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A Cultural Neuroscience Approach to the Biosocial Nature of the Human Brain

Shihui 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 of China; email: [email protected] 2Mind, Brain Imaging and Neuroethics, Institute of Mental Health Research University of Ottawa, Ottawa ON K1Z 7K4, Canada 3Department of Psychiatry, University of Cologne, Cologne 50924, Germany 4Institute of Neuroscience and Medicine, Cognitive Neuroscience (INM3), Research Center Juelich, Juelich 52425, Germany 5Department of Psychiatry, Yale University, New Haven, Connecticut 06519 6Department of Psychology, University of Michigan, Ann Arbor, Michigan 48109-1109

Annu. Rev. Psychol. 2013. 64:335–59 Keywords by University of Ottawa on 10/29/13. For personal use only. First published online as a Review in Advance on cultural neuroscience, culture, brain imaging, human brain, race September 17, 2012 Abstract

Annu. Rev. Psychol. 2013.64:335-359. Downloaded from www.annualreviews.org The Annual Review of Psychology is online at psych.annualreviews.org Cultural neuroscience (CN) is an interdisciplinary field that investigates This article’s doi: the relationship between culture (e.g., value and belief systems and prac- 10.1146/annurev-psych-071112-054629 tices shared by groups) and human brain functions. In this review we Copyright c 2013 by Annual Reviews. describe the origin, aims, and methods of CN as well as its conceptual All rights reserved framework and major findings. We also clarify several misunderstand- ings of CN research. Finally, we discuss the implications of CN findings for understanding human brain function in sociocultural contexts and novel questions that future CN research should address. By doing so, we hope to provide a clear picture of the CN approach to the human brain and culture and to elucidate the intrinsically biosocial nature of the functional organization of the human brain.

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CN touches upon concepts such as humans, Contents culture, and race that are often used in a wider context beyond and outside CN. Thus it is INTRODUCTION...... 336 important to take stock of the progress so far WHAT IS CULTURAL by providing a comprehensive overview of NEUROSCIENCE?...... 336 the field as well as addressing some questions Origin...... 337 and concerns that have been voiced about this KeyConcepts...... 338 endeavor. The goals of the current review are Aims...... 340 (a) to elucidate the origin, aims and concepts, Methods...... 342 empirical methods, and findings of CN studies; Findings...... 344 (b) to clarify misunderstanding of CN findings; WHAT IS CULTURAL (c) to discuss the implications of CN findings NEUROSCIENCE NOT? ...... 351 for understanding the biosocial nature of the IMPLICATIONS OF CULTURAL human brain; and (d ) to address key questions NEUROSCIENCE FINDINGS . . . 352 for future CN studies. FUTURE QUESTIONS FOR CULTURAL NEUROSCIENCE...... 353 WHAT IS CULTURAL CONCLUSION...... 354 NEUROSCIENCE? The term cultural neuroscience was initially introduced by Chiao & Ambady (2007, p. 238), who defined CN as “a theoretical and empirical INTRODUCTION approach to investigate and characterize the During the past few years, brain imaging mechanisms by which [the] hypothesized bidi- studies have uncovered variations in the neural rectional, mutual constitution of culture, brain, substrates of human cognition in different and genes occurs.” This definition was further cultural groups. The integration of theory and refined by Chiao (2010, p. 109), who described methods from social and cultural psychology, CN as an “interdisciplinary field bridging cognitive neuroscience, and other related cultural psychology, neurosciences and neuro- disciplines gave birth to a new field—cultural genetics that explains how the neurobiological neuroscience (CN). This new field investigates processes, such as genetic expression and brain how human brain functions are shaped by function, give rise to cultural values, practices

by University of Ottawa on 10/29/13. For personal use only. interactions between culture, the brain, and and beliefs as well as how culture shapes genes. CN has developed specific methods for neurobiological processes across macro- and uncovering cultural influences on human brain microtime scales.” These concepts resonate

Annu. Rev. Psychol. 2013.64:335-359. Downloaded from www.annualreviews.org functions, and it has provided a new perspective with recent efforts to integrate research findings on the functional organization of the human from neuroscience, genetics, developmental brain in sociocultural environments. The psychology, and sociology by highlighting growth of this new field can be seen in the in- the role of postnatal neuroplasticity in human creasing number of research articles and several development (e.g., Li 2003, Wexler 2006). In special issues dedicated to CN (e.g., Progress the remainder of this section we seek to provide in Brain Research in 2009; Social Cognitive and a description of CN, starting with its origin and Affective Neuroscience in 2010) in addition to aims. Next we define key theoretical concepts edited books (e.g., Han & Poppel¨ 2011) and its in CN. This is followed by an overview of coverage in textbooks (e.g., Ward 2012). common CN methodology. Finally, we end CN is receiving more and more attention the section by reviewing the empirical findings from researchers in different fields because of CN studies using different methods.

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Origin processes develop and are shaped (Kitayama CN emerges from the integration of differ- & Uskul 2011). The findings of cultural psychological research stimulate researchers ent branches of social sciences and natural sci- Independent ences and arises mainly from two disciplines, to investigate neural substrates of cultural self-construal: i.e., cultural psychology, which has provided diversity of human cognition and emotion. a tendency to view the the insight that cognitive, emotional, and mo- self as autonomous and bounded; an emphasis tivational tendencies and habits are shaped by Social cognitive neuroscience. Social cogni- tive neuroscience research investigates brain on affirming the culture, and neuroscience, which has demon- independence and strated that the brain is shaped by experience. mechanisms that allow human beings to uniqueness of the self understand the self and others and to effi- We here show that CN is based on four key ap- Interdependent proaches, i.e., cultural psychology, social cogni- ciently navigate social environments (Ochsner self-construal: tive neuroscience, the study of neuroplasticity, & Lieberman 2001). Early social cognitive a tendency to view the self as interconnected and the study of culture × gene interactions. neuroscience research focused on the neural substrates underpinning social cognition by and overlapping with combining brain imaging and social psycho- close others; an emphasis on affirming Cultural psychology. Although there is a logical paradigms. Most of these studies aimed close relationships and long history of the debate about the concept to uncover the neural mechanisms of social maintaining harmony of culture (Kroeber & Kluckhohn 1952), it is cognition and behavior without considering within them commonly acknowledged that cultural groups potential cultural differences. However, an have differentiated over thousands of years to important feature of social cognition and create what has been called cultural speciation behavior is context dependence. We are always (Goodall & Berman 1999). The resulting situationally embedded in a certain envi- cultural differences still exist in our globalized ronment, the “context,” which substantially world and are associated with certain differ- influences our perception of others and our ences in the ways people think and behave. understanding of the behavior of others. This These differences exist side by side with context dependency itself underlies substantial many cross-cultural commonalities. Cultural influences exerted by culture. In other words, differences in human behaviors are very well what social information is processed and how documented in anthropology (e.g., Haviland it is processed rely heavily on one’s interaction et al. 2008), and human development has been partners (in the case of dyadic interactions) viewed as a process of acquiring and embodying and, more broadly, on the social context in culture’s belief systems (Rogoff 2003). Cultural which the interactions occur. For example, cul-

by University of Ottawa on 10/29/13. For personal use only. differences in human mental processes and tural psychological studies have documented underlying cognitive mechanisms have been numerous variations in social cognitive pro- investigated extensively in cultural psychology cesses across different cultural contexts, such

Annu. Rev. Psychol. 2013.64:335-359. Downloaded from www.annualreviews.org during the past two decades. From this line as construal of the self (Markus & Kitayama of research, theoretical frameworks such as 1991, 2010), causal attribution of physical individualistic versus collectivistic values, inde- and social events (Choi et al. 1999), analytic pendent self-construals versus interdependent versus holistic attention (Masuda & Nisbett self-construals, and holistic versus analytic 2001), affective states that people ideally like cognitive tendencies have emerged to guide to feel (Tsai et al. 2006), and choice-induced empirical studies of cultural discrepancy in dissonance (Kitayama et al. 2004), among human cognition and emotion (Kitayama & many others. Due to the considerable evidence Cohen 2007, Nisbett et al. 2001, Varnum et al. for cultural divergence of human subjective 2010). Cultural psychology takes the view that experiences and psychological processes, human cognitive and affective processes vary as recently neuroscientists have shown increasing a function of cultural environments that provide interest in whether parallel differences in unique social contexts in which psychological neural mechanisms might also be present

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among people who were raised in different and personal experiences (Pascual-Leone et al. sociocultural contexts (Ambady & Bharucha 2005) and to adapt to social contexts during 2009; Ames & Fiske 2010; Chiao & Bebko development (Blakemore 2008). Given that 2011; Han & Northoff 2008, 2009; Kitayama human thoughts and behaviors differ substan- & Uskul 2011; Park & Gutchess 2006; Park tially across a variety of sociocultural contexts, & Huang 2010; Rule et al. 2012). On the the- it is not surprising that the human brain, the oretical basis alone, cultural influences on the source of human behaviors and the carrier of neural substrates underlying human cognitive human thoughts, is modulated by sociocultural and affective processes would seem highly environments and develops unique neural plausible given that it takes almost 20 years for mechanisms that help an individual to adapt a large portion of the brain to mature (Gogtay to culturally specific changes and pressures. et al. 2004). During this period the brain is Thus an intrinsic feature of the brain is its influenced by personal experiences in a specific sociocultural context dependence. cultural context. Thus in the beginning of the twenty-first century, researchers started Gene × environment interactions. The ba- to examine potential cultural differences in sic assumption of CN is that culture provides a human brain mechanisms involved in multiple framework for social behavior, communication, cognitive and affective processes by comparing and interaction that generates social values brain imaging results obtained from different and norms, assigns meaning to social events, cultural groups. There is now substantial interacts with biological variables (e.g., genes), evidence that individuals from different socio- and codetermines the functional organization cultural contexts show distinct patterns of brain of the brain. The CN approach investigates activity involved in cognition and behavior. mutual interactions between culture, the brain, and genome, consistent with culture-gene Neuroplasticity. Biological research has coevolution theory (Boyd & Richerson 1985, shown ample evidence for the intrinsic plas- Lumsden & Wilson 1981). This theory guides ticity of the human brain; that is, the brain research that explores how two complementary changes both structurally and functionally in and interacting evolutionary processes, i.e., response to the environment and experience genetic evolution and cultural evolution, (Shaw & McEachern 2001). For example, the influence human behavior. The CN approach occipital cortex, which is commonly involved aims to understand how sociocultural contexts in visual processing in sighted humans, can influence human behavior by examining

by University of Ottawa on 10/29/13. For personal use only. be engaged in auditory processing in blind cultural influences on underlying neural mech- individuals (Burton et al. 2002, Gougoux et al. anisms. CN views cultural differences in the 2009). Auditory deprivation results in the neural mechanisms underlying cognition as the

Annu. Rev. Psychol. 2013.64:335-359. Downloaded from www.annualreviews.org recruitment of the primary auditory cortex in product of the interaction between genes and the processing of vibrotactile stimuli (Levanen cultural environment, an idea that researchers et al. 1998) and sign language (Nishimura et al. in this field are beginning to test empirically. 1999) in deaf humans. The medial prefrontal cortex is engaged during self-reflection on visually but not aurally presented trait words in Key Concepts sighted humans, while the region is recruited Next we review some key concepts related to during self-reflection on aurally presented trait CN. We provide definitions and also highlight words in congenitally blind individuals (Ma conceptual distinctions between terms such as & Han 2011). These findings demonstrate an culture, nationality, and race, which some may intrinsic property of the brain, plasticity, which mistakenly use interchangeably. enables the nervous system to respond to en- Culture is obviously the most impor- vironmental pressures, physiological changes, tant concept in the field of CN. From

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anthropologists’ perspective, culture refers to hypothesized to be relevant to a particular a system of meanings used to make sense of pattern of neural activity. life (i.e., Kuper 1999). Although there are myr- Recent CN studies have directly measured iad definitions of culture [indeed, Kroeber & cultural values from different groups and Kluckhohn (1952) listed 164 definitions of cul- have assessed the relationship between these ture], culture is often used in three basic senses values and neural responses. For example, in social psychology (Chiu & Hong 2006). the Self-Construal Scale (Singelis 1994) has Material culture consists of all material arti- been used in several recent CN studies to facts produced by human beings. Social cul- evaluate individuals’ independent versus in- ture consists of social rules and social institu- terdependent self-construals (e.g., Chiao et al. tions. Subjective culture refers to shared ideas, 2009a, de Greck et al. 2012, Na & Kitayama values, beliefs, and behavioral scripts. These as- 2011, Sul et al. 2012). This approach seeks pects of culture, however, are dynamically re- to simultaneously capture both within- and lated within a given cultural tradition, locale, between-group variations instead of assuming, and/or community and together form a unique a priori, that people from two sociocultural social environment for the group of individuals. contexts must have different cultural values. It From the very beginning of their life, people en- therefore acknowledges individual differences gage in the complex composed of materials and in cultural values among those who grow up social rules or practices as well as folk beliefs of in the same sociocultural context. Moreover, their respective local communities, and by do- measuring cultural values allows researchers to ing so, they have their brains changed in such examine whether a group difference in brain a way that the resulting brain functions are at- activity is associated with a specific cultural tuned closely to the surrounding sociocultural value, whether cultural group differences environment. in brain activity are mediated by a specific Culture is different from nationality, which cultural value, and how individual differences is defined by social group membership based in brain activity are associated with variations on a shared nation state of origin. Although in cultural values within a given cultural group. the term culture emphasizes shared ideas, Cultural psychology views culture as a dy- values, beliefs, and practices, people of the namic knowledge system rather than a rigid set same nationality do not necessarily share the of stereotypes about a social group (Markus & same beliefs, values, or practices. Similar to Hamedani 2007). Culture represents a dynamic social psychology studies, most CN studies use concept of the social environment that is not

by University of Ottawa on 10/29/13. For personal use only. the term culture in the sense of a social group part of the innate biological condition of hu- whose members share social values, knowledge, mans. Humans are not born with propensities and practices. Some CN studies have recruited for any particular culture but rather with the

Annu. Rev. Psychol. 2013.64:335-359. Downloaded from www.annualreviews.org participants from two different cultural groups potential and the capacity to acquire and to cre- (e.g., Westerners and East Asians) based on ate culture (Harris 1999). Thus an individual cultural psychology findings that suggest that may change his/her cultural values and beliefs the two groups differ in specific cultural values as a result of experience (such as emigrating or specific cognitive processes. In some cases from his/her native country). People from the race and language are concomitants that also same cultural groups can be quite heteroge- differentiate two cultural groups. Other CN neous in terms of the values and beliefs they studies have investigated two cultural groups acquire. This is particularly true in contempo- who are from the same nation but are defined by rary societies where cultural exchanges occur religious or political beliefs. In such cases two often and rapidly. People in modern societies groups of participants share the same nation- are rarely monocultural because they are almost ality, race, and language but differ only in a set always exposed, often deeply, to other cultures’ of shared beliefs/values and practices that are practices and beliefs in multiple sociocultural

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contexts. Thus multiple cultural systems may alike) and argued that it is very difficult to ascer- become part of any single individual, and as tain the racial identity of individuals through a consequence, it is often required to switch their genes (Littlefield et al. 1982), recent large- to and fro between different cultural systems scale studies using genetic cluster analysis have during social interactions depending on spe- found correlations between self-reported racial cific contexts of social encounters (Hong et al. group membership and the genetic cluster of 2000). This dynamic model of culture allows racial groups (e.g., Paschou et al. 2010, Tang us to test whether brain activity underlying et al. 2005). However, these data do not speak human cognition can vary on a short temporal to the question of whether racial groups are scale as a function of recent use of one cultural psychologically different, nor do they address system or another (e.g., cultural priming, as is whether such differences (if they exist) are discussed in more detail when we turn to the solely or partially genetically driven. methodology of CN research). Taken together, Regardless of whether race has a biological CN studies view culture as a complex and component, race does have a number of dynamic external social environment in which complex sociopolitical implications and may the human brain is fostered and shaped. Rather be analyzed as a sociocultural construction. than considering the brain and its neuronal Moya & Markus (2011, p. 21) recently defined states by themselves, CN emphasizes the socio- race as “a dynamic set of historically derived cultural nature of the human brain and places and institutionalized ideas and practices that great weight on the influence of cultural values, (1) sorts people into ethnic groups according beliefs, and practices shared by a social group to perceived physical and behavioral human on functional organization of the human brain. characteristics; (2) associates differential value, Race is a way of categorizing human beings power, and privilege with these characteristics on the basis of external attributes, such as skin and establishes a social status ranking among tone and facial and body shapes, that differenti- the different groups; and (3) emerges when ate human populations. In many racial theories, groups are perceived (a) to pose a threat and in lay theories, racial groups also possess (political, economic, or cultural) to each different fixed and biologically determined psy- other’s world view or way of life; and/or (b) to chological traits and tendencies. Race is viewed justify the denigration and exploitation (past, as fixed both over the course of the lifespan and current, or future) of, and prejudice toward, across cultural contexts. People from the same other groups.” In addition, race has a strong racial group are thought to be homogenous evaluative component that has been often used

by University of Ottawa on 10/29/13. For personal use only. in terms of heritage and physical appearance. to qualify individuals from different races as However, in reality individuals classified as superior or inferior on the basis of untested or belonging to the same race do not necessarily discredited assumptions related to genetics.

Annu. Rev. Psychol. 2013.64:335-359. Downloaded from www.annualreviews.org share the same cultural values and experiences. For example, Native Chinese and Chinese Americans may be thought to belong to the Aims same racial group but may have distinct cultural The goal of CN studies is to investigate human values and beliefs and experiences. There has brain function and structure in diverse socio- long been a debate over whether racial differ- cultural contexts. Like cultural psychologists ences in psychological tendencies and behavior (Markus & Hamedani 2007), CN researchers exist and if so whether such differences are bio- have little interest in using brain activity as logically determined. As a starting point, there a way to classify people into groups. Instead, is the question of whether contemporary racial CN research investigates whether and how the categories can be genetically differentiated. Al- functional organization of the human brain though some have noted a great deal of genetic is shaped by culture and by the interaction similarities across races (humans are 99.9% between culture and genes on different

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time scales (Chiao & Ambady 2007, Han & specific neural substrates in the brain, CN stud- Northoff 2008). In addition, CN research aims ies have initially focused on whether different to investigate how neurobiological processes neural substrates may be engaged in a variety in the human brain contribute to the rise of cognitive processes among individuals in of divergent cultures in the world. Theories East Asian and Western cultural contexts. As built on CN findings will eventually help to a part of this line of research, CN researchers explain how cultural differences in human brain have found evidence for cultural differences in function mediate divergent social behaviors the neural mechanisms involved in visual per- across cultures while at the same time pointing ception (Goh et al. 2007, 2010; Gutchess et al. out the neural predispositions of psychosocial 2006; Jenkins et al. 2010), attention (Hedden commonalities across different cultures. CN et al. 2008, Lewis et al. 2008), causal attribution considers culture as a highly dynamic system of physical events (Han et al. 2011), seman- of continuous interaction and exchange among tic relationship processing (Gutchess et al. individuals. This system of social interaction 2010), musical processing (Nan et al. 2006, feeds back into social practices, values, and 2009), mental calculation (Tang et al. 2006), belief systems, thereby establishing circular, recognition of one’s own face (Sui et al. 2009), recursive, and reciprocal influences between self-reflection on personality traits (Chiao et al. interacting individuals and culture (Hacking 2009a,b; Wang et al. 2012; Zhu et al. 2007), 1999, Vogeley & Roepstorff 2009). perception of bodily expression (Freeman et al. Most current CN studies focus on cross- 2009), mental state reasoning (Adams et al. cultural differences in the neural substrates 2009, Kobayashi et al. 2006), empathy (de of human psychological processes including Greck et al. 2012), and in other domains. These cognition, emotion, and motivation (Ambady CN findings indicate that a fundamental aspect & Bharucha 2009; Han & Northoff 2008, 2009; of the functional organization of the human Kitayama & Uskul 2011). This line of research brain is its sensitivity to the sociocultural con- has mainly been stimulated by findings in texts in which individuals are brought up. How- cultural psychology that show cross-cultural ever, the final goal of CN research is not simply variation in multiple levels of psychological to show differences in brain activity across processes (Kitayama & Cohen 2007, Nisbett cultural groups. Rather, CN studies aim to pro- et al. 2001). By comparing behavioral perfor- vide a neuroscientific account of cross-cultural mances among individuals from Western (e.g., variation in human psychological functions European and American) and East Asian (e.g., and behaviors by discovering socioculturally

by University of Ottawa on 10/29/13. For personal use only. Chinese, Japanese, Korean) contexts, cultural patterned neural mechanisms and their devel- psychologists have shown evidence for dis- opment. Thus CN studies aim to reveal both tinct, culture-dependent cognitive processing culturally universal and culturally unique neu-

Annu. Rev. Psychol. 2013.64:335-359. Downloaded from www.annualreviews.org styles in perception ( Ji et al. 2000), attention ral processes by which human brains predispose (Kitayama et al. 2003, Masuda & Nisbett us to perceive self and others, communicate and 2001), memory (Wang & Conway 2004), interact with conspecifics, and guide actions. perspective taking (Wu & Keysar 2007), causal Recently, there has been an increasing attribution of events (Morris & Peng 1994, interest in how brain function is shaped by Peng & Knowles 2003), object categorization culture-gene interaction, based on culture-gene ( Ji et al. 2004), recognition of one’s own face coevolution theory (Boyd & Richerson 1985, (Liew et al. 2011b; Ma & Han 2009, 2010; Sui Lumsden & Wilson 1981). However, although et al. 2009), self-construal (Markus & Kitayama recent studies have shown that psychological 1991), and affect valuation (Tsai et al. 2006), tendencies and behavioral outcomes associated among many others. with specific genotypes are moderated by cul- Based on the fundamental hypothesis that ture (e.g., Kim et al. 2010a,b), direct evidence human psychological processes are mediated by that culture moderates the effect of genotype on

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the functional organization of the brain is still on whether and how two cultural groups differ lacking. Another line of research has focused in neural substrates of specific cognitive and on how the allelic frequencies of a genotype affective processes. A typical way to address Functional magnetic resonance imaging within a population may relate to cultural this issue is to compare functional magnetic (fMRI): a noninvasive differences in values. Chiao & Blizinsky (2010) resonance imaging (fMRI) or event-related method for recording examined the relationship between the cultural potential (ERP) brain data obtained from blood-oxygenation- phenotypes of individualism-collectivism and individuals who were raised in two different level-dependent allelic frequency of the serotonin transporter sociocultural contexts. signals that have high spatial resolution and functional polymorphism (5-HTTLPR) by One assumption of this approach is that, are used to examine assessing the prevalence of the short allele because participants from two cultural groups brain activations of 5-HTTLPR among different populations. differ in cultural knowledge, values, and/or associated with specific In a comparison of 29 countries, they found cognitive and affective processes, the un- stimuli or tasks that cultures that were high in collectivism derlying neural activity should be different Event-related contained a significantly greater proportion between the groups in a specific way. To potential (ERP): of short allele carriers and that increased address this assumption, CN research has synchronous activities of neuronal frequency of short allele carriers predicted taken cultural psychological research as a guide populations engaged in decreased anxiety and mood disorder preva- for its hypotheses about neural differences specific psychological lence. Further, the relationships between the between specific cultural groups. For example, processing, which are prevalence of short allele carriers within a behavioral research first showed that individ- time locked to population and the prevalence of anxiety and uals in Western cultures are more sensitive to stimulus events, can be recorded from mood disorders were mediated by collectivism. salient foreground objects compared to people electrodes over the Similarly, Way & Lieberman (2010) suggested in East Asian cultures, whereas individuals scalp, and have high that collectivism may have developed and in East Asian cultures are more inclined to temporal resolution persisted in populations with a high proportion focus their attention broadly on backgrounds of a functional polymorphism (A118G) in relative to people in Western cultures ( Ji et al. the μ-opioid receptor gene—a putative social 2000, Kitayama et al. 2003, Masuda & Nisbett sensitivity genotype that is compatible with 2001). Such findings lead to a reasonable collectivistic cultural groups. Caution is due hypothesis that neural substrates underlying because this body of evidence is entirely corre- visual perception of and attention to salient lational. Nevertheless, given that cultures may objects and contexts may show different interact with the 5-HTTLPR genotypes to patterns between individuals in the Western influence the prevalence of affective disorders and East Asian cultures (Goh et al. 2007,

by University of Ottawa on 10/29/13. For personal use only. such as anxiety and depression (Chiao & Gutchess et al. 2006, Hedden et al. 2008). Blizinsky 2010), it would be interesting to Similarly, evidence from cultural psychology study how the culture-gene interaction code- that the self is viewed as independent in

Annu. Rev. Psychol. 2013.64:335-359. Downloaded from www.annualreviews.org termines the intermediate endophenotype (i.e., Western cultures and interdependent in East neurobiological responsiveness) associated Asian cultures (Markus & Kitayama 1991, Sin- with culture-sensitive cognitive processes. gelis 1994) leads to the hypothesis that neural representation of the self and close others may overlap to a greater degree among East Asians Methods than among Westerners (Zhu et al. 2007). Although CN is a young field, CN researchers However, selection of participants from two have developed quite sophisticated method- different nations or sociocultural contexts does ologies by drawing on prior brain imaging not necessarily imply that the participants have and social and cultural psychology research. distinct cultural values (Oyserman et al. 2002). Methodological challenges include both the CN seeks to address this question by measuring design of psychological experiments and brain the value or self-construal dimensions that imaging techniques. Early CN studies focused are hypothesized to drive the relevant cultural

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differences in the neural process being studied. to reduce the effect of language processing to a Often, CN researchers directly assess the minimum degree. cultural values or self-construals of interest; Another elegant psychological paradigm this can be done using well-established ques- used by CN researchers is to prime cultural tionnaires developed by social and cultural identity or values before recording brain activ- psychologists. For example, the Self-Construal ity during a specific task. Such studies are based Scale (Singelis 1994) is widely used to evaluate on the assumption that individuals can acquire how people view themselves (either as indepen- more than one set of cultural knowledge and dent or interdependent) and has been shown can use different sets of cultural knowledge de- to differentiate cultural groups. Current CN pending on contextual cues (Hong et al. 2000). studies usually compare participants from two According to this dynamic constructivist model countries, usually a Western culture and an of culture, people who have been exposed to East Asian culture (e.g., British versus Chinese, multiple cultures may acquire multiple sets of American versus Japanese). Although previous cultural knowledge, and exposing individuals studies have demonstrated differences between to cultural symbols may activate specific these cultures in terms of values and cognitive cultural knowledge and result in mindsets and and affective processes, it is important to behaviors that are consistent with that culture. demonstrate that the participants recruited in For instance, after cultural priming, bicultural CN studies actually differ in these dimensions. individuals may switch between Western and Measurements of cultural values are also helpful East Asian mindsets that are consistent with the in situating CN studies as dealing with cultural most accessible cultural knowledge tradition differences rather than racial or national differ- (e.g., Hong et al. 2003). A number of studies ences. Equally important, measuring cultural have also shown that priming independence values allows for stronger inferences. By exam- and interdependence of self-construals in- ining whether individual differences in cultural fluences patterns of cognitive processes that values can predict individual differences in tend to differ between cultural groups (e.g., brain activity, researchers can probe the asso- Kuhnen¨ & Oyserman 2002, Lin & Han 2009; ciation between these values and patterns of for a review, see Oyserman & Lee 2008). brain activity. In addition, measuring cultural Similarly, CN studies take culture as a dynamic values in individual subjects makes it possible knowledge and meaning system, and they to assess whether cultural values mediate differ- manipulate cultural values as variables (Chiao ences in brain activity associated with specific et al. 2009b, Lin et al. 2008, Ng et al. 2010, Sui

by University of Ottawa on 10/29/13. For personal use only. tasks, which may be performed in different & Han 2007, Sui et al. 2012). CN studies using ways according to the respective degree of the cultural priming test for dynamic changes in cultural value, between two cultural groups. brain activity as a function of cultural values

Annu. Rev. Psychol. 2013.64:335-359. Downloaded from www.annualreviews.org Of course, when comparing participants and provide information that allows for causal from different sociocultural contexts, it is im- inference regarding the relationship between portant to control for potentially confounding cultural values and specific brain activity. variables such as gender, age, and education as CN studies often compare brain-imaging well as socioeconomic status. Language is an- results from multiple cultural groups, and thus other potential confound in cultural compar- several technical issues have to be considered isons if stimuli used in brain imaging studies are when designing CN research. It is ideal to scan based on verbal materials. This, however, can multiple cultural groups at the same experi- be controlled by using the native language for mental site in cross-cultural fMRI studies. This each cultural group so that the same language is can certainly avoid systematic, site-dependent used in an experimental condition and a control effects in fMRI sensitivity between the scanner condition. Such designs allow us to compare facilities, although between-subject differences the experimental and control conditions so as can account for nearly ten times more variance

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than site effects when participants are scanned intended to give an extensive review of current using the same type of scanner at two sites CN findings. Instead, we highlight a select set (Sutton et al. 2008). For those CN studies of recent CN studies in terms of their method- that scan multiple cultural groups at different ology in order to illustrate the intellectual sites, Chiao et al. (2010) has suggested several development of CN research. ways to reduce the probability of systematic, site-dependent effects in fMRI sensitivity. Distinct neural activity to cultural familiar/ First, both functional and anatomical MRI unfamiliar stimuli. One question that CN data should be collected using scanners from researchers are interested in is how human the same vendor with identical protocols. brain activity is tuned by culturally famil- Second, an interscanner reliability test can be iar/unfamiliar information. To address this, conducted by scanning a separate cohort of CN researchers simply recorded neural activity participants or phantom data at each scanner to culturally familiar/unfamiliar stimuli from facility, thus enabling one to quantify and sta- one cultural group. For example, to investigate tistically compare signal-to-noise ratio across the neural basis of musical phrase boundary scanner sites. Third, the presentation software processing during the perception of music from and hardware should be identical, calibrated native and nonnative cultures, Nan et al. (2008) and tested at each session, and scripts should used fMRI to record brain activity in German be written and implemented across the sites in musicians while they categorized phrased a culturally appropriate manner. Western and Chinese musical excerpts. They Relative to fMRI, electroencephalogram found that culturally familiar musical excerpts (EEG) is a simpler method for cross-cultural more strongly activated multiple brain regions comparisons of brain activity. Similar EEG including the superior frontal gyrus, the poste- recording systems can be found easily at differ- rior precentral gyrus, and the superior temporal ent recording sites. Portable EEG amplifiers gyrus, possibly reflecting enhanced sensorimo- are available and can be easily transferred tor integration. Culturally unfamiliar musical between different recording sites so that EEG excerpts, however, more strongly activated the data from different cultural groups can be posterior insula as well as the middle frontal and recorded using the same system. Analyzing angular gyri, possibly due to higher demands ERPs that are time locked to a stimulus or a on attention systems and higher loads on basic response is particularly helpful for uncovering auditory processing. Similarly, Demorest & the time course of neural responses to multiple Osterhout (2012) recorded ERPs while Amer-

by University of Ottawa on 10/29/13. For personal use only. cognitive processes such as recognition of one’s ican participants listened to melodies based in own face (Sui et al. 2009, 2012), inference of the Western folk tradition or North Indian one’s own and others’ personality traits (Mu classical music. ERPs showed that a long latency

Annu. Rev. Psychol. 2013.64:335-359. Downloaded from www.annualreviews.org & Han 2010, Na & Kitayama 2011), implicit positive activity was sensitive to the original and processing of vocal tone (Ishii et al. 2010), deviation form of the melodies, and this effect emotion regulation (Murata et al. 2012), and was more salient in the Western than in the musical processing (Nan et al. 2006, 2009). Indian context. The results suggest that people may generate specific expectancies when listen- ing to culturally familiar music, whereas they Findings may remain unable to develop such expectan- Several recent review articles have summarized cies when hearing culturally unfamiliar music. major findings from CN research (e.g., Am- Another example of this line of research bady & Bharucha 2009; Ames & Fiske 2010; examined whether observations of culturally Chiao & Bebko 2011; Han & Northoff 2008, familiar/unfamiliar symbolic gestures engage 2009; Kitayama & Uskul 2011; Park & Huang distinct neural subsystems. Liew and colleagues 2010; Rule et al. 2012). Thus this section is not (2011a) scanned Chinese participants while

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they perceived video clips in which a model One type of cultural modulation of brain showed culturally familiar/unfamiliar symbolic activity is that a specific neural activity is gestures. They found that culturally familiar significantly modulated by a particular task in Temporoparietal gestures increased activity in the posterior one cultural group but not in another cultural junction: the cortical cingulate cortex, the dorsal portion of the group. Based on the assumption that West- junction zone at the medial prefrontal cortex and the bilateral erners tend to attend to salient objects whereas border of the posterior temporoparietal junction. These brain regions East Asians are inclined to attend to a broad parts of the temporal constitute the neural circuit engaged in infer- perceptual and conceptual field (Nisbett et al. lobe and the inferior parts of the parietal ence of others’ intentions and beliefs (Frith & 2001, Nisbett & Masuda 2003), Jenkins et al. lobe, which has been Frith 2006). In contrast, unfamiliar gestures (2010) tested whether the neural activity in shown to be involved generated activity in the left inferior parietal the lateral occipital cortex to a target stimulus in belief reasoning and lobule, the left superior frontal gyrus, and the was more sensitive to background scenes in perspective taking bilateral superior parietal lobule. These brain East Asians than in Westerners. They scanned N400: a negative regions make up the neural network involved American and Chinese participants during per- potential that peaks in automatic motor simulations of observed ception of pictures consisting of a focal object around 400 ms after stimulus onset with the actions (Rizzolatti & Sinigaglia 2010). Appar- superimposed upon a background scene that maximum amplitude ently, the mentalizing system is engaged during was congruent (e.g., a deer in the woods) or in- over the parietal scale observation of culturally familiar gestures in congruent (e.g., a television in the desert) with site and is sensitive to order to understand others’ intentions or the target object. The target object was pre- semantic incongruity beliefs, whereas the mirror neuron system may sented on different novel scenes or on a single between stimuli be activated during perception of culturally un- repeated scene on four successive trials. Adap- familiar gestures so as to capture others’ mind tation magnitude was calculated by subtracting through automatic motor simulations of ob- the neural activity to objects on a repeated scene served actions. Taken together, these findings from that to objects on different novel scenes. indicate that long-term cultural experiences Jenkins et al. found that the neural activity in may result in specific neural mechanisms in the both the right and left lateral occipital cortex human brain that deal with culturally familiar showed significantly greater adaptation to information. This may allow the individual incongruent scenes than to congruent scenes, to quickly comprehend the meaning of social suggesting sensitivity of the occipital activity information in one’s own culture, predict to the background scenes. However, this effect others’ behavior, and take appropriate actions was evident in Chinese participants but not in in a specific cultural context. American participants. Similar results were ob-

by University of Ottawa on 10/29/13. For personal use only. served in another study that recorded ERPs to Cultural group differences in neural activ- target objects that were presented on semantic ity. The mainstream of CN studies focuses on congruent or incongruent background scenes

Annu. Rev. Psychol. 2013.64:335-359. Downloaded from www.annualreviews.org whether differences in cognitive processes be- (Goto et al. 2010). It was found that a negative tween two cultural groups revealed by cultural ERP component peaking at about 400 ms after psychology are associated with distinct pat- stimulus onset (N400), which has been shown terns of brain activity. There is now increasing to be sensitive to processing semantic relation- evidence that two cultural groups may employ ships (Kutas & Hillyard 1984), was enlarged by distinct neural mechanisms while performing target objects presented on semantically incon- seemingly identical cognitive and emotional gruent versus congruent background scenes; tasks. This is of fundamental importance to however, this effect was observed in Asian understanding brain-behavior relationships in Americans but not in European Americans. general as well as to CN. There are two pat- This pattern of cultural differences was terns of cultural group difference in the neural also observed in neural activity involved in activity involved in cognitive and affective high-level social cognition. To investigate processes. cultural differences in the neural mechanisms

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underlying causal attribution of physical and Westerners using fMRI during trait events, Han and colleagues (2011) first scanned judgments of oneself, a close other (i.e., one’s Chinese participants during causality versus mother), and a celebrity. They found that, Medial prefrontal cortex (MPFC): the motion direction judgments on animations of relative to trait judgments of a celebrity, trait medial region of the object collisions. Causality judgments asked judgments of oneself significantly activated the prefrontal cortex that participants to infer causes of physical events ventral region of the MPFC in both Chinese has been shown to be (i.e., changes of motion direction of a target and Westerners, suggesting a similar neural involved in social object after colliding with another object), and substrate of representation of oneself in the cognition, with the dorsal region engaged motion direction judgments required iden- two cultural groups. However, trait judgments in mental state tification of the motion direction of a target of one’s mother versus a celebrity activated the reasoning and the object after colliding with another object. same brain region in Chinese but not in West- ventral region engaged They showed that, relative to motion direction erners, suggesting shared neural representation in self-reflection judgments, causality judgments activated the of the self and a close other in Chinese but not medial prefrontal cortex (MPFC) and the left in Westerners. This finding reveals a neural parietal cortex (LPC). Moreover, they found model of cross-cultural variations in represen- that MPFC activity was sensitive to the demand tations of a close other in relation to the self. to infer causes of events, whereas LPC activity It is not always true that individuals from was modulated by the contextual complexity East Asian cultures show additional modu- of physical events. In a subsequent experiment, lations of neural activity in comparison with Han et al. (2011) scanned American and Chi- individuals from Western cultures. In a recent nese participants during causality versus motion fMRI study, de Greck et al. (2012) examined direction judgments on animations of object cultural differences in brain activity during collisions. They found that American and empathy with anger. They scanned Chinese Chinese participants showed similar MPFC and German participants during an intentional activity involved in causality judgments. empathy task and found empathy-related However, LPC activity elicited by causality neural activity that was specific for each judgments of physical events was evident in cultural group. Specifically, empathy for angry Chinese but not in Americans. Thus it can be faces activated the left dorsolateral prefrontal concluded that LPC activity associated with cortex in Chinese participants, but it activated the contextual processing is more sensitive the right temporoparietal junction, the right to cultural differences in causality perception inferior and superior temporal gyrus, and the than is MPFC activity engaged in inference of left middle insula in Germany participants.

by University of Ottawa on 10/29/13. For personal use only. causal relationships. These results implicate enhanced emotion reg- Zhu et al. (2007) tested cultural differences ulation during empathy with anger in Chinese in the neural activity underlying representation culture, in which the attitude toward harmony

Annu. Rev. Psychol. 2013.64:335-359. Downloaded from www.annualreviews.org of personality traits of oneself and a close is more valued (Kim & Markus 1999, Markus other. According to Markus & Kitayama & Kitayama 1991). However, in Germans, (1991, 2010), Western cultures encourage empathy with anger may be characterized by self-identity that is independent of social con- enhanced inference of others’ mind, given the texts and others, whereas East Asian cultures key role of the temporoparietal junction in emphasize fundamental social connections, mental state reasoning (Frith & Frith 2006). leading to an interdependent view of the self The aforementioned neuroimaging find- and partial overlap in representation of the self ings are consistent with the idea that cultural and close others. This proposition may predict practices may produce specific psychological shared neural mechanisms of representation of processes that are significant in one culture but the self and a close other in East Asian cultures not in another culture. One may consequently but not in Western cultures. To test this, associate culturally specific psychological Zhu and colleagues (2007) scanned Chinese process with particular underlying neural

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mechanisms that are observed in one culture modulation by the stimuli in the two cultural but not in others. groups. Americans showed greater activity in Another type of cultural modulation of these brain regions when perceiving dominant brain activity is that a specific neural activity is stimuli than when perceiving subordinate modulated by a particular task in two cultural stimuli, whereas the reverse pattern of neural groups but in opposite ways. These opposite activity in the same brain regions was evident patterns of neural activity may have nothing to among Japanese. Consistent with the fMRI do with the cultural familiarity of the stimuli, results, Americans self-reported a tendency but may instead reflect culturally specific toward more dominant behavior, whereas cognitive styles. Hedden et al. (2008) assessed Japanese self-reported a tendency toward more cultural differences in the neural activity un- subordinate behavior. Moreover, activity in the derlying attentional control by scanning East right caudate and MPFC correlated with be- Asians and Americans in context-dependent or havioral tendencies toward dominance versus context-independent judgment tasks. Partic- subordination. The findings suggest that func- ipants were presented with a series of stimuli, tional activity in the mesolimbic reward system each consisting of a vertical line inside a box. is modulated in different (and opposite) ways in The context-dependent task required judg- order to coordinate with cultural preferences ments of whether the box and line combination for dominant or subordinate behavior. of each stimulus matched the proportional Taken together, these brain-imaging find- scaling of the preceding combination. The ings indicate that the same neural substrates are context-independent judgment task required tuned to a particular task in opposite patterns judgments of whether the current line matched in different cultures. This may reflect the the previous line, regardless of the size of the effects of culturally specific cognitive styles or accompanying box. It was found that the neural values. These findings are in concordance with activity in the prefrontal and parietal cortices contemporary social psychological models of involved in the tasks showed an opposite pattern cultural differences in cognition and provide of activations in the two cultural groups; that is, possible neural accounts of previously observed Americans showed greater prefrontal and pari- cultural differences in psychological tendencies etal activity during the context-dependent than and behavior. As discussed below, in many of context-independent tasks, whereas East Asians these studies explicit effort has been made to exhibited stronger activity in the prefrontal link the cultural difference to underlying val- and parietal cortices during the context- ues, self-construals, and/or acculturation levels.

by University of Ottawa on 10/29/13. For personal use only. independent than context-dependent tasks. Hence, it is neither race nor nationality per se, The opposite pattern of neural activity was but rather pertinent psycho-cultural dimen- interpreted as reflecting enhanced sustained sions such as independence/interdependence,

Annu. Rev. Psychol. 2013.64:335-359. Downloaded from www.annualreviews.org attentional control during culturally nonpre- individualism/collectivism, or hierarchical/ ferred in comparison with preferred tasks. egalitarian orientations that modulate the Opposite patterns of neural activity in two brain activities that are observed. Thus the cultural groups may also arise from distinct concept of race may be regarded as irrelevant cultural values. To examine why American on the empirical level. Moreover, none of the culture tends to reinforce dominant behavior aforementioned CN findings can be simply at- whereas Japanese culture tends to reinforce tributed to differences in physical appearance, subordinate behavior, Freeman et al. (2009) if any, between different cultural groups. scanned American and Japanese individuals Other CN studies have investigated cultural during perception of body displays related influences on neural activity by comparing to dominance and subordination. The neu- subcultural groups within a single national ral activity in the bilateral caudate nucleus culture. For example, Han et al. (2008, 2010) and MPFC showed an opposite pattern of examined whether and how religious beliefs

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as cultural practice modulate the neural of brain activity across individuals is associated mechanisms underlying self-reflection. They with one’s self-construal. Self-construal styles scanned Christian and nonreligious Chinese can be estimated using the Self-Construal participants during personal trait judgments Scale (Singelis 1994), which assesses individual of the self and a celebrity and found that in differences in independent/interdependent nonreligious Chinese, self-judgments activated self-construals. Goto et al. (2010) found that the ventral MPFC, a region that is associated the modulation of the N400 amplitude to target with coding the self-relevance of stimuli (Han objects by semantically incongruent versus con- & Northoff 2009, Northoff et al. 2006). In gruent background scenes was stronger in Asian contrast, in Christian Chinese, self-judgments Americans than in European Americans. They activated the dorsal MPFC, a brain region that also showed evidence that smaller-magnitude is engaged in inference of others’ mental states N400 incongruity effects were associated with (Grezes` et al. 2004). The results suggest that higher independent self-construal scores across religious beliefs produce significant effects on the whole subject sample. the functional organization of the MPFC in Other CN studies found an association be- self-reflection independently of race (and also tween self-construal measurements and brain language and nationality). Similarly, using an activity that is directly related to the pro- ERP paradigm, Varnum et al. (2012) found cessing of self-related information. Chiao differences in neural responses indicating et al. (2009a) studied Japanese and Caucasian spontaneous trait inference when comparing Americans using a general self-referential task European Americans from working-class and (i.e., to judge whether a sentence can de- middle-class backgrounds that are parallel to scribe oneself in general) and a contextual self- differences between those observed between referential task (i.e., to judge whether a sentence European Americans and East Asians using the can describe oneself in a specific context). They same paradigm (Na & Kitayama 2011). found that MPFC activity during contextual versus general self-judgments was positively Association between cultural value and correlated with self-reported collectivism/ indi- brain activity. Increasingly, CN researchers vidualism. Similarly, Sul et al. (2012) examined have noticed that it is not enough to show the neural substrates underlying self-reflection cultural group differences in brain activity in- in Koreans with different cultural orientations volved in a specific task. It is also important and showed that interdependent self-construals to test whether neural activity varies across predicted stronger activation in the left superior

by University of Ottawa on 10/29/13. For personal use only. individuals with different cultural values and temporal gyrus related to personality trait judg- whether an observed cultural group difference ments. These findings provide evidence that in- in brain activity is mediated by a specific cultural dividual differences in brain activity can be as-

Annu. Rev. Psychol. 2013.64:335-359. Downloaded from www.annualreviews.org value. This line of research helps to further un- sociated with a specific cultural value. cover the mechanisms of cultural modulation of While some CN studies have shown cultural human brain activity. group differences in both brain activity and Even individuals from the same cultural a specific cultural value (e.g., de Greck et al. groups may differ in many culture-related 2012, Goto et al. 2010), other CN studies have values and behaviors. Thus it is a novel issue tried to address whether cultural values mediate whether observed brain activity in a specific task differences in neural activities that differentiate is associated with a cultural value across individ- between two cultural groups. This has been uals. For example, given the difference in self- tested using mediation analysis (MacKinnon construals between Western and East Asian et al. 2007), which can assess whether a cultures (Markus & Kitayama 1991), recent CN mediating variable transmits the effect of an studies have investigated whether the variation independent variable on a dependent variable.

348 Han et al. PS64CH13-Han ARI 8 November 2012 10:8

To examine cultural effects on neural responses Asian cultural contexts. They scanned Chinese to target objects and stimulus context, Lewis and Danish participants during judgments of et al. (2008) recorded ERPs from European and social, mental, and physical attributes of them- P3: a positive East Asian Americans while they responded to selves and public figures to assess cultural in- potential that peaks a target stimulus (the number 6) and ignored fluences on self-referential processing of per- around 300–400 ms frequent nontarget stimuli (three-character sonal attributes in different dimensions. Self- after stimulus onset words or numbers) and an infrequent nontar- construal scale measure first confirmed greater with the maximum get stimulus (the number 8). A cultural value endorsement of the cultural value of interde- amplitude over the parietal or frontal scale was measured using the Individualism and pendence in Chinese than in Danes. fMRI re- sites and is sensitive to Collectivism Attitude Scale (Triandis 1995). sults showed that judgments of self versus a stimulus probability Independent self-construal was measured public figure elicited greater activation in the and task relevance by calculating the average response on the MPFC in Danes than in Chinese regardless Individualism subscale, and interdependent of attribute dimensions for judgments. In con- self-construal was measured by calculating the trast, self-judgments of social attributes induced average response on the Collectivism subscale. greater activity in the temporoparietal junction They first showed that European Americans in Chinese than in Danes. Moreover, the tem- displayed relatively greater P3 amplitudes to poroparietal junction activity was correlated target events, whereas East Asian Americans with interdependence of self-construal across displayed relatively greater P3 amplitudes to all participants, being stronger in those with the infrequent nontarget stimulus (novelty greater endorsement of the cultural value of in- P3). They further found that culture predicted terdependence. Finally, the group difference in self-construal (the East Asian Americans were the temporoparietal junction activity was me- significantly more interdependent than the diated by the measure of interdependence of European Americans) and the P3 novelty self-construal. Thus the findings of differences amplitudes. Most importantly, the effects of in the brain activity are consistent at both the culture on the novelty P3 amplitudes were cultural group level and at the individual level. significantly reduced after including the mea- These CN studies not only demonstrated group surement of interdependent self-construal as differences in brain activity but also suggest a mediator. These results indicate that the that the variation in brain activity across cul- relationship between culture and novelty P3 tural groups is mediated by specific cultural was mediated by self-construal. values. Similarly, Na & Kitayama (2011) found that

by University of Ottawa on 10/29/13. For personal use only. the N400 to a trait adjective during a lexi- Modulations of neural activity by cultural cal decision task was enlarged when preceded priming. CN researchers have also used by a facial photo with trait-implying behavior priming in order to enable causal inference

Annu. Rev. Psychol. 2013.64:335-359. Downloaded from www.annualreviews.org that was semantically incongruent versus con- and to assess the degree to which culturally gruent with the target trait adjective. This ef- influenced patterns of neural function are fect was evident in European Americans but stable or malleable. This more proximal not in Asian Americans. Moreover, the N400 approach to the exploration of the relationship incongruity effect was significantly enhanced between culture and brain allows researchers to with increasing independent self-construal, and examine cultural influences on neural activity the cultural difference in the N400 incon- as dynamic processes operating on a short gruity effect was mediated by independent self- time scale. One approach is to apply iconic construal. Ma and colleagues (2012) recently cultural primes to bicultural participants. For investigated whether the brain activity engaged example, in a study with bicultural participants during self-reflection is different between in- living in Hong Kong, Ng et al. (2010) used dividuals who grow up in Western and East images of Chinese or Western cultural icons

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as cultural primes. Chinese cultural priming Thus the findings from priming studies in- decreased MPFC activity that differentiated dicate that cultural values dynamically shape between trait judgments of the self and mother, neural representations of the self and close Self-construal priming: a procedure whereas Western cultural priming produced others. that allows self- the opposite effect. Such dynamic variation of How does temporary access to other cultural construal to shift the neural correlates of the self is consistent frameworks interact with long-term cultural either to an with previously observed differences in the experiences to shape human brain activity? In interdependent or an neural representations of the self between an initial attempt to answer this question, Sui independent style in a given population (e.g., Chinese and Westerners that reflects the and colleagues (2012) recorded ERPs from by reading essays with chronic influences of cultural values and British and Chinese adults during judgments the instruction to practices. of orientations of one’s own and a friend’s faces focus one’s attention Another line of priming research has after they were primed with independent and on personal pronouns focused on manipulating cultural values that interdependent self-construals. They found either in the singular or plural form) are hypothesized to underlie group differences that priming an interdependent self-construal in neural function. For example, Sui & Han reduced the default anterior N2 in response to (2007) used self-construal priming (Gardner their own faces for British participants. By con- et al. 1999) with a group of Chinese partici- trast, priming an independent self-construal pants, asking them to search for independent or suppressed the default anterior N2 in response interdependent pronouns (e.g., “I” or “we”) in to their friend’s face for Chinese participants. essays. They showed that the right frontal activ- These findings illustrate how temporary and ity related to recognition of one’s own face was chronic cultural orientation may interact significantly reduced after the interdependent to shape neural responses. Chronic cultural versus independent self-construal priming. orientation may constrain the effect of cultural This finding indicates that the neural process priming on brain activity, reflecting a complex involved in recognition of one’s own face is pattern of interactions between short-term and shaped by dynamic variation of self-construals. long-term cultural experiences. Self-construal priming has also been shown In sum, CN findings are obtained using to modulate neural activity related to early three distinct sets of methods. Initial demon- perceptual processing (Lin et al. 2008). In this strations of cultural differences in brain activity study, Chinese participants were primed with focus on comparisons between individuals independent or interdependent self-construals from two cultural groups (e.g., East Asians before discriminating global or local features of and Westerners). This type of work suggests

by University of Ottawa on 10/29/13. For personal use only. hierarchical stimuli. The ERP results showed that sociocultural contexts may result in that independent self-construal priming different patterns of brain activity related to resulted in larger occipital P1 amplitudes to human cognition. These initial demonstrations

Annu. Rev. Psychol. 2013.64:335-359. Downloaded from www.annualreviews.org local targets than to global targets, whereas a re- are often followed by further analyses on verse pattern was observed after the interdepen- underlying value or self-construal dimensions, dent self-construal priming. Similarly, Chiao providing further evidence that the observed et al. (2009a) primed Asian Americans using the cultural group differences in brain activity are Sumerian Warrior Story Task and the Similar- associated with specific cultural values (e.g., ities and Differences with Family and Friends self-construals). A similar extension has also Task (Trafimow et al. 1991), which have been been attempted with priming (manipulating the shown to influence self-construal. They found salience of either culture or important cultural that priming individualistic values increased values) to test whether there are causal relation- activation in the ventral MPFC and posterior ships between culture and the neurocognitive cingulate cortex during general self-judgments processes involved in human cognition. In ad- relative to contextual self-judgments. Prim- dition to showing a causal role of cultural values ing collectivism led to the opposite pattern. and self-construals, the priming studies have

350 Han et al. PS64CH13-Han ARI 8 November 2012 10:8

demonstrated that culturally typical patterns of Another misunderstanding of CN arises brain activity are sometimes quite malleable. from the confusion between culture and race. While some CN studies have compared participants from cultural groups that are WHAT IS CULTURAL also purported to be racial groups, other NEUROSCIENCE NOT? CN studies have examined cultural effects The rapid development of CN has induced sev- such as differences in brain activity between eral misunderstandings of CN research. These religious groups and between social classes of misunderstandings relate to the origin of CN the same race. CN assumes that any difference research, the biological versus social nature of between these groups is primarily the result of the human brain, and the relationship between socialization and chronic cultural experiences. culture and race (e.g., Mateo et al. 2012). None of the aforementioned CN findings can One misunderstanding of CN research is to be simply explained by group differences in connect CN studies with anthropological ap- physical attributes (e.g., skin tone). CN studies proaches to explain the nature of culture, which have shown evidence that differences in brain produces the misimpression that the goal of activity between Westerners and East Asians CN studies is simply to find biological markers can be mediated by specific cultural values in the brain that differentiate cultural groups (e.g., independent versus interdependent and to demonstrate that any cultural differences self-construals). Cultural priming research in brain activity are determined biologically has demonstrated causal effects of culture on and are immutable. Such misunderstanding brain activity. These findings indicate the also arises from an ontologically dualistic importance of neuroplasticity in the study of opposition of the biological and cultural nature culture and demonstrate that culture is not of the human brain. Such a false dichotomy viewed as an analog for race in CN research. leads to a view of biology and culture as two The confusion between culture and race opposite accounts of the nature of the human may lead to allegations of racism against CN brain. CN actually has a nonreductionist research (e.g., CN findings “exert a tremendous view of the relationships between formative impact on the reproduction of stereotypes and biological and cultural properties of the human racism”; Mateo et al. 2012, p. 158). Racist brain. As previously noted, CN studies aim to accounts of human difference hold that human elucidate neuroplastic and culturally generated traits are biologically determined, fixed, and processes. This is fundamentally at odds with that members of racial groups are homogenous

by University of Ottawa on 10/29/13. For personal use only. cultural essentialism and hard-wired biological in these traits. In stark contrast to these beliefs, determinism. CN researchers generate specific CN researchers view a cultural group as a hypotheses about neurocognitive processes dynamic collection of individuals who share a

Annu. Rev. Psychol. 2013.64:335-359. Downloaded from www.annualreviews.org grounded in both behavioral findings from similar sociocultural context and whose mem- cultural psychology and brain imaging findings bers are affected by that context in divergent from cognitive neuroscience. These hypothe- ways (a view shared by cultural psychologists; ses limit the brain regions under investigation Heine 2012). CN research regards human and predict specific patterns of cultural group neurocognitive processes as being flexible and differences and individual differences in brain being continuously shaped by sociocultural en- activity. Thus CN research does not study vironments. CN findings demonstrate that an culture as a set of biologically determined individual’s brain is not doomed by biology to predispositions/constraints that can be used work in a specific way, but rather that the brain to rigidly categorize collections of people. is strongly shaped by long-term and short-term Instead, the CN approach emphasizes the flex- cultural experiences. It is sociocultural context ibility of the human brain that enables humans rather than race that matters. Knowing about to adapt to sociocultural environments. cultural differences in neurocognitive processes

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may discourage people from believing biologi- latory context dependence (Han & Northoff cally essentialist accounts of race and thus may 2008, Northoff 2012). Alternatively, the con- facilitate cross-cultural communication. In this stitution of any neural activity is dependent sense, CN studies should help to reduce rather upon the context; this amounts to what can be than facilitate the reproduction of stereotypes described as constitutive context dependence and racism. (Han & Northoff 2008, Northoff 2012). The distinction between these two conceptions of context dependence has far reaching IMPLICATIONS OF CULTURAL implications for the relationship between NEUROSCIENCE FINDINGS biological and social domains. In the case of CN findings have a host of both theoretical modulatory context dependence, neuronal and and practical implications. Here we list a few social activities interact with each other while of them. First, CN studies reveal the culturally remaining independent from each other in sensitive nature of the human brain and help us their respective constitution. The brain is then to understand how the human brain as a biolog- purely neuronal and thus biological, whereas ical organ is shaped by man-made sociocultural culture is social. This differs from the model of contexts. Human beings are different from constitutive context dependence, which posits other animals in that humans create the most that, if the constitution of the brain’s neuronal complicated and varied social environments. activity depends on the respective social We are also unique in our capacity for culture. context, a clear-cut distinction between the Although there are cultural universals, the spe- biological domain of the brain and the social cific contents of culture are greatly important. domain of culture is impossible. Rather than In comparison with other species, this is an being exclusively and completely biological, important advantage for the development of the brain and its neuronal activity must then be culture in a generalized sense as social commu- considered to be a hybrid of both biological and nities of conspecifics. Every person is fostered in social influences. In other words, our brains are a unique artificial environment, speaks his/her biosocial. The brain is then a relational organ mother language, behaves in accordance with that bridges the gap between the biological specific social rules, acts as a member of social world of the organism and the social world institutions, and interacts with people who of the environment and its culture (Northoff share specific cultural values with each other. 2012). Thus the human brain develops in a specific Third, CN findings help us to understand

by University of Ottawa on 10/29/13. For personal use only. sociocultural context during interactions with cultural differences in human behaviors. Tradi- others. Because there are large variations across tionally, there was usually a dominant culture in cultures, how to fit into one’s specific society a given society, and contact with out-groups was

Annu. Rev. Psychol. 2013.64:335-359. Downloaded from www.annualreviews.org and how to cooperate with others efficiently limited. Cognition and behavior adhere to the is a challenge for each person. CN studies cultural environment, and this may result in cul- indicate that the human brain has the capacity tural imprinting effects on the brain. In current to develop culture-specific neurocognitive societies, however, it is much easier for people processes that help an individual to function in to meet members of many different cultures. a specific sociocultural environment. One issue raised by CN studies is what kind Second, the context-dependent nature of of experiences during development may facili- the human brain can be understood in two tate the ability of individuals’ brains to fit into different senses. One possibility is that the their specific culture and to interact with indi- culturally different stimuli merely modulate viduals from other cultures. This is particularly already preexisting neural activity that, as such, important for those who emigrate to another remains independent of any contextual effects. culture. Does the brain adapt to a new culture, This amounts to what has been called modu- and if so, how quickly? CN studies, particularly

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those using cultural priming, suggest that even tural perspective, and it offers at the same time an adult’s brain is quite flexibly attuned to socio- a new way to enrich concepts of culture from a cultural environments. Thus cultural education neuroscience perspective. CN research has al- and experience may shape the brain in order to ready provided many important insights into fit into new cultural contexts. the mutual constitution of culture and the brain, Fourth, because CN studies help us to un- and it has raised intriguing questions for fu- derstand cultural differences in behavior, prac- ture researchers to address, some of which are tices, and psychological tendencies from a neu- discussed below. roscience perspective, CN findings may help to develop constructive ways to deal with mis- Brain and cultural value. Although there has understandings and conflicts between different been increasing evidence for cultural differ- cultural groups. In fact, CN studies not only ences in brain activity underlying multiple cog- show differences in brain activity mediating be- nitive and affective processes, future research havioral discrepancy in distinct sociocultural should further explore the association between contexts, but also highlight commonalities of cultural values and cultural group differences the human species across cultures. Understand- in brain activity. This may help to ascribe ing both cultural distinctiveness and cultural observed culture-specific neurocognitive pro- universality in the neural mechanisms under- cesses to the effect of specific cultural values lying human cognition and behavior may help or practices, and it may provide a more com- to reduce intergroup conflict and prejudice and plete account of variation in neurocognitive may provide insight in how best to facilitate in- processes as a function of cultural values. CN tergroup cooperation. studies have shown culturally specific patterns Finally, CN findings lead us to rethink the of brain activity. But it remains unclear why nature of culture. If the brain shows constitutive some neural processes are sensitive to cultural context dependence and can no longer be re- influences whereas others are not. A more con- garded as purely biological but rather as bioso- ceptual issue concerns how much the structure cial, it means conversely that the environment and organization of human culture in general, and thus culture also are not as purely social as and the extent to which specific cultures in par- is often assumed. Instead, culture is ingrained ticular, mirror the structure and organization by the neuronal structures and organization of of the brain. the brain’s neural activity via the latter’s consti- tutive context dependence. Hence rather than

by University of Ottawa on 10/29/13. For personal use only. being completely and exclusively social, culture Culture, genes, and biochemistry. CN must then be considered to be sociobiological. research has so far demonstrated cultural influ- The empirical results from CN clearly call ences on the brain in terms of activity changes

Annu. Rev. Psychol. 2013.64:335-359. Downloaded from www.annualreviews.org into question any simplified characterization of in brain regions or networks. However, CN the brain as merely biological and suggest that has yet to explore how culture may affect or a more sophisticated account of the brain as interact with biochemical substances of the biosocial better fits the evidence. This in turn brain. Cultural effects should be traced to the entails the need to redefine our concept of cul- neuronal and biochemical levels in order to ture and to reject any simplifications of culture understand the relationship between culture as an exclusively social construction. and micro-level neural processes. For instance, an important question for future researchers will be to determine whether and how certain FUTURE QUESTIONS FOR biochemical substances or neurotransmitters CULTURAL NEUROSCIENCE are sensitive to cultural experiences. Exploring CN research provides a new approach to the this question may help us to understand the understanding of the human brain from a cul- extent to which cultural influences on the brain

www.annualreviews.org • Cultural Neuroscience 353 PS64CH13-Han ARI 8 November 2012 10:8

operate not only on the functional neural level Further, is the association between genotype but also on the biochemical level. and mental illness similar across cultures, or is CN research also needs to be integrated it moderated by cultural context? Answering with genetic imaging. Although imaging stud- these questions may help to determine whether ies have shown both cultural and genetic the same treatments are appropriate for mental effects on brain function, there has been little disorders in different cultures. research that examines whether and how cul- ture and genes interact to affect neural pro- Brain and the creation of culture. Finally, cesses. One possible approach to this issue is to one of the main goals of CN research is to examine genotype and culture simultaneously address how the brain is involved in the cre- using brain imaging. Studies along this line will ation and maintenance of the cultures that exist offer a comprehensive description of how cul- today. This may seem like a research question ture and genes interact to shape the human that is beyond the grasp of CN. However, CN brain and may further challenge purely biolog- studies may create new paradigms to distribute ical accounts of the brain. values and practices in a small group of par- ticipants and examine whether and how such Brain and acculturation. The increasing manipulations may result in corresponding number of immigrants in the current society changes in brain activity. Such an approach may raises another important issue for CN research: provide the ability to directly test neuroscience Is there a sensitive period during develop- accounts of cultural differences. Further, such ment for acculturation of the brain? Recent be- an approach may provide insights into how havioral research suggests that, for individuals culturally influenced ways of thinking and feel- who immigrated before approximately age 14.5, ing are acquired and represented in the human identification with a new culture increased with brain. time living in the new culture (Cheung et al. 2011). However, for older immigrants, identifi- CONCLUSION cation with a new culture seemed not to change with the time in the new culture. Such findings This review presents a refined account of CN imply that there may be a sensitive period for in terms of origin, concept, method, findings, acculturation of the human brain. Among indi- and theory, and clarifies several misunderstand- viduals of different ages, this can be examined ings of the field. In sum, CN investigates the by investigating the variation of brain activity biosocial nature of the human brain by examin-

by University of Ottawa on 10/29/13. For personal use only. as a function of the time spent in a new culture. ing whether and how implicit and explicit pat- terns of beliefs, values, meanings, and practices Culture and abnormal brain function. CN in specific sociocultural contexts shape the neu- Annu. Rev. Psychol. 2013.64:335-359. Downloaded from www.annualreviews.org studies also raise issues concerning cross- ral mechanisms underlying human cognition, cultural differences in the prevalence (and emotion, and behavior. CN research has helped neural correlates) of psychiatric disorders such to provide a nuanced understanding of culture as schizophrenia and depression. Although by integrating methods from social and cultural these mental disorders appear to occur with psychology and neuroscience. The accumulat- similar lifetime prevalence in different cultures, ing findings in the field show strong evidence it remains unclear whether their symptomatic for cultural influences on the human brain and expression is influenced by cultural predispo- raise exciting new questions about the biosocial sitions (e.g., collectivism versus individualism) nature of human beings. We believe that the and whether their symptomatology is mediated continued growth and development of CN will by similar patterns of abnormal neural activity. promote cross-cultural understanding and pro- Do the same neurocognitive mechanisms medi- vide strong evidence against racist accounts of ate these psychiatric disorders across cultures? human difference.

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DISCLOSURE STATEMENT The authors are not aware of any affiliations, memberships, funding, or financial holdings that might 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 Program 2010CB833903) 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 Volkswagen Foundation.

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Annual Review of Psychology Volume 64, 2013 Contents

Prefatory Shifting Gears: Seeking New Approaches for Mind/Brain Mechanisms Michael S. Gazzaniga pppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppp1

Biological Bases of Behavior The Endocannabinoid System and the Brain Raphael Mechoulam and Linda A. Parker ppppppppppppppppppppppppppppppppppppppppppppppppppp21

Vision Synesthesia Jamie Ward ppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppp49

Scene Perception, Event Perception, Object Recognition Visual Aesthetics and Human Preference Stephen E. Palmer, Karen B. Schloss, and Jonathan Sammartino ppppppppppppppppppppppppp77

Attention and Performance Detecting Consciousness: A Unique Role for Neuroimaging Adrian M. Owen pppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppp109 by University of Ottawa on 10/29/13. For personal use only. Executive Functions Adele Diamond pppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppp135

Annu. Rev. Psychol. 2013.64:335-359. Downloaded from www.annualreviews.org Animal Learning and Behavior The Neuroscience of Learning: Beyond the Hebbian Synapse C.R. Gallistel and Louis D. Matzel pppppppppppppppppppppppppppppppppppppppppppppppppppppppp169

Evolutionary Psychology Evolutionary Psychology: New Perspectives on Cognition and Motivation Leda Cosmides and John Tooby ppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppp201 Origins of Human Cooperation and Morality Michael Tomasello and Amrisha Vaish pppppppppppppppppppppppppppppppppppppppppppppppppppp231

vi PS64-FrontMatter ARI 15 November 2012 14:20

Language and Communication Gesture’s Role in Speaking, Learning, and Creating Language Susan Goldin-Meadow and Martha Wagner Alibali ppppppppppppppppppppppppppppppppppppp257

Nonverbal and Verbal Communication The Antecedents and Consequences of Human Behavioral Mimicry Tanya L. Chartrand and Jessica L. Lakin ppppppppppppppppppppppppppppppppppppppppppppppppp285

Intergroup Relations, Stigma, Stereotyping, Prejudice, Discrimination Sexual Prejudice Gregory M. Herek and Kevin A. McLemore pppppppppppppppppppppppppppppppppppppppppppppp309

Social Neuroscience A Cultural Neuroscience Approach to the Biosocial Nature of the Human Brain Shihui Han, Georg Northoff, Kai Vogeley, Bruce E. Wexler, Shinobu Kitayama, and Michael E.W. Varnum pppppppppppppppppppppppppppppppppppppppp335

Organizational Climate/Culture Organizational Climate and Culture Benjamin Schneider, Mark G. Ehrhart, and William H. Macey pppppppppppppppppppppppp361

Industrial Psychology/Human Resource Management Employee Recruitment James A. Breaugh ppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppp389

Learning and Performance in Educational Settings Self-Regulated Learning: Beliefs, Techniques, and Illusions

by University of Ottawa on 10/29/13. For personal use only. Robert A. Bjork, John Dunlosky, and Nate Kornell ppppppppppppppppppppppppppppppppppppppp417

Teaching of Subject Matter Annu. Rev. Psychol. 2013.64:335-359. Downloaded from www.annualreviews.org Student Learning: What Has Instruction Got to Do With It? Hee Seung Lee and John R. Anderson pppppppppppppppppppppppppppppppppppppppppppppppppppppp445

Health Psychology Bringing the Laboratory and Clinic to the Community: Mobile Technologies for Health Promotion and Disease Prevention Robert M. Kaplan and Arthur A. Stone ppppppppppppppppppppppppppppppppppppppppppppppppppp471

Research Methodology Multivariate Statistical Analyses for Neuroimaging Data Anthony R. McIntosh and Bratislav Miˇsi´c ppppppppppppppppppppppppppppppppppppppppppppppppp499

Contents vii PS64-FrontMatter ARI 15 November 2012 14:20

Social Network Analysis: Foundations and Frontiers on Advantage Ronald S. Burt, Martin Kilduff, and Stefano Tasselli ppppppppppppppppppppppppppppppppppppp527

Indexes

Cumulative Index of Contributing Authors, Volumes 54–64 ppppppppppppppppppppppppppp549 Cumulative Index of Chapter Titles, Volumes 54–64 ppppppppppppppppppppppppppppppppppp554

Errata

An online log of corrections to Annual Review of Psychology articles may be found at http://psych.AnnualReviews.org/errata.shtml by University of Ottawa on 10/29/13. For personal use only. Annu. Rev. Psychol. 2013.64:335-359. Downloaded from www.annualreviews.org

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