A Cultural Neuroscience Approach to the Biosocial Nature ...cscn-pku.com/pubs/2013 Han et al Annu Rev Psych reprint.pdfPS64CH13-Han ARI 8 November 2012 10:8 A Cultural Neuroscience

PS64CH13-Han ARI 8 November 2012 10:8

A Cultural NeuroscienceApproach to the BiosocialNature of the Human BrainShihui Han,1 Georg Northoff,2 Kai Vogeley,3,4

Bruce E. Wexler,5 Shinobu Kitayama,6

and Michael E.W. Varnum1

1Department of Psychology, Peking University, Beijing, 100871, People’s Republic ofChina; email: [email protected], Brain Imaging and Neuroethics, Institute of Mental Health Research University ofOttawa, Ottawa ON K1Z 7K4, Canada3Department of Psychiatry, University of Cologne, Cologne 50924, Germany4Institute of Neuroscience and Medicine, Cognitive Neuroscience (INM3), ResearchCenter Juelich, Juelich 52425, Germany5Department of Psychiatry, Yale University, New Haven, Connecticut 065196Department of Psychology, University of Michigan, Ann Arbor, Michigan 48109-1109

Annu. Rev. Psychol. 2013. 64:335–59

First published online as a Review in Advance onSeptember 17, 2012

The Annual Review of Psychology is online atpsych.annualreviews.org

This article’s doi:10.1146/annurev-psych-071112-054629

Copyright c© 2013 by Annual Reviews.All rights reserved

Keywords

cultural neuroscience, culture, brain imaging, human brain, race

Abstract

Cultural neuroscience (CN) is an interdisciplinary field that investigatesthe relationship between culture (e.g., value and belief systems and prac-tices shared by groups) and human brain functions. In this review wedescribe the origin, aims, and methods of CN as well as its conceptualframework and major findings. We also clarify several misunderstand-ings of CN research. Finally, we discuss the implications of CN findingsfor understanding human brain function in sociocultural contexts andnovel questions that future CN research should address. By doing so,we hope to provide a clear picture of the CN approach to the humanbrain and culture and to elucidate the intrinsically biosocial nature ofthe functional organization of the human brain.

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Contents

INTRODUCTION . . . . . . . . . . . . . . . . . . 336WHAT IS CULTURAL

NEUROSCIENCE?. . . . . . . . . . . . . . . 336Origin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 337Key Concepts . . . . . . . . . . . . . . . . . . . . . . 338Aims . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 340Methods. . . . . . . . . . . . . . . . . . . . . . . . . . . 342Findings . . . . . . . . . . . . . . . . . . . . . . . . . . . 344

WHAT IS CULTURALNEUROSCIENCE NOT? . . . . . . . . 351

IMPLICATIONS OF CULTURALNEUROSCIENCE FINDINGS . . . 352

FUTURE QUESTIONSFOR CULTURALNEUROSCIENCE. . . . . . . . . . . . . . . . 353

CONCLUSION . . . . . . . . . . . . . . . . . . . . . 354

INTRODUCTION

During the past few years, brain imagingstudies have uncovered variations in the neuralsubstrates of human cognition in differentcultural groups. The integration of theory andmethods from social and cultural psychology,cognitive neuroscience, and other relateddisciplines gave birth to a new field—culturalneuroscience (CN). This new field investigateshow human brain functions are shaped byinteractions between culture, the brain, andgenes. CN has developed specific methods foruncovering cultural influences on human brainfunctions, and it has provided a new perspectiveon the functional organization of the humanbrain in sociocultural environments. Thegrowth of this new field can be seen in the in-creasing number of research articles and severalspecial issues dedicated to CN (e.g., Progressin Brain Research in 2009; Social Cognitive andAffective Neuroscience in 2010) in addition toedited books (e.g., Han & Poppel 2011) and itscoverage in textbooks (e.g., Ward 2012).

CN is receiving more and more attentionfrom researchers in different fields because

CN touches upon concepts such as humans,culture, and race that are often used in a widercontext beyond and outside CN. Thus it isimportant to take stock of the progress so farby providing a comprehensive overview ofthe field as well as addressing some questionsand concerns that have been voiced about thisendeavor. The goals of the current review are(a) to elucidate the origin, aims and concepts,empirical methods, and findings of CN studies;(b) to clarify misunderstanding of CN findings;(c) to discuss the implications of CN findingsfor understanding the biosocial nature of thehuman brain; and (d ) to address key questionsfor future CN studies.

WHAT IS CULTURALNEUROSCIENCE?

The term cultural neuroscience was initiallyintroduced by Chiao & Ambady (2007, p. 238),who defined CN as “a theoretical and empiricalapproach to investigate and characterize themechanisms by which [the] hypothesized bidi-rectional, mutual constitution of culture, brain,and genes occurs.” This definition was furtherrefined by Chiao (2010, p. 109), who describedCN as an “interdisciplinary field bridgingcultural psychology, neurosciences and neuro-genetics that explains how the neurobiologicalprocesses, such as genetic expression and brainfunction, give rise to cultural values, practicesand beliefs as well as how culture shapesneurobiological processes across macro- andmicrotime scales.” These concepts resonatewith recent efforts to integrate research findingsfrom neuroscience, genetics, developmentalpsychology, and sociology by highlightingthe role of postnatal neuroplasticity in humandevelopment (e.g., Li 2003, Wexler 2006). Inthe remainder of this section we seek to providea description of CN, starting with its origin andaims. Next we define key theoretical conceptsin CN. This is followed by an overview ofcommon CN methodology. Finally, we endthe section by reviewing the empirical findingsof CN studies using different methods.

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Origin

CN emerges from the integration of differ-ent branches of social sciences and natural sci-ences and arises mainly from two disciplines,i.e., cultural psychology, which has providedthe insight that cognitive, emotional, and mo-tivational tendencies and habits are shaped byculture, and neuroscience, which has demon-strated that the brain is shaped by experience.We here show that CN is based on four key ap-proaches, i.e., cultural psychology, social cogni-tive neuroscience, the study of neuroplasticity,and the study of culture × gene interactions.

Cultural psychology. Although there is along history of the debate about the conceptof culture (Kroeber & Kluckhohn 1952), it iscommonly acknowledged that cultural groupshave differentiated over thousands of years tocreate what has been called cultural speciation(Goodall & Berman 1999). The resultingcultural differences still exist in our globalizedworld and are associated with certain differ-ences in the ways people think and behave.These differences exist side by side withmany cross-cultural commonalities. Culturaldifferences in human behaviors are very welldocumented in anthropology (e.g., Havilandet al. 2008), and human development has beenviewed as a process of acquiring and embodyingculture’s belief systems (Rogoff 2003). Culturaldifferences in human mental processes andunderlying cognitive mechanisms have beeninvestigated extensively in cultural psychologyduring the past two decades. From this lineof research, theoretical frameworks such asindividualistic versus collectivistic values, inde-pendent self-construals versus interdependentself-construals, and holistic versus analyticcognitive tendencies have emerged to guideempirical studies of cultural discrepancy inhuman cognition and emotion (Kitayama &Cohen 2007, Nisbett et al. 2001, Varnum et al.2010). Cultural psychology takes the view thathuman cognitive and affective processes vary asa function of cultural environments that provideunique social contexts in which psychological

Independentself-construal:a tendency to view theself as autonomous andbounded; an emphasison affirming theindependence anduniqueness of the self

Interdependentself-construal:a tendency to view theself as interconnectedand overlapping withclose others; anemphasis on affirmingclose relationships andmaintaining harmonywithin them

processes develop and are shaped (Kitayama& Uskul 2011). The findings of culturalpsychological research stimulate researchersto investigate neural substrates of culturaldiversity of human cognition and emotion.

Social cognitive neuroscience. Social cogni-tive neuroscience research investigates brainmechanisms that allow human beings tounderstand the self and others and to effi-ciently navigate social environments (Ochsner& Lieberman 2001). Early social cognitiveneuroscience research focused on the neuralsubstrates underpinning social cognition bycombining brain imaging and social psycho-logical paradigms. Most of these studies aimedto uncover the neural mechanisms of socialcognition and behavior without consideringpotential cultural differences. However, animportant feature of social cognition andbehavior is context dependence. We are alwayssituationally embedded in a certain envi-ronment, the “context,” which substantiallyinfluences our perception of others and ourunderstanding of the behavior of others. Thiscontext dependency itself underlies substantialinfluences exerted by culture. In other words,what social information is processed and howit is processed rely heavily on one’s interactionpartners (in the case of dyadic interactions)and, more broadly, on the social context inwhich the interactions occur. For example, cul-tural psychological studies have documentednumerous variations in social cognitive pro-cesses across different cultural contexts, suchas construal of the self (Markus & Kitayama1991, 2010), causal attribution of physicaland social events (Choi et al. 1999), analyticversus holistic attention (Masuda & Nisbett2001), affective states that people ideally liketo feel (Tsai et al. 2006), and choice-induceddissonance (Kitayama et al. 2004), amongmany others. Due to the considerable evidencefor cultural divergence of human subjectiveexperiences and psychological processes,recently neuroscientists have shown increasinginterest in whether parallel differences inneural mechanisms might also be present

www.annualreviews.org • Cultural Neuroscience 337

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among people who were raised in differentsociocultural contexts (Ambady & Bharucha2009; Ames & Fiske 2010; Chiao & Bebko2011; Han & Northoff 2008, 2009; Kitayama& Uskul 2011; Park & Gutchess 2006; Park& Huang 2010; Rule et al. 2012). On the the-oretical basis alone, cultural influences on theneural substrates underlying human cognitiveand affective processes would seem highlyplausible given that it takes almost 20 years fora large portion of the brain to mature (Gogtayet al. 2004). During this period the brain isinfluenced by personal experiences in a specificcultural context. Thus in the beginning ofthe twenty-first century, researchers startedto examine potential cultural differences inhuman brain mechanisms involved in multiplecognitive and affective processes by comparingbrain imaging results obtained from differentcultural groups. There is now substantialevidence that individuals from different socio-cultural contexts show distinct patterns of brainactivity involved in cognition and behavior.

Neuroplasticity. Biological research hasshown ample evidence for the intrinsic plas-ticity of the human brain; that is, the brainchanges both structurally and functionally inresponse to the environment and experience(Shaw & McEachern 2001). For example, theoccipital cortex, which is commonly involvedin visual processing in sighted humans, canbe engaged in auditory processing in blindindividuals (Burton et al. 2002, Gougoux et al.2009). Auditory deprivation results in therecruitment of the primary auditory cortex inthe processing of vibrotactile stimuli (Levanenet al. 1998) and sign language (Nishimura et al.1999) in deaf humans. The medial prefrontalcortex is engaged during self-reflection onvisually but not aurally presented trait words insighted humans, while the region is recruitedduring self-reflection on aurally presented traitwords in congenitally blind individuals (Ma& Han 2011). These findings demonstrate anintrinsic property of the brain, plasticity, whichenables the nervous system to respond to en-vironmental pressures, physiological changes,

and personal experiences (Pascual-Leone et al.2005) and to adapt to social contexts duringdevelopment (Blakemore 2008). Given thathuman thoughts and behaviors differ substan-tially across a variety of sociocultural contexts,it is not surprising that the human brain, thesource of human behaviors and the carrier ofhuman thoughts, is modulated by socioculturalenvironments and develops unique neuralmechanisms that help an individual to adaptto culturally specific changes and pressures.Thus an intrinsic feature of the brain is itssociocultural context dependence.

Gene × environment interactions. The ba-sic assumption of CN is that culture provides aframework for social behavior, communication,and interaction that generates social valuesand norms, assigns meaning to social events,interacts with biological variables (e.g., genes),and codetermines the functional organizationof the brain. The CN approach investigatesmutual interactions between culture, the brain,and genome, consistent with culture-genecoevolution theory (Boyd & Richerson 1985,Lumsden & Wilson 1981). This theory guidesresearch that explores how two complementaryand interacting evolutionary processes, i.e.,genetic evolution and cultural evolution,influence human behavior. The CN approachaims to understand how sociocultural contextsinfluence human behavior by examiningcultural influences on underlying neural mech-anisms. CN views cultural differences in theneural mechanisms underlying cognition as theproduct of the interaction between genes andcultural environment, an idea that researchersin this field are beginning to test empirically.

Key Concepts

Next we review some key concepts related toCN. We provide definitions and also highlightconceptual distinctions between terms such asculture, nationality, and race, which some maymistakenly use interchangeably.

Culture is obviously the most impor-tant concept in the field of CN. From

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anthropologists’ perspective, culture refers toa system of meanings used to make sense oflife (i.e., Kuper 1999). Although there are myr-iad definitions of culture [indeed, Kroeber &Kluckhohn (1952) listed 164 definitions of cul-ture], culture is often used in three basic sensesin social psychology (Chiu & Hong 2006).Material culture consists of all material arti-facts produced by human beings. Social cul-ture consists of social rules and social institu-tions. Subjective culture refers to shared ideas,values, beliefs, and behavioral scripts. These as-pects of culture, however, are dynamically re-lated within a given cultural tradition, locale,and/or community and together form a uniquesocial environment for the group of individuals.From the very beginning of their life, people en-gage in the complex composed of materials andsocial rules or practices as well as folk beliefs oftheir respective local communities, and by do-ing so, they have their brains changed in sucha way that the resulting brain functions are at-tuned closely to the surrounding socioculturalenvironment.

Culture is different from nationality, whichis defined by social group membership basedon a shared nation state of origin. Althoughthe term culture emphasizes shared ideas,values, beliefs, and practices, people of thesame nationality do not necessarily share thesame beliefs, values, or practices. Similar tosocial psychology studies, most CN studies usethe term culture in the sense of a social groupwhose members share social values, knowledge,and practices. Some CN studies have recruitedparticipants from two different cultural groups(e.g., Westerners and East Asians) based oncultural psychology findings that suggest thatthe two groups differ in specific cultural valuesor specific cognitive processes. In some casesrace and language are concomitants that alsodifferentiate two cultural groups. Other CNstudies have investigated two cultural groupswho are from the same nation but are defined byreligious or political beliefs. In such cases twogroups of participants share the same nation-ality, race, and language but differ only in a setof shared beliefs/values and practices that are

hypothesized to be relevant to a particularpattern of neural activity.

Recent CN studies have directly measuredcultural values from different groups andhave assessed the relationship between thesevalues and neural responses. For example,the Self-Construal Scale (Singelis 1994) hasbeen used in several recent CN studies toevaluate individuals’ independent versus in-terdependent self-construals (e.g., Chiao et al.2009a, de Greck et al. 2012, Na & Kitayama2011, Sul et al. 2012). This approach seeksto simultaneously capture both within- andbetween-group variations instead of assuming,a priori, that people from two socioculturalcontexts must have different cultural values. Ittherefore acknowledges individual differencesin cultural values among those who grow upin the same sociocultural context. Moreover,measuring cultural values allows researchers toexamine whether a group difference in brainactivity is associated with a specific culturalvalue, whether cultural group differencesin brain activity are mediated by a specificcultural value, and how individual differencesin brain activity are associated with variationsin cultural values within a given cultural group.

Cultural psychology views culture as a dy-namic knowledge system rather than a rigid setof stereotypes about a social group (Markus &Hamedani 2007). Culture represents a dynamicconcept of the social environment that is notpart of the innate biological condition of hu-mans. Humans are not born with propensitiesfor any particular culture but rather with thepotential and the capacity to acquire and to cre-ate culture (Harris 1999). Thus an individualmay change his/her cultural values and beliefsas a result of experience (such as emigratingfrom his/her native country). People from thesame cultural groups can be quite heteroge-neous in terms of the values and beliefs theyacquire. This is particularly true in contempo-rary societies where cultural exchanges occuroften and rapidly. People in modern societiesare rarely monocultural because they are almostalways exposed, often deeply, to other cultures’practices and beliefs in multiple sociocultural


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contexts. Thus multiple cultural systems maybecome part of any single individual, and asa consequence, it is often required to switchto and fro between different cultural systemsduring social interactions depending on spe-cific contexts of social encounters (Hong et al.2000). This dynamic model of culture allowsus to test whether brain activity underlyinghuman cognition can vary on a short temporalscale as a function of recent use of one culturalsystem or another (e.g., cultural priming, as isdiscussed in more detail when we turn to themethodology of CN research). Taken together,CN studies view culture as a complex anddynamic external social environment in whichthe human brain is fostered and shaped. Ratherthan considering the brain and its neuronalstates by themselves, CN emphasizes the socio-cultural nature of the human brain and placesgreat weight on the influence of cultural values,beliefs, and practices shared by a social groupon functional organization of the human brain.

Race is a way of categorizing human beingson the basis of external attributes, such as skintone and facial and body shapes, that differenti-ate human populations. In many racial theories,and in lay theories, racial groups also possessdifferent fixed and biologically determined psy-chological traits and tendencies. Race is viewedas fixed both over the course of the lifespan andacross cultural contexts. People from the sameracial group are thought to be homogenousin terms of heritage and physical appearance.However, in reality individuals classified asbelonging to the same race do not necessarilyshare the same cultural values and experiences.For example, Native Chinese and ChineseAmericans may be thought to belong to thesame racial group but may have distinct culturalvalues and beliefs and experiences. There haslong been a debate over whether racial differ-ences in psychological tendencies and behaviorexist and if so whether such differences are bio-logically determined. As a starting point, thereis the question of whether contemporary racialcategories can be genetically differentiated. Al-though some have noted a great deal of geneticsimilarities across races (humans are 99.9%

alike) and argued that it is very difficult to ascer-tain the racial identity of individuals throughtheir genes (Littlefield et al. 1982), recent large-scale studies using genetic cluster analysis havefound correlations between self-reported racialgroup membership and the genetic cluster ofracial groups (e.g., Paschou et al. 2010, Tanget al. 2005). However, these data do not speakto the question of whether racial groups arepsychologically different, nor do they addresswhether such differences (if they exist) aresolely or partially genetically driven.

Regardless of whether race has a biologicalcomponent, race does have a number ofcomplex sociopolitical implications and maybe analyzed as a sociocultural construction.Moya & Markus (2011, p. 21) recently definedrace as “a dynamic set of historically derivedand institutionalized ideas and practices that(1) sorts people into ethnic groups accordingto perceived physical and behavioral humancharacteristics; (2) associates differential value,power, and privilege with these characteristicsand establishes a social status ranking amongthe different groups; and (3) emerges whengroups are perceived (a) to pose a threat(political, economic, or cultural) to eachother’s world view or way of life; and/or (b) tojustify the denigration and exploitation (past,current, or future) of, and prejudice toward,other groups.” In addition, race has a strongevaluative component that has been often usedto qualify individuals from different races assuperior or inferior on the basis of untested ordiscredited assumptions related to genetics.

Aims

The goal of CN studies is to investigate humanbrain function and structure in diverse socio-cultural contexts. Like cultural psychologists(Markus & Hamedani 2007), CN researchershave little interest in using brain activity asa way to classify people into groups. Instead,CN research investigates whether and how thefunctional organization of the human brainis shaped by culture and by the interactionbetween culture and genes on different

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time scales (Chiao & Ambady 2007, Han &Northoff 2008). In addition, CN research aimsto investigate how neurobiological processesin the human brain contribute to the riseof divergent cultures in the world. Theoriesbuilt on CN findings will eventually help toexplain how cultural differences in human brainfunction mediate divergent social behaviorsacross cultures while at the same time pointingout the neural predispositions of psychosocialcommonalities across different cultures. CNconsiders culture as a highly dynamic systemof continuous interaction and exchange amongindividuals. This system of social interactionfeeds back into social practices, values, andbelief systems, thereby establishing circular,recursive, and reciprocal influences betweeninteracting individuals and culture (Hacking1999, Vogeley & Roepstorff 2009).

Most current CN studies focus on cross-cultural differences in the neural substratesof human psychological processes includingcognition, emotion, and motivation (Ambady& Bharucha 2009; Han & Northoff 2008, 2009;Kitayama & Uskul 2011). This line of researchhas mainly been stimulated by findings incultural psychology that show cross-culturalvariation in multiple levels of psychologicalprocesses (Kitayama & Cohen 2007, Nisbettet al. 2001). By comparing behavioral perfor-mances among individuals from Western (e.g.,European and American) and East Asian (e.g.,Chinese, Japanese, Korean) contexts, culturalpsychologists have shown evidence for dis-tinct, culture-dependent cognitive processingstyles in perception ( Ji et al. 2000), attention(Kitayama et al. 2003, Masuda & Nisbett2001), memory (Wang & Conway 2004),perspective taking (Wu & Keysar 2007), causalattribution of events (Morris & Peng 1994,Peng & Knowles 2003), object categorization( Ji et al. 2004), recognition of one’s own face(Liew et al. 2011b; Ma & Han 2009, 2010; Suiet al. 2009), self-construal (Markus & Kitayama1991), and affect valuation (Tsai et al. 2006),among many others.

Based on the fundamental hypothesis thathuman psychological processes are mediated by

specific neural substrates in the brain, CN stud-ies have initially focused on whether differentneural substrates may be engaged in a varietyof cognitive processes among individuals inEast Asian and Western cultural contexts. Asa part of this line of research, CN researchershave found evidence for cultural differences inthe neural mechanisms involved in visual per-ception (Goh et al. 2007, 2010; Gutchess et al.2006; Jenkins et al. 2010), attention (Heddenet al. 2008, Lewis et al. 2008), causal attributionof physical events (Han et al. 2011), seman-tic relationship processing (Gutchess et al.2010), musical processing (Nan et al. 2006,2009), mental calculation (Tang et al. 2006),recognition of one’s own face (Sui et al. 2009),self-reflection on personality traits (Chiao et al.2009a,b; Wang et al. 2012; Zhu et al. 2007),perception of bodily expression (Freeman et al.2009), mental state reasoning (Adams et al.2009, Kobayashi et al. 2006), empathy (deGreck et al. 2012), and in other domains. TheseCN findings indicate that a fundamental aspectof the functional organization of the humanbrain is its sensitivity to the sociocultural con-texts in which individuals are brought up. How-ever, the final goal of CN research is not simplyto show differences in brain activity acrosscultural groups. Rather, CN studies aim to pro-vide a neuroscientific account of cross-culturalvariation in human psychological functionsand behaviors by discovering socioculturallypatterned neural mechanisms and their devel-opment. Thus CN studies aim to reveal bothculturally universal and culturally unique neu-ral processes by which human brains predisposeus to perceive self and others, communicate andinteract with conspecifics, and guide actions.

Recently, there has been an increasinginterest in how brain function is shaped byculture-gene interaction, based on culture-genecoevolution theory (Boyd & Richerson 1985,Lumsden & Wilson 1981). However, althoughrecent studies have shown that psychologicaltendencies and behavioral outcomes associatedwith specific genotypes are moderated by cul-ture (e.g., Kim et al. 2010a,b), direct evidencethat culture moderates the effect of genotype on


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Functional magneticresonance imaging(fMRI): a noninvasivemethod for recordingblood-oxygenation-level-dependentsignals that have highspatial resolution andare used to examinebrain activationsassociated with specificstimuli or tasks

Event-relatedpotential (ERP):synchronous activitiesof neuronalpopulations engaged inspecific psychologicalprocessing, which aretime locked tostimulus events, can berecorded fromelectrodes over thescalp, and have hightemporal resolution

the functional organization of the brain is stilllacking. Another line of research has focusedon how the allelic frequencies of a genotypewithin a population may relate to culturaldifferences in values. Chiao & Blizinsky (2010)examined the relationship between the culturalphenotypes of individualism-collectivism andallelic frequency of the serotonin transporterfunctional polymorphism (5-HTTLPR) byassessing the prevalence of the short alleleof 5-HTTLPR among different populations.In a comparison of 29 countries, they foundthat cultures that were high in collectivismcontained a significantly greater proportionof short allele carriers and that increasedfrequency of short allele carriers predicteddecreased anxiety and mood disorder preva-lence. Further, the relationships between theprevalence of short allele carriers within apopulation and the prevalence of anxiety andmood disorders were mediated by collectivism.Similarly, Way & Lieberman (2010) suggestedthat collectivism may have developed andpersisted in populations with a high proportionof a functional polymorphism (A118G) inthe μ-opioid receptor gene—a putative socialsensitivity genotype that is compatible withcollectivistic cultural groups. Caution is duebecause this body of evidence is entirely corre-lational. Nevertheless, given that cultures mayinteract with the 5-HTTLPR genotypes toinfluence the prevalence of affective disorderssuch as anxiety and depression (Chiao &Blizinsky 2010), it would be interesting tostudy how the culture-gene interaction code-termines the intermediate endophenotype (i.e.,neurobiological responsiveness) associatedwith culture-sensitive cognitive processes.

Methods

Although CN is a young field, CN researchershave developed quite sophisticated method-ologies by drawing on prior brain imagingand social and cultural psychology research.Methodological challenges include both thedesign of psychological experiments and brainimaging techniques. Early CN studies focused

on whether and how two cultural groups differin neural substrates of specific cognitive andaffective processes. A typical way to addressthis issue is to compare functional magneticresonance imaging (fMRI) or event-relatedpotential (ERP) brain data obtained fromindividuals who were raised in two differentsociocultural contexts.

One assumption of this approach is that,because participants from two cultural groupsdiffer in cultural knowledge, values, and/orcognitive and affective processes, the un-derlying neural activity should be differentbetween the groups in a specific way. Toaddress this assumption, CN research hastaken cultural psychological research as a guidefor its hypotheses about neural differencesbetween specific cultural groups. For example,behavioral research first showed that individ-uals in Western cultures are more sensitive tosalient foreground objects compared to peoplein East Asian cultures, whereas individualsin East Asian cultures are more inclined tofocus their attention broadly on backgroundsrelative to people in Western cultures ( Ji et al.2000, Kitayama et al. 2003, Masuda & Nisbett2001). Such findings lead to a reasonablehypothesis that neural substrates underlyingvisual perception of and attention to salientobjects and contexts may show differentpatterns between individuals in the Westernand East Asian cultures (Goh et al. 2007,Gutchess et al. 2006, Hedden et al. 2008).Similarly, evidence from cultural psychologythat the self is viewed as independent inWestern cultures and interdependent in EastAsian cultures (Markus & Kitayama 1991, Sin-gelis 1994) leads to the hypothesis that neuralrepresentation of the self and close others mayoverlap to a greater degree among East Asiansthan among Westerners (Zhu et al. 2007).

However, selection of participants from twodifferent nations or sociocultural contexts doesnot necessarily imply that the participants havedistinct cultural values (Oyserman et al. 2002).CN seeks to address this question by measuringthe value or self-construal dimensions thatare hypothesized to drive the relevant cultural

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differences in the neural process being studied.Often, CN researchers directly assess thecultural values or self-construals of interest;this can be done using well-established ques-tionnaires developed by social and culturalpsychologists. For example, the Self-ConstrualScale (Singelis 1994) is widely used to evaluatehow people view themselves (either as indepen-dent or interdependent) and has been shownto differentiate cultural groups. Current CNstudies usually compare participants from twocountries, usually a Western culture and anEast Asian culture (e.g., British versus Chinese,American versus Japanese). Although previousstudies have demonstrated differences betweenthese cultures in terms of values and cognitiveand affective processes, it is important todemonstrate that the participants recruited inCN studies actually differ in these dimensions.Measurements of cultural values are also helpfulin situating CN studies as dealing with culturaldifferences rather than racial or national differ-ences. Equally important, measuring culturalvalues allows for stronger inferences. By exam-ining whether individual differences in culturalvalues can predict individual differences inbrain activity, researchers can probe the asso-ciation between these values and patterns ofbrain activity. In addition, measuring culturalvalues in individual subjects makes it possibleto assess whether cultural values mediate differ-ences in brain activity associated with specifictasks, which may be performed in differentways according to the respective degree of thecultural value, between two cultural groups.

Of course, when comparing participantsfrom different sociocultural contexts, it is im-portant to control for potentially confoundingvariables such as gender, age, and education aswell as socioeconomic status. Language is an-other potential confound in cultural compar-isons if stimuli used in brain imaging studies arebased on verbal materials. This, however, canbe controlled by using the native language foreach cultural group so that the same language isused in an experimental condition and a controlcondition. Such designs allow us to comparethe experimental and control conditions so as

to reduce the effect of language processing to aminimum degree.

Another elegant psychological paradigmused by CN researchers is to prime culturalidentity or values before recording brain activ-ity during a specific task. Such studies are basedon the assumption that individuals can acquiremore than one set of cultural knowledge andcan use different sets of cultural knowledge de-pending on contextual cues (Hong et al. 2000).According to this dynamic constructivist modelof culture, people who have been exposed tomultiple cultures may acquire multiple sets ofcultural knowledge, and exposing individualsto cultural symbols may activate specificcultural knowledge and result in mindsets andbehaviors that are consistent with that culture.For instance, after cultural priming, biculturalindividuals may switch between Western andEast Asian mindsets that are consistent with themost accessible cultural knowledge tradition(e.g., Hong et al. 2003). A number of studieshave also shown that priming independenceand interdependence of self-construals in-fluences patterns of cognitive processes thattend to differ between cultural groups (e.g.,Kuhnen & Oyserman 2002, Lin & Han 2009;for a review, see Oyserman & Lee 2008).Similarly, CN studies take culture as a dynamicknowledge and meaning system, and theymanipulate cultural values as variables (Chiaoet al. 2009b, Lin et al. 2008, Ng et al. 2010, Sui& Han 2007, Sui et al. 2012). CN studies usingcultural priming test for dynamic changes inbrain activity as a function of cultural valuesand provide information that allows for causalinference regarding the relationship betweencultural values and specific brain activity.

CN studies often compare brain-imagingresults from multiple cultural groups, and thusseveral technical issues have to be consideredwhen designing CN research. It is ideal to scanmultiple cultural groups at the same experi-mental site in cross-cultural fMRI studies. Thiscan certainly avoid systematic, site-dependenteffects in fMRI sensitivity between the scannerfacilities, although between-subject differencescan account for nearly ten times more variance


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than site effects when participants are scannedusing the same type of scanner at two sites(Sutton et al. 2008). For those CN studiesthat scan multiple cultural groups at differentsites, Chiao et al. (2010) has suggested severalways to reduce the probability of systematic,site-dependent effects in fMRI sensitivity.First, both functional and anatomical MRIdata should be collected using scanners fromthe same vendor with identical protocols.Second, an interscanner reliability test can beconducted by scanning a separate cohort ofparticipants or phantom data at each scannerfacility, thus enabling one to quantify and sta-tistically compare signal-to-noise ratio acrossscanner sites. Third, the presentation softwareand hardware should be identical, calibratedand tested at each session, and scripts shouldbe written and implemented across the sites ina culturally appropriate manner.

Relative to fMRI, electroencephalogram(EEG) is a simpler method for cross-culturalcomparisons of brain activity. Similar EEGrecording systems can be found easily at differ-ent recording sites. Portable EEG amplifiersare available and can be easily transferredbetween different recording sites so that EEGdata from different cultural groups can berecorded using the same system. AnalyzingERPs that are time locked to a stimulus or aresponse is particularly helpful for uncoveringthe time course of neural responses to multiplecognitive processes such as recognition of one’sown face (Sui et al. 2009, 2012), inference ofone’s own and others’ personality traits (Mu& Han 2010, Na & Kitayama 2011), implicitprocessing of vocal tone (Ishii et al. 2010),emotion regulation (Murata et al. 2012), andmusical processing (Nan et al. 2006, 2009).

Findings

Several recent review articles have summarizedmajor findings from CN research (e.g., Am-bady & Bharucha 2009; Ames & Fiske 2010;Chiao & Bebko 2011; Han & Northoff 2008,2009; Kitayama & Uskul 2011; Park & Huang2010; Rule et al. 2012). Thus this section is not

intended to give an extensive review of currentCN findings. Instead, we highlight a select setof recent CN studies in terms of their method-ology in order to illustrate the intellectualdevelopment of CN research.

Distinct neural activity to cultural familiar/unfamiliar stimuli. One question that CNresearchers are interested in is how humanbrain activity is tuned by culturally famil-iar/unfamiliar information. To address this,CN researchers simply recorded neural activityto culturally familiar/unfamiliar stimuli fromone cultural group. For example, to investigatethe neural basis of musical phrase boundaryprocessing during the perception of music fromnative and nonnative cultures, Nan et al. (2008)used fMRI to record brain activity in Germanmusicians while they categorized phrasedWestern and Chinese musical excerpts. Theyfound that culturally familiar musical excerptsmore strongly activated multiple brain regionsincluding the superior frontal gyrus, the poste-rior precentral gyrus, and the superior temporalgyrus, possibly reflecting enhanced sensorimo-tor integration. Culturally unfamiliar musicalexcerpts, however, more strongly activated theposterior insula as well as the middle frontal andangular gyri, possibly due to higher demandson attention systems and higher loads on basicauditory processing. Similarly, Demorest &Osterhout (2012) recorded ERPs while Amer-ican participants listened to melodies based inthe Western folk tradition or North Indianclassical music. ERPs showed that a long latencypositive activity was sensitive to the original anddeviation form of the melodies, and this effectwas more salient in the Western than in theIndian context. The results suggest that peoplemay generate specific expectancies when listen-ing to culturally familiar music, whereas theymay remain unable to develop such expectan-cies when hearing culturally unfamiliar music.

Another example of this line of researchexamined whether observations of culturallyfamiliar/unfamiliar symbolic gestures engagedistinct neural subsystems. Liew and colleagues(2011a) scanned Chinese participants while

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they perceived video clips in which a modelshowed culturally familiar/unfamiliar symbolicgestures. They found that culturally familiargestures increased activity in the posteriorcingulate cortex, the dorsal portion of themedial prefrontal cortex and the bilateraltemporoparietal junction. These brain regionsconstitute the neural circuit engaged in infer-ence of others’ intentions and beliefs (Frith &Frith 2006). In contrast, unfamiliar gesturesgenerated activity in the left inferior parietallobule, the left superior frontal gyrus, and thebilateral superior parietal lobule. These brainregions make up the neural network involvedin automatic motor simulations of observedactions (Rizzolatti & Sinigaglia 2010). Appar-ently, the mentalizing system is engaged duringobservation of culturally familiar gestures inorder to understand others’ intentions orbeliefs, whereas the mirror neuron system maybe activated during perception of culturally un-familiar gestures so as to capture others’ mindthrough automatic motor simulations of ob-served actions. Taken together, these findingsindicate that long-term cultural experiencesmay result in specific neural mechanisms in thehuman brain that deal with culturally familiarinformation. This may allow the individualto quickly comprehend the meaning of socialinformation in one’s own culture, predictothers’ behavior, and take appropriate actionsin a specific cultural context.

Cultural group differences in neural activ-ity. The mainstream of CN studies focuses onwhether differences in cognitive processes be-tween two cultural groups revealed by culturalpsychology are associated with distinct pat-terns of brain activity. There is now increasingevidence that two cultural groups may employdistinct neural mechanisms while performingseemingly identical cognitive and emotionaltasks. This is of fundamental importance tounderstanding brain-behavior relationships ingeneral as well as to CN. There are two pat-terns of cultural group difference in the neuralactivity involved in cognitive and affectiveprocesses.

Temporoparietaljunction: the corticaljunction zone at theborder of the posteriorparts of the temporallobe and the inferiorparts of the parietallobe, which has beenshown to be involvedin belief reasoning andperspective taking

N400: a negativepotential that peaksaround 400 ms afterstimulus onset with themaximum amplitudeover the parietal scalesite and is sensitive tosemantic incongruitybetween stimuli

One type of cultural modulation of brainactivity is that a specific neural activity issignificantly modulated by a particular task inone cultural group but not in another culturalgroup. Based on the assumption that West-erners tend to attend to salient objects whereasEast Asians are inclined to attend to a broadperceptual and conceptual field (Nisbett et al.2001, Nisbett & Masuda 2003), Jenkins et al.(2010) tested whether the neural activity inthe lateral occipital cortex to a target stimuluswas more sensitive to background scenes inEast Asians than in Westerners. They scannedAmerican and Chinese participants during per-ception of pictures consisting of a focal objectsuperimposed upon a background scene thatwas congruent (e.g., a deer in the woods) or in-congruent (e.g., a television in the desert) withthe target object. The target object was pre-sented on different novel scenes or on a singlerepeated scene on four successive trials. Adap-tation magnitude was calculated by subtractingthe neural activity to objects on a repeated scenefrom that to objects on different novel scenes.Jenkins et al. found that the neural activity inboth the right and left lateral occipital cortexshowed significantly greater adaptation toincongruent scenes than to congruent scenes,suggesting sensitivity of the occipital activityto the background scenes. However, this effectwas evident in Chinese participants but not inAmerican participants. Similar results were ob-served in another study that recorded ERPs totarget objects that were presented on semanticcongruent or incongruent background scenes(Goto et al. 2010). It was found that a negativeERP component peaking at about 400 ms afterstimulus onset (N400), which has been shownto be sensitive to processing semantic relation-ships (Kutas & Hillyard 1984), was enlarged bytarget objects presented on semantically incon-gruent versus congruent background scenes;however, this effect was observed in AsianAmericans but not in European Americans.

This pattern of cultural differences wasalso observed in neural activity involved inhigh-level social cognition. To investigatecultural differences in the neural mechanisms


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Medial prefrontalcortex (MPFC): themedial region of theprefrontal cortex thathas been shown to beinvolved in socialcognition, with thedorsal region engagedin mental statereasoning and theventral region engagedin self-reflection

underlying causal attribution of physicalevents, Han and colleagues (2011) first scannedChinese participants during causality versusmotion direction judgments on animations ofobject collisions. Causality judgments askedparticipants to infer causes of physical events(i.e., changes of motion direction of a targetobject after colliding with another object), andmotion direction judgments required iden-tification of the motion direction of a targetobject after colliding with another object.They showed that, relative to motion directionjudgments, causality judgments activated themedial prefrontal cortex (MPFC) and the leftparietal cortex (LPC). Moreover, they foundthat MPFC activity was sensitive to the demandto infer causes of events, whereas LPC activitywas modulated by the contextual complexityof physical events. In a subsequent experiment,Han et al. (2011) scanned American and Chi-nese participants during causality versus motiondirection judgments on animations of objectcollisions. They found that American andChinese participants showed similar MPFCactivity involved in causality judgments.However, LPC activity elicited by causalityjudgments of physical events was evident inChinese but not in Americans. Thus it can beconcluded that LPC activity associated withthe contextual processing is more sensitiveto cultural differences in causality perceptionthan is MPFC activity engaged in inference ofcausal relationships.

Zhu et al. (2007) tested cultural differencesin the neural activity underlying representationof personality traits of oneself and a closeother. According to Markus & Kitayama(1991, 2010), Western cultures encourageself-identity that is independent of social con-texts and others, whereas East Asian culturesemphasize fundamental social connections,leading to an interdependent view of the selfand partial overlap in representation of the selfand close others. This proposition may predictshared neural mechanisms of representation ofthe self and a close other in East Asian culturesbut not in Western cultures. To test this,Zhu and colleagues (2007) scanned Chinese

and Westerners using fMRI during traitjudgments of oneself, a close other (i.e., one’smother), and a celebrity. They found that,relative to trait judgments of a celebrity, traitjudgments of oneself significantly activated theventral region of the MPFC in both Chineseand Westerners, suggesting a similar neuralsubstrate of representation of oneself in thetwo cultural groups. However, trait judgmentsof one’s mother versus a celebrity activated thesame brain region in Chinese but not in West-erners, suggesting shared neural representationof the self and a close other in Chinese but notin Westerners. This finding reveals a neuralmodel of cross-cultural variations in represen-tations of a close other in relation to the self.

It is not always true that individuals fromEast Asian cultures show additional modu-lations of neural activity in comparison withindividuals from Western cultures. In a recentfMRI study, de Greck et al. (2012) examinedcultural differences in brain activity duringempathy with anger. They scanned Chineseand German participants during an intentionalempathy task and found empathy-relatedneural activity that was specific for eachcultural group. Specifically, empathy for angryfaces activated the left dorsolateral prefrontalcortex in Chinese participants, but it activatedthe right temporoparietal junction, the rightinferior and superior temporal gyrus, and theleft middle insula in Germany participants.These results implicate enhanced emotion reg-ulation during empathy with anger in Chineseculture, in which the attitude toward harmonyis more valued (Kim & Markus 1999, Markus& Kitayama 1991). However, in Germans,empathy with anger may be characterized byenhanced inference of others’ mind, given thekey role of the temporoparietal junction inmental state reasoning (Frith & Frith 2006).

The aforementioned neuroimaging find-ings are consistent with the idea that culturalpractices may produce specific psychologicalprocesses that are significant in one culture butnot in another culture. One may consequentlyassociate culturally specific psychologicalprocess with particular underlying neural

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mechanisms that are observed in one culturebut not in others.

Another type of cultural modulation ofbrain activity is that a specific neural activity ismodulated by a particular task in two culturalgroups but in opposite ways. These oppositepatterns of neural activity may have nothing todo with the cultural familiarity of the stimuli,but may instead reflect culturally specificcognitive styles. Hedden et al. (2008) assessedcultural differences in the neural activity un-derlying attentional control by scanning EastAsians and Americans in context-dependent orcontext-independent judgment tasks. Partic-ipants were presented with a series of stimuli,each consisting of a vertical line inside a box.The context-dependent task required judg-ments of whether the box and line combinationof each stimulus matched the proportionalscaling of the preceding combination. Thecontext-independent judgment task requiredjudgments of whether the current line matchedthe previous line, regardless of the size of theaccompanying box. It was found that the neuralactivity in the prefrontal and parietal corticesinvolved in the tasks showed an opposite patternof activations in the two cultural groups; that is,Americans showed greater prefrontal and pari-etal activity during the context-dependent thancontext-independent tasks, whereas East Asiansexhibited stronger activity in the prefrontaland parietal cortices during the context-independent than context-dependent tasks.The opposite pattern of neural activity wasinterpreted as reflecting enhanced sustainedattentional control during culturally nonpre-ferred in comparison with preferred tasks.

Opposite patterns of neural activity in twocultural groups may also arise from distinctcultural values. To examine why Americanculture tends to reinforce dominant behaviorwhereas Japanese culture tends to reinforcesubordinate behavior, Freeman et al. (2009)scanned American and Japanese individualsduring perception of body displays relatedto dominance and subordination. The neu-ral activity in the bilateral caudate nucleusand MPFC showed an opposite pattern of

modulation by the stimuli in the two culturalgroups. Americans showed greater activity inthese brain regions when perceiving dominantstimuli than when perceiving subordinatestimuli, whereas the reverse pattern of neuralactivity in the same brain regions was evidentamong Japanese. Consistent with the fMRIresults, Americans self-reported a tendencytoward more dominant behavior, whereasJapanese self-reported a tendency toward moresubordinate behavior. Moreover, activity in theright caudate and MPFC correlated with be-havioral tendencies toward dominance versussubordination. The findings suggest that func-tional activity in the mesolimbic reward systemis modulated in different (and opposite) ways inorder to coordinate with cultural preferencesfor dominant or subordinate behavior.

Taken together, these brain-imaging find-ings indicate that the same neural substrates aretuned to a particular task in opposite patternsin different cultures. This may reflect theeffects of culturally specific cognitive styles orvalues. These findings are in concordance withcontemporary social psychological models ofcultural differences in cognition and providepossible neural accounts of previously observedcultural differences in psychological tendenciesand behavior. As discussed below, in many ofthese studies explicit effort has been made tolink the cultural difference to underlying val-ues, self-construals, and/or acculturation levels.Hence, it is neither race nor nationality per se,but rather pertinent psycho-cultural dimen-sions such as independence/interdependence,individualism/collectivism, or hierarchical/egalitarian orientations that modulate thebrain activities that are observed. Thus theconcept of race may be regarded as irrelevanton the empirical level. Moreover, none of theaforementioned CN findings can be simply at-tributed to differences in physical appearance,if any, between different cultural groups.

Other CN studies have investigated culturalinfluences on neural activity by comparingsubcultural groups within a single nationalculture. For example, Han et al. (2008, 2010)examined whether and how religious beliefs


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as cultural practice modulate the neuralmechanisms underlying self-reflection. Theyscanned Christian and nonreligious Chineseparticipants during personal trait judgmentsof the self and a celebrity and found that innonreligious Chinese, self-judgments activatedthe ventral MPFC, a region that is associatedwith coding the self-relevance of stimuli (Han& Northoff 2009, Northoff et al. 2006). Incontrast, in Christian Chinese, self-judgmentsactivated the dorsal MPFC, a brain region thatis engaged in inference of others’ mental states(Grezes et al. 2004). The results suggest thatreligious beliefs produce significant effects onthe functional organization of the MPFC inself-reflection independently of race (and alsolanguage and nationality). Similarly, using anERP paradigm, Varnum et al. (2012) founddifferences in neural responses indicatingspontaneous trait inference when comparingEuropean Americans from working-class andmiddle-class backgrounds that are parallel todifferences between those observed betweenEuropean Americans and East Asians using thesame paradigm (Na & Kitayama 2011).

Association between cultural value andbrain activity. Increasingly, CN researchershave noticed that it is not enough to showcultural group differences in brain activity in-volved in a specific task. It is also importantto test whether neural activity varies acrossindividuals with different cultural values andwhether an observed cultural group differencein brain activity is mediated by a specific culturalvalue. This line of research helps to further un-cover the mechanisms of cultural modulation ofhuman brain activity.

Even individuals from the same culturalgroups may differ in many culture-relatedvalues and behaviors. Thus it is a novel issuewhether observed brain activity in a specific taskis associated with a cultural value across individ-uals. For example, given the difference in self-construals between Western and East Asiancultures (Markus & Kitayama 1991), recent CNstudies have investigated whether the variation

of brain activity across individuals is associatedwith one’s self-construal. Self-construal stylescan be estimated using the Self-ConstrualScale (Singelis 1994), which assesses individualdifferences in independent/interdependentself-construals. Goto et al. (2010) found thatthe modulation of the N400 amplitude to targetobjects by semantically incongruent versus con-gruent background scenes was stronger in AsianAmericans than in European Americans. Theyalso showed evidence that smaller-magnitudeN400 incongruity effects were associated withhigher independent self-construal scores acrossthe whole subject sample.

Other CN studies found an association be-tween self-construal measurements and brainactivity that is directly related to the pro-cessing of self-related information. Chiaoet al. (2009a) studied Japanese and CaucasianAmericans using a general self-referential task(i.e., to judge whether a sentence can de-scribe oneself in general) and a contextual self-referential task (i.e., to judge whether a sentencecan describe oneself in a specific context). Theyfound that MPFC activity during contextualversus general self-judgments was positivelycorrelated with self-reported collectivism/ indi-vidualism. Similarly, Sul et al. (2012) examinedthe neural substrates underlying self-reflectionin Koreans with different cultural orientationsand showed that interdependent self-construalspredicted stronger activation in the left superiortemporal gyrus related to personality trait judg-ments. These findings provide evidence that in-dividual differences in brain activity can be as-sociated with a specific cultural value.

While some CN studies have shown culturalgroup differences in both brain activity anda specific cultural value (e.g., de Greck et al.2012, Goto et al. 2010), other CN studies havetried to address whether cultural values mediatedifferences in neural activities that differentiatebetween two cultural groups. This has beentested using mediation analysis (MacKinnonet al. 2007), which can assess whether amediating variable transmits the effect of anindependent variable on a dependent variable.

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To examine cultural effects on neural responsesto target objects and stimulus context, Lewiset al. (2008) recorded ERPs from European andEast Asian Americans while they responded toa target stimulus (the number 6) and ignoredfrequent nontarget stimuli (three-characterwords or numbers) and an infrequent nontar-get stimulus (the number 8). A cultural valuewas measured using the Individualism andCollectivism Attitude Scale (Triandis 1995).Independent self-construal was measuredby calculating the average response on theIndividualism subscale, and interdependentself-construal was measured by calculating theaverage response on the Collectivism subscale.They first showed that European Americansdisplayed relatively greater P3 amplitudes totarget events, whereas East Asian Americansdisplayed relatively greater P3 amplitudes tothe infrequent nontarget stimulus (noveltyP3). They further found that culture predictedself-construal (the East Asian Americans weresignificantly more interdependent than theEuropean Americans) and the P3 noveltyamplitudes. Most importantly, the effects ofculture on the novelty P3 amplitudes weresignificantly reduced after including the mea-surement of interdependent self-construal asa mediator. These results indicate that therelationship between culture and novelty P3was mediated by self-construal.

Similarly, Na & Kitayama (2011) found thatthe N400 to a trait adjective during a lexi-cal decision task was enlarged when precededby a facial photo with trait-implying behaviorthat was semantically incongruent versus con-gruent with the target trait adjective. This ef-fect was evident in European Americans butnot in Asian Americans. Moreover, the N400incongruity effect was significantly enhancedwith increasing independent self-construal, andthe cultural difference in the N400 incon-gruity effect was mediated by independent self-construal. Ma and colleagues (2012) recentlyinvestigated whether the brain activity engagedduring self-reflection is different between in-dividuals who grow up in Western and East

P3: a positivepotential that peaksaround 300–400 msafter stimulus onsetwith the maximumamplitude over theparietal or frontal scalesites and is sensitive tostimulus probabilityand task relevance

Asian cultural contexts. They scanned Chineseand Danish participants during judgments ofsocial, mental, and physical attributes of them-selves and public figures to assess cultural in-fluences on self-referential processing of per-sonal attributes in different dimensions. Self-construal scale measure first confirmed greaterendorsement of the cultural value of interde-pendence in Chinese than in Danes. fMRI re-sults showed that judgments of self versus apublic figure elicited greater activation in theMPFC in Danes than in Chinese regardlessof attribute dimensions for judgments. In con-trast, self-judgments of social attributes inducedgreater activity in the temporoparietal junctionin Chinese than in Danes. Moreover, the tem-poroparietal junction activity was correlatedwith interdependence of self-construal acrossall participants, being stronger in those withgreater endorsement of the cultural value of in-terdependence. Finally, the group difference inthe temporoparietal junction activity was me-diated by the measure of interdependence ofself-construal. Thus the findings of differencesin the brain activity are consistent at both thecultural group level and at the individual level.These CN studies not only demonstrated groupdifferences in brain activity but also suggestthat the variation in brain activity across cul-tural groups is mediated by specific culturalvalues.

Modulations of neural activity by culturalpriming. CN researchers have also usedpriming in order to enable causal inferenceand to assess the degree to which culturallyinfluenced patterns of neural function arestable or malleable. This more proximalapproach to the exploration of the relationshipbetween culture and brain allows researchers toexamine cultural influences on neural activityas dynamic processes operating on a shorttime scale. One approach is to apply iconiccultural primes to bicultural participants. Forexample, in a study with bicultural participantsliving in Hong Kong, Ng et al. (2010) usedimages of Chinese or Western cultural icons


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Self-construalpriming: a procedurethat allows self-construal to shifteither to aninterdependent or anindependent style in agiven population (e.g.,by reading essays withthe instruction tofocus one’s attentionon personal pronounseither in the singularor plural form)

as cultural primes. Chinese cultural primingdecreased MPFC activity that differentiatedbetween trait judgments of the self and mother,whereas Western cultural priming producedthe opposite effect. Such dynamic variation ofthe neural correlates of the self is consistentwith previously observed differences in theneural representations of the self betweenChinese and Westerners that reflects thechronic influences of cultural values andpractices.

Another line of priming research hasfocused on manipulating cultural values thatare hypothesized to underlie group differencesin neural function. For example, Sui & Han(2007) used self-construal priming (Gardneret al. 1999) with a group of Chinese partici-pants, asking them to search for independent orinterdependent pronouns (e.g., “I” or “we”) inessays. They showed that the right frontal activ-ity related to recognition of one’s own face wassignificantly reduced after the interdependentversus independent self-construal priming.This finding indicates that the neural processinvolved in recognition of one’s own face isshaped by dynamic variation of self-construals.Self-construal priming has also been shownto modulate neural activity related to earlyperceptual processing (Lin et al. 2008). In thisstudy, Chinese participants were primed withindependent or interdependent self-construalsbefore discriminating global or local features ofhierarchical stimuli. The ERP results showedthat independent self-construal primingresulted in larger occipital P1 amplitudes tolocal targets than to global targets, whereas a re-verse pattern was observed after the interdepen-dent self-construal priming. Similarly, Chiaoet al. (2009a) primed Asian Americans using theSumerian Warrior Story Task and the Similar-ities and Differences with Family and FriendsTask (Trafimow et al. 1991), which have beenshown to influence self-construal. They foundthat priming individualistic values increasedactivation in the ventral MPFC and posteriorcingulate cortex during general self-judgmentsrelative to contextual self-judgments. Prim-ing collectivism led to the opposite pattern.

Thus the findings from priming studies in-dicate that cultural values dynamically shapeneural representations of the self and closeothers.

How does temporary access to other culturalframeworks interact with long-term culturalexperiences to shape human brain activity? Inan initial attempt to answer this question, Suiand colleagues (2012) recorded ERPs fromBritish and Chinese adults during judgmentsof orientations of one’s own and a friend’s facesafter they were primed with independent andinterdependent self-construals. They foundthat priming an interdependent self-construalreduced the default anterior N2 in response totheir own faces for British participants. By con-trast, priming an independent self-construalsuppressed the default anterior N2 in responseto their friend’s face for Chinese participants.These findings illustrate how temporary andchronic cultural orientation may interactto shape neural responses. Chronic culturalorientation may constrain the effect of culturalpriming on brain activity, reflecting a complexpattern of interactions between short-term andlong-term cultural experiences.

In sum, CN findings are obtained usingthree distinct sets of methods. Initial demon-strations of cultural differences in brain activityfocus on comparisons between individualsfrom two cultural groups (e.g., East Asiansand Westerners). This type of work suggeststhat sociocultural contexts may result indifferent patterns of brain activity related tohuman cognition. These initial demonstrationsare often followed by further analyses onunderlying value or self-construal dimensions,providing further evidence that the observedcultural group differences in brain activity areassociated with specific cultural values (e.g.,self-construals). A similar extension has alsobeen attempted with priming (manipulating thesalience of either culture or important culturalvalues) to test whether there are causal relation-ships between culture and the neurocognitiveprocesses involved in human cognition. In ad-dition to showing a causal role of cultural valuesand self-construals, the priming studies have

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demonstrated that culturally typical patterns ofbrain activity are sometimes quite malleable.

WHAT IS CULTURALNEUROSCIENCE NOT?

The rapid development of CN has induced sev-eral misunderstandings of CN research. Thesemisunderstandings relate to the origin of CNresearch, the biological versus social nature ofthe human brain, and the relationship betweenculture and race (e.g., Mateo et al. 2012).

One misunderstanding of CN research is toconnect CN studies with anthropological ap-proaches to explain the nature of culture, whichproduces the misimpression that the goal ofCN studies is simply to find biological markersin the brain that differentiate cultural groupsand to demonstrate that any cultural differencesin brain activity are determined biologicallyand are immutable. Such misunderstandingalso arises from an ontologically dualisticopposition of the biological and cultural natureof the human brain. Such a false dichotomyleads to a view of biology and culture as twoopposite accounts of the nature of the humanbrain. CN actually has a nonreductionistview of the relationships between formativebiological and cultural properties of the humanbrain. As previously noted, CN studies aim toelucidate neuroplastic and culturally generatedprocesses. This is fundamentally at odds withcultural essentialism and hard-wired biologicaldeterminism. CN researchers generate specifichypotheses about neurocognitive processesgrounded in both behavioral findings fromcultural psychology and brain imaging findingsfrom cognitive neuroscience. These hypothe-ses limit the brain regions under investigationand predict specific patterns of cultural groupdifferences and individual differences in brainactivity. Thus CN research does not studyculture as a set of biologically determinedpredispositions/constraints that can be usedto rigidly categorize collections of people.Instead, the CN approach emphasizes the flex-ibility of the human brain that enables humansto adapt to sociocultural environments.

Another misunderstanding of CN arisesfrom the confusion between culture andrace. While some CN studies have comparedparticipants from cultural groups that arealso purported to be racial groups, otherCN studies have examined cultural effectssuch as differences in brain activity betweenreligious groups and between social classes ofthe same race. CN assumes that any differencebetween these groups is primarily the result ofsocialization and chronic cultural experiences.None of the aforementioned CN findings canbe simply explained by group differences inphysical attributes (e.g., skin tone). CN studieshave shown evidence that differences in brainactivity between Westerners and East Asianscan be mediated by specific cultural values(e.g., independent versus interdependentself-construals). Cultural priming researchhas demonstrated causal effects of culture onbrain activity. These findings indicate theimportance of neuroplasticity in the study ofculture and demonstrate that culture is notviewed as an analog for race in CN research.

The confusion between culture and racemay lead to allegations of racism against CNresearch (e.g., CN findings “exert a tremendousimpact on the reproduction of stereotypes andracism”; Mateo et al. 2012, p. 158). Racistaccounts of human difference hold that humantraits are biologically determined, fixed, andthat members of racial groups are homogenousin these traits. In stark contrast to these beliefs,CN researchers view a cultural group as adynamic collection of individuals who share asimilar sociocultural context and whose mem-bers are affected by that context in divergentways (a view shared by cultural psychologists;Heine 2012). CN research regards humanneurocognitive processes as being flexible andbeing continuously shaped by sociocultural en-vironments. CN findings demonstrate that anindividual’s brain is not doomed by biology towork in a specific way, but rather that the brainis strongly shaped by long-term and short-termcultural experiences. It is sociocultural contextrather than race that matters. Knowing aboutcultural differences in neurocognitive processes


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may discourage people from believing biologi-cally essentialist accounts of race and thus mayfacilitate cross-cultural communication. In thissense, CN studies should help to reduce ratherthan facilitate the reproduction of stereotypesand racism.

IMPLICATIONS OF CULTURALNEUROSCIENCE FINDINGS

CN findings have a host of both theoreticaland practical implications. Here we list a fewof them. First, CN studies reveal the culturallysensitive nature of the human brain and help usto understand how the human brain as a biolog-ical organ is shaped by man-made socioculturalcontexts. Human beings are different fromother animals in that humans create the mostcomplicated and varied social environments.We are also unique in our capacity for culture.Although there are cultural universals, the spe-cific contents of culture are greatly important.In comparison with other species, this is animportant advantage for the development ofculture in a generalized sense as social commu-nities of conspecifics. Every person is fostered ina unique artificial environment, speaks his/hermother language, behaves in accordance withspecific social rules, acts as a member of socialinstitutions, and interacts with people whoshare specific cultural values with each other.Thus the human brain develops in a specificsociocultural context during interactions withothers. Because there are large variations acrosscultures, how to fit into one’s specific societyand how to cooperate with others efficientlyis a challenge for each person. CN studiesindicate that the human brain has the capacityto develop culture-specific neurocognitiveprocesses that help an individual to function ina specific sociocultural environment.

Second, the context-dependent nature ofthe human brain can be understood in twodifferent senses. One possibility is that theculturally different stimuli merely modulatealready preexisting neural activity that, as such,remains independent of any contextual effects.This amounts to what has been called modu-

latory context dependence (Han & Northoff2008, Northoff 2012). Alternatively, the con-stitution of any neural activity is dependentupon the context; this amounts to what can bedescribed as constitutive context dependence(Han & Northoff 2008, Northoff 2012). Thedistinction between these two conceptionsof context dependence has far reachingimplications for the relationship betweenbiological and social domains. In the case ofmodulatory context dependence, neuronal andsocial activities interact with each other whileremaining independent from each other intheir respective constitution. The brain is thenpurely neuronal and thus biological, whereasculture is social. This differs from the model ofconstitutive context dependence, which positsthat, if the constitution of the brain’s neuronalactivity depends on the respective socialcontext, a clear-cut distinction between thebiological domain of the brain and the socialdomain of culture is impossible. Rather thanbeing exclusively and completely biological,the brain and its neuronal activity must then beconsidered to be a hybrid of both biological andsocial influences. In other words, our brains arebiosocial. The brain is then a relational organthat bridges the gap between the biologicalworld of the organism and the social worldof the environment and its culture (Northoff2012).

Third, CN findings help us to understandcultural differences in human behaviors. Tradi-tionally, there was usually a dominant culture ina given society, and contact with out-groups waslimited. Cognition and behavior adhere to thecultural environment, and this may result in cul-tural imprinting effects on the brain. In currentsocieties, however, it is much easier for peopleto meet members of many different cultures.One issue raised by CN studies is what kindof experiences during development may facili-tate the ability of individuals’ brains to fit intotheir specific culture and to interact with indi-viduals from other cultures. This is particularlyimportant for those who emigrate to anotherculture. Does the brain adapt to a new culture,and if so, how quickly? CN studies, particularly

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those using cultural priming, suggest that evenan adult’s brain is quite flexibly attuned to socio-cultural environments. Thus cultural educationand experience may shape the brain in order tofit into new cultural contexts.

Fourth, because CN studies help us to un-derstand cultural differences in behavior, prac-tices, and psychological tendencies from a neu-roscience perspective, CN findings may helpto develop constructive ways to deal with mis-understandings and conflicts between differentcultural groups. In fact, CN studies not onlyshow differences in brain activity mediating be-havioral discrepancy in distinct socioculturalcontexts, but also highlight commonalities ofthe human species across cultures. Understand-ing both cultural distinctiveness and culturaluniversality in the neural mechanisms under-lying human cognition and behavior may helpto reduce intergroup conflict and prejudice andmay provide insight in how best to facilitate in-tergroup cooperation.

Finally, CN findings lead us to rethink thenature of culture. If the brain shows constitutivecontext dependence and can no longer be re-garded as purely biological but rather as bioso-cial, it means conversely that the environmentand thus culture also are not as purely social asis often assumed. Instead, culture is ingrainedby the neuronal structures and organization ofthe brain’s neural activity via the latter’s consti-tutive context dependence. Hence rather thanbeing completely and exclusively social, culturemust then be considered to be sociobiological.The empirical results from CN clearly callinto question any simplified characterization ofthe brain as merely biological and suggest thata more sophisticated account of the brain asbiosocial better fits the evidence. This in turnentails the need to redefine our concept of cul-ture and to reject any simplifications of cultureas an exclusively social construction.

FUTURE QUESTIONS FORCULTURAL NEUROSCIENCE

CN research provides a new approach to theunderstanding of the human brain from a cul-

tural perspective, and it offers at the same timea new way to enrich concepts of culture from aneuroscience perspective. CN research has al-ready provided many important insights intothe mutual constitution of culture and the brain,and it has raised intriguing questions for fu-ture researchers to address, some of which arediscussed below.

Brain and cultural value. Although there hasbeen increasing evidence for cultural differ-ences in brain activity underlying multiple cog-nitive and affective processes, future researchshould further explore the association betweencultural values and cultural group differencesin brain activity. This may help to ascribeobserved culture-specific neurocognitive pro-cesses to the effect of specific cultural valuesor practices, and it may provide a more com-plete account of variation in neurocognitiveprocesses as a function of cultural values. CNstudies have shown culturally specific patternsof brain activity. But it remains unclear whysome neural processes are sensitive to culturalinfluences whereas others are not. A more con-ceptual issue concerns how much the structureand organization of human culture in general,and the extent to which specific cultures in par-ticular, mirror the structure and organizationof the brain.

Culture, genes, and biochemistry. CNresearch has so far demonstrated cultural influ-ences on the brain in terms of activity changesin brain regions or networks. However, CNhas yet to explore how culture may affect orinteract with biochemical substances of thebrain. Cultural effects should be traced to theneuronal and biochemical levels in order tounderstand the relationship between cultureand micro-level neural processes. For instance,an important question for future researcherswill be to determine whether and how certainbiochemical substances or neurotransmittersare sensitive to cultural experiences. Exploringthis question may help us to understand theextent to which cultural influences on the brain


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operate not only on the functional neural levelbut also on the biochemical level.

CN research also needs to be integratedwith genetic imaging. Although imaging stud-ies have shown both cultural and geneticeffects on brain function, there has been littleresearch that examines whether and how cul-ture and genes interact to affect neural pro-cesses. One possible approach to this issue is toexamine genotype and culture simultaneouslyusing brain imaging. Studies along this line willoffer a comprehensive description of how cul-ture and genes interact to shape the humanbrain and may further challenge purely biolog-ical accounts of the brain.

Brain and acculturation. The increasingnumber of immigrants in the current societyraises another important issue for CN research:Is there a sensitive period during develop-ment for acculturation of the brain? Recent be-havioral research suggests that, for individualswho immigrated before approximately age 14.5,identification with a new culture increased withtime living in the new culture (Cheung et al.2011). However, for older immigrants, identifi-cation with a new culture seemed not to changewith the time in the new culture. Such findingsimply that there may be a sensitive period foracculturation of the human brain. Among indi-viduals of different ages, this can be examinedby investigating the variation of brain activityas a function of the time spent in a new culture.

Culture and abnormal brain function. CNstudies also raise issues concerning cross-cultural differences in the prevalence (andneural correlates) of psychiatric disorders suchas schizophrenia and depression. Althoughthese mental disorders appear to occur withsimilar lifetime prevalence in different cultures,it remains unclear whether their symptomaticexpression is influenced by cultural predispo-sitions (e.g., collectivism versus individualism)and whether their symptomatology is mediatedby similar patterns of abnormal neural activity.Do the same neurocognitive mechanisms medi-ate these psychiatric disorders across cultures?

Further, is the association between genotypeand mental illness similar across cultures, or isit moderated by cultural context? Answeringthese questions may help to determine whetherthe same treatments are appropriate for mentaldisorders in different cultures.

Brain and the creation of culture. Finally,one of the main goals of CN research is toaddress how the brain is involved in the cre-ation and maintenance of the cultures that existtoday. This may seem like a research questionthat is beyond the grasp of CN. However, CNstudies may create new paradigms to distributevalues and practices in a small group of par-ticipants and examine whether and how suchmanipulations may result in correspondingchanges in brain activity. Such an approach mayprovide the ability to directly test neuroscienceaccounts of cultural differences. Further, suchan approach may provide insights into howculturally influenced ways of thinking and feel-ing are acquired and represented in the humanbrain.

CONCLUSION

This review presents a refined account of CNin terms of origin, concept, method, findings,and theory, and clarifies several misunderstand-ings of the field. In sum, CN investigates thebiosocial nature of the human brain by examin-ing whether and how implicit and explicit pat-terns of beliefs, values, meanings, and practicesin specific sociocultural contexts shape the neu-ral mechanisms underlying human cognition,emotion, and behavior. CN research has helpedto provide a nuanced understanding of cultureby integrating methods from social and culturalpsychology and neuroscience. The accumulat-ing findings in the field show strong evidencefor cultural influences on the human brain andraise exciting new questions about the biosocialnature of human beings. We believe that thecontinued growth and development of CN willpromote cross-cultural understanding and pro-vide strong evidence against racist accounts ofhuman difference.

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DISCLOSURE STATEMENT

The authors are not aware of any affiliations, memberships, funding, or financial holdings thatmight be perceived as affecting the objectivity of this review.

ACKNOWLEDGMENTS

S.H. and M.V. were supported by the National Basic Research Program of China (973 Program2010CB833903) and the National Natural Science Foundation of China (Project 30910103901,91024032, 81161120539). G.N. was supported by the HDRF/ISAN, the CIHR, and the CIHR-EJLB, as well as the Michael Smith Foundation. K.V. was supported by the German VolkswagenFoundation.

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PS64-FrontMatter ARI 15 November 2012 14:20

Annual Review ofPsychology

Volume 64, 2013 Contents

Prefatory

Shifting Gears: Seeking New Approaches for Mind/Brain MechanismsMichael S. Gazzaniga � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 1

Biological Bases of Behavior

The Endocannabinoid System and the BrainRaphael Mechoulam and Linda A. Parker � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � �21

Vision

SynesthesiaJamie Ward � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � �49

Scene Perception, Event Perception, Object Recognition

Visual Aesthetics and Human PreferenceStephen E. Palmer, Karen B. Schloss, and Jonathan Sammartino � � � � � � � � � � � � � � � � � � � � � � � � �77

Attention and Performance

Detecting Consciousness: A Unique Role for NeuroimagingAdrian M. Owen � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 109

Executive FunctionsAdele Diamond � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 135

Animal Learning and Behavior

The Neuroscience of Learning: Beyond the Hebbian SynapseC.R. Gallistel and Louis D. Matzel � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 169

Evolutionary Psychology

Evolutionary Psychology: New Perspectives on Cognitionand MotivationLeda Cosmides and John Tooby � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 201

Origins of Human Cooperation and MoralityMichael Tomasello and Amrisha Vaish � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 231

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Language and Communication

Gesture’s Role in Speaking, Learning, and Creating LanguageSusan Goldin-Meadow and Martha Wagner Alibali � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 257

Nonverbal and Verbal Communication

The Antecedents and Consequences of Human Behavioral MimicryTanya L. Chartrand and Jessica L. Lakin � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 285

Intergroup Relations, Stigma, Stereotyping, Prejudice, Discrimination

Sexual PrejudiceGregory M. Herek and Kevin A. McLemore � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 309

Social Neuroscience

A Cultural Neuroscience Approach to the Biosocial Natureof the Human BrainShihui Han, Georg Northoff, Kai Vogeley, Bruce E. Wexler,

Shinobu Kitayama, and Michael E.W. Varnum � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 335

Organizational Climate/Culture

Organizational Climate and CultureBenjamin Schneider, Mark G. Ehrhart, and William H. Macey � � � � � � � � � � � � � � � � � � � � � � � � 361

Industrial Psychology/Human Resource Management

Employee RecruitmentJames A. Breaugh � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 389

Learning and Performance in Educational Settings

Self-Regulated Learning: Beliefs, Techniques, and IllusionsRobert A. Bjork, John Dunlosky, and Nate Kornell � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 417

Teaching of Subject Matter

Student Learning: What Has Instruction Got to Do With It?Hee Seung Lee and John R. Anderson � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 445

Health Psychology

Bringing the Laboratory and Clinic to the Community: MobileTechnologies for Health Promotion and Disease PreventionRobert M. Kaplan and Arthur A. Stone � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 471

Research Methodology

Multivariate Statistical Analyses for Neuroimaging DataAnthony R. McIntosh and Bratislav Misic � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 499

Contents vii

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Social Network Analysis: Foundations and Frontiers on AdvantageRonald S. Burt, Martin Kilduff, and Stefano Tasselli � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 527

Indexes

Cumulative Index of Contributing Authors, Volumes 54–64 � � � � � � � � � � � � � � � � � � � � � � � � � � � 549

Cumulative Index of Chapter Titles, Volumes 54–64 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 554

Errata

An online log of corrections to Annual Review of Psychology articles may be found athttp://psych.AnnualReviews.org/errata.shtml

viii Contents

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A Cultural Neuroscience Approach to the Biosocial Nature ...cscn-pku.com/pubs/2013 Han et al Annu Rev Psych reprint.pdfPS64CH13-Han ARI 8 November 2012 10:8 A Cultural Neuroscience

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