Strikingly, several regions reviewed by Lindquist et al. are not only involved in the perception and experience of emotions, but are the very same neural regions most often implicated in social behaviors. Here, we argue that culture as the broadest form of social behavior depends upon – and evolved along with – general neural mechanisms underlying emotion. Specifically, we use cultural neuroscience findings on emotional vigilance and empathy to illustrate that general mechanisms, rather than specific emotional modules, support how we think, feel, and behave in our social worlds.
Implicating the basic processes of executive attention and core affect in social contexts, cultural neuroscience research reveals a relationship between collectivism and emotional vigilance. Specifically, participants primed with collectivistic values responded more quickly to fear-relevant stimuli in a dot-probe task (Chiao & Lee Reference Chiao and Lee2011). This is further supported by a cross-cultural study of Japanese, Japanese-American, and Caucasian-American participants, demonstrating that Japanese, relative to Americans of either race, show heightened amygdala activity in response to negative emotional stimuli (Chiao et al. Reference Chiao, Hariri, Harada, Mano, Hechtman, Komeda, Sadato, Parrish and Iidaka2011). Furthermore, bicultural Asian-Americans primed with collectivistic values show increased amygdala response to negative emotional stimuli, indicating that being in either an individualistic or collectivistic cultural mindset elicits a culturally congruent emotional effect (Hechtman et al. Reference Hechtman, Hariri, Harada, Mano, Sadato, Parrish, Iidaka and Chiao2011).
We interpret these findings as suggesting that collectivism necessitates increased emotional vigilance (in a constructionist's framework, core affect and executive attention). For example, collectivistic nations have historically had higher levels of environmental threat, such as increased pathogen prevalence, which may require vigilance for survival (Fincher et al. Reference Fincher, Thornhill, Murray and Schaller2008). Furthermore, collectivistic cultures typically place a strong emphasis on group membership and hierarchy (Fincher et al. Reference Fincher, Thornhill, Murray and Schaller2008). These tight constraints on appropriate social behavior may require increased attention to negative cues in one's environment, in order to correct one's behavior and bring behavior into line with social norms. Building on this evolutionary perspective, genetic factors may parallel and support culture's influence on emotion. For instance, recent evidence suggests that the serotonin transporter gene (5-HTTLPR) – known to regulate neurotransmission within the amygadala and other regions supporting emotional vigilance (Munafo et al. Reference Munafò, Brown and Hariri2008) – may have co-evolved along with cultural values of individualism–collectivism (Chiao & Blizinsky Reference Chiao and Blizinsky2010). Specifically, collectivistic nations have larger proportions of individuals who carry the S compared to the L allele of the 5-HTTLPR (Chiao & Blizinsky Reference Chiao and Blizinsky2010). S-allele carriers show increased amygdala response to negative stimuli (Hariri et al. Reference Hariri, Mattay, Tessitore, Kolachana, Fera and Goldman2002), paralleling the cultural effects. We therefore propose that genetic and cultural factors have exerted mutual evolutionary influence on neural systems underlying attention and core affect.
As implied by this culture-gene coevolutionary view, the aforementioned findings on amygdala and associated genes can be interpreted as mechanisms tuning people's basic emotional processes. This is consistent with a view of the amygdala as not merely supporting a discrete emotion category, but as a center that modulates other regions to create a nonspecific state of emotional vigilance (Davis & Whalen Reference Davis and Whalen2001). Similarly, since the amygdala typically serves an important social function (Adolphs Reference Adolphs2010), its activity likely enables appropriate cultural behaviors based on environmental context. Altogether, the amygdala functions as a modulator of emotional and attentional processes. A locationist view of emotion therefore fails to account for how the brain evolved and how it carries out its responses to sociocultural and environmental demands.
Besides moderating emotional vigilance, culture shapes other social cognitive processes involving core affect and conceptualization. Particularly, culture influences neural mechanisms underlying group coordination and helping behaviors, such as empathy and pro-sociality (Boyd & Richerson Reference Boyd and Richerson2009). When empathizing with others' feelings, people recruit the bilateral insula, the anterior cingulate cortex (ACC), and somatosensory cortices during both perception and experience of others' emotional states (Lamm et al. Reference Lamm, Decety and Singer2011). Additionally, neural response is enhanced within brain regions associated with mentalizing, such as the medial prefrontal cortex, which likely reflects the recruitment of self-related processes for understanding and sharing the emotional states of others (Frith & Frith Reference Frith and Frith2006).
Neural response within the empathic brain is modulated by a number of sociocultural factors, including social dominance orientation (SDO) (Chiao et al. Reference Chiao, Mathur, Harada and Lipke2009), implicit racial attitudes (IAT; Avenanti et al. Reference Avenanti, Sirigu and Aglioti2010; Bruneau & Saxe Reference Bruneau and Saxe2010), racial identification (MEIM; Mathur et al. Reference Mathur, Harada, Lipke and Chiao2010), and modern racism (MR; Gutsell & Inzlicht, in press). Specifically, people with strong social hierarchy preference (i.e., high SDO) show less ACC and insula activity when empathizing with others' pain. Additionally, people with stronger racial IAT biases show reduced sensorimotor response to out-group members (Avenanti et al. Reference Avenanti, Sirigu and Aglioti2010). Beyond ACC and insula activity, MEIM predicts extraordinary in-group empathy in African-Americans, indexed by increased activity in the medial prefrontal cortex (MPFC) (Mathur et al. Reference Mathur, Harada, Lipke and Chiao2010). Furthermore, MEIM is associated with default-mode network activity within cortical midline structures (Mathur et al., in press). Finally, Gutsell and Inzlicht (in press) demonstrate the association between MR and prefrontal asymmetry.
Similar to the proposed roles of the amygdala, neural mechanisms underlying empathy can be interpreted from an evolutionary perspective. In constructionist terms, processes of core affect (e.g., in the ACC and insula) and conceptualization (e.g., in the MPFC) give rise to empathy, which channels people's emotion towards others through re-experience of their feelings. Empathic processes, then, may have evolved to support pro-social behaviors in groups and, in turn, may have coevolved with intricate sociocultural structures. For example, sociocultural constraints on empathic feeling (e.g., SDO, racial IAT, MEIM, and MR) likely create (and are enabled by) differential feelings for others based on group and/or status. Here the modulation of various processes (e.g., core affect and conceptualization) supports both differential feelings towards, and differential concepts about, other people. Localist views of emotion cannot account for the multiple converging, socially shaped processes crucially involved in empathic feeling.
In sum, using cultural neuroscience work on emotional vigilance and empathy, we illustrate that both social and emotional behaviors are products of basic processes that are culturally and environmentally shaped. This process view, assuming evolutionary development of fundamental psychological building blocks, accounts for the vast overlap in social and emotional brain regions. Together, cultural neuroscience research, accounting for the evolution of culturally appropriate forms of emotion, supports constructionist, but not localist, views on emotion.
Strikingly, several regions reviewed by Lindquist et al. are not only involved in the perception and experience of emotions, but are the very same neural regions most often implicated in social behaviors. Here, we argue that culture as the broadest form of social behavior depends upon – and evolved along with – general neural mechanisms underlying emotion. Specifically, we use cultural neuroscience findings on emotional vigilance and empathy to illustrate that general mechanisms, rather than specific emotional modules, support how we think, feel, and behave in our social worlds.
Implicating the basic processes of executive attention and core affect in social contexts, cultural neuroscience research reveals a relationship between collectivism and emotional vigilance. Specifically, participants primed with collectivistic values responded more quickly to fear-relevant stimuli in a dot-probe task (Chiao & Lee Reference Chiao and Lee2011). This is further supported by a cross-cultural study of Japanese, Japanese-American, and Caucasian-American participants, demonstrating that Japanese, relative to Americans of either race, show heightened amygdala activity in response to negative emotional stimuli (Chiao et al. Reference Chiao, Hariri, Harada, Mano, Hechtman, Komeda, Sadato, Parrish and Iidaka2011). Furthermore, bicultural Asian-Americans primed with collectivistic values show increased amygdala response to negative emotional stimuli, indicating that being in either an individualistic or collectivistic cultural mindset elicits a culturally congruent emotional effect (Hechtman et al. Reference Hechtman, Hariri, Harada, Mano, Sadato, Parrish, Iidaka and Chiao2011).
We interpret these findings as suggesting that collectivism necessitates increased emotional vigilance (in a constructionist's framework, core affect and executive attention). For example, collectivistic nations have historically had higher levels of environmental threat, such as increased pathogen prevalence, which may require vigilance for survival (Fincher et al. Reference Fincher, Thornhill, Murray and Schaller2008). Furthermore, collectivistic cultures typically place a strong emphasis on group membership and hierarchy (Fincher et al. Reference Fincher, Thornhill, Murray and Schaller2008). These tight constraints on appropriate social behavior may require increased attention to negative cues in one's environment, in order to correct one's behavior and bring behavior into line with social norms. Building on this evolutionary perspective, genetic factors may parallel and support culture's influence on emotion. For instance, recent evidence suggests that the serotonin transporter gene (5-HTTLPR) – known to regulate neurotransmission within the amygadala and other regions supporting emotional vigilance (Munafo et al. Reference Munafò, Brown and Hariri2008) – may have co-evolved along with cultural values of individualism–collectivism (Chiao & Blizinsky Reference Chiao and Blizinsky2010). Specifically, collectivistic nations have larger proportions of individuals who carry the S compared to the L allele of the 5-HTTLPR (Chiao & Blizinsky Reference Chiao and Blizinsky2010). S-allele carriers show increased amygdala response to negative stimuli (Hariri et al. Reference Hariri, Mattay, Tessitore, Kolachana, Fera and Goldman2002), paralleling the cultural effects. We therefore propose that genetic and cultural factors have exerted mutual evolutionary influence on neural systems underlying attention and core affect.
As implied by this culture-gene coevolutionary view, the aforementioned findings on amygdala and associated genes can be interpreted as mechanisms tuning people's basic emotional processes. This is consistent with a view of the amygdala as not merely supporting a discrete emotion category, but as a center that modulates other regions to create a nonspecific state of emotional vigilance (Davis & Whalen Reference Davis and Whalen2001). Similarly, since the amygdala typically serves an important social function (Adolphs Reference Adolphs2010), its activity likely enables appropriate cultural behaviors based on environmental context. Altogether, the amygdala functions as a modulator of emotional and attentional processes. A locationist view of emotion therefore fails to account for how the brain evolved and how it carries out its responses to sociocultural and environmental demands.
Besides moderating emotional vigilance, culture shapes other social cognitive processes involving core affect and conceptualization. Particularly, culture influences neural mechanisms underlying group coordination and helping behaviors, such as empathy and pro-sociality (Boyd & Richerson Reference Boyd and Richerson2009). When empathizing with others' feelings, people recruit the bilateral insula, the anterior cingulate cortex (ACC), and somatosensory cortices during both perception and experience of others' emotional states (Lamm et al. Reference Lamm, Decety and Singer2011). Additionally, neural response is enhanced within brain regions associated with mentalizing, such as the medial prefrontal cortex, which likely reflects the recruitment of self-related processes for understanding and sharing the emotional states of others (Frith & Frith Reference Frith and Frith2006).
Neural response within the empathic brain is modulated by a number of sociocultural factors, including social dominance orientation (SDO) (Chiao et al. Reference Chiao, Mathur, Harada and Lipke2009), implicit racial attitudes (IAT; Avenanti et al. Reference Avenanti, Sirigu and Aglioti2010; Bruneau & Saxe Reference Bruneau and Saxe2010), racial identification (MEIM; Mathur et al. Reference Mathur, Harada, Lipke and Chiao2010), and modern racism (MR; Gutsell & Inzlicht, in press). Specifically, people with strong social hierarchy preference (i.e., high SDO) show less ACC and insula activity when empathizing with others' pain. Additionally, people with stronger racial IAT biases show reduced sensorimotor response to out-group members (Avenanti et al. Reference Avenanti, Sirigu and Aglioti2010). Beyond ACC and insula activity, MEIM predicts extraordinary in-group empathy in African-Americans, indexed by increased activity in the medial prefrontal cortex (MPFC) (Mathur et al. Reference Mathur, Harada, Lipke and Chiao2010). Furthermore, MEIM is associated with default-mode network activity within cortical midline structures (Mathur et al., in press). Finally, Gutsell and Inzlicht (in press) demonstrate the association between MR and prefrontal asymmetry.
Similar to the proposed roles of the amygdala, neural mechanisms underlying empathy can be interpreted from an evolutionary perspective. In constructionist terms, processes of core affect (e.g., in the ACC and insula) and conceptualization (e.g., in the MPFC) give rise to empathy, which channels people's emotion towards others through re-experience of their feelings. Empathic processes, then, may have evolved to support pro-social behaviors in groups and, in turn, may have coevolved with intricate sociocultural structures. For example, sociocultural constraints on empathic feeling (e.g., SDO, racial IAT, MEIM, and MR) likely create (and are enabled by) differential feelings for others based on group and/or status. Here the modulation of various processes (e.g., core affect and conceptualization) supports both differential feelings towards, and differential concepts about, other people. Localist views of emotion cannot account for the multiple converging, socially shaped processes crucially involved in empathic feeling.
In sum, using cultural neuroscience work on emotional vigilance and empathy, we illustrate that both social and emotional behaviors are products of basic processes that are culturally and environmentally shaped. This process view, assuming evolutionary development of fundamental psychological building blocks, accounts for the vast overlap in social and emotional brain regions. Together, cultural neuroscience research, accounting for the evolution of culturally appropriate forms of emotion, supports constructionist, but not localist, views on emotion.