relief; however, the relief did not come as a result of all aspects of painbeing diminished. Rather, only the subjective distress appears to have abated. Patientswould report that the sensory aspects of pain continued, but it no longer seemed tobother them (Foltz & White, 1968). Alternatively, there is a reported case of a patientwith damage to somatosensory cortex. When painful stimulation was applied to thebody region represented by the damaged part of somatosensory cortex, the patientstimulation as distressing (Nagasako, Oaklander, & Dworkin, 2003). So although inten-roimaging study in which they used hypnotic suggestion to alter the perceivedunpleasantness of painful stimulation without changing the perceived intensity.pain distress were highly correlated with changes in dACC activity. The changes werenot correlated with activity in somatosensory cortex, suggesting that this region,Until recently little research has examined neural correlates of social pain in humans; however, some studies from the animal literature suggest a role for dACC inbecause similar neural regions in different animals do not always serve the same func-
macaques, not squirrel monkeys. However, other squirrel monkeys in this study hadtions. Moreover, the monkeys were still capable of making different kinds of vocal-Electrical stimulation studies support this conclusion as well. Over half a centuryago, Smith (1945; Jurgens & Muller-Preuss, 1977) observed that electrical stimulationTo examine neural correlates of social exclusion in humans, and begin to assess
scanner. Numerous manipulations have been used successfully by social psychologists(see Williams, Forgas, & von Hippel, in press).ing task, so they will be working on their own (Leary et al., 1995; Twenge et al., 2001;if they have close friends and loved ones now (Baumeister & De Wall, in press).Williams and Sommer (1997) used the most direct and overt manipulation of socialaggressiveness, and increased conformity to group norms (Leary, in press).neuroimaging study. Fortunately, the ball-tossing manipulation has been convertedplayers, and produces the same effects as the in-person version (Williams, Cheung, &report feeling social pain as a result of the experience (Zadro & Williams, 1998).onance imaging (fMRI) study.Subjects in our study of social exclusion (Eisenberger, Lieberman, & Williams, 2003)at Johns Hopkins University, involves scanning many subjects simultaneously whileneural patterns of the different subjects can be analyzed. We showed subjects a write-Nature Neuroscience
to do the same kind of procedure in our laboratory, but that we were still in the tech-or the other (an animation of the game can be seen at www.scn.ucla.edu).however, our subjects knew that they were not being intentionally excluded. This con-dition conceptually replicated the study by Zadro and Williams (1998), in which sub-link so that they would be able to play. Subjects played with the other two playerspercent probability. In the third and nal scan (explicit exclusion), subjects were fullyOur primary analysis comparing brain activations showed greater activity in dACC,pain, however, dACC and right ventral prefrontal activity were both strongly related
results, consistent with other results (Petrovic et al., 2002; Wager et al., 2004), suggestposition, cluster size, and intensity. This may be somewhat surprising given that sub-this condition. However, it should be recalled that Zadro and Williams subjects expe-system. Similarly, the need to detect potential exclusion may be so important orin this case that it is not truly exclusionary.less likely to try to self-regulate the negative experience. Consistent with this view, notive to inclusion, even at very liberal statistical thresholds. Given that right ventral
negatively evaluated attitude objects (Crockett, Eisenberger, & Lieberman, 2004; Cun-rience, it appears that in our study, subjects were not engaged in these sorts of mentalexcluded, self-regulatory and attributional mechanisms were not engaged. One caveataway our cover story.The results thus far, from our own work as well as from the broader physical and socialphysical pain. The next obvious question to ask is why. What is the function of theto for an answer.If clinical and psychopathology researchers are asked, they will probably respondstates such as pain, anxiety, and distress. Each of these states serves important func-tions in motivating adaptive and appropriate behavior (Mandler, 1975). Thus, theseresearchers focus on phenomenological contributions of the ACC.If cognitive researchers are asked, they will probably respond by saying that thesuggested that dACC is activated when a discrepancy exists between ones goals andones prepotent responses. For instance, during the Stroop task when a person is showncolor words in different color ink (e.g., R-E-D written in blue ink), ones goal is to sayand noties lateral prefrontal cortex that top-down control processing is necessaryresearchers in this eld agree that the dACCs role is limited to detecting conict anddirectly.
tions underlying that activity.We suggest that it might be protable to consider the possibility that the dACC workswith one another. Any alarm system, whether it is a clock alarm or a smoke alarm,must integrate two functions to work effectively. First, it must be able to detect theroom crosses some threshold for unacceptability. Second, the alarm must be able tolling with smoke and possibly re. Conceptually, these functions are separable (e.g.,for the alarm to function properly.two processes operate together, they might function as a unitary alarm system, detect-ing conicts and making noise to attract the persons attention.may be the phenomenological consequence of the detection of discrepancy. It is pos-would still be integrated with one another, but this might help explain why per-
accounts. However, two studies partially investigated this question by examiningstudy (Derbyshire, Vogt, & Jones, 1998) noted that the regions in dACC activated bya pain and Stroop task were adjacent to one another, but also sometimes overlapping.tive task, however, the task did not involve conict detection and thus is not entirelyconict-detection processes covary with one another? If so, this would suggest thatperforming its own process independently. Two studies examined the covariationdisorder (OCD), which is characterized by distress and worry, were scanned while addition, a trend was observed, although not signicant, such that patients with moreA second neuroimaging study (Eisenberger, Lieberman, & Satpute, in press) investi-
consider an alternative formulation. After a search of the literature, although admit-the conict is represented symbolically or nonsymbolically.Symbolically and nonsymbolically represented conict should vary in a number ofthe conict or of the source of the conict.According to this view, when conict isThese two kinds of conict processing should also vary in the kind of computationalnetworks representing the combined outputs of multiple interconnected inputs natu-various inputs. When inputs are coherent and consistent with one another, the
The resources of awareness and attention are limited (Miller, 1956) such that only aprocessed serially. Thus, although many conicts may be nonsymbolically processed,We propose that symbolically represented conict is processed primarily in rACC,whereas nonsymbolically represented conict is processed primarily in dACC. Withinputs can nicely model standard oddball, gono-go, and Stroop effects. Alternatively,and the nature of the error. Using a Talaraich y-coordinate of 30 to divide rostral fromactivate dACC (Braver et al., 2001; Bush et al., 2003; Carter et al., 2000; Weissman et al., 2003). Alternatively, error-detection tasks, which involve symbolic conict rep-resentations, tend to activate rACC (Garavan et al., 2003; Kiehl, Liddle, & Hopnger,outcomes and what actually occurs (Frijda, 1986; Lazarus, 1991; Mandler, 1975). Mostnegative emotions (anger, fear, sadness) are thought to have a specic Intentionalis. Anxiety, however, is a negative affect that is distinguished from fear in that it lackssymbolic-nonsymbolic theory of the ACC. Sadness, anger, and fear reliably activateet al., 1999; Liotti et al., 2000; Mayberg et al., 1999; Shin et al., 2000). Anxiety,however, tends to activate dACC (Kimbrell et al., 1999; Liotti et al., 2000). Alongsimilar lines, perception of discrete emotional expressions activates rACC (Ueda et al.,2003) whereas perception of ambiguous emotional expressions activates dACC
Finally, pain distress can also be divided along symbolic and nonsymbolic lines. Typ-ically, pain is a nonsymbolic bottom-up process (animals without symbolic capacitiesoften associated with dACC activation (Hsieh, Stone-Elander, & Ingvar, 1999; Ploghauset al., 1999; Sawamoto et al., 2000; Tolle et al., 1999). However, when pain is anti-rACC. They do, however, seem to be organized such that more symbolic forms of cog-and place. If one is aware of the source of ones distress, one may choose to delayresponding (one may choose to ignore the distress caused by ones boss while the bossI have lunch with my bosss boss next week). Nonsymbolic conict simply producesWe suggest that because of the role of social attachment in mammals, the social painaddition, the dACC may have been one of the primary sites in which this overlapevolved, such that today, this region produces similar experiences of distress inresponse to both physical and social injuries. More generally, we propose that theredirects our attention and motivation toward dealing with the source of the threat.Finally, we attempted to integrate our model with what previously was hypothesized
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...We benign implications of categorization, such as carrying an umbrella when perceptionlikely. As Allport recognized, though, categorization becomes potentially more prob-lematic when applied to the perception of people. This is because we often have strongactivated after categorization. Allport noted, The category saturates all that it con-verbal behavior, response latencies), but with the continued development of neuro-questions from a more explicitly neuroscientic perspective. In this chapter, we reviewthe line of research we have pursued using event-related brain potentials (ERPs) tostudy aspects of social perception.Using ERPs to Study Social Perceptionsuch presentation of a visual stimulus or commission of a behavior. They can be10The Social Neuroscience of Stereotyping and Prejudice: UsingEvent-Related Brain Potentials to Study Social PerceptionTiffany A. Ito, Geoffrey R. Urland, Eve Willadsen-Jensen, and Joshua Correll
peaks at around 100msec after stimulus onset. Other naming schemes refer to the pre-ERPs are attractive for the study of social perception for several reasons. First, theyviding a large corpus of research to draw on in linking observed electrical activity that occur very quickly after stimulus onset, and allow for inferences about the might be unaware. As the research reviewed in this chapter will illustrate, perceivingand there is no reason to expect that perceivers are explicitly aware of all aspects willingness to report internal states accurately, ERPs are useful in measuring sociallysensitive topics such as processes related to stereotyping and prejudice. Finally,distribution of observed activity can be used to obtain estimates of neuroanatomicalception. Two main associated issues are how social category information relevant toclassifying individuals into meaningful social groups is perceived, and how this infor-mation, in conjunction with stereotypes, inuences behavior.Perception of Social Category InformationA very simply model of social perception is as follows: if we categorize an individualT. A. Ito and colleagues
earliest aspect, involving encoding cues indicative of social category membership, isless well understood. It has generally been assumed that an individuals membershipin different social categories is perceived relatively automatically (Brewer, 1988;Bruner, 1957; Fiske & Neuberg, 1990; Macrae & Bodenhausen, 2000; Stangor et al.,1992), but it was not known how quickly this information could be perceived andwhether the perceptions could be easily modied.encoded when perceiving faces, and whether attention to these factors requires per-ceivers to attend to these dimensions explicitly (Ito & Urland, 2003). To do so, we hadtive-going component with a mean latency of 122msec after face onset, a positive-going component with a mean latency of 176msec, a negative-going component witha mean latency of 256msec, and a positive-going component with a mean latency of485msec. We refer to these components based on their polarity and latency as N100,, respectively.ERP responses early in processing. Specically, racial category information affectedattending to gender.Gender category information had effects slightly later, beginning in the P200. Asof target gender, P200s were larger to males than to females. As with race effects, thisoccurred for participants explicitly attending to both race and gender, and also for
105010128960321281024Amplitude (mV)P200N200Black targetsN100105010128960321281024Amplitude (mV)Latency (ms)Female targets
N100, P200, and N200 effects as a function of target race (top) and gender (bottom). Waveformscategorizable individuals, by T. A. Ito and G. R. Urland, 2003, Journal of Personality and Social
orienting to relevant and/or salient features (Czigler & Geczy, 1996; Eimer, 1997; Kenemans, Kok, & Smulders, 1993; Naatanen & Gaillard, 1983; Wijers et al., 1989).We can also ask what the direction of target race and gender effects reects. This is an issue we are still investigating, but we do have preliminary hypotheses. In which can occur as early as 100msec (Pizzagalli, Regard, & Lehmann, 1990; Smith etto whites may represent greater, more individuated processing of racial in-groupsistent with the large body of research showing that whites and racial in-groupmembers are spontaneously processed more deeply than other racial groups (Anthony,Cooper, & Mullen, 1992; Levin, 2000). The meaning of larger N200s to females is lessclear. Processing differences in favor of female over male targets have been obtained(Lewin & Herlitz, 2002; McKelvie, 1981; McKelvie et al., 1993; OToole et al., 1998),We are therefore continuing our examination of what these particular patterns of
attention, we have examined a component linked with working memory processes insensitivity led to the belief that P300 amplitude reects updates to working memory1981). To examine the effect that social category information has on working memory,Consistent with past P300 research on nonsocial stimuli (Donchin, 1981), P300swere larger when a target individuals social category membership differed from thatblacks. This shows, not surprisingly, that working memory processes are sensitive tothe social category dimension along which categorization was explicitly occurring.However, we also obtained evidence of implicit categorization effects on workingmemory. P300 amplitude also increased when a target picture differed from individ-larger to females than males. Both explicit and implicit working memory effects canrace, and the bottom panel from participants attending to gender. In both panels, datawas done to examine the degree to which low-level perceptual features such as lumi-were responding to social cues per se, rather than to general perceptual features (lumi-nance) that happen to covary with the social cues.Together, these studies provide relatively direct evidence that when perceiversT. A. Ito and colleagues
Race Categorization TaskGender Categorization Task
P300 effects as a function of a target individuals race and gender relative to the race and gender ofand the bottom panel shows responses from participants categorizing by gender. Waveforms arerizable individuals, by T. A. Ito and G. R. Urland, 2003, Journal of Personality and Social Psychology
gender information are both activated at very early stages in processing. That race andautomatic (Brewer, 1988; Fiske & Neuberg, 1990; Macrae & Bodenhausen, 2000).However, it is the case that all participants were performing some type of social-category information, even dimensions that were not directly task relevant.Boundary Conditions on Attention to Race and GenderTasks that focus attention on a targets individual characteristics and foster person-based (as opposed to category-based) processing have proved successful in reducingstereotyping and prejudice (Fiske & Neuberg, 1990). Similarly, focusing attention atlower levels of analysis, for example, on a nonsocial physical cue such as a circle on afor three different reasons: (1) encoding of category membership has been attenuated,thereby blocking activation and application of stereotypes and prejudice; (2) category(3) category membership is encoded and stereotypes-prejudice are activated, but theirapplication is attenuated. To the degree that categorization is afforded unconditionalautomatic status (Brewer, 1988; Fiske & Neuberg, 1990; Macrae & Bodenhausen, 2000),option 1 would seem unlikely. To assess this, and to examine the effects obtained in Itoegory cues. The vegetable task was chosen because it decreases stereotype activation andthought to reect greater negativity toward the out-group (Wheeler & Fiske, 2005). WeWe observed the same four components as in Ito and Urland (2003), with veryT. A. Ito and colleagues
attending to social category. Specically, sensitivity of the P200, N200, and P300 wasreplicated even under these very different processing goals. This indicates that direct-aspects of racial and gender perception observed under conditions of explicit atten-tion to race and gender, further supporting the obligatory nature of social categoryof processing, after at least elementary aspects of social category information haveing of racial cues observed in more visually simple contexts (Ito & Urland, 2003). Itof attentional load in reducing visual attention (for a review, see Rees & Lavie, 2001).in all cases. Even when effects are delayed to the P200, they occurr within 190msecLooking across these studies examining the initial perception of racial and gendercues, the ease with which social category information can be encoded is consistentlydemonstrated. Access to very quick aspects of processing afforded by ERPs reveals thatearly perceptual aspects of social categorization are relatively obligatory, driven moreby properties of the individual being perceived than by goals and intentions of theperceiver. Such encoding occurs even when attention is explicitly directed to anothersocial category dimension (Ito & Urland, 2003), when targets are processed in a
that attention to category membership need not invariantly result in stereotyping andprejudice (cf. Wolsko et al., 2000).Understanding how social category information is processed is an important piece ofthe puzzle that is social perception, but it is also important to understand how thestored information perceivers have about social groups (stereotypes, prejudice) comesto inuence behavior. We found that ERPs are also well suited to addressing this issue.We particularly focused on understanding how race inuences the detection ofTimothy Thomas, Anthony Dwain Lee). In some cases, ofcers reported fearing thatTo assess this experimentally, Correll et al. (2002) developed a simulation in whichunarmed targets by pressing another. Behavioral results showed a consistent biasbetween blacks and violence-aggression grew stronger.T. A. Ito and colleagues
to be activated easily and quickly in this task. They may, however, conict with othervated representations of hes black and Ive shot him compared with the repre-sentations of hes white and Ive shot him. Similarly, less conict seems likelybetween the representations of hes white and I did not shoot him than betweenhes black and I did not shoot him.modeling has located the source of the ERN to medial prefrontal areas (Dehaene,Posner, & Tucker, 1994).after error commission. We therefore examined ERNs in response to errors as partici-
(580600msec after stimulus onset) to ensure a relatively large number of errors. Replicating Correll et al.s initial behavioral studies, the pattern of errors participantsERNs were observed after erroneous responses, but their amplitude differed depend-the explicit point contingencies of the game. To motivate performance, a cumulativescore based on performance was displayed after every trial. Correct detection of anincorrect detection of an unarmed target cost twenty, and incorrect detection of anquences. Within this point contingency, failing to shoot an armed target is the mostdiffer. Thus, consistent with stereotypes, shooting someone who is not associatedT. A. Ito and colleagues
These initial results implicate cognitive control processes, in particular, failure toof cognitive control should be observed. To examine this, we conducted a second studyParticipants were given a slightly longer window within which to respond (850msec),detected conict (Curtin & Fairchild, 2003; West & Alain, 1999). Source modeling ofThis occurred rst in the N100, which peaked at a mean latency of 159msec, and
continued for all subsequent components. This indicates that although searching forgun status very early in processing. Although correct decisions to shoot and notshoot occur much later (at around 550650msec), these effects reect the early pointat which perception begins to differentiate between armed and unarmed individuals.The second effect observed in the ERP waveforms was a stereotype-consistent differ-whites. Said differently, starting as early as 200msec, the safeness of unarmed whiteswas being perceived, but unarmed blacks were being processed in a manner similar toimplementation of cognitive control (Curtin & Fairchild, 2003; West & Alain, 1999),The purpose of conict detection and executive control is to regulate behavior. To the degree that the ERP effects we observed reect the operation of behavior-regulation mechanisms, we would expect ERP effects to predict subsequent behavior.T. A. Ito and colleagues
point in processing that predicts behavior. Contrary to expectations, N400 effects didnot predict behavior, which therefore leaves us to question whether they did reectAllport wrote his seminal book on prejudice from which this chapters opening quo-psychology is really not so different from the eld Allport knew. We still grapple withmany of the same issues, and we still seek interdisciplinary perspectives to addressthem. The two core issues addressed in this chapterhow social category informationis processed and how racial stereotypes inuence behaviorwould be very much atinquiry.Consistent with Allports concern over the ease and uency of social categorization,and subsequent suggestions that social category information is processed automati-cally (Brewer, 1988; Fiske & Neuberg, 1990; Macrae & Bodenhausen, 2000), ourresearch with ERPs reveals perceivers sensitivity to racial and gender cues. Perceiversorient to this information early in perception and process it in a relatively obligatorystereotypes and prejudice (Ito & Urland, 2003, 2005). Moreover, early social percep-Allport (1954/1979) noted that the social category saturates all that it contains withthe same ideational and emotional avor, (p. 21), but how exactly is this achieved?
Integration of neuroscience into the investigation of social perception suggests thatwith automatically encoded social category cues, for instance, may be simultaneouslythese numerous sources of representations are consistent with each other, behavioralior successfully. This analysis suggests that understanding behavior thought to beinuenced by a targets social category membership requires consideration of activatedwhich cognitive control is being implemented. To some degree, cognitive control canbe inferred if a perceivers behavior and some likely activated representations areknown, but ERPs provide a means for observing these processes as they occur on-linerelated to perceptions of threat and danger, differences in conict monitoring and press; Ito & Correll, unpublished data). To some extent, emphasis on behavior regu-lation in understanding bias is not new. Other researchers noted that motivations control offer a more detailed model of the processes involved in regulation. Moreover,differencesThere are many challenging aspects in studying social perception, such as perceiversT. A. Ito and colleagues
Supported by NIMH grants R21 MH66739 and R03 MH61327 to Tiffany A. Ito, aNational Science Foundation Graduate Research Fellowship to Geoffrey R. Urland, and1.Although the P300 had a latency longer than 300msec, we use the P300 name because its2.The N100, P200, and N200 amplitudes were insensitive to the preceding stimuli and differedAllport, G. W. (1954/1979). The Nature of PrejudiceAmodio, D. M., Harmon-Jones, E., Devine, P. G., Curtin, J. J., Hartley, S. L., & Covert, A. E. (2004).Anthony, T., Copper, C., & Mullen, B. (1992). Cross-racial facial identication: A social cognitiveBotvinick, M. M., Carter, C. S., Braver, T. S., Barch, D. M., & Cohen, J. D. (2001). Conict mon-Brewer, M. C. (1988). A dual process model of impression formation. In R. Wyer & T. Scrull (Eds.),Bruner, J. S. (1957). On perceptual readiness. Carter, C. S., Braver, T. S., Barch, D. M., Botvinick, M. M., Noll, D., & Cohen, J. D. (1998). Ante-Correll, J., Park, B., Judd, C. M., & Wittenbrink, B. (2002). The police ofcers dilemma: UsingJournal of Personality and Social Correll, J., Urland, G. R., & Ito, T. A. (in press). Shooting straight from the brain: Early attentionJournal of Experimental Social PsychologyCurtin, J. J. & Fairchild, B. A. (2003). Alcohol and cognitive control: Implications for regulationJournal of Abnormal Psychology
Czigler, I. & Geczy, I. (1996). Event-related potential correlates of color selection and lexical deci-sion: Hierarchical processing or late selection? International Journal of PsychophysiologyDehaene, S., Posner, M. I., & Tucker, D. M. (1994). Localization of a neural system for error detec-...Surprise? Eimer, M. (1997). An event-related potential (ERP) study of transient and sustained visual atten-BiologicalPsychologyFabiani, M., Gratton, G., & Coles, M. G. H. (2000). Event-related brain potentials. In J. T.Cacioppo, L. G. Tassinary, & G. G. Berntson (Eds.), ,2nd ed. (ppPerspectives of Event-Related Potentials Research Fiske, S. T. & Neuberg, S. L. (1990). A continuum of impression formation, from category-basedGehring, W. J., Coles, M. G. H., Meyer, D. E., & Donchin, E. (1995). A brain potential manifes-Perspectives of Event-Related Potentials ResearchGehring, W. J., Goss, B., Coles, M. G. H., Meyer, D. E., & Donchin, E. (1993). A neural systemGehring, W. J., Gratton, G., Coles, M. G. H., & Donchin, E. (1992). Probability effects on stim-Journal of Experimental PsychologyHuman Perception andPerformanceGilovich, T. & Medvec, V. H. (1995). The experience of regret: What, when, and why. Greenwald, A. G., Oakes, M. A., & Hoffman, H. G. (2003). Targets of discrimination: Effects ofJournal of Experimental PsychologyHolroyd, C. B., Larsen, J. T., & Cohen, J. D. (2004). Context dependence of the event-relatedIto, T. A., Thompson, E., & Cacioppo, J. T. (2004). Neural mechanisms of social perception: TheT. A. Ito and colleagues
Ito, T. A. & Urland, G. R. (2003). Race and gender on the brain: Electrocortical measures of Journal of Personality and SocialIto, T. A. & Urland, G. R. (2005). The inuence of processing objectives on the perception offaces: An ERP study of race and gender perception. and Behavioral NeuroscienceJames, M. S., Johnstone, S. J., & Hayward, W. G. (2001). Event-related potentials, conguralJournal of PsychophysiologyKenemans, J. L., Kok, A., & Smulders, F. T. Y. (1993). Event-related potentials to conjunctions ofElectroencephalographyandClinicalNeurophysiologyLevin, D. T. (2000). Race as a visual feature: Using visual search and perceptual social perceptionJournal of Experimental Psychology129Lewin, C. & Herlitz, A. (2002). Sex differences in face recognitionWomens faces make the dif-Liotti, M., Woldorff, M. G., Perez, R., & Mayberg, H. S. (2000). An ERP study of the temporalNeuropsychologiaMacDonald III, A. W., Cohen, J. D., Stenger, V. A., & Carter, C. S. (2000). Dissociating the role ofMacrae, C. N. & Bodenhausen, G. V. (2000). Social cognition: Thinking categorically about others.AnnualReviewofPsychologyMacrae, C. N., Bodenhausen, G. V., Milne, A. B., Thorn, T. M. J., & Castelli, L. (1997). On theJournal of Experi-McKelvie, S. J. (1981). Sex differences in memory for faces. Journal of PsychologyMcKelvie, S. J., Standing, L., St. Jean, D., & Law, J. (1993). Gender differences in recognitionmemory for faces and cars: Evidence for the interest hypothesis. Journal of Personality and Social PsychologyNaatanen, R. & Gaillard, A. W. K. (1983). The orientating reex and the N2 deection of theevent-related potential (ERP). In A. W. K. Gaillard & W. Ritter (Eds.), Tutorials in ERP Research(pp. 119141). New York: North-Holland.
Nieuwenhuis, S., Yeung, N., Van Den Wildenberg, W., & Ridderinkhof, K. R. (2003). Electro-ict and trial type frequency. Affective andBehavioral NeuroscienceOToole, A. J., Deffenbacher, K. A., Valentin, D., McKee, K., & Abdi, H. (1998). The perception ofMemory and Cogni-Payne, B. K. (2001). Prejudice and perception: The role of automatic and controlled processes inmisperceiving a weapon. Journal of Personality and Social Psychologyright and left brain hemispheres: An ERP study. NeuroreportPlant, E. A. & Devine, P. G. (1998). Internal and external motivation to respond without preju-Journal of Personality and Social PsychologyNeuropsychologiarelated brain activity, judgments of response accuracy, and types of errors. Journal of Experimen-Human Perception and PerformanceSmith, N. K., Cacioppo, J. T., Larsen, J. T., & Chartrand, T. L. (2003). May I have your attention,NeuropsychologiaStangor, C., Lynch, L. Duan, C., & Glass, B. (1992). Categorization of individuals on the basis ofJournal of Personality and Social PsychologyWest, R. & Alain, C. (1999). Event-related neural activity associated with the Stroop task. tive Brain ResearchWheeler, M. E. & Fiske, S. T. (2005). Controlling racial prejudice: Social cognitive goals affectWijers, A., Mulder, G., Okita, T., Mulder, L. J. M., & Scheffers, M. (1989). Attention to color: An analysis of selection, controlled search, and motor activation, using event-related potentials.Wolsko, C. V., Park, B., Judd, C. M., & Wittenbrink, B. (2000). Framing interethnic ideology:Journal of Personality and Social PsychologyT. A. Ito and colleagues
Successful human social interaction relies on each individuals ability to understandother peoples intentions, beliefs, and desires (Baron-Cohen, 1988). An importantthey feel. We can infer this in many ways, for example, from a persons tone of voiceprimates use these expressions as a primary means of communicating emotions(Darwin, 1872).Ambady, 2002a). Cultural differences in display rules indicating when it is appropri-ability. From the beginning of life, infants show proclivity to express facial emotions,11Race and Emotion: Insights from a Social Neuroscience PerspectiveNalini Ambady, Joan Y. Chiao, Pearl Chiu, and Patricia Deldin
(Widen & Russell, 2003). Even children who are blind and deaf and have no visual orauditory notion of what it is like to communicate an emotion create spontaneousthrough the face. Darwin (1872) rst noted over 100 years ago that humans share very similar to a human expression of fear (Knapp & Hall, 1997). Finally, people fromvarious cultures are able to recognize emotional expressions such as happiness, fear,anger, sadness, disgust, and surprise at above-chance levels of accuracy (Ekman, 1992,studies show that the amygdala plays a role in the perception, detection, and subse-that medial frontal gyri are critically involved in recognizing angry expressions (Phanemotional content in faces is a rapid process that occurs very early in the perceptualoccurs as early as 160msec after stimulus presentation (Pizzagalli et al., 2002). Perceiv-ing fearful facial expressions, in particular, modulates neural responses in frontocentralregions even earlier, at approximately 120msec (Eimer & Holmes, 2002). Thus, itappears that the neural processes associated with emotion recognition occur early.In summary, substantial evidence indicates that the human capacity to express andrecognize emotions through the face has evolutionary roots, is shared across humancultures, and has dedicated neural machinery.
internal attributes such as how someone feels, but also external features (age, gender,race) that shape a persons identity by inuencing how one sees oneself and howlevels of accuracy (Ekman, 1992). However, they recognize emotions most accuratelyAmbady, 2002b; gure 11.1). Several factors are thought to moderate this recognitionadvantage, including differences in attitude toward ones own cultural group rela-(Elfenbein & Ambady, 2002a). Moreover, cultural standards for when it is appropriate
Angry
Figure 11.1
However, given the impact of emotion on social perception and interaction (Hess,Barry, & Kleck, 2000; Keltner, Ellsworth, & Edwards, 1993; Vrana & Rollock, 1998,attitudes and stereotypical judgments (Jackson et al., 2001; Asuncion & Mackie, 1996;Bodenhausen, Kramer, & Susser, 1994; Bodenhausen, Sheppard, & Kramer, 1994;Lambert et al., 1997), and evidence that physiological reactivity of perceivers duringsocial interaction varies according to both racial and emotional contexts (Vrana &targets of such judgment (Vaes et al., 2003). Indeed, little previous work to our knowl-edge has examined perceivers responses to in-group and out-group members express-We conducted functional magnetic resonance imaging (fMRI) and event-related(ERP) studies in our laboratory that emphasize not only the impactsocial behavior.To examine how racial group membership affects brain processes during emotionCaucasian participants (4 men) while they explicitly identied fear, anger, and neutralwomen. Participants were shown each facial expression for 750msec and respondedwithin 2500msec, pressing an appropriate button to indicate which emotion the facewas expressing. We predicted that participants would recognize all expressions but
nized at better-than-chance accuracy levels; however, Caucasian participants recog-nized neutral faces better than fearful and angry faces. They were also best atAmerican faces, specically, fear and anger.to Asian-American and African-American anger. These ndings suggest that neuralthey viewed expressions and judged whether or not each face was angry, fearful, orneutral. We hypothesized that race of the facial target would inuence basic structuralface processing approximately 170msec after stimulus onset and this would be observ-able in the amplitude of the VPP, an ERP that is critical to face processing (Bentin et al.,1996; Jeffreys, 1989, 1996). Moreover, we predicted that the emotional expression beingprocessed would affect the extent to which race inuenced neural processing. To detected angry expressions most accurately in African-American and Caucasian Caucasian faces relative to African-American and Asian-American faces. Finally, neutralmore, angry expressions yielded the most positive amplitude for African-American Taken together, these neuroimaging and ERP data suggest that race affects brainprocesses involved in recognizing of fear and anger. First, regions important in the
the race of the expressor. Second, behavioral and neural evidence suggests that not allHowever, Asian-American and African-American anger faces are processed more other, thus leading to better recognition of emotions expressed by those groups relative to African-Americans (Elfenbein & Ambady, 2002a). Furthermore, socialgroups often vary in social status relative to each other. This observation was however, social psychologists and sociologists have also applied this subordina-tion hypothesis to racial groups as they vary not only in socioeconomic status minority out-groups are stigmatized by the majority. Of importance, neuroimagingTogether, our data are among the rst to indicate that the emotional expression ofonly the behavioral but also the physiological level. As we describe below, using judice. Specically, theories of prejudice emphasize that a combination of factors and that emotion clearly inuences social behavior, an investigation of the interac-
2000; Richeson, et al., 2003). A series of studies from our laboratory examined corti-(Chiu, Ambady, & Deldin, 2004).basis of their responses on the modern racism scale; McConahay, Hardee, & Batts,
S1Target Face250 ms4750 ms
w, w, b, r) presented for 250msec followed 4750msec later by a corresponding target face (happywhite, angry white, etc.). Participants were asked to respond yes or no according to their prefer-
stimulus that requires anticipation of a target stimulus (Walter et al., 1964; Picton &to respond to the target stimulus (Low & McSherry, 1968; Forth & Hare, 1989; Hamon& Seri, 1987). Moreover, the presence of the early CNV is generally thought to be a1988; Shiffrin & Schneider, 1977). Several groups demonstrated the sensitivity of theKlorman & Ryan, 1980; Yee & Miller, 1988; Regan & Howard, 1995).for low-prejudiced individuals (gure 11.3). Specically, low-prejudiced individualsshowed an increased CNV not only to angry out-group stimuli, but also in anticipa-tion of angry faces more generally, compared with happy faces. Longer behavioralresponse latencies of the low-prejudiced group in evaluating angry black targets furtherindicate this enhanced processing. Together, these data support, and extend to include
mVms
15085018508.0000.000High
S1, indicating the subsequent presentation of an angry black American face (S1
of angry black targets compared with all other targets, supporting theories that indi-viduals high in explicit prejudice may be characterized by a decreased tendency, shorter behavioral response latencies of the high-prejudiced group to angry blacktargets further reects absence of effortful suppression of prejudiced behavior. Inpation of happy white targets; this suggests a greater recruitment of cognitive resourcesmembers may contribute to the expression of prejudice (Brewer, 1999; Miller & Brewer,tants of race perception and race bias. In the one fMRI study (Phelps et al., 2000) thatcortical activity to angry black targets. At rst glance, these data may seem at oddswith those of Phelps et al. However, most of their participants scored below 2 on thegroup. Indeed, within this group, the data show greatest cortical resources to angryet al., 2003). Briey, participants with high scores on the implicit association test of
nitive resources to monitoring their behavior toward out-group members (BlascovichRicheson et al. Clearly, it would be of theoretical interest to explicate further the na-these groups were enhanced or attenuated by simply varying the anticipated andactual valence of facial expression from angry to happy. The pattern of null group dif-differences to angry out-group stimuli, is striking and suggests at least that prejudiceand stereotyping are not unitary phenomena and may be malleable. It should beemphasized, however, that no analyses within race or emotion alone yielded signi-Preliminary data from two other ERP studies from our laboratory supplement theseresources and cognitive processes when confronted with racial stereotypes and viola-tion of these race-based expectancies. Briey, we measured N400 and late positive com-that conrmed or violated racial stereotype-based expectancies (Jamaals favoritesport is lacrosse, versus Jamaals favorite sport is basketball). The N400 is typically
level information processing (Halgren, 1990; Kutas & Van Petten, 1994; Osterhout &the ease with which new information is integrated into ones representation of theCoulson, King, & Kutas, 1998; Gunter, Stowe, & Mulder, 1997). Our data indicate thatcies; lack of between-prejudice differences). Notably, although the representation ofin-group and out-group stereotypes, respectively, may be equivalent between high-stereotypes appear to be quite different. Specically, low-prejudiced individuals exhib-(Chiu et al., unpublished observations).out-group emotional faces. Preliminary analyses indicate that racial and emotional differentiation occur as early as 120msec after stimulus presentation (as evidenced byto angry faces compared with happy faces), and effects of target gender are seen even80msec; female targets elicited more negative voltage than male targets). Ofevident at all stages of processing, beginning as early as 80msec after stimulus presentation. Although data analyses from this study are still under way, preliminaryand emotion on person perception and social behavior (Chiu et al., unpublished observations).Studies in our laboratory emphasize not only the impact of emotion on the per-
converging measures to clarify the impact of race and emotion on social behavior. In-temporal manifestation of cortical resources and specic cognitive processes that maymeasures. We believe that these may provide both converging evidence for existingtion of prejudice depends how individuals process this information. Moreover, thespecicity of our between-group ndings to the early, but not the late, component ofmotoric preparation to the late CNV in anticipation of angry black targets is evidencedThe brains functional anatomy places important constraints on psychological theories of racial and emotional processing, and their relationship in social behavior.phenomena that may not otherwise not be assessed, and allows an assessment of
emotion on the perception of members of in-groups and out-groups but also, weon social behavior.social behavior. Indeed, ndings from our laboratory are among the rst to indicateand low on explicit measures of racial prejudice. Together, these data underscore bothsocial behavior. These investigations are, however, a rst step toward understandingto how we perceive and interact with each other.Supported by NSF PECASE grant to N.A. and NSF predoctoral fellowship to J.Y.C. Wethank Hillary Anger-Elfenbein, Y. Susan Choi, Heather Gray, Abigail Marsh, Ashli1.As indexed by geographical distance and self-report.2.An extensive and rapidly growing literature in cognitive psychophysiology suggests that event-ated with social behavior. The amplitude and latency of these voltages changes are thought tosensitive to experimenter effects and self-presentation biases. Moreover, since ERPs are consid-
3.Whole-brain analyses were conducted using a general linear model in SPM99. All contrastCurrent Opinion in NeurobiologyAdolphs, R., Tranel, D., & Damasio, A. R. (2003). Dissociable neural systems for recognizing emo-Baron-Cohen, S. (1988). Without a theory of mind one cannot participate in a conversation. Bentin, S., Allison, T., Puce, A., Perez, A., & McCarthy, G. (1996). Electrophysiological studies offace perception in humans. Journal of Cognitive NeuroscienceBlascovich, J., Wyer, N., Swart, L., & Kibler, J. (1997). Racism and racial categorization. Journal ofBodenhausen, G., Kramer, G., & Susser, K. (1994). Happiness and stereotypic thinking in socialJournal of Personality and Social PsychologyBodenhausen, G. & Macrae, N. (1998). Stereotype activation and inhibition. In R. Wyer (Ed.),Bodenhausen, G., Sheppard, L., & Kramer, G. (1994). Negative affect and social judgment. European Journal of Social PsychologyBrewer, M. (1999). The psychology of prejudice: Ingroup love or outgroup hate? Journal of SocialElectroencephalography and Clinical Neurophysiol-Cacioppo, J., Berntson, G., Lorig, T., Norris, C., Rickett, E., & Nusbaum, H. (2003). Just becauseyoure imaging the brain doesnt mean you can stop using your head: A primer and set of rstJournal of Personality and Social PsychologyChiu, P., Ambady, N., & Deldin, P. (2004). Contingent negative variation to emotional in- andJournal of Cognitive Neuro-
Coulson, S., King, J. W., & Kutas, M. (1998). ERPs and domain specicity: Beating a straw horse.Language and Cognitive Processescondition to elicit a stimulus-preceding negativity. Darwin, C. (1872). The Expression of Emotion in Man and Animals. New York: Appleton.Devine, P. (1989). Stereotypes and prejudice: Their automatic and controlled components. JournalDevine, P., Monteith, M., Zuwerink, J., & Elliot, A. (1991). Prejudice with and without com-Journal of Personality and Social Psychology...Surprise? Eimer, M. & Holmes, A. (2002). An ERP study on the time course of emotional face processing.NeuroreportEkman, P. & Oster, H. (1979). Facial Expressions of Emotion. Ekman, P. (1992). Are there basic emotions? Ekman, P. (1994). Strong evidence for universals in facial expressions: A reply to Russells mis-Elfenbein, H. A. & Ambady, N. (2002a). On the universality and cultural specicity of emotionElfenbein, H. A. & Ambady, N. (2002b). Is there an ingroup advantage in emotion recognition?Fabiani, M., Gratton, G., & Coles, M. G. H. (2000). Event-Related Brain Potentials. In J. T.Cacioppo & L. G. Tassinary et al. (Eds.), York, NY: Cambridge University Press.Fazio, R. H., Jackson, J. R., Dunton, B. C., & Williams, C. J. (1995). Variability in automatic acti-Journal of Personality
Fiske, S., Cuddy, A., Glick, P., & Xu, J. (2002). A model of (often mixed) stereotype content: Com-petence and warmth respectively follow from perceived status and competition. Journal of Fiske, S. & Neuberg, S. L. (1990). A continuum of impression formation, from category-based toFox, E., Lester, V., Russo, R., Bowles, R. J., Pichler, A., & Dutton, K. (2000). Facial expressions ofemotion: Are angry faces detected more efciently? Gehring, W., Karpinksi, A., & Hilton, J. (2003). Thinking about interracial interactions. NatureNeuroscienceGolby, A. J., Gabrieli, J. D. E., Chiao. J. Y., & Eberhardt, J. L. (2001). Differential responses in theNature NeuroscienceGunter, T. C., Stowe, L. A., & Mulder, G. (1997). When syntax meets semantics. reading. In A. B. Scheibel & A. F. Wechsler (Eds.), Neurobiology of Higher Cognitive Function. UCLAForum in Medical Sciences, No. 29 (pp. 103150). New York, NY: Guilford Press.Activitas Nervosa SuperiorHarmer, C. J., Thilo, K. V., Rothwell, J. C., & Goodwin, G. M. (2001). Transcranial magnetic stimulation of medial-frontal cortex impairs the processing of angry facial expressions. NatureNeuroscienceHart, A. J., Whalen, P. J., Shin, L. M., McInerney, S. C., Fischer, H., & Rauch, S. L. (2000). NeuroreportHess, U., Barry, S., & Kleck, R. (2000). The inuence of facial emotion displays, gender, and Journal of Nonverbal BehaviorThe managed heart: Commercialization of human feelingHugenberg, K. & Bodenhausen, G. (2003). Facing prejudice: Implicit prejudice and the percep-cross-cultural research.
Jackson, L., Lewandowski, D., Fleury, R., & Chin, P. (2001). Effects of affect, stereotype consis-Journal of Social PsychologyBrain ResearchVisual CognitionKeltner, D., Ellsworth, P., & Edwards, K. (1993). Beyond simple pessimism: Effects of sadness andanger on social perception. Journal of Personality and Social PsychologyBelmont, CA: Wadsworth.Kutas, M. & Van Petten, C. K. (1994). Psycholinguistics electried: Event-related brain potentialJournal of Personality and Social PsychologyLow, M. & McSherry, J. (1968). Further observations of psychological factors involved in CNVElectroencephalography and Clinical NeurophysiologyMcConahay, J., Hardee, B., & Batts, V. (1981). Has racism declined in America? It depends onJournal of Conict Resolutionena, and theories. In L. P. Lipsitt (Ed.), Advances in Infancy ResearchNorwood, NJ: Ablex.Miller, N. & Brewer, M. (1986). Categorization effects on ingroup and outgroup perception. In J.Prejudice, Discrimination, and RacismMonteith, M., Devine, P., & Zuwerink, J. (1993). Self-directed versus other-directed affect as aJournal of Personality and Social PsychologyNelson, C. A., Morse, P. A., & Leavitt, L. A. (1979). Recognition of facial expressions by seven-Oster, H. & Eleman, P. (1978). Facial behavior in child development. In W. A. Collins (Ed.), Osterhout, L. & Holcomb, P. J. (1995). Event related potentials and language comprehension. InElectrophysiology of Mind: Event-related Brain Potentials andCognition. Oxford psychology series, No. 25
Phan, K. L., Wager, T., Taylor, S. F., & Liberzon, I. (2002). Functional neuroanatomy of emotion:NeuroimagePhelps, E., OConnor, K., Cunningham, W., Funayama, E., Gatenby, J., Gore, J., et al. (2000). Journal of Cognitive NeurosciencePicton, T. & Hillyard, S. (1988). Endogenous components of the event-related brain potential. InT. Picton (Ed.), Elsevier.R. J. (2002). Affective judgments of faces modulate early activity (approximately 160ms) withinNeuroimagePlant, A. & Devine, P. (1998). Internal and external motivation to respond without prejudice.Journal of Personality and Social PsychologyRicheson, J., Baird, A., Gordon, H., Heatherton, T., Wyland, C., Trawalter, S., et al. (2003). AnNature NeuroscienceRockstroh, B., Elbert, T., Lutzenberger, W., & Birbaumer, N. (1979). Slow cortical potentials underconditions of uncontrollability. Memory and CognitionShrifn, R. & Schneider, W. (1977). Controlled and automatic human information processing. II.Perceptual learning, automatic attending, and a general theory. Vaes, J., Paladino, M., Castelli, L., Leyens, J., & Giovanazzi, A. (2003). On the behavioral conse-Journal of Personality and Social PsychologyVrana, S. R. & Rollock, D. (1998). Physiological response to a minimal social encounter: Effectsof gender, ethnicity, and social context. Vrana, S. R. & Rollock, D. (2002). The role of ethnicity, gender, emotional content, and contex-tual differences in physiological, expressive, and self-reported emotional responses to imagery.
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understanding of the human brain. Initially, this growth relied on detailed models ofof brain signals observed in humans (Cohen, Noll, & Schneider, 1993). However, aswe move from investigating the neural mechanisms of simple perceptual, motor, andior differs fundamentally from the types of behavior typically observed in laboratoryanimals. Evidence suggests, however, that even in the study of complex social behav-although the content and complexity of stimuli may vary widely. By using animalClassical conditioning was rst described by Pavlov over a century ago, but more12Animal Models of Human Attitudes: Integrations Across
(fMRI), amygdala activation was observed during fear conditioning (Buchel et al.,1998; LaBar et al., 1998). Moreover, the strength of that activation is correlated withfear, humans with damage to this region fail to demonstrate a CR as assessed with SCRHuman beings, however, can also be asked to report on their experience of emotion.expression of conditioned fear, these same patients are able to report explicitly on theevents that constitute fear conditioning. For example, patient SP, who suffers fromblue square after a few pairings with the shock, an indication of conditioned fear. SP,however, never demonstrated a SCR to the blue square, even though her SCR to themild shock was normal. When shown her data indicating a lack of conditioned fear,very beginning, except for the very rst one when I was surprised. That was my reactionI knewit was going to happen. So I learned from the very beginning that is was going to happen: blue-distinct neural circuits. This discovery has obvious implications for dissociations that
are observed in the behavioral responses of humans to complex situations in whichIn particular, animal data on fear conditioning indicating a dissociation betweenof attitudes, whereas implicit measures of attitudes reveal large preferences. Or, explicitthe implicit association test (IAT), subjects see items that belong to one of four cate-(joy, love, friend versus devil, vomit, agony). For half the trials, the task involves clas-bad using another key. For the remainder of the trials, the opposite pairing isanother. The difference in response latencies in these two conditions constitutes theIAT effect and is regarded to be a measure of evaluative strength of association regard-ing these race groups. Responses on the IAT (Greenwald & Banaji, 1995) using largeown group compared to white Americans, but their IAT data show a much weakerAfrican-Americans, as opposed to 72 percent of whites, favor their own group.damage to either the hippocampus or amygdala. Similarly, these models offered a start-ing point to infer the mechanisms that underlie the dissociations in attitudes observedin normal humans. Moreover, human data on implicit-explicit dissociations in raceis the meaning of the associative strength observed in response latencies that reveal(the measure reects an effect of familiarity, not attitude; the measure taps somethingularly useful concerned a specic objection regarding the construct validity of the IAT.
The claim was that the IAT response was not a measure of evaluation or affect at all;portion of ones responses to any object.the IAT while also shedding light on the dissociation itself. To test this hypothesis, weof conditioned fear, might also mediate implicit expressions of race bias as measuredby both the IAT and a physiological assessment (Phelps et al., 2000). To the extentthat this is observed, behavioral data resulting from the IAT can be assumed to bewhile they observed pictures of Black and White unfamiliar male faces and correlatedobserved brain activity with two behavioral measures, the IAT and startle eyeblink.repeated. Afterward, they were given the two implicit measures of race bias; the IAT,Racism Scale (MRS; McConahay, 1986). The IAT effect was computed as a differencewere words that again, could be clearly classied as good or bad (love, joy, friend versusagony, vomit, devil).response is an eyeblink, the strength of which can be assessed in the laboratory by(Lang, Bradley, & Cuthbert, 1990). In our assessment of implicit race bias, subjectsVariability among the White American subjects was seen in the amygdala response
cant. We had expected overall greater amygdala activation to Black than to Whiteamygdala activation and the IAT, startle reex, and MRS. When amygdala activationIAT (exploratory analysis generated correlation maps indicating brain regions where behav-amygdala correlated with both the IAT and startle eyeblink measures of bias (gureThis study revealed a variety of results that can be the basis of future research. First,
was signicantly correlated with the strength of implicit race bias as assessed by the IAT (left)
observed amygdala activation in White Americans to Black versus White faces that arepresented subliminally, so quickly that subjects are unaware of their presentation andas fear. It also is implicated in fear extinction, in which a CS is no longer paired witha UCS and, eventually, the CS fails to elicit a CR. During initial extinction learning,extinction to specic individual members of a race group who are familiar. A furtherTo assess the effect of familiarity and exposure on the neural systems of race bias,male faces were not unfamiliar, but rather were well known gures from the popularmedia with relatively positive public images (Martin Luther King, John F. Kennedy,Denzel Washington, Harrison Ford). With these stimuli, we failed to nd consistentcategory as whole continues to elicit them.In the Phelps et al. (2000) study, we did not explore responses to the entire brain,or inhibition of the amygdala response. However, Cunningham et al. (2004) founding (Phelps et al., 2004). However, this was observed only when subjects were aware
bias. The precise regions of the PFC observed in these studies differ from thoseobserved in extinction learning (Phelps et al., 2004), but they are consistent with& Tropp, 2000). These results are consistent with the hypothesis that exposure,Thus far, we have argued that the neural systems of classical fear conditioning mightbe similar to the neural systems of culturally acquired race bias. However, a funda-A primary means of social learning in humans is languagea symbolic form of com-
investigate the verbal communication of fears is instructed fear. This is similar to fearmay, in the future, predict a possible aversive consequence is unlikely to elicit a strongemotional response at the time this information is learned. However, it is possible thatsubjects actually received a shock at any time. Afterward, all subjects indicated aware-SCR response to the blue square was increased, a physiological indication of fear.show a physiological indication of instructed fear, although they were able to report
Another powerful means of social learning is observation (Bandura, Ross, & Ross,observation has been demonstrated in infants viewing the emotional reactions of theirmothers (Gerull & Rapee, 2002), as well as in monkeys that observed reactions of otheriological expression of fear acquired through social (instruction and observation) andnonsocial (classical conditioning) means (Olsson & Phelps, 2004). In the observation(observational CS) that was paired with a mild shock to the wrist (observational UCS).the video. The subjects were then presented the observational CS, but never actually
Activation to instructed fear. A group composite map (left) and selected individual subjects (right)
physiological fear expression when the stimulus (conditioned, instructed, or observa-tional CS) that was paired with the aversive event was presented supraliminally, sofear conditioning and observational fear resulted in the physiological expression ofpresented. When an instructed CS was presented subliminally, there was no indica-tion of a fear response (Olsson & Phelps, 2004). Thus, observing another individualsvational fear suggests overlapping neural circuits. Although the acquisition of verballyexperience of observing another individuals aversive reaction to a UCS paired with aCS evokes an amygdala response. To investigate this, an fMRI study (Olsson et al.,2004) examined amygdala activation when observing a confederate receiving a mildvation occurred to both observing a confederate receive a shock and expecting toexpression of observational fear learning, similar to fear conditioning.Studies on the neural systems of instructed and observational fear learning indicateacquisition (observation) and expression (observation and verbal instruction) of preferences, which are learned through social communication and observation, to
that this is not the case. Monkeys raised in the laboratory that never had experience) is not and serves as a base-line, suggest that instances of some categories can serve as prepared stimuli. A specicprepared (Ebert et al., 2005). We asked to what extent can social groups that vary infaces in White and Black American subjects. We hypothesized that an outgroup face,from each race group served as a CS(UCS) during acquisition, while the other served as a CSan indication of fear conditioning. Regardless of subjects race, both Black and Whitestimuli showed evidence of acquisition of a CR. However, the CR to an out-
be more durable than an acquired negative response to an ingroup race member. UsingAs the research outlined in this chapter indicates, animal models can be quite usefullearning and memory that may be extensions of the processes that are common to conscious awareness, intention, and control to shape behavior. A human being, tion, even when hungry, to dine with a friend, to deciding to recode stimuli inhibition that are common across species (Phelps et al., 2004) is unclear. It is clear,however, that animal models can be useful tools in our investigations of the neural systems underlying human social behavior. They serve as the starting point
and NIMH and Rockefeller Foundations Bellagio Study Center to M. R. Banaji. WeBandura, A., Ross, D., & Ross, S. A. (1961). Transmission of aggression through imitation of aggres-Journal of Abnormal Social PsychologyBechara, A., Tranel, D., Damasio, H., Adolphs, R., Rockland, C., & Damasio, A. R. (1995). Doubleconditioning: An event-related fMRI study. NeuronCahill, L., Weinberger, N. M., Roozendaal, B., & McGaugh, J. L. (1999). Is the amygdala a locusNeuronCohen, J. D., Noll, D. C., & Schneider, W. (1993). Functional magnetic resonance imaging:Overview and methods for psychological research. Behavioral Research Methods, Instruments andCunningham, W. A., Johnson, M. K., Raye, C. L., Gatenby, J. C., Gore, J. C., & Banaji, M. R.Journal of Neuropsy-chiatry and Clinical Neuroscienceand Social Psychology.Falls, W. A., Miserendino, M. J., & Davis, M. (1992). Extinction of fear-potentiated startle: Journal of Neuroscienceaffective modulation of startle in humans: Effects of unilateral temporal lobectomy. Journal ofCognition NeuroscienceGerull, F. C. & Rapee, R. M. (2002). Mother knows best: Effects of maternal modelling on theBehaviour Research and Therapy
Hart, A. J., Whalen, P. J., Shin, L. M., McInerney, S. C., Fischer, H., & Rauch, S. L. (2000). Dif-Neurore-portJournal of Experimental PsychologyHygge, S. & Ohman, A. (1978). Modeling processes in the acquisition of fears: Vicarious electro-Journal Personal Social PsychologyKapp, B. S., Frysinger, R. C., Gallagher, M., & Haselton, J. R. (1979). Amygdala central nucleusKing, N. J., Eleonora, G., & Ollendick, T. H. (1998). Etiology of childhood phobias: Current statusof Rachmans three pathways theory. Behavior Research and TherapyLaBar, K. S., Gatenby, J. C., Gore, J. C., LeDoux, J. E., & Phelps, E. A. (1998). Human amygdalaactivation during conditioned fear acquisition and extinction: A mixed-trial fMRI study. NeuronLaBar, K. S., LeDoux, J. E., Spencer, D. D., & Phelps, E. A. (1995). Impaired fear conditioning fol-Journal of NeuroscienceLang, P. J., Bradley, M. M., & Cuthbert, B. N. (1990). Emotion, attention, and the startle reex.. New York: Simon & Schuster.Synaptic Self: How Our Brains Become Who We Are. New York: Viking.McConahay, J. P. (1986). Modern racism, ambivalence, and the modern racism scale. In J. F.Prejudice, Discrimination and RacismMineka, S. & Cook, M. (1993). Mechanisms involved in the observational conditioning of fear.Journal of Experimental PsychologyMineka, S., Davidson, M., Cook, M., & Keir, R. (1984). Observational conditioning of snake fearJournal of Abnormal PsychologyNeuroscienceLetterOchsner, K. N., Bunge, S. A., Gross, J. J., & Gabrieli, J. D. (2002). Rethinking feelings: An fMRIJournal of Cognitive NeuroscienceOhman, A. & Mineka, S. (2001). Fears, phobias, and preparedness: Toward an evolved module
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Marcus Aurelius was a thinking persons Roman emperor, and his many observations about society, leadership, life, and death that ll this diarylike textis one whose importance might be overlooked by the casual reader. Aurelius observed,insight that the world is what we make of it, and that our mental machinery con-structs our experience of the world. Second, they betray Aureliuss role as Romes com-more generally, to cripple even the heartiest of soldiers. Aurelius sought to understandhow one might be protected from the consequences of injury, both psychic and A full appreciation of Aureliuss insights may have never been more timely. Ourmodern landscape may lack Roman emperors and Roman legions, but it is full of every-day battles both personal and societal. We pursue professional success. We wrestle withromantic relationships. We confront conicted feelings. We intervene in intergroupconicts. We wage war on terrorism. Over two millennia ago a philosophical generalSwung one way, it serves your enemy, causing distress and pain. Swung another, itserves your champion, cutting a swath through fear, through grief, through false belief.Indeed, your mind is the best weapon you can carry into any battle, for with it you13Characterizing the Functional Architecture of Affect Regulation:
multidisciplinary research can provide answers to these questions that move beyondchology, researchers have begun constructing theories that link the experience of neg-together comprise a functional architecture for affect regulation.Although the terms emotion and affect often are used interchangeably by psycholo-iology and behavior, a useful distinction between them can be made. Emotion refers1999; Feldman Barrett, Ochsner, & Gross, in press; Lazarus, 1991). Emotions arethat trigger a response whose trajectory can be described and demarcated by an endexternal events and internal goals, wants, and needs (Scherer, Schorr, & Johnstone,conditions and consciously perceived referents, as well as moment-to-momenting to regulate a valenced response. As considered below, thinking or cognition canbe used to regulate ones positive or negative perception of another person or attitudeobject more generally, as well as to regulate ones own personal experience of emotion. The second is that neuroscience research is increasingly suggesting that aboth person perception and personal experience (Anderson et al., 2003; Anderson &
interpretations that lead an individual to perceive another person in a favorable, appraisals that lead an individual to feel happy, sad, angry, or glad. In keeping withAureliuss recognition of the pernicious consequences of psychological distress, inSince the late 1990s, evidence from both human and animal research has implicatedral lobes medial wall, in a number of functions intimately related to affective evalu-The amygdala seems to play a special role in the automatic, rapid, early, and evenbased on the discovery that a subcortical pathway from sensory organs to amygdala,2000). This suggestive nding was not directly tested, however, until functional(Morris, Ohman, & Dolan, 1999). More recent studies show that the amygdalasdala is important for consolidating explicit memory for emotionally arousing events.The Functional Architecture of Affect Regulation
epinephrine release in the amygdala eliminated the memory enhancement typicallyfound for the emotional components of a story. Subsequent imaging studies linkedtographs to subsequent memory for them (Hamann et al., 1999).More broadly, amygdala activation has been observed in response to a wide varietyperception of aversive or threatening stimuli in a positive direction, leading toimpaired recognition of fear faces, a tendency to perceive as friendly people norma-tively appearing to be unfriendly, and a tendency to classify normatively unpleasantimages as more pleasant (Adolphs, Tranel, & Damasio, 1998). As these data indicate,the amygdala may play a special role in perceiving social stimuli, a role further sup-ported by the nding that amygdala lesions impair perception of subtle expressions& Tranel, 2002), and that for Caucasian participants, amygdala activation to African-(Abercrombie et al., 1998), and accompanies evoked symptoms of anxiety, phobia, andBeyond the amygdala, a number of other richly interconnected brain structures playrepresenting the unpleasant properties of physical pain, and may more generally serveEisenberger & Lieberman, 2004; Ochsner et al., 2001; Peyron, Laurent, & Garcia-Larrea,
to alter a stimulus-reinforcer association once it is learned, and single-unit recordingevaluation that may bias perception of social targets and be a rst step in the gener-to understand the nature of ones despair or desire, and thereby be free from its grasp.Whether it is biblical tales of the very rst family and their inhibitory failures, a Romanemperors meditations on the epistemology of pain, the legend of Siddhartha and hisBuddhist renunciation of worldly wealth, stories of concentration camp survival, orgossip about the indiscretions of contemporary politicians, humans are always seekingcan impact emotion will survive empirical test, and what mechanisms are involved?constrain theories. Until recently, information about the brain bases of affect and The Functional Architecture of Affect Regulation
bases of simple behaviors (for discussion, see Ochsner, 2004; Ochsner & Lieberman,domains of neuroscience research relevant to questions about affect regulation. Cog-that at whatever stage they might intervene, control processes used to regulate areemotion and the specic kind of regulatory strategy employed (Ochsner & Gross, 2004,in retrieving information from semantic memory, holding information in mind, andmaintaining representations of strategic (e.g. regulatory) goals (Miller & Cohen, 2001).one important caveat: most studies investigated only the regulatory dynamicsThe use of attention to control the ow of sensory input long has been studied in thecontext of visual object recognition and vision more generally. Neuroimaging studiesby attention. To the extent that a brain regions activation does not vary as a func-
tion of the allocation of attentional resources, computations carried out by that regionconscious, on-line, attentional monitoring). Thus far, for simple nonaffective visualKanwisher, 2001). In the context of effectively charged stimuli, the question becomesditions of either full or diminished attention to the face stimuli. Two studies foundthat the amygdalas response to faces does not vary as a function of attentionalresource deployment. In the rst study (Vuilleumier et al., 2001), researchers askedtively specialized for processing face stimulidid vary as a function of attention, asanother, under conditions of full and divided attention. Each type of grayscale stim-the face or a feature of the house on each trial. As was the case for Vuilleumier et al.,The Functional Architecture of Affect Regulation
diminished, whereas the amygdalas response to disgust faces actually increased asattentional allocation to them diminished. This suggests that the tuning curve of theamygdala actually broadens when fewer attentional resources are available to identifydalas response prole changes in different circumstances.In direct contrast to these ndings, two studies observed amygdala modulation inticipants with fearful, happy, and neutral faces in the center of the screen and hadresponse to neutral faces. Similar results were obtained in a study employing a verysciously perceived neutral face of the same gender, which participants were instructedto view passively. During supraliminal presentations, fear faces activated the rightamygdala and disgust faces activated the insula bilaterally, but both activations dis-tion of affectively charged stimuli is necessary for the affective salience of the stimulussible leakage of affectively charged stimuli into awareness. This was very unlikely insion on subliminal trials was at chance. A problem for this account, however, is pre-sented by research of Cunningham, Johnson et al. (2004), who found greater amygdalasubliminally, but that this difference disappeared when faces were presented supral-
iminally. Although Cunningham et al. did not verify unawareness of subliminal stim-ulus presentations as rigorously as did Philips et al., which according to PhilipssPhilips et al.s account cannot explain failureregulate their responses to black faces when they could consciously perceive them (soresponses to consciously perceived fearful faces, which in experimental contexts maystimuli is observed. To the extent that low-level visual features simply are not beingencoded as one fails to attend to and perceive those features, it is not likely that theinformation necessary to discriminate affectively relevant information would be avail-the extent to which they can attend to the pain. Typically, reports of pain affect dropobserve this phenomenon. Participants experienced either a nonpainful cool or aThe Functional Architecture of Affect Regulation
observed, although it is not clear whether these regions were involved in active regu-neutral trials, and an activation interaction was observed between the presence ofTwo regions associated with cognitive control and awareness of affectrightdemonstrated that distraction can modulated thalamic responses, and Tracey et al.brainstem region known as the periacqueductal gray (PAG), which is important fordescending modulation of pain inputs from the spinal cord. Although Tracey et al.focused only on PAG activation, and so could not speak to the issue of which regionsmight have been involved in sending cortically based top-down regulatory instruc-tions, others observed PAG-prefrontal interactions that speak to this issue (Valet et al.,2004). Valet et al. employed a task essentially similar to that employed by Bantick et al., and like that group, observed that rostral medial prefrontal regions were moredistraction in the absence of pain. Although they did not report Banticks negativerelated thalamic nuclei as well as the PAG inversely correlated with activation of rostral
tive evaluation of performing a secondary task or diverting spatial attention, two otheried semantic, perceptual or emotional stimulus dimension, such as judging theinteractions, as discussed below.(1991) to describe our ability to alter the trajectory of a continuing emotional responsegoals, wants, or needs. In an early study, he showed that providing a stress-reducingquences of, and context for, reappraisal (see Gross, 1998, for review), until recentlyattend to and be aware of their emotional responses, but not to try and change them;ipants could reappraise an otherwise sad photograph of four women crying outside ofa church as involving a wedding rather than a funeral, which would be an occasionto aversive compared with neutral photographs dropped signicantly, as did subjec-The Functional Architecture of Affect Regulation
detached observers (Beauregard, Levesque, & Bourgouin, 2001). Right amygdala andinate among these possibilities. In a subsequent study, these investigators used aThinking is used to regulate feeling not only in the service of decreasing feelings,but in the service of increasing them as well. Indeed, when we worry, make ourselvesfacto up-regulate, or at least maintain, our negative emotion that might otherwisewith greater N). Schaefer et al. (2002) obtained consistent results by observing bilat-
mine which regulatory mechanisms are responsible for these effects.inferred straightforwardly from perceptual features.ulus presented at the top of the screen (Hariri, Bookheimer, & Mazziotta, 2000). In theperceptual condition, all three stimuli were faces that expressed anger or fear. In theexpression labels (i.e. the words angry and afraid). Right amygdala activation wasgreater for matching perceptually, whereas right ventral lateral prefrontal activationof negatively valenced stimuli, including the perception of African-American faces(Lieberman et al., unpublished observations) and aversive photographs (Hariri et al.,were noted when participants matched to labels compared with percepts. As a group,problem for this account, however, is that the perceptual condition in which moreamygdala activation is observed includes three faces, whereas the labeling conditionBe that as it may, explicit labeling of affective properties of stimuli down-regulateswere angry faces, neutral faces, or blank control stimuli (Nomura et al., 2004). Partic-ipants judged whether the consciously perceived target face seemed to be angry, happy,or neutral. In general, greater right amygdala activation was observed for trials withThe Functional Architecture of Affect Regulation
angry primes, and right amygdala activation was correlated with the tendency to judgethe target face as angry. Right ventral lateral PFC showed precisely the oppositeangry expressions. In the second, decreased amygdala activation was observed byTaylor et al. (2003) when participants rated the valence of aversive and neutral pho-tographs compared with viewing them passively. In the third, Critchley et al. (2000)had participants view happy, angry, or neutral faces, and judge either face gender orwhether or not each face was emotionally expressive. In general, perception of emotional compared with neutral faces activated the left amygdala, and strikingly,number of studies failed to observe modulation of amygdala responses when partici-For example, Gorno-Tempini et al. (2001) had participants judge either the genderor emotional expression of happy, disgust, or neutral faces. Although left amygdalaPFC. It should be noted, however, that because the authors did not report the gendermight have produced greater amygdala activation, as was observed by Critchley et al.(2000). This ambiguity of analysis was not a problem for a study in which happy, sad,expressive (Winston, ODoherty, & Dolan, 2003). Although increasing intensity ofexpression activated the amygdala bilaterally, in contrast to results of Critchley et al.(2000), amygdala activation did not vary with judgment type.ertheless expressing a high degree of anger), Winston et al. (2002) had participants
untrustworthy faces was observed that once again did not vary as a function of judg-ment type. Finally, Cunningham et al. (2003) had participants judge whether photosdid not vary as a function of judgment type.will take place? One possibility, suggested earlier in the context of attentional control,is that regulatory interactions between PFC and amygdala take place when one hasthe explicit motive or goal to regulate ones evaluative responses to stimulus dimen-instructed to control ones evaluation, participants may do so spontaneously, andsession, participants rated each word for affective intensity, good-bad valence, anddala activation. Critically, when making good-bad judgments, the motive to controlThe overarching question motivating this chapter has been, what are the neurocog-stock of what the preceding review suggests as an answer, or answers, to this question,The Functional Architecture of Affect Regulation
praisal, and alterations of construal more generally, can change the way we affectivelyevaluate a stimulus. Careful behavioral observation and empirical experimentation, as1.Interactions between prefrontal and cingulate systems that implement control pro-2.The simplest alternative explanation for the effects of cognitive control on affec-tive processingnamely, that engaging in cognitive processing, or thinking, dimin-with respect to variations in attention (Anderson et al., 2003) and judgment (Winston3.Down-regulation of negative affect by cognitively changing the meaning of a stim-nonconscious (and likely unregulated) perception of black faces (Cunningham et al.,4.Mechanisms mediating regulation of experience and regulation of perception-judgment may be highly overlapping. Virtually all studies investigating reappraisalthe way in which they made judgments about it. Virtually all studies involving the
made no reference to experience in any way, instructing participants simply to attendright PFC-OFC regions to mediate their regulatory impact on an affective outcomevation in both cases. It should be noted, however, that one reason regulation ofexperience and perception may recruit similar mechanisms is because most tasksinvolved both types of regulatory processing, and failed to discriminate them cleanly.Research on the neurocognitive mechanisms of affect regulation has only been aprimary focus of functional imaging research for the past ve to ten years, so it is1.Although some of the primary neural players in the cognitive regulation of affecthave been identied, the precise functional nature of their regulatory interactions withtypes of regulatory mechanisms: (1) processes involved in selecting appropriate,ations in input to the amygdala that either (a) cut off or limit the ow of perceptualsemantic and/or perceptual inputs that feed forward to the amygdala, indicating thetive evaluation. It remains for future research that manipulates the psychological2.A second related question concerns the ways in which different types of strategiesmay involve different types of regulatory interactions. Distraction compared withThe Functional Architecture of Affect Regulation
strategy. Strategies that involve an inward focus on oneself and ones emotional expe-regulation of pain experience (Bantick et al., 2002; Frankenstein et al., 2001; Valet 3.To what extent must the conscious goal or motive be present in order for regula-tory effects on the amygdala to be observed? Studies that explicitly gave participantsspontaneous tendency to regulate (Cunningham et al., 2004) consistently observedexplicit regulatory goals less consistently observed amygdala modulation. The simplefact that in everyday life thinking often inuences feeling without our having therequires the conscious goal to change how one feels or perceives the world. What kindof processing goal is necessary is not clear, however.4.Are the same control mechanisms and regulatory interactions involved in atten-tional control, cognitive change, and response modulation? To address this question,experiment, using the same participants and stimuli for each type of regulatory inter-explicitly behavior-focused affect-regulation strategy, such as suppression of emotionexpressive behavior.5.To what extent are regulatory dynamics such as those described here also involvedof as explicitly regulatory (for discussion, see Ochsner & Gross, in press)? Stimulus-
6.To what extent are mechanisms mediating affect regulation similar to, or differentvisual-spatial, and auditory information? In other words, to what extent are systemsworking memory (DEsposito, Postle, & Rypma, 2000), or response inhibition (Braver7.How do individual differences in the tendency to generate affective responses ofbroad question may include specic questions about the development of regulatoryin normal healthy populations. Our models of the functional architecture of affecttribute to emotional responding and regulatory success in different individualsThe capacity Marcus Aurelius identied over two millennia ago, for our estimates ofability to adapt to stressful circumstances. Clearly, a complete understanding of mech-very useful kind of explanation speaks to several levels of analysis, linking socioemo-verge on theories consistent with numerous methodologies is essential and necessary.With any luck, in the next decade our estimate of the progress we have made in char-acterizing the functional architecture of affect regulation will produce a greaterThe Functional Architecture of Affect Regulation
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In the distant future I see open elds for far more important researches. Psychology will be basedon a new foundation, that of the necessary acquirement of each mental power and capacity byCharles Darwin, When Charles Darwin wrote those words in 1859, at the end of his famous book, hewas presciently advocating the extension of evolutionary theory as applied to mor-tionary psychology have indeed taken up the baton, albeit with agendas that not evenDarwin could have anticipated. The detailed claims of these disciplines remaindebated, but their central question is as clear and urgent as ever. What distinguishesan answer; however, it is equally easy to point out some specic facts that providepurchase. Take the observation by Richard Passingham (1982) that, Our species isearth. That observation is patently correct. by natural selection as Darwin had in mind. So how has this dramatic change beenThe answer, of course, lies with a different form of evolution than the one based on14What Is Special about Social Cognition?
we nd Michael Tomasello (1999) writing, The basic fact is thus that human beingsare able to pool their cognitive resources in ways that other species are not...madepossible by a single very special form of social cognition, namely, the ability of human mind special remains popular. One idea has been that intragroup or intergroupbehavior. Both tactical deception (Whiten & Byrne, 1997) and social cooperativity areour distinctive cognitive abilities (Dunbar, 1998)?We humans think of other people as having minds, having experiences, feelings,do. Broadly, this capacity has been dubbed theory of mind, and there has been con-thing similar (Povinelli & Vonk, 2003; Premack & Woodruff, 1978; Tomasello, Call, &Hare, 2003). This is an issue on which Tomasello himself recently changed his mind.Despite its prima facie plausibility, the idea still leaves unresolved the question of thischapter. Granted that our social cognitive abilities are special: are they the reason ourThere is no shortage of examples of the latter possibility. Our ability to conceive ofother points of view. Thus, we have the mental time travel hypothesis (Suddendorfand imagining the future. Conceiving of ourselves from alternative points of view,ior, our interactions with other people. Of course, in some sense nearly all informa-
tion is relevant for social behavior, making social cognition in that broad sense toogeneral a term to be useful. Researchers generally restrict the term to that informationdirectly perceived from, or inferred about, other people. A further restriction focusesSocial cognition, then, concerns perception of, attention to, memory for, and think-processing. Examples include processing other peoples faces and voices, judging theirpersonality, predicting their likely behavior, and planning our own interactions withthem. In one typical example, it begins with the sensory processing of social infor-mation (visual perception of someones face or body posture); proceeds to formationand behavior. Although pervasive in the sense that it has the potential to inuenceessentially all aspects of cognition and behavior, social cognition draws on a circum-varies considerably. Even among primates, a highly social group of mammals, largedifferences range from the essentially solitary existence of orang-utans to closeinteractions; but we also regularly drive our car, memorize phone numbers, and planThis distinction was formalized in the concept of modularity. First detailed by thephilosopher Jerry Fodor (1983), modularity has come to mean several things, and itsWoodward, 2004). Without going into detail, we can itemize the following attributes
Are Best understood, from the view of cognitive psychology, as a distinct class ofare not informationally encapsulated, to use Fodors original term.of social cognition is considered together, certainly components appear stronglymodular. Pheromonal signaling by way of the vomeronasal system (a pathway thatmodular. Language appears highly modular. All of these are components, but theis, pheromone reception and speech perception might both be modular insofar as
evolutionary psychologys controversial idea that the mind is massively modularthrough and through. That is the Wason selection task (Barkow, Cosmides, & Tooby,Debate continues about this particular example, but the Wason selection task aside,plenty of counterexamples exist in which certain quite circumscribed aspects of socialTwo good examples of current study in cognitive neuroscience are the social cogni-and lesions in both can result in disproportionate impairments in social behavior.and how we generate empathy. Again, the basic mechanism appears to be derivativeto more general aspects of motor control and imagery.(Broadmans area 10) are disproportionately larger in humans than in any other ape(Semendeferi et al., 2001). It is thought that this region of the brain subserves long-competences that Tomasello, Corballis, and others claimed to exist only in humans.
that these cells serve rapid signaling of social information during error detection.in social behavior. Their social decision making is poor, their social relationships withof the standard cognitive IQ (Bar-On et al., 2003; cf. Bechara this volume). In tryingbehavior after sustaining such a lesion (Barrash, Tranel, & Anderson, 2000). Severalemotional experience, apathy, inappropriate affect, and lack of planning.cortex. Some of these paradigms are specically social, others are not. Two regions ofrejection, for instance. Yet reasons for activation in social experiments are presumedgulate cortex is activated by the pain of social rejection (Eisenberger, Lieberman, &Williams, 2003) or by observing someone else in pain (Singer et al., 2004), yet playsa general role in pain processing. In fact, it has a very general role in detecting any
by observing others express disgust, as well as by experiencing pain, disgust, and otherThe second structure, the amygdala, presents us with a parallel story. In the 1930s,cial behaviors such as approach tendencies toward novel objects (Emery et al., 2001).et al., 2000; Calder et al., 1996; Young et al., 1996), complemented by functionalnition of expressions of fear, a conclusion supported also by some functional imagingwas raised by observing that some patients with complete bilateral amygdala damage
impaired in the same way, on the same tasks, or on the same emotions (Adolphs, 1999;The amygdalas role is not limited to making judgments about basic emotions, buthuman primates (Kling & Brothers, 1992; Kluver & Bucy, 1937; Rosvold, Mirsky, &lesions, and by using more sophisticated ways of assessing social behavior (Emery &Amaral, 1999; Emery et al., 2001), and they have been shown also in humans. Build-called theory of mind abilities: a collection of abilities whereby we attribute internal2000; Fine, Lumsden, & Blair, 2001). Three sets of studies from our laboratory cor-In one study, we asked subjects to judge how much they would trust, or how muchthey would want to approach, an unfamiliar person (Adolphs, Tranel, & Damasio,they performed normally when judging people who looked very trustworthy andndings were corroborated by functional imaging in normal individuals (Winston to look untrustworthy, amygdala activation increases, compared with activation whenthey look at people judged to look trustworthy. Moreover, this pattern of activationments about trustworthiness or making an unrelated judgment. Thus, the amygdalassocial judgments, may be fairly automatic, rapid, and obligatory.Heider, who designed short video clips depicting geometric shapes moving on a whiteotherwise look social), normal subjects immediately make social attributions to such
easier, and subjects normally cannot help but to see the stimuli in social terms. Byhow to go about answering this question, but one study (Adolphs, Tranel, & Baron-jealousy, pride, or embarrassment, necessarily require a social context and require aconcept of a social self that is situated within a social group. Two subjects with com-it is to guide responses to food, water, or electric shock and the nonsocial stimuli withwhich these reinforcers can be associated. However, in primates, and especially inworse and more pervasive than their impairments in making simpler emotional judg-
social behavior. Perhaps the human amygdala does not play a role in social cognitionnonmotivational property.) Preliminary arguments can be made both in favor of and against this view.ity of social groups (Barton & Aggleton, 2000). A different theory apparently againstthe idea comes from the observation that monkeys with selective bilateral amygdalacontra Kluver and Bucy, a normal repertoire of social behaviors, if those can be trig-gies that depend on very many different neural structures, of which the amygdala isone. One strategy relies on recruiting basic motivational and emotional circuitry, butlanguage and on declarative memory stores, may not rely on the amygdala.Rather than trying to establish that the above data argue for the domain specicity ofon a host of cognitive resources, and to ask instead in which way social and nonso-cial information processing support one another. A rich example adumbrated abovewhat it is like to be another person whom we are observing. Interest in the mecha-selves from the observation of another persons behavior. The ability may derive from
basic motor control adaptations, and yet may have expanded vastly in the service ofthe need to predict social behavior accurately.other peoples emotional states, at least in part, through some kind of articulated emu-lation (see Grush, 2004, for a review). Premotor cortices are engaged when we observeothers behaving emotionally, as are somatosensory cortices and insula. One interpre-tation of these ndings is that we engage some of the same machinery during emu-supports this idea: observing other people express emotions results in some mirroringof the emotional state in the viewer. In this case it seems that the emulator is the samewhen we model another persons emotion would be not only economical, but sug-neurally.Typically, of course, emulation of emotion should be less than the real thing. Thusthe external environment and an organisms interaction with it. Three interestinglikely to be a very dynamic, iterative process. It seems unlikely that we obtain all thelator, perhaps to different depths of detail, to approximate the answer we seek. Second,are hierarchically structured in some way. One could imagine emotion emulation
modeling required, much as visual imagery involves different levels of the visual hier-archy, depending on the grain of the imagery. And third, we can well imagine extend-ing the modeling outside the bounds of the body. To obtain social information, wemay query not only our own bodies, but other peoples. Clearly, this is the case in ageneral sense: we probe other peoples reactions to initial and often subtle behaviorsrules to models entirely internal to the brain or encompassing varying degrees of thecourse that draws on both. Social cognition is too broad a capacity, and makes contactsulated, impenetrable module. Yet it is reasonable to think that it comprises compu-social computations are relatively involuntary, automatic, and below the level of ourIn redening the question, it may be necessary to abandon our predilection fornot conversely. The assumption is that we could impair social knowledge while sparingthan another. Why not suppose that social cognition is the basic adaptive package,social and nonsocial cognition. Is one reducible to the other? Well, some components
to the question of this chapter: the level of analysis of a system. At a very molar other people rather than objects. At a very microscopic level, social cognition is question. Psychologically, behaviorally, aspects of it are indeed likely to be quitebiologically, it will be more difcult to nd similar evidence, since the special behav-in a much broader context within which they contribute to behavior. That context,that a particular social cognitive ability, or a particular aspect of social behavior,are for social cognition. Rather, we should consider social cognition as emergingNeuroscientistAdolphs, R., Tranel, D., & Baron-Cohen, S. (2002). Amygdala damage impairs recognition of socialJournal of Cognitive Neuroscience
Adolphs, R., Tranel, D., & Damasio, A. R. (1998). The human amygdala in social judgment. NatureAdolphs, R., Tranel, D., Damasio, H., & Damasio, A. (1994). Impaired recognition of emotion NatureAdolphs, R., Tranel, D., Hamann, S., Young, A., Calder, A., Anderson, A., et al. (1999). Recogni-NeuropsychologiaAllman, J. M. (2002). Two phylogenetic specializations of the human brain. NeuroscientistAmaral, D. G., Capitanio, J. P., Jourdain, M., Mason, W. A., Mendoza, S. P., & Prather, M. (2003).NeuropsychologiaAnderson, A. K., Spencer, D. D., Fulbright, R. K., & Phelps, E. A. (2000). Contribution of NeuropsychologyBarkow, J. H., Cosmides, L., & Tooby, J. (Eds.). (1992). The Adapted Mind: Evolutionary Psychologyand the Generation of Culture.New York: Oxford University Press.Baron-Cohen, S., Ring, H. A., Bullmore, E. T., Wheelwright, S., Ashwin, C., & Williams, S. C. R.(2000). The amygdala theory of autism. Neuroscience and Biobehavioral ReviewsBar-On, R., Tranel, D., Denburg, N., & Bechara, A. (2003). Exploring the neurological substrateBarrash, J., Tranel, D., & Anderson, S. W. (2000). Acquired personality disturbances associatedDevelopmental Neuropsychology, 2nd ed. (pp. 479508). New York: Oxford University Press.Breiter, H. C., Etcoff, N. L., Whalen, P. J., Kennedy, W. A., Rauch, S. L., Buckner, R. L., et al. (1996).NeuronCalder, A. J., Young, A. W., Rowland, D., Perrett, D. I., Hodges, J. R., & Etcoff, N. L. (1996). Facialemotion recognition after bilateral amygdala damage: Differentially severe impairment of fear.Cognitive NeuropsychologyTrends in Cognitive Sciences
Cowie, F. & Woodward, J. F. (2004). Mental modules did not evolve by natural selection. In C.Great Debates in Philosophy: Philosophy of Science.New York: Blackwell.the body. Nature Reviews NeuroscienceDescartes Error: Emotion, Reason, and the Human Brain.New York:Darwin, C. (1859). London: John Murray.ior in the free ranging rhesus monkey. Dunbar, R. (1998). The social brain hypothesis. Evolutionary AnthropologyEisenberger, N. I., Lieberman, M. D., & Williams, K. D. (2003). Does rejection hurt? An fMRIEmery, N. J. & Amaral, D. G. (1999). The role of the amygdala in primate social cognition. In Cognitive Neuroscience of EmotionEmery, N. J., Capitanio, J. P., Mason, W. A., Machado, C. J., Mendoza, S. P., & Amaral, D. G.Behavioral NeuroscienceFine, C., Lumsden, J., & Blair, R. J. R. (2001). Dissociation between theory of mind and exec-Fodor, J. A. (1983). Grush, R. (2004). The emulation theory of representation: Motor control, imagery and percep-Hamann, S. B., Stefanacci, L., Squire, L. R., Adolphs, R., Tranel, D., Damasio, H., et al. (1996).Naturesocial knowledge and perception. Proceedings of the National Academy of Sciences of the United StatesHeider, F. & Simmel, M. (1944). An experimental study of apparent behavior. American Journal ofKling, A. S. & Brothers, L. A. (1992). The amygdala and social behavior. In J. P. Aggleton (Ed.),The Amygdala: Neurobiological Aspects of Emotion, Memory, and Mental Dysfunction.New York:WileyLiss.Kluver, H. & Bucy, P. C. (1937). Psychic blindness and other symptoms following bilateral tem-American Journal of Physiology
Kluver, H. & Bucy, P. C. (1939). Preliminary analysis of functions of the temporal lobes inArchives of Neurology and PsychiatryMorris, J. S., Frith, C. D., Perrett, D. I., Rowland, D., Young, A. W., Calder, A. J., et al. (1996). ANatureNimchinsky, E. A., Gilissen, E., Allman, J. M., Perl, D. P., Erwin, J. M., & Hof, P. R. (1999). A neu-Proceedings of the National Academy ofPaus, T. (2001). Primate anterior cingulate cortex: Where motor control, drive and cognitionNature Reviews NeurosciencePovinelli, D. J. & Vonk, J. (2003). Chimpanzee minds: Suspiciously human? Trends in CognitivePremack, D. & Woodruff, G. (1978). Does the chimpanzee have a theory of mind? Rapcsak, S. Z., Galper, S. R., Comer, J. F., Reminger, S. L., Nielsen, L., Kaszniak, A. W., et al. (2000).NeurologyRosvold, H. E., Mirsky, A. F., & Pribram, K. (1954). Inuence of amygdalectomy on social behav-Journal of Comparative and Physiological PsychologySchmolck, H. & Squire, L. R. (2001). Impaired perception of facial emotions following bilateralNeuropsychologySemendeferi, K., Armstrong, E., Schleicher, A., Zilles, K., & Van Hoesen, G. W. (2001). PrefrontalAmerican Journal of Physical Anthro-Singer, T., Seymour, B., ODoherty, J., Kaube, H., Dolan, R. J., & Frith, C. D. (2004). Empathy forpain involves the affective but not sensory components of pain. Stone, V. E., Cosmides, L., Tooby, J., Kroll, N., & Knight, R. T. (2002). Selective impairment ofProceedings ofSuddendorf, T. & Corballis, M. (1997). Mental time travel and the evolution of the human mind.Tomasello, M. (1999). Cambridge: Harvard University Press.Tomasello, M., Call, J., & Hare, B. (2003). Chimpanzees understand psychologial statesTheTrends in Cognitive Science
Whalen, P. J., Shin, L. M., McInerney, S. C., Fischer, H., Wright, C. I., & Rauch, S. L. (2001). Afunctional MRI study of human amygdala responses to facial expressions of fear versus anger.Whiten, A., & Byrne, R. W. (Eds.). (1997). Whiten, A., Goodall, J., McGrew, W. C., Nishida, T., Reynolds, V., Sugiyama, Y., et al. (1999). NatureWinston, J. S., Strange, B. A., ODoherty, J., & Dolan, R. J. (2002). Automatic and intentionalNature NeuroscienceYoung, A. W., Hellawell, D. J., Van de Wal, C., & Johnson, M. (1996). Facial expression process-ing after amygdalotomy. Neuropsychologia
William James, in his monumental two-volume work devotes two insightful chapters to the brain and its relationship to behavior. The pre-...complexof the relations of mind and brain must show how the elementary ingredients of theformer correspond to the elementary functions of the latter (page 28, vol. 1).by no less a neuroscientist than Sir Charles Sherrington (1906) who wrote, ...physi-ology and psychology, instead of prosecuting their studies, as some now recommend,more strictly apart one from another than at present, will nd it serviceable for eachary hitherto (page 385). Although it is clear that progress was made from the time ofpart of the twentieth century.15Social Neuroscience: A PerspectiveMarcus E. Raichle
(1928; for translation, see Brugger, 1997). For many years such a vision seemed merefantasy.to realize Pavlovs vision of monitoring human brain function in a safe yet increas-ingly detailed and quantitative way.neuroscience (for a review, see Posner & Raichle, 1994).Charitable Trusts initiated their program in cognitive neuroscience, were faculty posi-nity, however, comes with a clear requirement. Social scientists and neuroscientistsa very multidisciplinary work in progressin its development. It must be kept in mind that this eld is itself in a very activesurvey, see Raichle, 2003b).To begin, it is useful to consider the intended goal of functional brain imaging. Thismay seem self-evident to most. Yet, interpretations frequently stated or implied aboutMarcus E. Raichle
version of nineteenth-century phrenology (for alternative views, Posner, 2003; Raichle,2003b). In this regard, it is worth noting that functional imaging researchers them-tion of the human brain from both microscopic and macroscopic points of view,discovery of modern imaging technology (1909 to be exact). He said, Indeed, recentlytheories have abounded which, like phrenology, attempt to localize complex mentalactivity such as memory, will, fantasy, intelligence or spatial qualities such as appre-ciation of shape and position to circumscribed cortical zones. He went on, These...One cannot think of their taking place in any other way thanous elementary activities....in each particular case [these] supposed elementary func-...Such activities are...always the result...of the function of a largenumber of suborgans distributed more or less widely over the cortical surface...(Garey, 1994, pp. 254255).With this prescient admonition in mind, the task of functional brain imaging wouldseem to be clear. First, identify the network of regions of the brain and its relation-way in the functional imaging community and is complemented by a long history ofneurophysiological and neuroanatomical studies in laboratory animals. Second, anddenitely more challenging, identify the elementary operations performed withinbe seen in different tasks, indicating the presence of elementary operations that are
manner in which the brain allocates its nite processing resources to the accom-this observation comes from studies in invertebrate neurobiology, in which ensem-bles of neurons were observed to recongure their relationships depending on the taskat hand, thus multiplying the potential of nite resources to serve the needs of theorganism (Marder & Weimann, 1991). It would not be too surprising if this rather sensible strategy had been conserved in an expanded version in the vertebrate brain.Obviously, success in pursuing information of this type will require very careful taskmight be expected in particular areas (i.e., how elementary is elementary when seekingresults and discussions about the specicity of face perception in the human brain (forcontrasting views, see Duchaine et al., 2004; Grill-Spector, Knouf, & Kanwisher, 2004;Hanson, Matsuka, & Haxby, 2004; Haxby et al., 2001). We must be careful in adjudi-related to emotion are suspected (Bush, Luu, & Posner, 2000; Drevets et al., 1997;Mayberg et al., 1999; Simpson et al., 2001). Extensions of detailed cytoarchitectonicsmaps from monkey to human (Gusnard et al., 2003; Ongur, Ferry, & Price, 2003) illus-role in future research.within the brain and circulatory and metabolic signals that are observed with func-tional imaging techniques (Lauritzen, 2001; Logothetis et al., 2001). To put this workMarcus E. Raichle
associated with the input to cells that are reected in very complex electrical signals(often called local eld potentials) generated by millions of tiny but very importantmeasure are different. Whereas they may at times correlate nicely, they are not causallyboth neurophysiologists and social neuroscientists have necessary but not sufcienttive neuroscience and functional brain imaging, an early worry was that individualacross individuals. The results were stunning (Fox et al., 1988; for a historical review,roscience, and with great success. However, for all who have examined such data in
chiatry, we are poised to make major advances in this area that should certainly be ofstones associated with the maturation of the human brain. Missing, however, is sat-isfactory understanding of the maturation process within systems of the human brain.Whether it is the development of attention, language, memory, management of dis-tress, or personality development more generally, we need information about brainonly the challenge of safely and accurately accessing the necessary information inapproach. Tools available to cognitive neuroscientists are becoming available thatallow safe access to the information in humans. Various imaging techniques applica-pioneering investigators (Dehaene-Lambertz, Dehaene, & Hertz-Pannier, 2002; McKinstry et al., 2002; Miller et al., 2003; Munakata, Casey, & Diamond, 2004; Finally, it will be critically important to maintain a sense of proportion when it comeschanges observed in functional imaging studies are very small. Although this highMarcus E. Raichle
1999), it has only recently received the attention that it deserves (for a recent briefreview, see Raichle & Gusnard, 2002), primarily because of the realization that mostnor an observed output? Chapter 3 presents experimental observations that provideinteresting avenues for future research in this important area.one to the view, long espoused by Llins (1974, 2001; Llins & Pare, 1991), that thebrain operates intrinsically with sensory information, often in an impoverished form,modulating rather than informing the system. As William James (1890) observed,whilst part of what we perceive comes through our senses from the object before us,another part (and it may be the larger part) always comes...out of our own headhave appeared (Kuhl, 2003; Meltzoff & Decety, 2003). As well, an experiment in theabsence of sensory input, the cat visual cortex appeared to be creating, in its sponta-neous activity, representations of anticipated visual stimuli (Kenet et al., 2003). Theseand other observations led us to endorse the general view that the brain develops andactivity is an internal representation of that model with which sensory informationTaking a lesson from the immense success of cognitive neuroscience, those interestedbrain imaging. These new researchers must not only understand the major questions
issues of interest to social scientists. We will all be the beneciaries of the success of1.Like most things in science, the introduction of brain imaging had important antecedentsVergleichende lokalisationlehre der grosshirnrindeBrugger, P. (1997). Pavlov on neuroimaging. Journal of Neurology, Neurosurgery and PsychiatryBush, G., Luu, P., & Posner, M. I. (2000). Cognitive and emotional inuences in anterior cingu-Trends in Cognitive SciencesCacioppo, J. T. & Berntson, G. G. (1992). Social psychological contributions to the decade of theClark, D. D. & Sokoloff, L. (1999). Circulation and energy metabolism of the brain. In G. J. Siegel,B. W. Agranoff, R. W. Albers, S. K. Fisher, & M. D. Uhler (Eds.), Basic Neurochemistry. Molecular,Brain Work: The Coupling of Function, Metabolism and BloodDehaene-Lambertz, G., Dehaene, S., & Hertz-Pannier, L. (2002). Functional imaging of speechperception in children. Drevets, W. C., Price, J. L., Simpson, J. R., Jr., Todd, R. D., Reich, T., Vannier, M. W., et al. (1997).NatureDuchaine, B. C., Dingle, K., Butterworth, E., & Nakayama, K. (2004). Normal greeble learning inNeuronFox, P. T., Mintun, M. A., Rieman, E. M., & Raichle, M. E. (1988). Enhanced detection of focalJournal of Cerebral Blood Flow and MetabolismGarey, L. J. (1994). Brodmanns Localization in the Cerebral Cortex(L. J. Garey, Trans.). London:Grill-Spector, K., Knouf, N., & Kanwisher, N. (2004). The fusiform face area subserves face per-ception, not generic within-category indentication. Nature NeuroscienceMarcus E. Raichle
Gusnard, D. A., Ollinger, J. M., Shulman, G. L., Cloninger, C. R., Price, J. L., Van Essen, D. C., et al. (2003). Persistence and brain circuitry. Proceedings of the National Academy of Sciences of Hanson, S. J., Matsuka, T., & Haxby, J. V. (2004). Combinatorial codes in ventral temporal lobeNeuroimageHaxby, J. V., Gobbini, M. I., Furey, M. L., Ishai, A., Schouten, J. L., & Pietrini, P. (2001). Distrib-Illes, J., Krischen, M. P., & Gabrieli, J. D. E. (2003). From neuroimaging to neuroethics. NatureNeuroscienceJames, W. (1890). (pp. 9799). New York: Henry Holt.Kenet, T., Bibitchkov, D., Tsodyks, M., Grinvald, A., & Ariell, A. (2003). Spontaneously emergingNatureKuhl, P. K. (2003). Human speech and birdsong: Communication and the social brain. Proceed-ow. Journal of Cerebral Blood Flow and MetabolismLlins, R. (1974). La forme et la fonction des cellules nerveuses. RechercheI of the Vortex: From Neurons to SelfNeuroscienceLogothetis, N. K., Pauls, J., Augath, M., Trinath, T., & Oeltermann, A. (2001). Neurophysiologi-NatureMarder, E. & Weimann, J. M. (1991). Modulatory control of multiple task processing in the stom-atogastric nervous system. In J. Kien, C. McCrohan, & B. Winlow (Eds.), Neurobiology of MotorProgram Selection: New Approaches to Mechanisms of Behavioral Choice(pp. 319). Manchester, U.K.:Mayberg, H. S., Liotti, M., Brannan, S. K., McGinnis, S., Mahurin, R. K., Jerabek, P. A., et al. (1999).American Journal of PsychiatryMcKinstry, R. C., Mathur, A., Miller, J. H., Ozcan, A., Snyder, A. Z., Schefft, G. L., et al. (2002).Cerebral CortexMeltzoff, A. N. & Decety, J. (2003). What imitation tells us about social cognition: A rapproche-Philosophical Transactions
Miller, J. H., McKinstry, R. C., Philip, J. V., Mukherjee, P., & Neil, J. J. (2003). Diffusion-tensorAmerican Journal of RoentgenologyMunakata, Y., Casey, B. J., & Diamond, A. (2004). Developmental cognitive neuroscience: ProgressTrends in Cognitive SciencesNichols, M. J. & Newsome, W. T. (1999). The neurobiology of cognition. NatureJ. S. Werner (Eds.), The Visual NeurosciencesOngur, D., Ferry, A. T., & Price, J. L. (2003). Architectonic subdivisions of the human orbital andJournal of Comparative NeurologyPosner, M. I. (2003). Imaging a science of mind. Trends in Cognitive SciencesPosner, M. I. & Raichle, M. E. (1994). . New York: Freeman.Raichle, M. E. (2000). A brief history of human functional brain mapping. In A. W. Toga & J. C.Journal of Neuro-Journalof NeuroscienceRaichle, M. E. & Gusnard, D. A. (2002). Appraising the brains energy budget. Proceedings of theSchlagger, B. L., Browm, T. T., Lugar, H. M., Visscher, K. M., Miezen, F. M., & Petersen, S. E. (2002).Schwartz, W. J., Smith, C. B., Davidsen, L., Savaki, H., Sokoloff, L., Mata, M., et al. (1979). The Integrative Action of the Nervous System.New Haven: Yale UniversitySimpson, J. R. J., Drevets, W. C., Snyder, A. Z., Gusnard, D. A., & Raichle, M. E. (2001). Emotion-induced changes in human medial prefrontal cortex. II. During anticipatory anxiety. ProceedingsUttal, W. R. (2001). The New PhrenologyMarcus E. Raichle
California Institute of TechnologyTufts UniversityHarvard UniversityUniversity of TexasGary G. BerntsonJoan Y. ChiaoHarvard UniversityHarvard UniversityWashington UniversityTiffany A. ItoMalia F. MasonJason P. MitchellDartmouth College and HarvardNew York UniversityMarcus E. RaichleWashington University School ofHarvard University
Valerie E. StoneEve Willadsen-Jensen
Adaptability, 29, 30, 33Allport, Gordon W., 189, 203Anxiety, 180Aurelius, Marcus, 245Bar-On emotional quotient inventory. Emotional quotient inventoryBelief-desire psychology, 110112
Boxological models of behavior, 136alsospecic topicsCategory error, 3Computer metaphor, xiCortical modularity. Crying. Laughing and cryingDarwin, Charles, 269Distributed circuits, 36Emotional quotient inventory (EQ-i), 2729,insular-somatosensory cortex and, 2022neuroanatomy, 1416primary and secondary inducers of, 18, 21,Empathy, 161. Social categoryEvolutionary context of social cognition, 103,
theory of mind and, 115118Faux Pas Recognition Task, 117Fear. Flexibility, 29, 30, 33Gender. Identifying people and their behavior, 9092Implicit association test (IAT), 231232Insular-somatosensory cortex, lesions of,Intentionality, 179, 182n1James, William, 287, 293other, 132activity, 139141Language research, 131132, 134Laughing and crying, pathologic, 2526Left temporoparietal junction (LTPJ), 89, 93,
history, 23Memory, 30, 31, 33working, and social category information, 194inferring another persons, 6869theory of mind and, 112122Metarepresentation-recursion and theory ofModularity, 133134Neural circuits and networks, 135, 136person-perception processes and, 160161self-perception processes and, 158159theory of mind and, 117120Pavlov, Ivan, 287288Perception, 30, 31, 3334Periacqueductal gray (PAG), 254LTPJ and, 9798Pretend play, 109Proximity, corollary of, 7Pseudobulbar palsy, 2526
boundary conditions on attention to,Reexivity. Representing people and their behavior, 9092as neural substrate theory of mind, 9293agents and their behavior, 9092Self-perception, accuracy of, 158159Self-perception processesSelf-regulation, role of self-perception andperson perception in, 157161Social category information, perception of,boundary conditions on attention to raceand gender, 196198See alsospecic topicsevolutionary story for the link, 169170Social perception. See alsospecic topicsERPs used to study, 189190person-perception processes and, 160self-perception processes and, 158,
Somatosensory cortex. somatosensory cortexSpeech perception, 142143, 145146. Stereotyping and behavior, 198199,Social category informationTemporoparietal junction (TPJ), 120, 121. Theory of mind (ToM), 85, 98, 270. pretend play, 109neuroscience research on, 114122Theory of mind (ToM) stories/tasks, alsospecic regionsguiding social behavior, 7375inferring another persons current mentalTime. Tomasello, Michael, 270Tone discrimination, 138139Ventral striatum, 248Ventromedial prefrontal (VM) region, 2225,Vertex positive potential (VPP), 213, 214Visual system, 175Williams syndrome, 75