gene × environment interaction in social cognition 26.pdf · gene × environment interaction in...

26Gene × Environment Interaction in Social

Cognition

J o a n Y . C h i a o , B o b b y K . C h e o n , G e n n a M . B e b k o , R o b e r t W . L i v i n g s t o n ,

& Y i n g - Y i H o n g

INTRODUCTION

In an ideal world the scientist should find a method to prevent the most severe forms of autism but allow the milder forms to survive. After all, the really social people did not invent the first stone spear. It was probably invented by an Aspie who chipped away at rocks while the other people socialized around the campfire. Without autism traits we might still be living in caves. (Temple Grandin, 1996)

How can we explain the nature and origin of social cognition? One of the most remarkable hallmarks of human and non-human primate living is the ability to successfully coexist over centuries in incredibly complex social groups of varying size, from small-scale hunter-gatherer tribes, ranging from a few to a few hundred people, to large-scale settled horticultural tribes, ranging from a few hundred to a few thousand people. According to the social brain hypothesis, this versatility in social living arrangements is possible due to humans having evolved an unusu-ally large brain with increased cognitive capacities (Brothers, 2001; Dunbar, 1998). Supporting this view, a number of quantitative studies have shown that, among primates, the relative volume of the neocortex is positively correlated with a range of

markers of social group complexity, including the average size of a social group, number of females in the group, grooming group size, frequency of coalitions, prevalence of social play, prevalence of deception, and frequency of social learning (Dunbar & Shultz, 2007). These findings illustrate the evolutionary role of the brain mechanisms involved in facilitating social cognition as well as genes that facilitate their transmission across generations (see Chapter 23).

Whereas the capacity for social cognition is largely ubiquitous across individuals and social groups, the extent to which people are adept at understanding other people varies across indi-viduals and cultures. These differences in under-standing can be seen in individuals with autism, who have difficulty reading the minds of others, to individuals with Williams syndrome, who dis-play hypersociality (see Chapter 21). Understand-ing how and why such a wide range of social-cognitive ability exists in society through-out human history requires an understanding of how genetic and environmental factors, such as cultural values, practices, and beliefs, give rise to social cognition and its underlying neural sub-strates, as well as the evolutionary basis for such social-cognitive abilities.

Here we adopt the cultural neuroscience framework for understanding how genetic and

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GENE × ENVIRONMENT INTERACTION IN SOCIAL COGNITION 517

environmental influences shape social cognition across cultural contexts (Chiao & Ambady, 2007; Chiao, 2009; Chiao, 2011; see Figure 26.1). The idea that complex behavior results from the dynamic interaction of genes and cultural environ-ment is not new (Caspi & Moffitt, 2007; Johnson, 1997; Li, 2003); however, cultural neuroscience represents a novel empirical approach to demon-strating bidirectional interactions between culture and biology by integrating theory and methods from cultural psychology (Kitayama & Cohen, 2007), neuroscience (Gazzaniga, Ivry, & Mangun, 2002), and neurogenetics (Canli & Lesch, 2006; Green et al., 2008, Hariri, Drabant, & Weinberger, 2006). Similar to other interdisciplinary fields, cultural neuroscience aims to explain a given mental phenomenon in terms of a synergistic product of mental, neural, and genetic events. Cultural neuroscience shares overlapping research goals with social neuroscience, in particular, as understanding how neurobiological mechanisms facilitate cultural transmission involves investigat-ing primary social processes that enable humans to learn from one another, such as imitative learn-ing. However, cultural neuroscience is also unique from related disciplines in that it focuses explic-itly on ways that mental and neural events vary as a function of culture and genetic traits in some meaningful way (Figure 26.2). Additionally, cul-tural neuroscience illustrates how cultural and genetic traits may alter neurobiological and psy-chological processes beyond those that facilitate social experience and behavior, such as perception and cognition.

Over the past decade, social-cognitive neuro-scientists have been uncovering with rapid preci-sion how our unusually large human brain has enabled complex social behavior by mapping net-works of brain structures to complex social func-tions (Lieberman, 2010; Ochsner & Lieberman, 2001). Convergent social neuroscience evidence to date indicates that a core network of brain regions underlie the capacity for social cognition or thinking about other people, including the extrastriate cortex, superior temporal gyrus, medial prefrontal cortex, temporoparietal junc-tion, amygdala and lateral prefrontal regions, and anterior insula and secondary somatosensory cortex (Figure 26.3). Regions of extrastriate cortex, such as the fusiform gyrus, and superior temporal gyrus, are involved in the ability to rec-ognize personal identity, from perceptual cues such as the face (Kanwisher, McDermott, & Chun, 1997) and the voice (Belin et al., 2000). Medial prefrontal cortex (MPFC) and temporopa-rietal junction (TPJ) play important roles in the ability to infer other people’s thoughts, feelings, and intentions through either simulated (Mitchell, 2009) or conceptual (Saxe, 2006) processing of

social cues in the environment. Limbic circuitry, such as the amygdala, facilitate an emotional (Pessoa & Adolphs, 2010) and motivational (Cunningham & Zelazo, 2007) response to threat-ening or ambiguous signals in the environment, whereas prefrontal cortex, including orbital and lateral regions, are associated with the regulation of this emotional response in the context with social and cultural norms (Olsson & Ochsner, 2008; Quirk & Beer, 2006). Finally, anterior

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Figure 26.1 Illustration of the cultural neuroscience framework (adapted from Chiao, 2011).

Figure 26.2 Illustration of different models of gene-to-behavior pathways. (a) The direct linear approach to gene and behavior. (b) The endophenotype approach, which assumes that the gene-to-behavior pathway is mediated by proximate or intermediate phenotypes such as neural activity. (c) The gene-by-environment approach, which assumes interaction between genetic and environmental factors influences neural activity and behavior. (d) The culture-by-gene approach, which identifies specific cultural traits and genes that interact and mutually influence neural activity and subsequent behavior (Chiao, 2011).

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THE SAGE HANDBOOK OF SOCIAL COGNITION518

insula and secondary somatosensory cortex are brain regions associated with responding to the perception and sharing of the experience of social (Eisenberger, 2010), emotional (Mathur et al., 2010), and physical pain (Apkarian, Baliki, & Geha, 2009; Lamm, Decety, & Singer, 2011).

Some of these brain regions underlie related social-cognitive abilities in non-human primates as well. For instance, dominant male rhesus macaques show increased right amygdala and right superior temporal sulcus response during mate competition, suggesting shared neural circuitry associated with social vigilance across species (Rilling, Winslow, & Kilts, 2004). Additionally, some components of the social-cognitive brain are engaged during childhood, and to a greater extent, compared to adults. For instance, children showed heightened amygdala activation in response to emotional faces relative to adults (Hoehl & Striano, 2010). Even in early infancy, the superior temporal sulcus can distinguish between vocal and non-vocal sounds, setting the stage for neural discrimination of auditory cues facilitating con-specific communication (Belin & Grosbras, 2010). The existence of a core network of brain regions across species and engaged early in development suggests that the human ability to navigate the social world, to distinguish between one’s self and others, between a person’s intentions from their thoughts and desires, to understand one as

part of a group, and to infer consciousness of others from both perceptual and conceptual infor-mation is reliant, at least in part, on innate bio-logical machinery that facilitates transmission of social-cognitive processes across phylogeny and ontogeny.

GENETIC BASIS OF SOCIAL COGNITION

The notion of social cognition as arising from innate machinery is supported by evidence from twin studies showing that several foundational social skills, such as empathy, motivation, and decision making, are between 15 and 50% herita-ble (Ebstein et al., 2010), or shared to a greater extent between monozygotic (MZ) compared to dizygotic (DZ) twins. MZ twins share nearly 100% of their genome, whereas DZ twins only share 50%. Assuming that both MZ and DZ twins are raised in a similar shared environment, differ-ences in correlation of behavior between MZ and DZ twins can be attributed to a genetic influence (see also Constantino & Todd, 2000).

Furthermore, behavioral and neurogenetic stud-ies of social behavior in human and non-human primates have made significant progress in associ-ating specific functional polymorphisms (SNPs) or genes with specific social-cognitive processes

Figure 26.3 Illustration of network of brain regions involved in understanding the mental states of self and others (adapted from Hein & Singer, 2008). MFC, medial prefrontal cortex; ACC, anterior cingulate cortex; AI, anterior insula; SII, secondary somatosensory cortex; TP, temporal poles; STS, superior temporal sulcus; TPJ, temporoparietal junction.



underlying interpersonal processes. For instance, the serotonin transporter gene (SLC6A4) is typi-cally associated with emotional (Hariri et al., 2002) and social-cognitive processes (Canli & Lesch, 2007). People who carry the short (S) allele of the serotonin transporter gene are more likely to show greater emotional reactivity and amygdala response to emotional information compared to those who carry the long (L) allele (Munafo, Brown, & Hariri, 2008). Empathy, or the ability to feel what others are feeling, is asso-ciated with the oxytocin receptor gene (OXTR). Recent evidence shows that the rs2254298A allele of OXTR is significantly associated with larger bilateral amygdala, but not hippocampal, volume (Inoue et al., 2010). Monoamine oxidase A (MAOA) genotype has been associated with antisocial behavior across a number of studies (Taylor & Kim-Cohen, 2007). In addition, recent behavioral genetics further suggests evidence of a genetic basis for intergroup processes as well. For example, in a large-sample twin study, Lewis and Bates (2010) revealed that MZ twins displayed significantly more similar levels of intergroup bias relative to DZ twins, and both shared genes and environment contributed to this process. Hence, social cognition underlying both interpersonal and intergroup processes arises, at least in part, due to genes regulating brain regions underlying emo-tional and social processes.

CULTURAL BASIS OF SOCIAL COGNITION

Whereas emerging behavioral and neurogenetics work shows that social cognition is due at least in part to genetic factors, a plethora of cultural psy-chology (Chapter 22) and cultural neuroscience evidence (Chiao & Bebko, 2010; Han & Northoff, 2009) indicates that environmental factors, such as cultural values, practices, and beliefs, have a tremendous influence on a range of core skills underlying the human capacity to understand and think about other people.

Self and other knowledge

One of the most robust ways that values, such as individualism and collectivism, influence human behavior is in self-construal, or how people think about themselves in relation to others. Individual-ists think of themselves as autonomous from others, while collectivists think of themselves as highly interconnected with others (Markus & Kitayama, 1991; Triandis, 1995). Recent cultural neuroscience evidence indicates that the brain basis of the self is modulated by cultural values

of individualism and collectivism (Chiao et al., 2009; Han & Northoff, 2008). For instance, research has shown Caucasians, but not Chinese, showed greater neural activity within the MPFC during evaluation of personality traits of one’s self relative to a close other (i.e., mother), suggest-ing cultural variation in MPFC response during self-evaluation (Zhu et al., 2007; see Figure 26.4). More recent evidence has demonstrated that cultural values (i.e., individualism−collectivism), rather than cultural affiliation (i.e., East Asian−Westerners) per se, modulate neural response during self-evaluation. In another cross-cultural neuroimaging study, people in both Japan and the United States who endorsed individualistic values showed greater MPFC activity for general relative to contextual self-descriptions, whereas people who endorsed collectivistic values demonstrated greater MPFC activation for contextual relative to general self-descriptions (Chiao et al., 2009; see Figure 26.5). Supporting this view, another study using cultural priming (Hong et al., 2000) showed that even temporarily heightening awareness of individualistic and collectivistic values in bicul-tural individuals (i.e., bicultural Asian-Americans) modulates MPFC and posterior cingulate cortex (PCC) in a similar manner (Chiao et al., 2010; see Figure 26.6). In addition to cultural values modu-lating neural responses during explicit self-processing, a further neuroimaging study shows that dorsal, but not ventral, regions of MPFC are modulated by cultural priming of individualism and collectivism when thinking about one’s self in an implicit manner (Harada, Li, & Chiao, 2010; see Figure 26.7). Such findings suggest that cultural values dynamically shape neural rep-resentations during the evaluation, rather than the detection, of self-relevant information. Taken together, these studies provide convergent evi-dence that environmental factors, such as cultural values of individualism−collectivism, shape the psychological and neural basis of the self.

In addition to cultural values of individualism−collectivism, religious beliefs may also play an important role in modulating neural responses underlying social cognition. One set of neuro-imaging studies examining the neural substrates of religiousity found that activity within Theory of Mind (ToM) regions, including left precuneus, left temporoparietal junction, and left middle frontal gyrus, was correlated with the degree of one’s religiousity (Kapogiannis et al., 2009). Additionally, religious practices, such as pray-ing, also modulate neural responses within ToM regions. For instance, compared to formalized prayer and secular cognition, improvised praying activated the temporopolar region, medial pre-frontal cortex, temporoparietal junction, and pre-cuneus (Schjoedt et al., 2009). Finally, religious



beliefs affect neural representations of the self. Whereas atheists typically recruit ventral MPFC during self-evaluation, religious individuals show greater response within dorsal MPFC, suggesting that religious beliefs promote greater evalua-tion, rather than representation, of one’s self (Han et al., 2008). Hence, the human ability to possess

religious beliefs and exercise religious practices relies on ToM and mentalizing brain regions that facilitate the representation and evaluation of own and others (e.g., human, God) mental states.

Although the lion’s share of cultural neuro-science research on knowledge of self and others has been conducted with human neuroimaging

Figure 26.5 In both Japan and the United States, the degree of collectivistic or individualis-tic cultural values predicts neural response within medial prefrontal cortex (MPFC) to contextual or general self-judgments, respectively (adapted from Chiao et al., 2009).

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methodology, a couple of recent studies have examined the effect of culture on electrophysio-logical indices of social cognition. In one study, Lewis and colleagues (2008) measured event-related potentials (ERPs) while participants

completed the oddball task, where they are shown visual stimuli in either a frequent or infrequent (i.e., oddball stimulus) manner. Results demon-strated that European-American participants showed greater novelty P3, or late positive

Figure 26.6 Dynamic cultural influences on neural response to self-judgments. (a−c) Bicultural individuals primed with collectivistic cultural values show greater medial prefrontal cortex (MPFC) and posterior cingulate cortex (PCC) response to explicit contextual self-judgments, whereas bicultural individuals primed with individualistic cultural values show greater MPFC and PCC response to explicit general self-judgments (adapted from Chiao et al., 2010). (d−e) Degree of cultural priming predicts neural response to culturally congruent self-judgments within both MPFC and PCC.

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potential, amplitude for target events, whereas East Asians showed greater P3 amplitude for odd-ball events. Another study by Iishi and colleagues (2009) found that amplitude of the N400, a late negative potential, was significantly larger when individuals perceived incongruent relative to congruent information and the degree of late negativity activity was reliably predicted by chronic social orientation (e.g., interdependence)

for females. Both electrophysiological studies demonstrate the effect of cultural values of individualism−collectivism on how people respond to information that is either congruent or incongruent to one another. Hence, cultural values of individualism−collectivism not only affect how people represent knowledge about self and others but also respond to congruent or incongruent informational cues in the environment.

Figure 26.7 Dynamic neural response within ventral and dorsal regions of medial prefrontal cortex (VMPFC and DMPFC) during implicit self-judgments (Harada, Li, & Chiao, 2010). (a) Neural response during self-relevant and father-relevant trials compared to control trials for both individualistic and collectivistic priming conditions. (b) Mean parameter estimates within VMPFC and DMPFC during implicit self-judgments.

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Interpersonal perception

Minute perceptual cues from the body, such as the eye region of the face, can convey a wealth of information about what people are thinking and feeling. Recent neuroimaging evidence indicates cultural variation in neural responses when infer-ring the internal states of others, particularly from the eye region (Adams et al., 2010). Native Japanese and US Caucasian participants per-formed the “Reading the Mind in the Eyes” Test (RME), a measure of mental state decoding from visual stimuli only depicting an individual’s eyes (Baron-Cohen, Wheelwright, Hill, Raste, & Plumb, 2001). In the study participants were more accurate at decoding the mental state of members of one’s own culture relative to members of another culture, and activity within the poste-rior superior temporal sulcus (pSTS) increased during the same-culture mental state decoding relative to other-culture mental state decoding (see Figure 26.8). Additionally, the intracultural advan-tage was significantly negatively correlated with pSTS activity during other-culture mental state decoding such that as pSTS activity increased, the intracultural advantage decreased. This correla-tion was not significant for same-culture mental state decoding from the eyes, suggesting that the intracultural advantage may be due to less pSTS recruitment during other-culture mental state decoding. These findings support the universal recruitment of pSTS in ToM, while at the same time revealing culturally modulated pSTS recruit-ment underlying the intracultural advantage in ToM. Another recent study found that activity within the mesolimbic system responds more for culturally congruent dominant and submissive facial cues (Freeman et al., 2009). Individuals from egalitarian cultures, such as the United States, show greater mesolimbic response to dom-inant facial cues, whereas individuals from hierar-chical cultures, such as Japan, show greater mesolimbic response to submissive facial cues. Taken together, these studies highlight how cul-tural variation in attribution styles may modulate neural activity underlying processes related to interpersonal perception.

Emotion recognitionCulture affects how people prefer to experience, express, recognize, and regulate their emotions (Mesquita & Leu, 2007). East Asians prefer to experience low-arousal relative to high-arousal positive emotions (Tsai, 2007) and are more likely to suppress their emotions relative to Westerners (Butler, Lee, & Gross, 2007). Additionally, both East Asians and Westerners demonstrate cultural specificity in emotion recognition, whereby they show greater recognition for emotions expressed

by their own cultural group members relative to members of other cultural groups (Elfenbein & Ambady, 2002). Recent cultural neuroscience of emotion research has shown cultural specificity effects within a number of brain regions involved in emotion recognition. Moriguchi and colleagues (2005) found greater activation in the posterior cingulate, supplementary motor cortex, and amy-gdala in Caucasians, relative to Japanese, who showed greater activity within the right inferior frontal, premotor cortex, and left insula when participants were asked to explicit recognize emo-tions from the face. Chiao and colleagues (2008) examined neural responses in adults living in either the United States or Japan and found that across cultures people exhibit greater bilateral amygdala response to fear faces expressed by own-culture relative to other-culture members (Chiao et al., 2008, Figure 26.9). Another recent neuroimaging study comparing neural responses during emotion recognition in Asians and Europeans found a significant negative correlation between duration of stay and amygdala response, such that amygdala response during emotion rec-ognition was higher in individuals who were recent immigrants to the region, suggesting that experience alters neural responses to emotional expressions (Derntl et al., 2009). Taken together, this research indicates that activity within the human amygdala is modulated by cultural group membership. An important question for future research will be to determine whether neural mechanisms that support other facets of emotion, such as experience and regulation, are affected by culture.

EmpathyEmpathy is the capacity to share the emotional states of others (Batson, Duncan, Ackerman, Buckley, & Birch, 1981; Preston & de Waal, 2002). The perception−action model of empathy indicates that empathy is a key motivator (Decety & Grèzes, 2006) and the proximate mechanism (de Waal, 2008) of altruistic behavior, whereby an individual perceives and shares in the distress of another person, and acts to reduce his or her suffering (Preston & de Waal, 2002). Prior social neuroscience research indicates that empathy is a multi-component process that includes affect shar-ing, cognitive perspective taking, and cognitive appraisal (Decety & Jackson, 2004; Hein & Singer, 2008; Lamm, Batson, & Decety, 2007; Olsson & Ochsner, 2008). Empathy for pain is supported by neuroanatomical circuits underlying both affective and cognitive processes (Decety & Jackson, 2004; Hein & Singer, 2008; Lamm et al., 2007; Olsson & Ochsner, 2008). A distinct neural matrix, including bilateral anterior insula (AI) and anterior cingulate cortex (ACC) (Decety &



Jackson, 2004; Hein & Singer, 2008; Olsson & Ochsner, 2008) is thought to underlie the affective components of empathy. AI and ACC code the autonomic and affective dimension of pain and, in particular, the subjective experience of empathy when perceiving pain or distress in others (Decety & Jackson, 2004; Hein & Singer, 2008; Olsson & Ochsner, 2008).

Recent evidence indicates that empathic neural response is modulated by culture. For instance, a recent neuroimaging study by Xu and colleagues (2009) examined whether or not cultural group membership modulates neural response during the perception of pain in others. Chinese and Caucasian participants were scanned while observ-ing Chinese and Caucasian targets either in physi-cally painful (e.g., needlestick) or neutral (e.g., Q-tip probe) scenes. All participants showed greater ACC and AI response to painful relative to neutral scenes; however, they also showed greater ACC response to in-group relative to out-group members (see Figure 26.10). Furthermore, recent transcranial magnetic stimulation (TMS) findings

by Avenanti and colleagues (2010) show that both Black and White participants showed greater muscle-specific corticospinal inhibition when watching a needle penetrate the hand, but only when the hand was a person of the same race, indicating an in-group bias in the activation of pain representations within the perceiver’s senso-rimotor system. Importantly, however, both groups of participants showed increased neural response to violet hands, highlights a pivotal role for cul-ture in changing how and when humans share and respond to the suffering of same and other races (see Figure 26.11). Taken together, these findings demonstrate that cultural group membership affects neural responses to perceived physical pain of others and suggest a neural precursor to group selection in altruistic behavior (Wilson, 2006).

Theory of MindAnother key social cognitive process is theory of mind (ToM), or the ability to understand and rep-resent the psychological state of others (Wellman,

Figure 26.8 Graphs depict regions of left and right posterior superior temporal sulcus (pSTS) activation for same- versus other-culture mental state decoding (Adams et al., 2010). Neural response within STS is heightened for same- versus other-culture mental state decoding across both cultural groups.



Cross, & Watson, 2001). Normally developing children demonstrate ToM starting at 4 years of age, while younger children and children with autism typically fail to demonstrate ToM (Baron-Cohen, Leslie, & Frith, 1985). Such developmen-tal findings provide evidence for ToM as a universal developmental process (Fodor, 1983; Leslie, Friedman, & German, 2004; Scholl & Leslie, 1999) with an underlying biological basis (Frith & Frith, 2001; Scholl & Leslie, 1999). While some cross-cultural studies support the universality of ToM, other studies suggest ToM may be culturally and linguistically dependent (for review, see Kobayashi, Glover, & Temple, 2006). For example, variation in cultural attribu-tion styles may influence ToM performance in Asian children (Naito, 2003) who are raised in a culture that attributes behavior to external and contextual causes rather than to internal causes, as in American−European cultures (Masuda & Nisbett, 2001; Nisbett, 2003). Similarly, speaking a non-English language with few mental state verbs may negatively influence children’s per-formance on ToM tasks (Vinden, 1996).

Neuroimaging provides further evidence for both the universal (Saxe, 2006; Saxe & Kanwisher, 2003) and culturally specific influences on ToM

processes (Kobayashi, Glover, & Temple, 2006). A number of prior neuroimaging studies of ToM conducted on individuals from Western populations have found greater activity within the right temporoparietal junction (rTPJ), specifically when participants read stories about another per-son’s thoughts (Saxe, 2006; Saxe & Kanwisher, 2003). Recently, Kobayashi and colleagues (2006) used functional magnetic resonance imaging (fMRI) to examine cultural and linguistic influ-ences on neural activity underlying ToM in American English-speaking monolinguals and Japanese-English late bilinguals. Neural activity was recorded using fMRI while participants com-pleted second-order false-belief ToM stories in both English and Japanese languages. Universally recruited brain regions associated with ToM processing included the right MPFC, right ante-rior cingulate cortex (ACC), right MFG/DLPFC (middle frontal gyrus/dorsal-lateral prefrontal cortex), and TPJ. In the American English-speaking monolinguals, culturally modulated neural activity underlying ToM was observed in the right insula, bilateral temporal poles, and right MPFC relative to the Japanese-English bilinguals, while the Japanese-English late bilinguals showed culturally modulated neural activity in the right

Figure 26.9 Cultural specificity in bilateral amygdala response to fear faces (adapted from Chiao et al., 2008). (a) Examples of Japanese and Caucasian-American fear faces. (b) Bilateral amygdala. (c, d) Participants show greater left (c) and right (d) amygdala response to fear expressed by members of one’s own cultural group.

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orbitofrontal gyrus (OFG) and right inferior frontal gyrus (IFG) associated with ToM process-ing relative to the American English-speaking monolinguals. Greater insular and TP activity in the American English-speaking monolinguals suggest that ToM in American culture emphasizes integrating sensory modalities with limbic input, whereas greater OFG and IFG activity in the Japanese-English late bilinguals suggest ToM in Japanese culture may rely more on emotional mentalizing. Taken together, these findings dem-onstrate universality and cultural diversity in neural mechanisms underlying theory of mind.

Intergroup perception

Recent evidence from social cognition and social neuroscience has demonstrated that diverse mech-anisms are involved in the perception and process-ing of group membership, and, in particular, has highlighted the role of social context in modulat-ing neural response to in-group and out-group targets. Early work on the social neuroscience of interracial perception demonstrated greater amygdala reactivity when viewing the faces of

out-group members relative to in-group members (Hart et al., 2000; Phelps et al., 2000), which reflected, in part, unconscious racial bias (Cunningham et al., 2004; Phelps et al., 2000; Richeson et al., 2003; Wheeler & Fiske, 2005).

Biases in intergroup perception and processing may be represented through other patterns of neural reactivity not directly related to evaluation or detection of threat. For instance, an individual with damage to the bilateral amygdala is still capable of exhibiting negative racial biases (Phelps, Cannistraci, & Cunningham, 2003), sug-gesting that other neural correlates may be involved. Though negative biases, evaluations, or stereotypes may be spontaneously activated, such automatic processes may be followed by control-led inhibitory processes (Devine, 1989; Shelton, 2003). Such regulatory activity may be reflected by responses in the ACC, which may detect inappropriate potential responses, and the dorsal-lateral prefrontal cortex (DLPFC), a region involved in inhibitory processes (Amodio et al., 2004; Cunningham et al., 2004). Supporting this view, Richeson and colleagues (2003) demon-strated that implicit racial bias was associated with increased reactivity within the ACC and

Figure 26.10 Increased activations in the anterior cingulate cortex (ACC) and the frontal/insula cortex when participants perceived racial in-group faces (Xu et al., 2009). LFC, lateral frontal cortex; SMA, supplementary motor area.



DLPFC when White participants viewed Black faces during neuroimaging. Moreover, the level of cognitive depletion experienced by White participants following face-to-face interracial interaction with a Black target corresponded to greater reactivity in the DLPFC when viewing Black faces during neuroimaging. These find-ings suggest that neural responses underlying intergroup bias are modulated by the degree of self-monitoring or regulation exerted by partici-pants when processing in-group and out-group members.

Finally, intergroup processes may be reflected not only in the activation and suppression of potentially negative evaluations but also the with-holding of activity in regions that may be involved in prosocial responding. One critical contribution from social neuroscience to social cognition has been the demonstration that cognitive processes recruit qualitatively different neural processes within social and non-social contexts. For exam-ple, the MPFC is typically recruited when one thinks about or infers the traits and characteristics of others, but less so when similar cognitions or

inferences are applied to understanding non-social phenomena (Amodio & Frith, 2006; Mitchell, Macrae, & Banaji, 2005). This “social” profile of cognitive activity may be attenuated when per-ceiving out-group relative to in-group members. For instance, when viewing highly stigmatized out-group members, participants show lower levels of MPFC response, suggesting stigma-tized out-group members are processed in a de-individuated fashion more similar to aversive objects rather than people (Harris & Fiske, 2006, 2007).

GENE-BY-ENVIRONMENT INTERACTION IN SOCIAL COGNITION

Understanding how and when environmental fac-tors such as culture shape social-cognitive brain function is a laudable and necessary stepping stone to the larger project of understanding how the interaction of genetic and environmental factors give rise to a range of social-cognitive

Figure 26.11 Empathic sensorimotor contagion as a function of race (Avenanti, Sirigu, & Aglioti, 2010; Chiao & Mathur, 2010). (a) Black and White participants observe a needle penetrating a specific muscle in a Black, White, or Violet hand. (b) For Black and White participants, motor-evoked potential (MEP) inhibition is greater for in-group relative to out-group hands. (c) Unconscious racial bias predicts degree of in-group bias in empathic sensorimotor contagion.

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abilities (Canli & Lesch, 2007; Caspi et al., 2010). Groundbreaking work by Suomi and colleagues, for instance, has revealed a gene-by-environment interaction in aggression levels of non-human primates (Suomi, 1994). Specifically, levels of high-risk aggression in male macaques carrying the S allele of the serotonin transporter gene (SLC6A4) were significantly higher if they were also exposed to early adversity in the form of peer rearing (Schwandt et al., 2010). Environmental influence on the expression of the serotonin trans-porter gene has similarly been found in humans. In an early seminal study, Caspi and colleagues (2003) found that humans carrying at least one S allele were significantly more likely to experience depressive episodes compared to those carrying the L allele, but only when exposed to major life stress, such as threat of loss of a significant other (see also Risch et al., 2009). This model of gene-by-environment interaction associated with the serotonin transporter gene has similarly been found at the neural level in humans. For instance, Canli and colleagues (2005) recently showed that life stress interacts with the effect of serotonin transporter genotype on both amygdala and hippocampal resting activation in humans. Taken together, these studies indicate the importance of understanding how environmental factors interact with genes in the production of socio-emotional behavior, and their underlying neural bases.

Intriguingly, recent research has begun to dem-onstrate culture−gene interaction in production and maintenance of complex behavior. In one recent study, Kim and colleagues (2010) found an interaction between the 5-HTR1A genotype and culture, such that Koreans were more likely to attend to the field compared to European-Americans, and this cultural difference in locus of attention was moderated by the serotonin 1A receptor polymorphism (5-HTR1A). Specifically, European-Americans carriers of the G allele recently showed increased attention to focal objects, whereas Korean carriers of the G allele showed increased attention to the context. In another recent study, Kim and colleagues (2010) found that distressed American carriers of the GG/AG genotype of the oxytocin receptor poly-morphism (OXTR) rs53576 reported seeking more emotional social support, compared with those with the AA genotype, whereas Korean partici-pants did not differ significantly by genotype. By contrast, OXTR groups did not differ significantly in either cultural group when not in distress. Taken together, these findings illustrate specific functional polymorphisms that are sensitive to input from the social environment − specifically cultural norms − in both basic visual perception and emotional support seeking.

FUTURE DIRECTIONS IN GENE-BY-ENVIRONMENT INTERACTION IN SOCIAL COGNITION

How and why does social cognition emerge from gene-by-environment and culture−gene interac-tion? One possible theory, alluded to at the begin-ning of this chapter, is that culture−gene coevolutionary forces have shaped the social-cognitive brain (Boyd & Richerson, 1985). By this view, environmental pressures, such as the pres-ence of infectious diseases (Fincher et al., 2006), made it important throughout human history to have the cultural capacity to infer the mental states of conspecifics and having this trait was adaptive and selected for across successive generations, giving human and non-human primates the ability to coexist with each other and coordinate tasks essential to group survival. Genes would then enable the refinement of cognitive and neural architecture underlying social cognitive capacities across successive generations. Finally, any geo-graphic variation of environmental pressures driv-ing social cognition would then lead to cultural variation in the psychological and biological mechanisms underlying how to understand each other’s thoughts, intentions, and feelings (see Chapter 22). An important puzzle for future research is to understand how culture−gene coev-olution may have shaped mechanisms in the social mind and brain differently across cultural con-texts, due to diversity of selection pressures across geographical regions.

Supporting this view, recent cross-national culture−gene association studies have identified specific cultural and genetic factors associated with social cognition. Chiao and Blizinsky (2010) found that cultural values of individualism and collectivism are associated with the serotonin transporter across nations (see Figure 26.12). Collectivistic cultures were significantly more likely to be composed of individuals carrying the S allele of the 5-HTTLPR across 29 nations. Furthermore, across nations, global pathogen prevalence was positively associated with collec-tivistic social norms, due, at least in part, to selec-tion of the S allele of the serotonin transporter gene. Way and Lieberman (2010) have similarly found that additional genes important to social cognition and social sensitivity, such as MAOA -uVNTR and opioid (OPRM1 A118G) receptor genes, vary cross-nationally and appear to confer similar adaptive benefits, such as reduced preva-lence of mood disorders.

Importantly, recent cross-cultural behavioral genetics studies have similarly revealed the impor-tance of genetic and cultural factors underlying social cognition. Recent cross-cultural behavioral



Figure 26.12 Culture−gene coevolution of individualism−collectivism and the serotonin transporter gene (5-HTTLPR) (Chiao & Blizinsky, 2010). (a) Color map of frequency distribu-tion of IND-COL from Hofstede (2001). (b) Color map of frequency distribution of S alleles of 5-HTTLPR. (c) Collectivistic nations showed higher prevalence of S allele carriers.

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genetics work further suggests that the G allele is associated with different behavioral phenotypes, depending on culture and environment (Kim et al., 2010, see Figure 26.13). The oxcytocin receptor polymorphism (OXTR) contains a polymorphism in the third intron of OXTR that has two known allelic variants, the G allele and the A allele. In Western populations, people carrying the G allele of OXTR, relative to those with the A allele, show increased maternal sensitivity (Bakermans-Kranenburg & van Ijzendoorg, 2008), empathy (Rodrigues et al., 2009), and positive emotion (Lucht et al., 2009). Distressed Americans who carry the G allele are more likely to seek emo-tional support relative to those who carry the A allele, whereas equally distressed Koreans who carry the G allele do not (Kim et al., 2010). Given the allelic frequency variation of the OXTR poly-morphism, it is possible that carrying the A rather than the G allele in East Asian cultures is advanta-geous and thus is selected for disproportionately within the geographic region and confers differen-tial functional utility across cultures, which is observable in brain and behavior.

Despite some early clues regarding the nature and origin of social cognition, many questions remain. For instance, what kinds of evolutionary selection pressures have led to diversity in cultural and genetic factors underlying social cognition? How might the human brain construct and con-strain a diversity of social cognitive styles across cultures? Future work adopting a cultural neuro-science approach to social cognition may be helpful for illuminating these questions (Chiao & Bebko, 2010; Han & Northoff, 2009).

IMPLICATIONS OF GENE-BY-ENVIRONMENT INTERACTION MODELS OF SOCIAL COGNITION

Theory

To date, theories of social cognition have focused primarily on the role of environmental factors, such as the situation and culture, and how they



contribute to the extent to which people infer the mental states of others. However, less well under-stood is how both genetic and environmental factors affect mental-state inference and social behavior across cultures. Given that frequency of allelic variation in genes associated with social cognition, such as the serotonin transporter gene and oxcytocin receptor polymorphism, varies across geographic regions and is known to interact with environmental factors, such culture, an important direction for future research in social cognition is to examine how both genetic and environmental factors contribute to social cogni-tion across cultures.

Application

Disruptions in social cognition, including autism, Williams syndrome, prosopagnosia, social phobia, and schizophrenia, underlie some of the most complex and challenging mental health disorders around the globe. While these mental health disor-ders may simply reflect naturally occurring indi-vidual variation in social cognition that may even confer adaptive benefits to the population as a whole, individuals with autism or Williams syn-drome often suffer detriments to the quality of their social life and will seek treatment as a means of reducing the emotional burden of living with the disorder. However, the cost of treating mental health disorders is often exorbitant for individuals and institutions alike. Ganz (2006) estimates that caring for an autistic person alone can cost about $3.2 million over his or her lifetime and that caring for all people with autism over their

lifetimes costs an estimated $35 billion per year within the United States. Similar to other kinds of complex atypical behavior, such as mood disorders, the source and nature of these social-cognitive disorders likely results from complex interactions between genetic and environmental factors. Developing a more precise model of gene-by-environment interaction in social cognition may enable us to discover how to effectively treat such complex health disorders as well as create a richer basic science understanding of how social-cognitive mechanisms in the mind and brain give rise to the fundamental human capacity for social living.

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