The Neuroscience of Stereotyping
The Neuroscience of Stereotyping
The Neuroscience of Stereotyping
Social motivations, such as the desire to affiliate or com- characteristics ascribed to a social group, such as personal
Social motivations
Motives that operate in social pete with others, rank among the most potent of human traits (for example, unintelligent) or circumstantial attrib-
contexts and satisfy basic, drives1. Not surprisingly, the capacity to discern ‘us’ from utes (for example, poor)5. Although they are distinguish-
often universal, goals and ‘them’ is fundamental in the human brain. Although this able by content and process, prejudices and stereotypes
aspirations, such as to affiliate computation takes just a fraction of a second2,3, it sets the often operate in combination to influence social behav-
(for example, form relationships
and communities) or to achieve
stage for social categorization, stereotypes, prejudices, inter- iour 6. Moreover, both forms of bias can operate implicitly,
dominance (for example, within group conflict and inequality, and, at the extremes, war such that they may be activated and influence judgements
a social hierarchy). and genocide. Thus, although prejudice stems from a and behaviours without conscious awareness7–9.
mechanism of survival, built on cognitive systems that Despite the persistence of prejudices and stereotypes
Stereotypes
‘structure’ the physical world, its function in modern in contemporary society, their effects on behaviour are
Conceptual attributes
associated with a group and its society is complex and its effects are often deleterious. often countered by people’s egalitarian personal beliefs
members (often through over- For the neuroscientist, the domain of prejudice pro- and pro-social norms7. Guided by these beliefs and
generalization), which may vides a unique context for examining neural mechanisms norms, people frequently engage self-regulatory pro-
refer to trait or circumstantial of the human mind that guide complex behaviour. Social cesses to mitigate the effects of bias on their behaviour.
characteristics.
prejudices are scaffolded by basic-level neurocognitive Hence, a theoretical analysis of prejudice and stereo-
Prejudices structures, but their expression is guided by personal typing is incomplete without a consideration of these
Evaluations of or affective goals and normative expectations, played out in dyadic regulatory processes. Here, self-regulation refers to
responses towards a social and intergroup settings; this is truly the human brain the process of acting in an intentional manner, often
group and its members based
in vivo. Although probing the neural basis of prejudice through mechanisms of cognitive control.
on preconceptions.
is a challenging endeavour — in which the rigours of The neuroscientific research conducted on prejudice
reductionism are balanced with the richness of context and stereotyping over the past decade suggests that these
— it offers neuroscientists the opportunity to connect complex forms of human behaviour involve different
their knowledge to some of society’s most pressing prob- interacting networks of neural structures. In this article, I
lems, such as discrimination, intergroup conflict and describe the functions of key structures in each network,
disparities in health and socioeconomic status. including both their broader neurocognitive functions
New York University,
In this article, I review research on the role of the brain and their specific roles in prejudice and stereotyping.
Department of Psychology,
6 Washington Place, in social prejudice and stereotyping. The term prejudice is This article extends previous reviews on this topic —
New York, New York 10003, used broadly to refer to preconceptions — often negative which were guided by a social psychological analysis10
USA. — about groups or individuals based on their social, racial or emphasized a particular neuroimaging method11,12 —
e-mail: david.amodio@nyu. or ethnic affiliations4. Within the field of social psychol- by providing a comprehensive overview of the literature
edu
doi:10.1038/nrn3800
ogy, prejudice refers more specifically to evaluations (that from a neural-systems perspective. Although many of
Published online is, attitudes) and emotional responses towards a group and the conclusions drawn from this emerging literature rely
4 September 2014 its members. Stereotypes, by comparison, are generalized heavily on reverse inference from neuroimaging data,
Self-regulation these inferences are strengthened by converging theory many findings in this literature concern basic mechanisms
The process of responding in and behavioural data from the extensive psychological of social cognition that, to varying extents, underlie other
an intentional manner, often literature on intergroup bias and self-regulation13,14. forms of bias, such as those based on ethnicity, gender,
involving the inhibition or The majority of the research reviewed here concerns sexual preference and nationality.
overriding of an alternative
response tendency.
racial prejudice — a form of prejudice with clearly defined
social categories linked to identifiable physical attrib- Neural basis of prejudice
utes (BOX 1). In particular, prejudice of white Americans In the modern social psychology literature, prejudice
towards black people (that is, individuals of African or is defined as an attitude towards a person on the basis
Caribbean descent) has deep historical roots and contem- of his or her group membership. Prejudice may reflect
porary relevance to social issues, and the majority of stud- preference towards ingroup members or dislike of out-
ies have examined prejudice in this context. Nevertheless, group members, and it is typically imbued with affect,
with emotions ranging from love and pride to fear, dis-
gust and hatred15,16. Consequently, research on the neu-
Box 1 | Seeing race: the role of visual perception ral basis of prejudice has primarily focused on neural
structures involved in emotion and motivation, such as
Social interactions often begin with the perception of a face. Mounting evidence reveals the amygdala, insula, striatum and regions of orbital and
that social motivations can alter the way a face is seen, which presumably reflects the
ventromedial frontal cortices (FIG. 1). Although they are
modulatory influences of signals from the temporal cortex, prefrontal cortex and
amygdala to the fusiform gyrus148. This insight suggests that prejudices and stereotypes often examined independently, these structures appear
may alter early face processing. to form a core network for the experience and expression
Early functional MRI (fMRI) research demonstrated greater fusiform activity (see the of prejudice.
figure, panel a) in response to faces of one’s own racial group (that is, the ingroup) — an
effect that was associated with better recognition of ingroup faces than outgroup faces50. Amygdala. Research on the neural basis of prejudice
Research examining the N170 component of the event-related potential (ERP), which has most frequently examined the amygdala, a complex
indexes the degree of initial configural face encoding at just ~170 ms, revealed enhanced subcortical structure located bilaterally in the medial
processing of ingroup versus outgroup faces (see the figure, panel b), even when groups temporal lobes (FIG. 1). Although the amygdala is some-
were defined arbitrarily3. This finding is consistent with fMRI data showing that faces of times described as a neural locus of emotion (for exam-
‘coalition members’ elicited greater activity in the fusiform gyrus than did other faces,
ple, fear), it in fact comprises approximately 13 distinct
regardless of race149. Hence, social group membership, even when defined on the basis of
minimal categories, promotes greater visual encoding. These findings dovetail with nuclei that, in conjunction, perform multiple functions
behavioural research showing that biased visual representations of outgroup members to support adaptive behaviour 17 (FIG. 2).
facilitate discriminatory actions towards them150,151. The amygdala receives direct (or nearly direct) affer-
In the context of race, outgroup members are often viewed as threatening and ents from all sensory organs into its lateral nucleus, ena-
therefore may elicit vigilant attention. Indeed, larger N170 ERP amplitudes in response to bling it to respond very rapidly to immediate threats in
viewing black versus white faces (equated in luminance) have been observed in subjects advance of more elaborative processing of a stimulus18.
with stronger implicit prejudice152 and in subjects who were made to feel anxious about Within the amygdala, the central nucleus (CeA) has
appearing biased42. These and other findings suggest that the visual processing of race is been implicated in Pavlovian (classical) fear condition-
malleable and depends on social motivations and contexts153–158. ing in both rats and humans19–22, and signals emerging
Neural representations of race (black versus white), as determined by multivoxel
from the CeA activate hypothalamic and brainstem
pattern analysis (MVPA), have been observed in the fusiform gyrus, and these neural
representations have been associated with behavioural indices of implicit prejudice structures to induce arousal, attention, freezing and
and stereotyping49,52,159,160. It is notable that MVPA has also identified race preparation for fight or flight — a response that is often
representation in the medial occipital cortex; however, because full-colour photos characterized as ‘fear’. By comparison, output from the
were used in these studies, the effect may reflect differences in luminance associated basal nucleus guides appetitive and instrumental responses
with skin tone rather than the race of the people depicted. Nevertheless, the broader via projections to the ventral striatum22,23. Both the fear-
body of findings suggests that social category cues modulate the early visual related and appetitive functions of the amygdala involve
processing of ingroup and outgroup members’ faces in ways that support the motivation and attention, but to different ends, and they
perceivers’ biased or egalitarian social goals. probably correspond to different aspects of a prejudice-
Panel a of the figure is from REF. 50, Nature Publishing Group. Panel b of the figure is based response. In humans, the amygdala is integral
reprinted from J. Exp. Soc. Psychol., 49, Ratner, K. G. & Amodio, D. M., Seeing “us versus
to the processing of fear in facial expressions as well as
them”: minimal group effects on the neural encoding of faces, 298–301, Copyright (2013),
with permission from Elsevier. other salient social cues24. Given the amygdala’s ability
to respond rapidly to potential social threat, researchers
a b interested in the neural substrate of implicit prejudice
4 first looked to this brain structure.
Ingroup
Outgroup A pair of early functional MRI (fMRI) studies exam-
2
ined the amygdala activity of white research subjects in
Amplitude (µV)
Time Time
Evaluation > stereotype Stereotype > evaluation members in pain71. In a conceptually related study of
gender bias, men who reported highly sexist attitudes
exhibited lower mPFC activity when viewing sexual-
ized images of female (but not male) bodies than men
with less sexist views — a pattern consistent with idea
that sexual objectification involves a form of dehuman-
ization82. Hence, the mPFC’s role in prejudice seems to
be marked by an absence of activity, which may reflect
a lack of humanization and empathy regarding disliked
Decoding accuracy
Decoding accuracy
56 56
or disrespected outgroup members.
54 54
52 52
A neural network for prejudice. Prejudice is a complex
50 50 social cognitive process that seems to be supported by
48 48 a network of neural structures (FIG. 1). The amygdala
46 46 supports threat-based associations, which are thought
Judgement type Judgement type
to underlie the most common form of implicit preju-
Stereotype Evaluation dice, and it is also involved in initial responses to sali-
Figure 3 | Neural representation of racial bias in ent positive or negative cues, including cues regarding
Nature Reviews
affect-based and stereotype-based | Neuroscience
judgements. group membership. Thus, depending on the situation
A multivoxel pattern analysis approach revealed a unique and nuclei of interest, amygdala activity may be associ-
representation (that is, decoding accuracy) of race in the ated with social threat or with social reward. Activity in
orbital frontal cortex when subjects judged images of the anterior insula supports the subjective experience of
black and white males according to an evaluative negative affect (which often accompanies a prejudiced
dimension (who is more likely to be a friend?), and a response), whereas the mPFC is involved in mental-
unique representation of race in the medial prefrontal izing and perspective taking, which may be engaged
cortex when judging black and white males on a more strongly towards ingroup than outgroup members.
stereotypical trait dimension (who is more interested in
Neural projections from the amygdala and insula to the
athletics?)49. Reprinted from Neuropsychologia, 50,
Gilbert, S. J., Swencionis, J. K. & Amodio, D. M., Evaluative ventral mPFC may support the integration of affective
versus trait representation in intergroup social responses with mentalizing and empathy processes.
judgments: distinct roles of anterior temporal lobe and Finally, appetitive responses such as positive attitudes
prefrontal cortex, 3600–3611, Copyright (2012), with and approach-related behavioural tendencies, which are
permission from Elsevier. often expressed towards ingroup members, are primar-
ily supported by the striatum. These brain regions may
function in concert to support the learning, experience
interconnections with the ACC, the insula, the OFC and and expression of prejudice.
the dorsolateral PFC (dlPFC), as well as other structures62.
In the context of social cognition, mPFC activity Neural basis of stereotyping
has been primarily associated with the formation of In contrast to prejudice, which reflects an evaluative or
impressions about other people, especially impressions emotional component of social bias, stereotypes repre-
that require mentalizing — the process of considering sent the cognitive component — the conceptual attrib-
a person’s unique perspective and motives (that is, utes linked to a particular group as defined by a culture
engaging in theory of mind)80. Because the mPFC is or society. This process involves the encoding and stor-
typically activated during judgements about other peo- age of stereotype concepts, the selection and activa-
ple (as opposed to inanimate objects)77, mPFC activ- tion of these concepts into working memory and their
ity, particularly in ventral, perigenual regions (FIG. 1), application in judgements and behaviours5,83. As such,
has been interpreted by some theorists as reflecting a stereotyping involves cortical structures that support
‘humanization’ process and, by extension, empathy 67,81. more general forms of semantic memory, object mem-
Hence, a lack of mPFC activity in response to a social ory, retrieval and conceptual activation, such as the tem-
target may indicate a form of prejudice that is charac- poral lobes and inferior frontal gyrus (IFG), as well as
terized by a lack of humanization (that is, dehumani- regions that are involved in impression formation, such
zation) and empathy. Indeed, the ventral mPFC has as the mPFC10,84,85. Although there is substantial overlap
been shown to be more highly activated when a sub- between these structures and those implicated in preju-
ject views members of esteemed groups associated with dice, as described above, these structures appear to form
pride and admiration than when a subject views mem- the core of a stereotyping network in the brain that may
bers of low-status groups associated with disgust and operate separately from a prejudice network (FIG. 4).
disregard (for example, homeless people)67. Moreover,
in a study in which Chinese and Caucasian subjects Temporal lobe. Stereotypes reflect conceptual associa-
viewed images of people being exposed to a painful or tions between social groups and a particular set of attrib-
non-painful stimulus (that is, a needle penetrating the utes — associations that are thought to reside in semantic
cheek versus a Q‑tip touch), mPFC and ACC activity memory 83. As such, stereotype associations are posited
was elicited only in response to seeing racial ingroup to involve regions of the lateral temporal lobe that
Dorsal mPFC Lateral temporal lobe that the ATL is necessary for stereotype representation.
Impression formation Semantic and episodic memory Thus, knowledge of social stereotypes appears to reside
in the ATL.
lateral PFC with the basal ganglia and motor cortex sup- the coherence of these proposed functional networks for
port the coordination of complex actions that are guided prejudice and stereotyping and their interaction has yet
by working memory and high-level cognition 73,110. to be tested, their existence is consistent with known
Stereotypes are a form of social cognition that guide anatomical connectivity (which has been primarily
behaviour, and indeed the process of applying stereo observed in non-human animals) and is supported by
types in judgement and behaviour has been shown to decades of theory and behavioural research in the social
specifically involve activity in the IFG111. psychology literature4,5.
Whereas the retrieval of conceptual knowledge typi-
cally involves the left IFG, activity in the right IFG has Regulation of prejudice and stereotyping
been observed in research subjects who were judging In an era of increasing diversity, international relations,
whether gender-stereotyped traits applied to a series of global communication and awareness of civil rights
male and female individuals (as compared with traits that issues, intergroup biases are often deemed to be both per-
were unrelated to gender stereotypes)111. Given other evi- sonally and socially unacceptable. Preferences based on
dence that the right IFG has a role in domain-general racial and ethnic categories that may have been adaptive
response inhibition112, it is possible that activation of in less complex societies are no longer so. Fortunately,
the right IFG during stereotype judgement tasks reflects the human mind is adept at self-regulation, and although
an individual’s efforts to inhibit the influence of stereo- stereotypes and prejudices may come to mind automati-
types on behaviour. This pattern of lateralized function cally in intergroup contexts, their expression can often be
— stereotype retrieval and implementation on the left moderated. Neuroscience research on the mechanisms
and response inhibition on the right — suggests a useful supporting the control of intergroup responses incorpo-
distinction in the processes through which stereotypes rates existing domain-general models of cognitive con-
are applied in behaviour. trol into broader models that consider the influence of
social factors. For example, the impetus for the control
A neural network for stereotyping. The research described of racial bias may arise from internal cues (for example,
above suggests that a network of neural structures sup- the personal rejection of prejudice) or external cues (for
ports stereotyping processes (FIG. 4). The ATL is believed example, social pressure to respond without prejudice),
to represent stereotype-related knowledge, and it provides and engagement in control is frequently associated with
input to the mPFC, possibly also during the online for- social emotions such as social anxiety or guilt 43. A neural
mation of impressions about an individual. In this way, model of prejudice control should account for these dif-
social stereotypes ‘stored’ in the ATL may influence trait ferent impetuses and emotion effects. In this way, neuro-
impression processes associated with dorsal mPFC activ- science research on prejudice has inspired an expanded
ity. The application of stereotypes to behaviour seems to view of the neural and psychological processes involved
involve regions of the lateral PFC that are associated with in control.
goal representation and response inhibition. Together,
the structures in this putative network may support the Anterior cingulate cortex. The ACC has been widely
storage, activation and behavioural expression of social implicated in the monitoring and detection of response
stereotypes. conflict 116,117. In particular, the dorsal region of the ACC
(FIG. 5) is often activated during cognitive control tasks,
Interacting networks such as the Stroop or Flanker tasks, on trials involving
The framework described above suggests separate net- a high degree of conflict between one’s desired response
works for prejudice and stereotyping, but in most cases and a countervailing tendency 118,119. Conflict monitor-
these two processes operate in concert. Thus, although ing theory posits that as the conflict signal in the ACC
they are largely rooted in distinct neural systems, their rises, the ACC increasingly engages dlPFC regions that
effects converge in higher-level cognition and behav- function to implement goal-directed behaviour 120. This
ioural expression. Neuroscience studies suggest several model is consistent with the ACC’s connectivity with
places at which this convergence may occur, although PFC regions involved in high-level goal representa-
this is primarily based on studies of connectivity in non- tion and with the PFC’s connectivity with the striatum,
human animals. For example, the anatomical connec- through which top-down control is implemented in
tivity of the amygdala and OFC with the ATL, via the behaviour 73,110,121.
Event-related potential
uncinate fasciculus92, is consistent with behavioural evi- In a social context, cognitive control is needed to
(ERP). An electrical signal dence that affective responses may influence the activa- curb the unwanted influence of implicit stereotypes and
produced by summated tion of stereotype concepts, and vice versa113,114. Similarly, prejudices on behaviour 7,122. Building on conflict moni-
postsynaptic potentials of signals from several structures that are involved in both toring theory, it has been proposed that the control of
cortical neurons in response to
prejudice and stereotyping — including the amygdala, implicit bias requires the detection of a conflict between
a discrete event, such as a
stimulus or a response in an insula, striatum, OFC and ATL — converge in regions of a biased tendency and one’s goal to act without bias123.
experimental task. Typically the mPFC, where information seems to be integrated in Support for this proposal was provided by a study that
recorded from the scalp in support of elaborate person representations62,63. Finally, assessed ACC activity, which was indexed by the error-
humans, ERPs can be the joint influences of prejudiced affect and stereotype related negativity component of the event-related potential
measured with extremely high
temporal resolution and can be
concepts on behaviour are likely to converge in the stria- (ERP). In this study, subjects performed a task that
used to track rapid, real-time tum, which receives inputs from the amygdala, OFC, lat- required them to inhibit the automatic expression of
changes in neural activity. eral PFC and ATL, as well as other regions73,115. Although racial stereotypes on some trials but not others (the
dACC
feedback about their bias. In two fMRI studies in which
Conflict processing subjects completed IAT measures of racial attitudes, false
mPFC
IAT feedback to the subjects indicating that he or she
Representation of showed racial prejudice elicited heightened ACC activity,
interpersonal cues and this degree of activity was associated with feelings
rACC of guilt — a self-regulatory emotion that promotes pro-
Monitoring social behaviours129,130. Together, these findings suggest
external cues that the ACC supports the detection of one’s unwanted
social biases and the engagement of cognitive control in
dlPFC order to avoid the expression of bias.
Response selection
Lateral prefrontal cortex. As noted above, the PFC is
IFG
Response inhibition associated with working memory, response selection and
the representation of high-level goals131,132, and it governs
most goal-directed responses in humans. Lateral PFC
regions (FIG. 5), in particular, coordinate the control of
action and attention; most findings indicate that activity
Figure 5 | Regulation network. Neural structures supporting
NaturetheReviews
regulation of
| Neuroscience in the left lateral PFC is linked to the implementation of
intergroup responses. Conflicts between a biased tendency and either internal goals or action, whereas activity in the right lateral PFC is linked
external cues (for example, social norms) are processed in the dorsal anterior cingulate to action inhibition (in right-handed individuals)133,134.
cortex (dACC) and rostral ACC (rACC), respectively. The medial prefrontal cortex (mPFC) On the basis of research in cognitive neuroscience, lateral
is involved in perspective taking and mentalizing, and activation in this region provides
PFC regions have been proposed as primary substrates of
further representation of interpersonal cues to guide regulatory processing. Intergroup
response goals are represented in the lateral PFC and implemented in behaviour in cognitive control of prejudice27,37,64.
coordination with the striatum and motor cortex. dlPFC, dorsolateral PFC; IFG, inferior Until recently, fMRI studies of responses towards
frontal gyrus. ingroup versus outgroup faces were not designed to
elicit or assess cognitive control. That is, most stud-
ies used tasks in which pictures of racial ingroup and
weapons identification task; see BOX 2). ACC activity outgroup members were viewed passively by subjects.
was selectively greater on trials requiring stereotype Interestingly, these studies consistently revealed activ-
inhibition, and the degree of subjects’ ACC activity on ity in regions of the PFC, most often the right IFG, in
these trials predicted their success at controlling the response to explicit presentations of black faces com-
expression of stereotypes in task behaviour 123. This find- pared with white faces37,45,48. Although this finding is
ing has been replicated and extended in several studies difficult to interpret in terms of control, the established
using different tasks and with alternative ERP indices of role of the right IFG in response inhibition suggests that
ACC activity (for example, the error-related negativity exposure to black faces in these tasks may have spon-
and response-locked N2)124–127. These studies revealed, taneously elicited a form of inhibitory control, perhaps
for example, that ACC activation in response to stereo- owing to subjects’ concern about appearing prejudiced.
type conflict occurs implicitly and without deliberation, In addition, right IFG activity has been reported to cor-
is observed on a range of cognitive control tasks that relate negatively with amygdala activity in response to
require the inhibition of stereotype-based responses and viewing black faces. This could suggest a potential regu-
is associated with an individual’s motivation to respond latory circuit for prejudice control37,45; however, as these
without prejudice. data are merely correlational and as there is little direct
In several fMRI studies, ACC activation has also been connectivity between the IFG and amygdala135, it is more
observed in white American subjects while they viewed likely that right IFG activity reflects a form of response
images of black (versus white) faces. However, this acti- inhibition rather than the direct downregulation of
vation typically occurred in the absence of an opportu- amygdala activity.
nity for cognitive control (that is, during tasks that did The role of lateral PFC activity in the cognitive con-
not require control, such as passive face viewing)37,45,48. trol of race biased behaviour was examined directly
Thus, it is difficult to interpret ACC activations in these in an electroencephalography (EEG) study, in which
studies in terms of a control process. Nevertheless, subjects completed a task that assessed the behavioural
these findings suggest that exposure to black faces may inhibition of stereotypes (the weapons identification
spontaneously elicit conflict detection processes. In task)136. Greater dlPFC activity was found to be associ-
one notable exception — a study in which fMRI was ated with better behavioural control (as modelled using
recorded during performance on a racial attitude IAT, the process dissociation procedure), indicating a direct
which requires cognitive control — ACC activity was link between the dlPFC and control of stereotyping.
associated with the ability to identify the correct (that Furthermore, the relation between dlPFC activity and
is, non-biased) task response64. This finding is concep- stereotype control was mediated by greater attentional
tually consistent with evidence from ERP studies that orienting to black faces than white faces, as indexed
link ACC activity to conflict processing and the engage- by the P2 component of the ERP. This pattern sug-
ment of top-down control128. Similar patterns have been gested that PFC activity tuned perceptual attention to
observed when people are confronted with explicit relevant stimuli, which in turn facilitated behavioural
control (BOX 2). Findings consistent with this idea have that these two forms of bias are subserved by different
been reported in studies using EEG, fMRI and brain learning and memory systems — a clue that interventions
lesion approaches in combination with behavioural to reduce prejudice and stereotyping may require differ-
tasks designed to assess elements of prejudice con- ent approaches10,28. Implicit prejudice has been linked to
trol51,64,126,137,138. Together, these studies have begun to fear conditioning involving the amygdala, whereas ste-
identify the specific pathways through which the PFC reotype associations appear to reflect conceptual learn-
guides the control of intergroup responses. ing systems involving the temporal cortex and PFC.
Importantly, learning and expression differ consider-
Medial prefrontal cortex. As noted in previous sections, ably between these systems: fear conditioning may be
the mPFC contributes to aspects of both stereotyping acquired in a single trial and expressed primarily through
and prejudice. However, this region is large, heterogene- behavioural freezing, anxiety and heightened vigilance18,
ous and widely interconnected, and emerging theories whereas conceptual associations require many exposures
and research suggest that its function may be closely for acquisition and are expressed through high-level rep-
tied to regulatory processes as well. Amodio and Frith62 resentations of impressions and goal-directed actions140.
proposed that, given its role in mentalizing, the mPFC Social cognition studies have begun to adopt interven-
supports the regulation of behavioural responses accord- tion strategies that are consistent with this analysis, using
ing to social cues. Early evidence from ERP data sug- tasks in which images of racial outgroup members are
gested that activity in the mPFC and/or rostral ACC repeatedly paired with positive images and appetitive
was uniquely associated with behavioural control that responses or with counter-stereotypical concepts, in an
is guided by external social cues, whereas activity in effort to selectively target the affective or semantic mem-
the dorsal ACC was associated with internally con- ory systems underlying implicit prejudices and stereo-
trolled behaviour 124. In addition, ventral portions of the types, respectively 29,30,141. By considering the operations
mPFC are interconnected with the OFC and amygdala, of these different neural systems, researchers are gaining
and through these connections it may support the top- a better understanding of how and under what condi-
down modulation of emotional responses139. Hence, the tions different forms of bias are activated, expressed and
mPFC may support the regulation of intergroup affect, potentially extinguished.
such as threat or contempt, although this hypothesis Despite some success in reducing behavioural and
remains to be tested. Considered broadly, emerging evi- physiological expressions of implicit bias in the labo-
dence regarding mPFC function suggests that it has a ratory 29,30,141,142, most forms of implicit learning are
larger role in cognitive control than previously thought, resistant to extinction140,143. Implicit racial biases are
particularly in the context of regulating complex social particularly difficult to change in a cultural milieu that
responses. constantly reinforces racial prejudices and stereotypes
(for example, in mainstream media). Thus, although
A network for the regulation of prejudice and stereotyping. attempts to undo learned intergroup associations are
Self-regulation is critical for the adaptive expression laudable, such strategies may be ineffective for reducing
of social behaviour. This is especially true with regard the expression of bias in behaviour outside the labora-
to stereotyping and prejudice given the potential tory. Instead, interventions that enhance the cognitive
(unwanted) influence of implicit biases on behaviour. control of behaviour should be more effective. Such
The putative network of neural regions involved in the control-based strategies may not reduce prejudice in
regulation of intergroup responses includes ACC and the mind, but they can prevent its effect on potential
PFC regions that have been implicated in existing mod- victims. Over time, control-driven changes in behaviour
els of cognitive control, as well as additional regions that may become habitual, and prejudiced and stereotypical
facilitate control in social contexts (FIG. 5). Specifically, associations in the mind may weaken7,144. Neuroscience
although the dorsal ACC and lateral PFC may carry out models suggest that control-based interventions should
the domain-general functions of detecting conflict and focus on (at least) two separate processes: those for mon-
implementing top-down control, the mPFC and rostral itoring unwanted racially biased tendencies and those
ACC are important for guiding control that is based on involved in the top-down control of behaviour. Strategies
social cues, such as norms against expressing prejudice, to enhance ACC-mediated conflict-monitoring processes
and intergroup emotions. include interventions that increase people’s awareness of
the potential for bias, increase attention to specific cues
From brain to society indicating that control may be needed (for example, the
Research on the neural basis of prejudice occupies a appearance of an outgroup member in an interaction)
special position at the interface of the natural and social and increase the sensitivity of conflict monitoring systems
sciences, and as such it is uniquely situated to bring (for example, by activating cognitive conflict prior to an
neuroscientific advances to bear on real-world social intergroup response)145,146. For the effective control of
issues. To date, the neuroscientific analysis of prejudice behaviour, conflict monitoring processes must be paired
has advanced theories of how prejudices are formed, with top-down response plans. To this end, psychological
expressed and potentially controlled, and these can be research has shown that goal strategies that link a spe-
used to inform interventions aimed at reducing discrimi- cific cue (for example, ‘if I meet a black person’) with a
nation. For example, research linking implicit prejudice pre-planned response (for example, ‘I will ignore his or
and stereotyping to different neural substrates suggests her race’ or ‘I will respond more carefully’) are especially
effective at facilitating the control of implicit stereotypes effects of social factors on neural function, but the neu-
in behaviour 147. By helping to inspire and explicate strate- roscience of prejudice is a particularly rich topic as it
gies such as these, the neuroscience of prejudice is already considers the roles of personal attitudes and motivations,
beginning to inform policy and interventions aimed at social norms and social emotions as they relate to com-
reducing prejudice in society. plex interpersonal behaviours. If the human brain has
Beyond its implications for social issues regarding evolved to support survival and prosperity in a complex
intergroup relations, research on the neural basis of social environment, then a research approach that con-
prejudice provides a context for understanding neural siders this range of factors will be needed to truly under-
function as it relates to the real-world lives of human stand neural function. Research on the neural basis of
beings. Many areas of social neuroscience consider the prejudice is an important step in this direction.
1. Brewer, M. B. The psychology of prejudice: ingroup 23. Rolls, E. T. & Rolls, B. J. Altered food preferences after familiar black and white faces. J. Cogn. Neurosci. 26,
love and outgroup hate? J. Soc. Issues 55, 429–444 lesions in the basolateral region of the amygdala in the 1992–2004 (2014).
(1999). rat. J. Comp. Physiol. Psychol. 83, 248–259 (1973). 40. Olsson, A., Ebert, J. P., Banaji, M. R. & Phelps, E. A.
2. Ito, T. A. & Urland, G. R. Race and gender on the 24. Adolphs, R. Fear, faces, and the human amygdala. The role of social group in the persistence of learned
brain: electrocortical measures of attention to the race Curr. Opin. Neurobiol. 18, 166–172 (2008). fear. Science 309, 785–787 (2005).
and gender of multiply categorizable individuals. 25. Phelps, E. A. et al. Performance on indirect measures 41. Richeson, J. A. & Trawalter, S. The threat of appearing
J. Pers. Soc. Psychol. 85, 616–626 (2003). of race evaluation predicts amygdala activation. prejudiced and race-based attentional biases. Psychol.
3. Ratner, K. G. & Amodio, D. M. Seeing “us versus J. Cogn. Neurosci. 12, 729–738 (2000). Sci. 19, 98–102 (2008).
them”: minimal group effects on the neural encoding An early fMRI investigation of neural correlates of 42. Ofan, R. H., Rubin, N. & Amodio, D. M. Situation-
of faces. J. Exp. Soc. Psychol. 49, 298–301 (2013). implicit prejudice, focusing on the amygdala. based social anxiety enhances the neural processing
4. Allport, G. W. The Nature of Prejudice (Addison- Relative differences in amygdala activity in of faces: evidence from an intergroup context. Soc.
Wesley, 1954). response to black compared with white faces were Cogn. Affect. Neurosci. http://dx.doi.org/10.1093/
5. Fiske, S. T. in The Handbook of Social Psychology associated with a behavioural index of implicit scan/nst087 (2013).
Vol. 2, 4th edn (eds Gilbert, D. T., Fiske, S. T. & prejudice and startle responses to black versus 43. Plant, E. A. & Devine, P. G. Internal and external
Lindzey, G.) 357–411 (McGraw Hill, 1998). white faces. motivation to respond without prejudice. J. Pers. Soc.
6. Amodio, D. M. & Devine, P. G. Stereotyping and 26. Hart, A. J. et al. Differential response in the human Psychol. 75, 811–832 (1998).
evaluation in implicit race bias: evidence for amygdala to racial outgroup versus ingroup face 44. Stephan, W. G. & Stephan, C. W. Intergroup anxiety.
independent constructs and unique effects on stimuli. Neuroreport 11, 2351–2355 (2000). J. Soc. Issues 41, 157–175 (1985).
behavior. J. Pers. Soc. Psychol. 91, 652–661 (2006). 27. Amodio, D. M., Harmon-Jones, E. & Devine, P. G. 45. Lieberman, M. D., Hariri, A., Jarcho, J. M.,
7. Devine, P. G. Stereotypes and prejudice: their Individual differences in the activation and control of Eisenberger, N. I. & Bookheimer, S. Y. An fMRI
automatic and controlled components. J. Pers. Soc. affective race bias as assessed by startle eyeblink investigation of race-related amygdala activity in
Psychol. 56, 5–18 (1989). responses and self-report. J. Pers. Soc. Psychol. 84, African-American and Caucasian-American individuals.
8. Fazio, R. H., Jackson, J. R., Dunton, B. C. & 738–753 (2003). Nature Neurosci. 8, 720–722 (2005).
Williams, C. J. Variability in automatic activation as an The first reported difference in amygdala activity in 46. Wheeler, M. E. & Fiske, S. T. Controlling racial
unobtrusive measure of racial attitudes: a bona fide response to black compared with white (and Asian) prejudice: social-cognitive goals affect amygdala and
pipeline? J. Pers. Soc. Psychol. 69, 1013–1027 faces. By using the startle-eyeblink method to stereotype activation. Psychol. Sci. 16, 56–63 (2005).
(1995). assess amygdala activity related to the CeA, the 47. Van Bavel, J. J., Packer, D. J. & Cunningham, W. A.
9. Greenwald, A. G. & Banaji, M. R. Implicit social results link implicit prejudice more directly to a The neural substrates of in‑group bias: a functional
cognition: attitudes, self-esteem, and stereotypes. Pavlovian conditioning mechanism of learning and magnetic resonance imaging investigation. Psychol.
Psychol. Rev. 102, 4–27 (1995). behavioural expression. Sci. 11, 1131–1139 (2008).
10. Amodio, D. M. The social neuroscience of intergroup 28. Amodio, D. M. & Ratner, K. G. A memory systems This study compared the independent effects of race
relations. Eur. Rev. Soc. Psychol. 19, 1–54 (2008). model of implicit social cognition. Curr. Dir. Psychol. and coalition (team membership) on amygdala
11. Ito, T. A. & Bartholow, B. D. The neural correlates of Sci. 20, 143–148 (2011). responses to faces and showed that when team
race. Trends Cogn. Sci. 13, 524–531 (2009). 29. Olson, M. A. & Fazio, R. H. Reducing automatically- membership was salient, the amygdala responded to
In a field dominated by fMRI studies, this review activated racial prejudice through implicit evaluative coalition (subject’s team or other team) and not race
presents important research on ERP approaches to conditioning. Pers. Soc. Psychol. Bull. 32, 421–433 (black or white). This finding demonstrated that the
probing the neural underpinnings of sociocognitive (2006). amygdala response to race is not inevitable but rather
processes involved in prejudice and stereotyping. 30. Kawakami, K., Phills, C. E., Steele, J. R. & Dovidio, J. F. corresponds to the subject’s particular task goal.
12. Kubota, J. T., Banaji, M. R. & Phelps, E. A. The (Close) distance makes the heart grow fonder: 48. Richeson, J. A. et al. An fMRI investigation of the
neuroscience of race. Nature Neurosci. 15, 940–948 improving implicit racial attitudes and interracial impact of interracial contact on executive function.
(2012). interactions through approach behaviors. J. Pers. Soc. Nature Neurosci. 6, 1323–1328 (2003).
13. Amodio, D. M. Can neuroscience advance social Psychol. 92, 957–971 (2007). 49. Gilbert, S. J., Swencionis, J. K. & Amodio, D. M.
psychological theory? Social neuroscience for the 31. Amodio, D. M. & Hamilton, H. K. Intergroup anxiety Evaluative versus trait representation in intergroup
behavioral social psychologist. Soc. Cogn. 28, effects on implicit racial evaluation and social judgments: distinct roles of anterior temporal
695–716 (2010). stereotyping. Emotion 12, 1273–1280 (2012). lobe and prefrontal cortex. Neuropsychologia 50,
14. Poldrack, R. A. Can cognitive processes be inferred 32. Chekroud, A. M., Everett, J. A. C., Bridge, H. & 3600–3611 (2012).
from neuroimaging data? Trends Cogn. Sci. 10, 59–63 Hewstone, M. A review of neuroimaging studies of Using fMRI, the authors distinguished neural
(2006). race-related prejudice: does amygdala response reflect processes involved in stereotyping and prejudiced
15. Mackie, D. M. & Smith, E. R. in The Social Self: threat? Front. Hum. Neurosci. 8, 179 (2014). attitudes. Stereotype-based judgements of black
Cognitive, Interpersonal, and Intergroup Perspectives 33. Ronquillo, J. et al. The effects of skin tone on race- compared with white people uniquely involved the
(eds Forgas, J. P. & Williams, K. D.) 309–326 related amygdala activity: an fMRI investigation. Soc. mPFC; affect-based judgements uniquely involved
(Psychology Press, 2002). Cogn. Affect. Neurosci. 2, 39–44 (2007). the OFC.
16. Cottrell, C. A. & Neuberg, S. L. Different emotional 34. Richeson, J. A., Todd, A. R., Trawalter, S. & Baird, A. A. Eye- 50. Golby, A. J., Gabrieli, J. D. E., Chiao, J. Y. &
reactions to different groups: a sociofunctional threat- gaze direction modulates race-related amygdala activity. Eberhardt, J. L. Differential fusiform responses to
based approach to “prejudice”. J. Pers. Soc. Psychol. Group Process. Interg. Relat. 11, 233–246 (2008). same- and other-race faces. Nature Neurosci. 4,
88, 770–789 (2005). 35. Freeman, J. B., Schiller, D., Rule, N. O. & Ambady, N. 845–850 (2001).
17. Swanson, L. W. & Petrovich, G. D. What is the The neural origins of superficial and individuated The first demonstration of race effects in visual
amygdala? Trends Neurosci. 21, 323–331 (1998). judgments about ingroup and outgroup members. processing, using fMRI. Differences in fusiform
18. LeDoux, J. E. Emotion circuits in the brain. Annu. Rev. Hum. Brain Mapp. 31, 150–159 (2010). responses to black and white faces predicted the
Neurosci. 23, 155–184 (2000). 36. Forbes, C. E., Cox, C. L., Schmader, T. & Ryan, L. degree of ‘own race bias’ in memory in white subjects.
19. Davis, M. Neural systems involved in fear and anxiety Negative stereotype activation alters interaction 51. Knutson, K. M., Mah, L., Manly, C. F. & Grafman, J.
measured with fear-potentiated startle. Am. Psychol. between neural correlates of arousal, inhibition, and Neural correlates of automatic beliefs about gender
61, 741–756 (2006). cognitive control. Soc. Cogn. Affect. Neurosci. 7, and race. Hum. Brain Mapp. 28, 915–930 (2007).
20. Fendt, M. & Fanselow, M. S. The neuroanatomical and 771–781 (2012). 52. Brosch, T., Bar-David, E. & Phelps, E. A. Implicit race
neurochemical basis of conditioned fear. Neurosci. 37. Cunningham, W. A. et al. Separable neural bias decreases the similarity of neural representations
Biobehav. Rev. 23, 743–760 (1999). components in the processing of black and white of black and white faces. Psychol. Sci. 24, 160–166
21. LeDoux, J. E., Iwata, J., Cicchetti, P. & Reis, D. J. faces. Psychol. Sci. 15, 806–813 (2004). (2013).
Different projections of the central amygdaloid nucleus 38. Telzer, E. H., Humphreys, K., Shapiro, M. & 53. Schreiber, D. & Iacoboni, M. Huxtables on the brain:
mediate autonomic and behavioral correlates of Tottenham, N. L. Amygdala sensitivity to race is not an fMRI study of race and norm violation. Polit.
conditioned fear. J. Neurosci. 8, 2517–2529 (1988). present in childhood but emerges in adolescence. Psychol. 33, 313–330 (2012).
22. Holland, P. C. & Gallagher, M. Amygdala circuitry in J. Cogn. Neurosci. 25, 234–244 (2013). 54. Whalen, P. J. Fear, vigilance, and ambiguity: initial
attentional and representational processes. Trends 39. Cloutier, J., Li, T. & Correll, J. The impact of childhood neuroimaging studies of the human amygdala. Curr.
Cogn. Sci. 3, 65–73 (1999). experience on amygdala response to perceptually Dir. Psychol. Sci. 7, 177–188 (1998).
55. Said, C. P., Baron, S. & Todorov, A. Nonlinear 80. Frith, C. D. & Frith, U. Interacting minds—a biological 106. Buckner, R. L. & Tulving, E. in Handbook of
amygdala response to face trustworthiness: basis. Science 286, 1692–1695 (1999). Neuropsychology Vol. 10 (eds Boller, F. & Grafman, J.)
contributions of high and low spatial frequency 81. Cikara, M., Bruneau, E. G. & Saxe, R. Us and them: 439–466 (Elsevier, 1995).
information. J. Cogn. Neurosci. 21, 519–528 (2009). intergroup failures of empathy. Curr. Dir. Psychol. Sci. 107. Demb, J. B. et al. Semantic encoding and retrieval in
56. Cunningham, W. A., Van Bavel, J. J. & Johnsen, I. R. 20, 149–153 (2011). the left inferior prefrontal cortex: a functional MRI
Affective flexibility: evaluative processing goals shape 82. Cikara, M., Eberhardt, J. L. & Fiske, S. T. From agents study of task difficulty and process specificity.
amygdala activity. Psychol. Sci. 19, 152–160 to objects: sexist attitudes and neural responses to J. Neurosci. 15, 5870–5878 (1995).
(2008). sexualized targets. J. Cogn. Neurosci. 23, 540–551 108. Thompson-Schill, S. L. Neuroimaging studies of
57. Phelps, E. A., Cannistraci, C. J. & Cunningham, W. A. (2011). semantic memory: inferring “how” from “where”.
Intact performance on an indirect measure of race bias 83. Hamilton, D. L. & Sherman, J. W. in Handbook of Neuropsychologia 41, 280–292 (2003).
following amygdala damage. Neuropsychologia 41, Social Cognition Vol. 2, 2nd edn (eds Wyer, R. S. Jr 109. Miller, E. K., Freedman, D. J. & Wallis, J. D. The
203–208 (2003). & Srull, T. K.) 1–68 (Erlbaum, 1994). prefrontal cortex: categories, concepts and cognition.
58. Bechara, A., Damasio, H. & Damasio, A. R. Emotion, 84. Olson, I. R., McCoy, D., Klobusicky, E. & Ross, L. A. Phil. Trans. R. Soc. Lond. B 357, 1123–1136 (2002).
decision making and the orbitofrontal cortex. Cereb. Social cognition and the anterior temporal lobes: 110. Balleine, B. W., Delgado, M. R. & Hikosaka, O. The
Cortex 10, 295–307 (2000). a review and theoretical framework. Soc. Cogn. Affect. role of the dorsal striatum in reward and decision-
59. O’Doherty, J., Kringelbach, M. L., Rolls, E. T., Neurosci. 8, 123–133 (2013). making. J. Neurosci. 27, 8161–8165 (2007).
Hornak, J. & Andrews, C. Abstract reward and 85. Quadflieg, S. & Macrae, C. N. Stereotypes and 111. Mitchell, J. P., Ames, D. L., Jenkins, A. C. &
punishment representations in the human stereotyping: what’s the brain got to do with it? Eur. Banaji, M. R. Neural correlates of stereotype
orbitofrontal cortex. Nature Neurosci. 4, 95–102 Rev. Soc. Psychol. 22, 215–273 (2011). application. J. Cogn. Neurosci. 21, 594–604
(2001). A comprehensive review of neuroscience research (2009).
60. Rushworth, M. F. S., Noonan, M. P., Boorman, E. D., on social stereotyping processes. 112. Aron, A. R., Robbins, T. W. & Poldrack, R. A. Inhibition
Walton, M. E. & Behrens, T. E. Frontal cortex and 86. Gabrieli, J. D. Cognitive neuroscience of human and the right inferior frontal cortex. Trends Cogn. Sci.
reward-guided learning and decision-making. Neuron memory. Annu. Rev. Psychol. 49, 87–115 (1998). 8, 170–177 (2004).
70, 1054–1069 (2011). 87. Martin, A. The representation of object concepts in 113. Bodenhausen, G. V., Sheppard, L. & Kramer, G. P.
61. Beer, J. S., Heerey, E. H., Keltner, D., Scabini, D. & the brain. Annu. Rev. Psychol. 58, 25–45 (2007). Negative affect and social perception: the differential
Knight, R. T. The regulatory function of self-conscious 88. Patterson, K., Nestor, P. J. & Rogers, T. T. Where do impact of anger and sadness. Eur. J. Soc. Psychol. 24,
emotion: insights from patients with orbitofrontal you know what you know? The representation of 45–62 (1994).
damage. J. Pers. Soc. Psychol. 85, 594–604 (2003). semantic knowledge in the human brain. Nature Rev. 114. DeSteno, D., Dasgupta, N., Bartlett, M. Y. &
62. Amodio, D. M. & Frith, C. D. Meeting of minds: the Neurosci. 8, 976–987 (2007). Cajdric, A. Prejudice from thin air: the effect of
medial frontal cortex and social cognition. Nature Rev. 89. Quadflieg, S. et al. Exploring the neural correlates of emotion on automatic intergroup attitudes. Psychol.
Neurosci. 7, 268–277 (2006). social stereotyping. J. Cogn. Neurosci. 21, Sci. 15, 319–324 (2004).
63. Ongür, D. & Price, J. L. The organization of networks 1560–1570 (2009). 115. Choi, E. Y., Yeo, B. T. & Buckner, R. L. The organization
within the orbital and medial prefrontal cortex of rats, 90. Olson, I. R., Plotzker, A. & Ezzyat, Y. The enigmatic of the human striatum estimated by intrinsic
monkeys and humans. Cereb. Cortex 10, 206–219 temporal pole: a review of findings on social and functional connectivity. J. Neurophysiol. 108,
(2000). emotional processing. Brain 130, 1718–1731 2242–2263 (2012).
64. Beer, J. S. et al. The Quadruple Process model (2007). 116. Bush, G., Luu, P. & Posner, M. L. Cognitive and
approach to examining the neural underpinnings of 91. Zahn, R. et al. Social concepts are represented in the emotional influences in anterior cingulate cortex.
prejudice. Neuroimage 43, 775–783 (2008). superior anterior temporal cortex. Proc. Natl Acad. Trends Cogn. Sci. 4, 215–222 (2000).
65. Craig, A. D. How do you feel? Interoception: the sense Sci. USA 104, 6430–6435 (2007). 117. Mansouri, F. A., Tanaka, K. & Buckley, M. J. Conflict-
of the physiological condition of the body. Nature Rev. 92. de Schotten, M. T., Dell’Acqua, F., Valabregue, R. & induced behavioral adjustment: a clue to the executive
Neurosci. 3, 655–666 (2002). Catani, M. Monkey to human comparative anatomy of functions of the prefrontal cortex. Nature Rev.
66. Craig, A. D. How do you feel — now? The anterior the frontal lobe association tracts. Cortex 48, 82–96 Neurosci. 10, 141–152 (2009).
insula and human awareness. Nature Rev. Neurosci. (2012). 118. Gerhing, W. J., Goss, B., Coles, M. G. H., Meyer, D. E.
10, 59–70 (2009). 93. Contreras, J. M., Banaji, M. R. & Mitchell, J. P. & Donchin, E. A neural system for error detection and
67. Harris, L. T. & Fiske, S. T. Dehumanizing the lowest of Dissociable neural correlates of stereotypes and other compensation. Psychol. Sci. 4, 385–390 (1993).
the low: neuro-imaging responses to extreme forms of semantic knowledge. Soc. Cogn. Affect. 119. Botvinick, M., Nystrom, L., Fissell, K., Carter, C. &
outgroups. Psychol. Sci. 17, 847–853 (2006). Neurosci. 7, 764–770 (2012). Cohen, J. Conflict monitoring versus selection-for-
68. Singer, T. et al. Empathy for pain involves the affective An fMRI study on the neural underpinnings of action in anterior cingulate cortex. Nature 402,
but not sensory components of pain. Science 303, stereotyping that distinguishes the role of trait 179–181 (1999).
1157–1162 (2004). inference processes from that of object knowledge 120. Botvinick, M. M., Braver, T. S., Barch, D. M.,
69. Singer, T. et al. Empathic neural responses are representation. Carter, C. S. & Cohen, J. D. Conflict monitoring and
modulated by the perceived fairness of others. Nature 94. Gallate, J., Wong, C., Ellwood, S., Chi, R. & Snyder, A. cognitive control. Psychol. Rev. 108, 624–652
439, 466–469 (2006). Noninvasive brain stimulation reduces prejudice (2001).
70. Lamm, C., Meltzoff, A. N. & Decety, J. How do we scores on an implicit association test. 121. Shackman, A. J. et al. The integration of negative
empathize with someone who is not like us? Neuropsychology 25, 185–192 (2011). affect, pain and cognitive control in the cingulate
A functional magnetic resonance imaging study. 95. Milne, E. & Grafman, J. Ventromedial prefrontal cortex cortex. Nature Rev. Neurosci. 12, 154–167 (2011).
J. Cogn. Neurosci. 22, 362–376 (2010). lesions in humans eliminate implicit gender 122. Amodio, D. M. & Devine, P. G. in Self Control in
71. Xu, X., Zuo, X., Wang, X. & Han, S. Do you feel my pain? stereotyping. J. Neurosci. 21, 1–6 (2001). Society, Mind and Brain (eds Hassin, R. R., Ochsner,
Racial group membership modulates empathic neural 96. Mitchell, J. P., Macrae, C. N. & Banaji, M. R. K. N. & Trope, Y.). 49–75 (Oxford Univ. Press, 2010).
responses. J. Neurosci. 29, 8525–8529 (2009). Dissociable medial prefrontal contributions to 123. Amodio, D. M. et al. Neural signals for the detection
In a demonstration of racial bias in empathy, mPFC judgments of similar and dissimilar others. Neuron 50, of unintentional race bias. Psychol. Sci. 15, 88–93
and ACC activity linked to empathy was observed 655–663 (2006). (2004).
only when subjects viewed members of their own 97. Quadflieg, S. et al. Stereotype-based modulation of This study demonstrated the role of the ACC in the
racial group being exposed to painful stimuli. person perception. Neuroimage 57, 549–557 control of racial bias. Using ERPs, the authors
72. Cikara, M. & Fiske, S. T. Bounded empathy: neural (2011). found an increase ACC activity to stereotype-based
responses to outgroup targets’ (mis)fortunes. J. Cogn. 98. Forbes, C. E. et al. Identifying temporal and causal conflict that predicted enhanced control of bias in
Neurosci. 23, 3791–3803 (2011). contributions of neural processes underlying the behaviour.
73. Alexander, G. E., DeLong, M. R. & Strick, P. L. Parallel Implicit Association Test. (IAT). Front. Hum. Neurosci. 124. Amodio, D. M., Kubota, J. T., Harmon-Jones, E. &
organization of functionally segregated circuits linking 6, 320 (2012). Devine, P. G. Alternative mechanisms for regulating
basal ganglia and cortex. Annu. Rev. Neurosci. 9, 99. Mitchell, J. P. Social psychology as a natural kind. racial responses according to internal versus external
357–381 (1986). Trends Cogn. Sci. 13, 246–251 (2009). cues. Soc. Cogn. Affect. Neurosci. 1, 26–36 (2006).
74. Knutson, B., Adams, C. S., Fong, G. W. & Hommer, D. 100. Gilbert, S. J. et al. Distinct regions of medial rostral 125. Amodio, D. M., Devine, P. G. & Harmon-Jones, E.
Anticipation of monetary reward selectively recruits prefrontal cortex supporting social and nonsocial Individual differences in the regulation of intergroup
nucleus accumbens. J. Neurosci. 21, RC159 (2001). functions. Soc. Cogn. Affect. Neurosci. 2, 217–226 bias: the role of conflict monitoring and neural
75. O’Doherty, J. et al. Dissociable roles of ventral and (2007). signals for control. J. Pers. Soc. Psychol. 94, 60–74
dorsal striatum in instrumental conditioning. Science 101. Baetens, K., Ma, N., Steen, J. & van Overwalle, F. (2008).
304, 452–454 (2004). Involvement of the mentalizing network in social and 126. Bartholow, B. D., Dickter, C. L. & Sestir, M. A.
76. Stanley, D. A. et al. Race and reputation: perceived non-social high construal. Soc. Cogn. Affect. Neurosci. Stereotype activation and control of race bias:
racial group trustworthiness influences the neural 9, 817–824 (2014). cognitive control of inhibition and its impairment
correlates of trust decisions. Phil. Trans. R. Soc. B 102. Saxe, R. Uniquely human social cognition. Curr. Opin. by alcohol. J. Pers. Soc. Psychol. 90, 272–287
367, 744–753 (2012). Neurobiol. 16, 235–239 (2006). (2006).
77. Mitchell, J. P., Heatherton, T. F. & Macrae, C. N. 103. Frith, C. D. & Frith, U. Implicit and explicit processes 127. Correll, J., Urland, G. R. & Ito, T. A. Event-related
Distinct neural systems subserve person and object in social cognition. Neuron 60, 503–510 (2008). potentials and the decision to shoot: the role of threat
knowledge. Proc. Natl Acad. Sci. USA 99, 104. Uddin, L. Q., Iacoboni, M., Lange, C. & Keenan, J. P. perception and cognitive control. J. Exp. Soc. Psychol.
15238–15243 (2002). The self and social cognition: the role of cortical 42, 120–128 (2006).
78. Van Overwalle, F. Social cognition and the brain: a midline structures and mirror neurons. Trends Cogn. 128. Gonsalkorale, K., Sherman, J. W., Allen, T. J.,
meta-analysis. Hum. Brain Mapp. 30, 829–858 Sci. 11, 153–157 (2007). Klauer, K. C. & Amodio, D. M. Accounting for
(2009). 105. Blaxton, T. A. et al. Functional mapping of human successful control of implicit racial bias: the roles of
79. Krueger, F., Barbey, A. K. & Grafman, J. The medial memory using PET: comparisons of conceptual and association activation, response monitoring, and
prefrontal cortex mediates social event knowledge. perceptual tasks. Can. J. Exp. Psychol. 50, 42–56 overcoming bias. Pers. Soc. Psychol. Bull. 37,
Trends Cogn. Sci. 13, 103–109 (2009). (1996). 1534–1545 (2011).
129. Fourie, M. M., Thomas, K. G. F., Amodio, D. M., training on the negation of stereotypic associations remembering others: an electro-physiological study.
Warton, C. M. R. & Meintjes, E. M. Neural correlates on stereotype activation. J. Pers. Soc. Psychol. 78, Brain Res. 1388, 123–133 (2011).
of experienced moral emotion: an fMRI investigation 871–888 (2000). 154. Caldara, R., Rossion, B., Bovet, P. & Hauert, C. A.
of emotion in response to prejudice feedback. Soc. 142. Mallan, K. M., Sax, J. & Lipp, O. V. Verbal Event-related potentials and time course of the ‘other-
Neurosci. 9, 203–218 (2014). instruction abolishes fear conditioned to racial out- race’ face classification advantage. Neuroreport 15,
130. Van Nunspeet, F., Ellemers, N., Derks, B. & group faces. J. Exp. Soc. Psychol. 45, 1303–1307 905–910 (2004).
Nieuwenhuis, S. Moral concerns increase attention (2009). 155. Caldara, R. et al. Face versus non-face object
and response monitoring during IAT performance: ERP 143. Bouton, M. E. Conditioning, remembering, and perception and the ‘other-race’ effect: a
evidence. Soc. Cogn. Affect. Neurosci. 9, 141–149 forgetting. J. Exp. Psychol. Anim. B 20, 219–231 spatiotemporal event-related potential study. Clin.
(2014). (1994). Neurophysiol. 114, 515–528 (2003).
131. Fuster, J. M. The prefrontal cortex—an update: time is 144. Devine, P. G., Plant, E. A., Amodio, D. M., Harmon- 156. Ito, T. A. & Urland, G. R. The influence of processing
of the essence. Neuron 2, 319–333 (2001). Jones, E. & Vance, S. L. The regulation of explicit and objectives on the perception of faces: an ERP study of
132. Badre, D. & D’Esposito, M. Is the rostro–caudal axis implicit racial bias: the role of motivations to respond race and gender perception. Cogn. Affect. Behav.
of the frontal lobe hierarchical? Nature Rev. Neurosci. without prejudice. J. Pers. Soc. Psychol. 82, 835–848 Neurosci. 5, 21–36 (2005).
10, 659–669 (2009). (2002). 157. Walker, P. M., Silvert, L., Hewstone, M. & Nobre, A. C.
133. Koechlin, E., Ody, C. & Kouneiher, F. The architecture 145. Kleiman, T., Hassin, R. R. & Trope, Y. The control-freak Social contact and other-race face processing in the
of cognitive control in the human prefrontal cortex. mind: stereotypical biases are eliminated following human brain. Soc. Cogn. Affect. Neurosci. 3, 16–25
Science 302, 1181–1185 (2003). conflict-activated cognitive control. J. Exp. Psychol. (2008).
134. Aron, A. R. The neural basis of inhibition in cognitive Gen. 143, 498–503 (2014). 158. Wiese, H., Stahl, J. & Schweinberger, S. R. Configural
control. Neuroscientist 13, 214–228 (2007). 146. Monteith, M. J., Ashburn-Nardo, L., Voils, C. I. & processing of other-race faces is delayed but not
135. Ghashghaei, H. T., Hilgetag, C. C. & Barbas, H. Czopp, A. M. Putting the brakes on prejudice: on the decreased. Biol. Psychol. 81, 103–109 (2009).
Sequence of information processing for emotions development and operation of cues for control. J. Pers. 159. Contreras, J. M., Banaji, M. R. & Mitchell, J. P.
based on the anatomic dialogue between prefrontal Soc. Psychol. 83, 1029–1050 (2002). Multivoxel patterns in fusiform face area differentiate
cortex and amygdala. Neuroimage 34, 905–923 147. Mendoza, S. A., Gollwitzer, P. M. & Amodio, D. M. faces by sex and race. PLoS ONE 8, e69684 (2013).
(2007). Reducing the expression of implicit stereotypes: 160. Ratner, K. G., Kaul, C. & Van Bavel, J. J. Is race
136. Amodio, D. M. Coordinated roles of motivation and reflexive control through implementation intentions. erased? Decoding race from patterns of neural
perception in the regulation of intergroup responses: Pers. Soc. Psychol. Bull. 36, 512–523 (2010). activity when skin color is not diagnostic of group
frontal cortical asymmetry effects on the P2 event- 148. Bar, M. et al. Top-down facilitation of visual boundaries. Soc. Cogn. Affect. Neurosci. 8, 750–755
related potential and behavior. J. Cogn. Neurosci. 22, recognition. Proc. Natl Acad. Sci. USA 103, 449–454 (2013).
2609–2617 (2010). (2006). 161. Greenwald, A. G., Poehlman, T. A., Uhlmann, E. &
This study demonstrated the role of the PFC in the 149. Van Bavel, J. J., Packer, D. J. & Cunningham, W. A. Banaji, M. R. Understanding and using the Implicit
behavioural control of racial bias and showed that Modulation of the fusiform face area following Association Test: III. meta-analysis of predictive
this effect involves changes in early perceptual minimal exposure to motivationally relevant faces: validity. J. Pers. Soc. Psychol. 97, 17–41 (2009).
attention to racial cues. The study provided initial evidence of in‑group enhancement (not out-group 162. Payne, B. K. Prejudice and perception: the role of
evidence for a ‘motivated perception’ account of disregard). J. Cogn. Neurosci. 23, 3343–3354 automatic and controlled processes in misperceiving
self-regulation. (2011). a weapon. J. Pers. Soc. Psychol. 81, 181–192
137. Amodio, D. M., Devine, P. G. & Harmon-Jones, E. 150. Ratner, K. G., Dotsch, R., Wigboldus, D., (2001).
A dynamic model of guilt: implications for motivation van Knippenberg, A. & Amodio, D. M. Visualizing 163. Greenwald, A. G., McGhee, D. E. & Schwartz, J. K. L.
and self-regulation in the context of prejudice. minimal ingroup and outgroup faces: implications Measuring individual differences in implicit cognition:
Psychol. Sci. 18, 524–530 (2007). for impressions, attitudes, and behavior. J. Pers. the Implicit Association Test. J. Pers. Soc. Psychol. 74,
138. Gozzi, M., Raymont, V., Solomon, J., Koenigs, M. & Soc. Psychol. 106, 897–911 (2014). 1464–1480 (1998).
Grafman, J. Dissociable effects of prefrontal and 151. Krosch, A. K. & Amodio, D. M. Economic scarcity
anterior temporal cortical lesions on stereotypical alters the perception of race. Proc. Natl Acad. Sci. USA
Acknowledgements
gender attitudes. Neuropsychologia 47, 2125–2132 111, 9079–9084 (2014).
Work on this article was supported by a National Science
(2009). 152. Ofan, R. H., Rubin, N. & Amodio, D. M. Seeing race:
Foundation grant (BCS 0847350). The author thanks mem-
139. Milad, M. R. & Quirk, G. J. Neurons in medial N170 responses to race and their relation to
bers of the NYU Social Neuroscience Laboratory, J. Freeman,
prefrontal cortex signal memory for fear extinction. automatic racial attitudes and controlled
K. Ratner and the three anonymous reviewers for their helpful
Nature 420, 70–74 (2002). processing. J. Cogn. Neurosci. 23, 3152–3161
feedback on earlier versions of this article.
140. Sloman, S. A. The empirical case for two systems of (2011).
reasoning. Psychol. Bull. 119, 3–22 (1996). 153. Brebner, J. L., Krigolson, O., Handy, T. C.,
141. Kawakami, K., Dovidio, J. F., Moll, J., Hermsen, S. & Quadflieg, S. & Turk, D. J. The importance of skin Competing interests statement
Russin, A. Just say no (to stereotyping): effects of color and facial structure in perceiving and The author declares no competing interests.