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The Neuroscience of Prejudice and Stereotyping

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Abstract

Despite global increases in diversity, social prejudices continue to fuel intergroup conflict, disparities and discrimination. Moreover, as norms have become more egalitarian, prejudices seem to have 'gone underground', operating covertly and often unconsciously, such that they are difficult to detect and control. Neuroscientists have recently begun to probe the neural basis of prejudice and stereotyping in an effort to identify the processes through which these biases form, influence behaviour and are regulated. This research aims to elucidate basic mechanisms of the social brain while advancing our understanding of intergroup bias in social behaviour.
Social motivations, such as the desire to affiliate or com-
pete with others, rank among the most potent of human
drives1. Not surprisingly, the capacity to discern ‘us’ from
‘them’ is fundamental in the human brain. Although this
computation takes just a fraction of a second2,3, it sets the
stage for social categorization, stereotypes, prejudices, inter-
group conflict and inequality, and, at the extremes, war
and genocide. Thus, although prejudice stems from a
mechanism of survival, built on cognitive systems that
‘structure’ the physical world, its function in modern
society is complex and its effects are often deleterious.
For the neuroscientist, the domain of prejudice pro-
vides a unique context for examining neural mechanisms
of the human mind that guide complex behaviour. Social
prejudices are scaffolded by basic-level neurocognitive
structures, but their expression is guided by personal
goals and normative expectations, played out in dyadic
and intergroup settings; this is truly the human brain
invivo . Although probing the neural basis of prejudice
is a challenging endeavour — in which the rigours of
reductionism are balanced with the richness of context
— it offers neuroscientists the opportunity to connect
their knowledge to some of society’s most pressing prob-
lems, such as discrimination, intergroup conflict and
disparities in health and socioeconomicstatus.
In this article, I review research on the role of the brain
in social prejudice and stereotyping. The term prejudice is
used broadly to refer to preconceptions — often negative
— about groups or individuals based on their social, racial
or ethnic affiliations4. Within the field of social psychol-
ogy, prejudice refers more specifically to evaluations (that
is, attitudes) and emotional responses towards a group and
its members. Stereotypes, by comparison, are generalized
characteristics ascribed to a social group, such as personal
traits (for example, unintelligent) or circumstantial attrib-
utes (for example, poor)5. Although they are distinguish-
able by content and process, prejudices and stereotypes
often operate in combination to influence social behav-
iour6. Moreover, both forms of bias can operate implicitly,
such that they may be activated and influence judgements
and behaviours without conscious awareness7–9.
Despite the persistence of prejudices and stereotypes
in contemporary society, their effects on behaviour are
often countered by peoples egalitarian personal beliefs
and pro-social norms7. Guided by these beliefs and
norms, people frequently engage self-regulatory pro-
cesses to mitigate the effects of bias on their behaviour.
Hence, a theoretical analysis of prejudice and stereo-
typing is incomplete without a consideration of these
regulatory processes. Here, self-regulation refers to
the process of acting in an intentional manner, often
through mechanisms of cognitive control.
The neuroscientific research conducted on prejudice
and stereotyping over the past decade suggests that these
complex forms of human behaviour involve different
interacting networks of neural structures. In this article, I
describe the functions of key structures in each network,
including both their broader neurocognitive functions
and their specific roles in prejudice and stereotyping.
This article extends previous reviews on this topic —
which were guided by a social psychological analysis10
or emphasized a particular neuroimaging method11,12
by providing a comprehensive overview of the literature
from a neural-systems perspective. Although many of
the conclusions drawn from this emerging literature rely
heavily on reverse inference from neuroimaging data,
Social motivations
Motives that operate in social
contexts and satisfy basic,
often universal, goals and
aspirations, such as to affiliate
(for example, form relationships
and communities) or to achieve
dominance (for example, within
a social hierarchy).
Stereotypes
Conceptual attributes
associated with a group and its
members (often through over-
generalization), which may
refer to trait or circumstantial
characteristics.
Prejudices
Evaluations of or affective
responses towards a social
group and its members based
on preconceptions.
The neuroscience of prejudice and
stereotyping
David M.Amodio
Abstract | Despite global increases in diversity, social prejudices continue to fuel intergroup
conflict, disparities and discrimination. Moreover, as norms have become more egalitarian,
prejudices seem to have ‘gone underground’, operating covertly and often unconsciously,
such that they are difficult to detect and control. Neuroscientists have recently begun to
probe the neural basis of prejudice and stereotyping in an effort to identify the processes
through which these biases form, influence behaviour and are regulated. This research aims
to elucidate basic mechanisms of the social brain while advancing our understanding of
intergroup bias in social behaviour.
New York University,
Department of Psychology,
6 Washington Place,
New York, New York 10003,
USA.
e-mail: david.amodio@nyu.
edu
doi:10.1038/nrn3800
Published online
4 September 2014
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a
0
–7.5
L R
Amplitude (µV)
4
2
0
–2
–4
–6
–200 0 200 400
Time (ms)
Ingroup
Outgroup
N170Face onset
b
μV
Self-regulation
The process of responding in
an intentional manner, often
involving the inhibition or
overriding of an alternative
response tendency.
these inferences are strengthened by converging theory
and behavioural data from the extensive psychological
literature on intergroup bias and self-regulation13,14.
The majority of the research reviewed here concerns
racial prejudice — a form of prejudice with clearly defined
social categories linked to identifiable physical attrib-
utes (BOX1). In particular, prejudice of white Americans
towards black people (that is, individuals of African or
Caribbean descent) has deep historical roots and contem-
porary relevance to social issues, and the majority of stud-
ies have examined prejudice in this context. Nevertheless,
many findings in this literature concern basic mechanisms
of social cognition that, to varying extents, underlie other
forms of bias, such as those based on ethnicity, gender,
sexual preference and nationality.
Neural basis of prejudice
In the modern social psychology literature, prejudice
is defined as an attitude towards a person on the basis
of his or her group membership. Prejudice may reflect
preference towards ingroup members or dislike of out-
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-
ral basis of prejudice has primarily focused on neural
structures involved in emotion and motivation, such as
the amygdala, insula, striatum and regions of orbital and
ventromedial frontal cortices (FIG.1). Although they are
often examined independently, these structures appear
to form a core network for the experience and expression
of prejudice.
Amygdala. Research on the neural basis of prejudice
has most frequently examined the amygdala, a complex
subcortical structure located bilaterally in the medial
temporal lobes (FIG.1). Although the amygdala is some-
times described as a neural locus of emotion (for exam-
ple, fear), it in fact comprises approximately 13 distinct
nuclei that, in conjunction, perform multiple functions
to support adaptive behaviour17 (FIG.2).
The amygdala receives direct (or nearly direct) affer-
ents from all sensory organs into its lateral nucleus, ena-
bling it to respond very rapidly to immediate threats in
advance of more elaborative processing of a stimulus18.
Within the amygdala, the central nucleus (CeA) has
been implicated in Pavlovian (classical) fear condition-
ing in both rats and humans19–22, and signals emerging
from the CeA activate hypothalamic and brainstem
structures to induce arousal, attention, freezing and
preparation for fight or flight — a response that is often
characterized as ‘fear’. By comparison, output from the
basal nucleus guides appetitive and instrumental responses
via projections to the ventral striatum22,23. Both the fear-
related and appetitive functions of the amygdala involve
motivation and attention, but to different ends, and they
probably correspond to different aspects of a prejudice-
based response. In humans, the amygdala is integral
to the processing of fear in facial expressions as well as
other salient social cues24. Given the amygdalas ability
to respond rapidly to potential social threat, researchers
interested in the neural substrate of implicit prejudice
first looked to this brain structure.
A pair of early functional MRI (fMRI) studies exam-
ined the amygdala activity of white research subjects in
response to blocked presentations of black and white
faces25,26. Although neither study found that amygdala
activity to faces varied significantly as a function of ‘race,
their results were suggestive: one study showed that the
relative difference in subjects’ amygdala activity to black
versus white faces was correlated with a behavioural
indicator of implicit prejudice (BOX2) and with rela-
tive differences in the startle eyeblink response to black
Box 1 | Seeing race: the role of visual perception
Social interactions often begin with the perception of a face. Mounting evidence reveals
that social motivations can alter the way a face is seen, which presumably reflects the
modulatory influences of signals from the temporal cortex, prefrontal cortex and
amygdala to the fusiform gyrus148. This insight suggests that prejudices and stereotypes
may alter early face processing.
Early functional MRI (fMRI) research demonstrated greater fusiform activity (see the
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.
Research examining the N170 component of the event-related potential (ERP), which
indexes the degree of initial configural face encoding at just ~170 ms, revealed enhanced
processing of ingroup versus outgroup faces (see the figure, panel b), even when groups
were defined arbitrarily3. This finding is consistent with fMRI data showing that faces of
‘coalition members’ elicited greater activity in the fusiform gyrus than did other faces,
regardless of race149. Hence, social group membership, even when defined on the basis of
minimal categories, promotes greater visual encoding. These findings dovetail with
behavioural research showing that biased visual representations of outgroup members
facilitate discriminatory actions towards them150,151.
In the context of race, outgroup members are often viewed as threatening and
therefore may elicit vigilant attention. Indeed, larger N170 ERP amplitudes in response to
viewing black versus white faces (equated in luminance) have been observed in subjects
with stronger implicit prejudice152 and in subjects who were made to feel anxious about
appearing biased42. These and other findings suggest that the visual processing of race is
malleable and depends on social motivations and contexts153–158.
Neural representations of race (black versus white), as determined by multivoxel
pattern analysis (MVPA), have been observed in the fusiform gyrus, and these neural
representations have been associated with behavioural indices of implicit prejudice
and stereotyping49,52,159,160. It is notable that MVPA has also identified race
representation in the medial occipital cortex; however, because full-colour photos
were used in these studies, the effect may reflect differences in luminance associated
with skin tone rather than the race of the people depicted. Nevertheless, the broader
body of findings suggests that social category cues modulate the early visual
processing of ingroup and outgroup members’ faces in ways that support the
perceivers’ biased or egalitarian social goals.
Panel a of the figure is from REF.50, Nature Publishing Group. Panel b of the figure is
reprinted from J.Exp. Soc. Psychol., 49, Ratner,K.G. & Amodio,D.M., Seeing “us versus
them”: minimal group effects on the neural encoding of faces, 298–301, Copyright (2013),
with permission from Elsevier.
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Amygdala
Early threat or
reward processing Striatum
Instrumental approach response
Insula
Visceral subjective emotion
Ventral mPFC
Empathy and mentalizing
OFC
Affective judgements
Configural face encoding
The visual encoding of a face in
terms of its basic structural
characteristics (for example,
the eyes, nose, mouth and the
relative distances between
these elements). Configural
encoding may be contrasted
with featural encoding, which
refers to the encoding of
feature characteristics that
make an individual’s face
unique.
Instrumental responses
Actions performed to achieve a
desired outcome (that is,
goal-directed responses).
versus white faces25. The other study, which also included
black subjects26, showed that amygdala responses habitu-
ated more slowly to racial outgroup faces. Together, these
studies identified the amygdala as a candidate substrate
of implicit prejudice.
To examine the timing and function of the amygdala
response to race more precisely, a later study used the
startle eyeblink method to index CeA-dependent amyg-
dala activity at very brief intervals following the presen-
tation of a face image27. This study revealed significantly
greater startle activity in response to black faces relative to
white or Asian faces — an effect that varied with subjects
self-reported motivations to respond without prejudice.
By demonstrating this differential response to race with
an index associated with CeA activity, this study more
directly implicated fear conditioning as a mechanism
underlying implicit prejudice. This link suggested that
the extensive literature on fear conditioning can, to some
extent, inform our understanding of implicit prejudice,
specifically regarding how this form of bias may be
learned, expressed and potentially extinguished28–31.
The role of the amygdala in implicit prejudice has
been examined in many subsequent studies. Much of
this research suggests that amygdala activation reflects
an immediate (or implied) threat response to racial
outgroup members10,12,32. For example, in white sub-
jects viewing images of black faces, amygdala activa-
tion is greater in response to faces with darker rather
than lighter skin tone33; when the eyegaze of the target
face is direct rather than averted34; when judgements of
faces are made on the basis of superficial information35;
and in contexts evoking interracial threat36. Moreover,
some evidence suggests that the amygdala response is
stronger when ingroup and outgroup faces are presented
very briefly, presumably because the brief presentation
precludes the regulation of this response37. By contrast,
familiarity with racial outgroup members is associated
with an attenuated difference in the amygdala response
to outgroup versus ingroup faces, both in children and
adults25,38–40. Together, these findings corroborate social
psychology theories of implicit prejudice as reflecting a
form of threat processing and suggest new links between
implicit prejudice, Pavlovian fear conditioning and
affective processes.
It is also possible that the amygdala response in some
studies reflects not a direct threat from an outgroup
member but rather the threat of appearing prejudiced
in the presence of others who may disapprove of bias.
Indeed, in white subjects, anxiety about appearing prej-
udiced to others has been shown to enhance eyegaze fix-
ations and early visual processing of black faces41,42, and
low-prejudice individuals who worried about appear-
ing prejudiced to others showed larger startle eyeblink
responses to black versus white faces compared with
low-prejudice individuals without this concern27. This
possibility — that amygdala activity in response to racial
outgroups is due to the threat of appearing prejudiced to
others — is consistent with findings from many social
psychology studies43,44 but has not been tested directly.
More recent studies have emphasized that the
amygdala response to an ingroup or outgroup mem-
ber depends on a perceiver’s goals: when exposure to
images of people from a different racial group is com-
bined with an unrelated secondary task (for example, to
detect the appearance of a small dot on the image), race
no longer drives the amygdala response45,46. In fact, in a
study in which the subject’s goal was to identify white
and black individuals in terms of coalition (for example,
whether each individual belonged to ones own sports
team, irrespective of race), it was coalition, and not
race, that drove the amygdala response47. Specifically,
amygdala activity was highest in response to the sub-
ject’s own team members. Still other studies have found
no differences in amygdala activity in response to dif-
ferent racial groups, presumably because the study
designs focused subjects’ attention on task features
other than race48–53. Although these findings may seem
to contradict other research linking amygdala activity
to threat, they are consistent with a broader model of
amygdala function, which proposes that it responds to
motivationally relevant cues — aversive or rewarding
— to guide adaptive behaviours22,23,54–56.
To date, the research literature suggests that there
are three main patterns of amygdala function with
respect to intergroup responses. One pattern reflects a
learned threat response to racial outgroups, which is
ostensibly rooted in fear conditioning. A second, but
still speculative, pattern may reflect the threat expe-
rienced by a perceiver who worries about appearing
prejudiced in the eyes of others when viewing faces of
racial outgroup members. Both of these patterns prob-
ably represent activity of the CeA, given its known
role in fear conditioning and anxiety. A third pattern
seems to reflect instrumental (that is, goal-directed)
responses, suggesting approach-related motivation
and attention towards members of the ingroup (which
can be based on race or other social categories). This
instrumental response probably reflects output from the
basal nucleus, given the involvement of this nucleus in
Figure 1 | Prejudice network. An interactive set of neural structures that underlie
components of a prejudiced response. The amygdala is involved in the rapid processing
of social category cues, including racial groups, in terms of potential threat or reward.
Approach-related instrumental responses are mediated by the striatum. The insula
supports visceral and subjective emotional responses towards social ingroups or
outgroups. Affect-driven judgements of social outgroup members rely on the orbital
frontal cortex (OFC) and may be characterized by reduced activity in the ventral medial
prefrontal cortex (mPFC), a region involved in empathy and mentalizing.
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LA
BA
CeA
ITC
Sensory input
Visual
Auditory
Somatosensory
Olfactory
Gustatory
Ventral striatum
(instrumental actions)
Hypothalamus and
brainstem (SNS, hormones)
Neuromodulatory systems
(arousal)
PFC (regulation)
PAG (freezing)
Implicit bias
Prejudiced or
stereotype-based perceptions
or responses that operate
without conscious awareness.
Deliberative judgements
Judgements that result from
thoughtful considerations
(often involving cognitive
control) as opposed to rapid,
gut-level, ‘snap’ judgements.
goal-directed behaviour. Together, these findings iden-
tify the amygdala as a major substrate of different forms
of implicit prejudice. However, it is important to note
that behavioural expressions of bias, such as in social
interactions or on a laboratory task (for example, the
implicit association test (IAT)), may reflect other pro-
cesses — such as conceptual associations, intentions and
cognitive control — in addition to an amygdala-based
response57. As the contributions of different amyg-
dala nuclei become better understood, and with more
refined behavioural assessments of implicit bias, the role
of the amygdala in prejudice and other social processes
will become increasinglyclear.
Orbital frontal cortex. The orbital frontal cortex (OFC)
(FIG.1), which is often considered to include the inferior
ventral medial prefrontal cortex (mPFC), is associated
with the processing of affective cues, contingency-based
learning, evaluation and decision making58–60. In the social
domain, the OFC supports the monitoring of social cues
and subsequent adjustment of ones behaviour61. This
function is crucial in intergroup situations involving
social norms, in which responses may be influenced by
others’ expectations62. Moreover, the OFC is anatomi-
cally interconnected with brain regions involved in all
sensory modalities and with structures that are known
to represent emotional and reward processes (such as
the basal nuclei of the amygdala and striatum) and social
knowledge (such as the medial frontal cortex and tempo-
ral poles)63. In comparison with the amygdala, the OFC
seems to support more complex and flexible evaluative
representations that are more directly applicable to the
intricacies of social behaviour.
To date, relatively few studies have examined the role
of the OFC in prejudice, most likely because the field
has primarily focused on comparatively basic responses
to racial group members (for example, through passive
viewing) rather than the kind of complex evaluative
processes that are known to involve the OFC. However,
findings from these studies are generally consistent
with the OFC’s proposed role in complex evaluations of
people based on group membership, beyond its poten-
tial role in implicit racial attitudes64. For example, OFC
activity has been associated with subjects’ deliberative
judgements regarding the prospect of befriending black,
relative to white, individuals49 (FIG.3). OFC activity has
also been associated with subjects’ preference for mem-
bers of their own team independently of race, indicating
that the OFC may have a broader role in group-based
evaluation47. Given its role in the regulation of social
behaviour61, the OFC is likely to emerge as an impor-
tant substrate of more elaborated forms of intergroup
evaluation.
Insula. The insula (FIG.1a) is a large cortical region
that runs medial to the temporal lobes, adjacent to
the frontal cortex, and broadly functions to represent
somatosensory states (including visceral responses)
and emotions related to such states (such as disgust)65.
Posterior insula regions are thought to provide primary
representation of interoceptive signals, whereas ante-
rior regions support the cognitive re-representation
of these signals. This re-representation in the anterior
insula provides an interface with the anterior cingulate
cortex (ACC) and PFC, which are involved in subjec-
tive awareness of emotion and cognitive control66. It
is the anterior insula, rather than the posterior insula,
that is most frequently associated with aspects of social
cognition and social emotion.
Although the insula is rarely of focal interest in neu-
roimaging studies of prejudice, its activity is frequently
associated with responses to racial outgroup versus
ingroup members in experimental tasks33,45,48. This find-
ing has been interpreted as reflecting a negative visceral
reaction, such as disgust, to racial outgroups67, and it has
been specifically associated with white subjects’ implicit
negative attitudes towards black people51,64. Thus, the
insula seems to contribute to the subjective affect that
is often experienced as part of a prejudiced response. It
could be speculated that the representation of this affec-
tive response in the anterior insula may — through its
connections with the ACC and PFC — facilitate the abil-
ity to detect and regulate ones behaviour on the basis of
a prejudicial affective response.
It is notable that the insula is also implicated in pro-
social emotions, such as empathy, towards liked indi-
viduals68–70. For example, insula activity was found to
increase when subjects viewed another person being
exposed to a painful stimulus, but only if that person
was of the same racial group71. Similarly, another study
observed insula activity when members of liked, but not
disliked, outgroups were harmed67. Both findings sug-
gest that empathy-related activity in the insula depends
on the victim’s social affiliation. In an interesting twist,
Figure 2 | The amygdala and its role in prejudice. Amygdala activity is
frequently observed in individuals while they view members of racial outgroups,
but it has also been found in response to viewing members of one’s own group
independently of race47. This mixed finding may reflect the different functions of
nuclei within the amygdala. The figure depicts three amygdala nuclei that
probably contribute to these two forms of prejudice: sensory inputs enter via the
lateral nucleus of the amygdala (LA) and, depending on the context and nature of
the stimuli, this signal is directed to the central nucleus of the amygdala (CeA),
which supports a threat response, or to the basal nucleus of the amygdala (BA),
which supports an instrumental response18. Because of the inhibitory nature of
within-amygdala projections, activating signals involve connections through
intercalated masses (ITCs). PAG, periaqueductal grey; PFC, prefrontal cortex;
SNS, sympathetic nervous system.
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600 ms
600 ms 600 ms
600 ms
a
Happy
+Awful
+
b
‘Compatible’ block
c‘Incompatible’ block
Time Time
+ +
Black or
negative
White or
positive
Black or
positive
White or
negative
Happy Black or
negative
White or
positive Awful Black or
positive
White or
negative
1 s 1 s
1 s 1 s
Fixation Prime Target
200 ms
200 ms
200 ms
200 ms
Response Fixation Prime Target Response
200 ms
200 ms
200 ms
200 ms
‘Pleasant’ or
‘unpleasant’?
‘Pleasant’ or
‘unpleasant’?
‘Gun’ or ‘tool’? ‘Gun’ or ‘tool’?
insula activity has also been observed when a disliked
outgroup member is rewarded — a case of outgroup envy
— and the degree of this activation predicted subjects
intention to harm that individual72. Although our under-
standing of insula function in social contexts is still devel-
oping, these findings highlight a role of visceral responses
to other people that has been largely overlooked in past
social-cognition research but that may nonetheless be
crucial for guiding intergroup social behaviour.
Striatum. The striatum is a component of the basal gan-
glia that comprises the caudate nucleus and putamen
(FIG.1). This structure is broadly involved in instrumental
learning and reward processes, including the coordina-
tion of goal-directed and habit-based responses through
bidirectional connections with the PFC (via the caudate
nucleus) and with motor areas (via the putamen), respec-
tively73. Findings from functional neuroimaging research
on economic bargaining and reinforcement learning sug-
gest that striatal activation is associated with the compu-
tation of value (that is, value placed on a potential action)
and anticipated outcomes74,75.
Consistent with a role of the striatum in reward pro-
cessing, fMRI studies of social perception have revealed
increased striatal activity in response to viewing pic-
tures of ingroup versus outgroup members47. In a study
in which white subjects completed an IAT that assessed
preferences for black versus white individuals, caudate
activity was stronger when subjects viewed white faces
compared with black faces, and this difference was asso-
ciated with an implicit preference for white ingroup
members64. In an economic bargaining game, the degree
of trust shown by white subjects towards a black part-
ner was associated with striatal activity76. These initial
findings suggest that the striatum has a role in guiding
positive intergroup interactions through instrumental
and approach-related responses.
Medial prefrontal cortex. The medial frontal cortex —
which encompasses Brodmann area8 (BA8), BA9 and
BA10 along the medial wall of the frontal cortex, superior
and anterior to the ACC — has emerged as a particularly
important structure for the processing of social informa-
tion62,77–79. This highly associative region has prominent
Box 2 | Measuring implicit prejudice and stereotyping
Unlike explicit racial beliefs, implicit attitudes and stereotypes reflect
associations in the mind that operate without conscious awareness9.
Implicit attitudes associated with race are formed through direct
or indirect exposure to members of these racial groups in negative
(or sometimes positive) contexts. Such implicit racial associations
are typically assessed using computerized priming tasks; the
priming effect is considered to be ‘implicit’ because subjects may be
unaware that they possess racial associations or may be otherwise
unaware of how their racial associations affect their task responses.
Racial bias assessed by implicit measures such as these has been
shown to predict a wide range of behavioural forms of
discrimination161.
In an example of a sequential priming task, subjects view and classify
target words as either ‘pleasant’ or ‘unpleasant’ (see the figure, part a).
Each target word is preceded by a prime stimulus that represents a social
category: for example, white and black faces. Implicit prejudices are
revealed in task performance: among white Americans, negative (versus
positive) words are often classified more quickly following black faces
than following white faces8.
A variant used to assess implicit stereotype associations is the weapons
identification task, in which white and black face stimuli (primes) are
followed by images of handguns and handtools162 (see the figure, part b).
Black primes typically facilitate the categorization of guns and interfere
with the categorization of tools, reflecting the stereotype of black
Americans as dangerous. Because this task creates stereotype-based
interference (on black-face prime–tool trials), it is also used to elicit and
index the cognitive control of stereotyping.
In the implicit association test (IAT), subjects view a series of stimuli, such
as white and black faces and positive and negative words163 (see the figure,
part c). During ‘compatible’ trials, white faces and positive words are
categorized using one key, whereas black faces and negative words are
categorized with a different key. During ‘incompatible’ trials, categories
are rearranged: white faces and negative words are categorized with one
key, and black faces and positive words with the other key. A tendency to
respond more quickly on compatible than incompatible blocks is taken to
indicate an anti-black and/or pro-white attitude. The IAT effect represents
the difference in average response latency between these two trial blocks,
with higher scores indicating stronger implicit prejudice.
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Stereotype > evaluation
56
54
52
50
48
46
Decoding accuracy
Judgement type
56
54
52
50
48
46
Decoding accuracy
Judgement type
Stereotype Evaluation
Evaluation > stereotype
interconnections with the ACC, the insula, the OFC and
the dorsolateral PFC (dlPFC), as well as other structures62.
In the context of social cognition, mPFC activity
has been primarily associated with the formation of
impressions about other people, especially impressions
that require mentalizing — the process of considering
a persons unique perspective and motives (that is,
engaging in theory of mind)80. Because the mPFC is
typically activated during judgements about other peo-
ple (as opposed to inanimate objects)77, mPFC activ-
ity, particularly in ventral, perigenual regions (FIG.1),
has been interpreted by some theorists as reflecting a
‘humanization’ process and, by extension, empathy67,81.
Hence, a lack of mPFC activity in response to a social
target may indicate a form of prejudice that is charac-
terized by a lack of humanization (that is, dehumani-
zation) and empathy. Indeed, the ventral mPFC has
been shown to be more highly activated when a sub-
ject views members of esteemed groups associated with
pride and admiration than when a subject views mem-
bers of low-status groups associated with disgust and
disregard (for example, homeless people)67. Moreover,
in a study in which Chinese and Caucasian subjects
viewed images of people being exposed to a painful or
non-painful stimulus (that is, a needle penetrating the
cheek versus a Q-tip touch), mPFC and ACC activity
was elicited only in response to seeing racial ingroup
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
or disrespected outgroup members.
A neural network for prejudice. Prejudice is a complex
social cognitive process that seems to be supported by
a network of neural structures (FIG.1). The amygdala
supports threat-based associations, which are thought
to underlie the most common form of implicit preju-
dice, and it is also involved in initial responses to sali-
ent positive or negative cues, including cues regarding
group membership. Thus, depending on the situation
and nuclei of interest, amygdala activity may be associ-
ated with social threat or with social reward. Activity in
the anterior insula supports the subjective experience of
negative affect (which often accompanies a prejudiced
response), whereas the mPFC is involved in mental-
izing and perspective taking, which may be engaged
more strongly towards ingroup than outgroup members.
Neural projections from the amygdala and insula to the
ventral mPFC may support the integration of affective
responses with mentalizing and empathy processes.
Finally, appetitive responses such as positive attitudes
and approach-related behavioural tendencies, which are
often expressed towards ingroup members, are primar-
ily supported by the striatum. These brain regions may
function in concert to support the learning, experience
and expression of prejudice.
Neural basis of stereotyping
In contrast to prejudice, which reflects an evaluative or
emotional component of social bias, stereotypes repre-
sent the cognitive component — the conceptual attrib-
utes linked to a particular group as defined by a culture
or society. This process involves the encoding and stor-
age of stereotype concepts, the selection and activa-
tion of these concepts into working memory and their
application in judgements and behaviours5,83. As such,
stereotyping involves cortical structures that support
more general forms of semantic memory, object mem-
ory, retrieval and conceptual activation, such as the tem-
poral lobes and inferior frontal gyrus (IFG), as well as
regions that are involved in impression formation, such
as the mPFC10,84,85. Although there is substantial overlap
between these structures and those implicated in preju-
dice, as described above, these structures appear to form
the core of a stereotyping network in the brain that may
operate separately from a prejudice network (FIG.4).
Temporal lobe. Stereotypes reflect conceptual associa-
tions between social groups and a particular set of attrib-
utes — associations that are thought to reside in semantic
memory83. As such, stereotype associations are posited
to involve regions of the lateral temporal lobe that
Figure 3 | Neural representation of racial bias in
affect-based and stereotype-based judgements.
A multivoxel pattern analysis approach revealed a unique
representation (that is, decoding accuracy) of race in the
orbital frontal cortex when subjects judged images of
black and white males according to an evaluative
dimension (who is more likely to be a friend?), and a
unique representation of race in the medial prefrontal
cortex when judging black and white males on a
stereotypical trait dimension (who is more interested in
athletics?)49. Reprinted from Neuropsychologia, 50,
Gilbert,S.J., Swencionis,J.K. & Amodio,D.M., Evaluative
versus trait representation in intergroup social
judgments: distinct roles of anterior temporal lobe and
prefrontal cortex, 36003611, Copyright (2012), with
permission from Elsevier.
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Dorsal mPFC
Impression formation
IFG
Stereotype activation
ATL
Social knowledge
Lateral temporal lobe
Semantic and episodic memory
underpin semantic knowledge10,85–89 (FIG.4). In particu-
lar, the anterior temporal lobe (ATL) is associated with
the representation of social knowledge, such as attributes
that describe people but not inanimate objects84,90,91. The
dorsal part of the ATL, which is implicated more spe-
cifically in the representation of social objects (that is,
people), is densely interconnected with the regions of
the mPFC that are associated with trait judgement and
impression formation92. This suggests that social infor-
mation represented in the ATL is selected into the mPFC
to support the process of social cognition.
Not surprisingly, the ATL is frequently implicated
in studies of stereotype representation. In one fMRI
study examining the neural basis of stereotyping, sub-
jects considered either social or non-social categories
(for example, men versus women or violins versus
guitars) and judged which category was more likely to
be characterized by a particular feature (for example,
enjoys romantic comedies or has six strings). A contrast
of brain activity between social and non-social condi-
tions revealed that ATL activity was uniquely activated
during stereotype-relevant judgements of social catego-
ries93. A different fMRI study used multivoxel pattern
analysis (MVPA) to examine neural activity represent-
ing judgements of black and white individuals on the
basis of stereotype traits (athleticism) versus evaluations
(potential for friendship)49. Results showed that when
subjects made trait judgements, a behavioural index of
implicit stereotyping correlated with ATL activity, and
when they made evaluative judgements, a behavioural
index of implicit racial attitudes correlated with activity
in the same part of the ATL. Consistent with these find-
ings, the disruption of ATL activity by transcranial mag-
netic stimulation attenuated the behavioural expression
of implicit gender stereotype associations94, suggesting
that the ATL is necessary for stereotype representation.
Thus, knowledge of social stereotypes appears to reside
in theATL.
Medial prefrontal cortex. As discussed above, the mPFC
is consistently involved in the representation of an indi-
vidual’s traits, preferences and mental states during
impression formation77,80. Although relevant to aspects
of prejudice, the mPFC is more directly involved in
stereotyping.
To date, the neural substrates of stereotyping have
mainly been examined within the domains of gender
and political orientation. These studies have linked
mPFC activity, typically in dorsal regions, with the acti-
vation of gender-related and political concepts during
behavioural tasks such as the IAT51,89,95–98. The mPFC
has been implicated in the domain of racial stereotyping
as well, during tasks that require subjects to infer per-
sonal traits of individuals from racial minority groups
(for example, African Americans)35. In an fMRI study
designed to distinguish the neural representation of ste-
reotype-based judgements of black versus white people
from judgements based on affective responses, MVPA
results identified the rostral dorsal mPFC as the only
region representing stereotype judgements49.
Although the mPFC has been linked to stereotyping,
its precise role in this process remains a point of inquiry.
Some authors have conceptualized the anterior mPFC
as a repository of social knowledge79,99 or as a region
that integrates information about social knowledge
with goals in order to coordinate social behaviour49,62,93.
Researchers are beginning to investigate these alternative
functions100,101. Nevertheless, in either case, the mPFC
seems to be centrally involved in the stereotype-based
processing of people.
It is notable that the mPFC is often considered to
function as part of a social-cognition (or mentalizing)
network, together with the temporoparietal junction,
superior temporal sulcus, precuneus and ATL78,102104. As
discussed, the mPFC and ATL have been directly linked
to social stereotyping processes, whereas the other
regions seem to be primarily associated with theory-
of-mind processing, action understanding and self-
consciousness — processes that are less directly relevant
to stereotyping and prejudice. Hence, the set of regions
involved in functional networks proposed for one psy-
chological function (for example, making mental state
inferences) may not cohere in the context of another
(for example, stereotyping) despite the fact that both
functions represent aspects of social cognition. In this
case, an involvement of the mPFC in stereotyping does
not necessarily implicate other components of networks
associated with mentalizing and social cognition.
Lateral prefrontal cortex. The lateral PFC — more spe-
cifically, the regions often referred to as theIFG (FIG.4)
(BA44, BA45 and BA47) — is associated with the selec-
tion of concepts into working memory to support goal-
directed action87,105109. William James famously observed
that ‘thinking is for doing’, and the left IFG, in particular,
reflects this notion: strong reciprocal connections of the
Figure 4 | Stereotyping network. Neural structures that
underlie components of intergroup stereotyping. Semantic
information stored in the lateral temporal lobe —
especially representations of stereotype-related
knowledge about people and social groups in the anterior
temporal lobe (ATL) — is recruited into the dorsal medial
prefrontal cortex (mPFC) to support the formation of
impressions (that is, stereotypes) and, in conjunction, into
the inferior frontal gyrus (IFG) to support goal-directed
actions that are guided by these stereotypes.
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Event-related potential
(ERP). An electrical signal
produced by summated
postsynaptic potentials of
cortical neurons in response to
a discrete event, such as a
stimulus or a response in an
experimental task. Typically
recorded from the scalp in
humans, ERPs can be
measured with extremely high
temporal resolution and can be
used to track rapid, real-time
changes in neural activity.
lateral PFC with the basal ganglia and motor cortex sup-
port the coordination of complex actions that are guided
by working memory and high-level cognition73,110.
Stereotypes are a form of social cognition that guide
behaviour, and indeed the process of applying stereo-
types in judgement and behaviour has been shown to
specifically involve activity in the IFG111.
Whereas the retrieval of conceptual knowledge typi-
cally involves the left IFG, activity in the right IFG has
been observed in research subjects who were judging
whether gender-stereotyped traits applied to a series of
male and female individuals (as compared with traits that
were unrelated to gender stereotypes)111. Given other evi-
dence that the right IFG has a role in domain-general
response inhibition112, it is possible that activation of
the right IFG during stereotype judgement tasks reflects
an individual’s efforts to inhibit the influence of stereo-
types on behaviour. This pattern of lateralized function
— stereotype retrieval and implementation on the left
and response inhibition on the right — suggests a useful
distinction in the processes through which stereotypes
are applied in behaviour.
A neural network for stereotyping. The research described
above suggests that a network of neural structures sup-
ports stereotyping processes (FIG.4). The ATL is believed
to represent stereotype-related knowledge, and it provides
input to the mPFC, possibly also during the online for-
mation of impressions about an individual. In this way,
social stereotypes ‘stored’ in the ATL may influence trait
impression processes associated with dorsal mPFC activ-
ity. The application of stereotypes to behaviour seems to
involve regions of the lateral PFC that are associated with
goal representation and response inhibition. Together,
the structures in this putative network may support the
storage, activation and behavioural expression of social
stereotypes.
Interacting networks
The framework described above suggests separate net-
works for prejudice and stereotyping, but in most cases
these two processes operate in concert. Thus, although
they are largely rooted in distinct neural systems, their
effects converge in higher-level cognition and behav-
ioural expression. Neuroscience studies suggest several
places at which this convergence may occur, although
this is primarily based on studies of connectivity in non-
human animals. For example, the anatomical connec-
tivity of the amygdala and OFC with the ATL, via the
uncinate fasciculus92, is consistent with behavioural evi-
dence that affective responses may influence the activa-
tion of stereotype concepts, and vice versa113,114. Similarly,
signals from several structures that are involved in both
prejudice and stereotyping — including the amygdala,
insula, striatum, OFC and ATL — converge in regions of
the mPFC, where information seems to be integrated in
support of elaborate person representations62,63. Finally,
the joint influences of prejudiced affect and stereotype
concepts on behaviour are likely to converge in the stria-
tum, which receives inputs from the amygdala, OFC, lat-
eral PFC and ATL, as well as other regions73,115. Although
the coherence of these proposed functional networks for
prejudice and stereotyping and their interaction has yet
to be tested, their existence is consistent with known
anatomical connectivity (which has been primarily
observed in non-human animals) and is supported by
decades of theory and behavioural research in the social
psychology literature4,5.
Regulation of prejudice and stereotyping
In an era of increasing diversity, international relations,
global communication and awareness of civil rights
issues, intergroup biases are often deemed to be both per-
sonally and socially unacceptable. Preferences based on
racial and ethnic categories that may have been adaptive
in less complex societies are no longer so. Fortunately,
the human mind is adept at self-regulation, and although
stereotypes and prejudices may come to mind automati-
cally in intergroup contexts, their expression can often be
moderated. Neuroscience research on the mechanisms
supporting the control of intergroup responses incorpo-
rates existing domain-general models of cognitive con-
trol into broader models that consider the influence of
social factors. For example, the impetus for the control
of racial bias may arise from internal cues (for example,
the personal rejection of prejudice) or external cues (for
example, social pressure to respond without prejudice),
and engagement in control is frequently associated with
social emotions such as social anxiety or guilt43. A neural
model of prejudice control should account for these dif-
ferent impetuses and emotion effects. In this way, neuro-
science research on prejudice has inspired an expanded
view of the neural and psychological processes involved
in control.
Anterior cingulate cortex. The ACC has been widely
implicated in the monitoring and detection of response
conflict116,117. In particular, the dorsal region of the ACC
(FIG.5) is often activated during cognitive control tasks,
such as the Stroop or Flanker tasks, on trials involving
a high degree of conflict between one’s desired response
and a countervailing tendency118,119. Conflict monitor-
ing theory posits that as the conflict signal in the ACC
rises, the ACC increasingly engages dlPFC regions that
function to implement goal-directed behaviour120. This
model is consistent with the ACC’s connectivity with
PFC regions involved in high-level goal representa-
tion and with the PFC’s connectivity with the striatum,
through which top-down control is implemented in
behaviour73,110,121.
In a social context, cognitive control is needed to
curb the unwanted influence of implicit stereotypes and
prejudices on behaviour7,122. Building on conflict moni-
toring theory, it has been proposed that the control of
implicit bias requires the detection of a conflict between
a biased tendency and ones goal to act without bias123.
Support for this proposal was provided by a study that
assessed ACC activity, which was indexed by the error-
related negativity component of the event-related potential
(ERP). In this study, subjects performed a task that
required them to inhibit the automatic expression of
racial stereotypes on some trials but not others (the
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rACC
Monitoring
external cues
dACC
Conflict processing
Nature Reviews | Neuroscience
mPFC
Representation of
interpersonal cues
dlPFC
Response selection
IFG
Response inhibition
weapons identification task; see BOX2). ACC activity
was selectively greater on trials requiring stereotype
inhibition, and the degree of subjects’ ACC activity on
these trials predicted their success at controlling the
expression of stereotypes in task behaviour123. This find-
ing has been replicated and extended in several studies
using different tasks and with alternative ERP indices of
ACC activity (for example, the error-related negativity
and response-locked N2)124127. These studies revealed,
for example, that ACC activation in response to stereo-
type conflict occurs implicitly and without deliberation,
is observed on a range of cognitive control tasks that
require the inhibition of stereotype-based responses and
is associated with an individual’s motivation to respond
without prejudice.
In several fMRI studies, ACC activation has also been
observed in white American subjects while they viewed
images of black (versus white) faces. However, this acti-
vation typically occurred in the absence of an opportu-
nity for cognitive control (that is, during tasks that did
not require control, such as passive face viewing)37,45,48.
Thus, it is difficult to interpret ACC activations in these
studies in terms of a control process. Nevertheless,
these findings suggest that exposure to black faces may
spontaneously elicit conflict detection processes. In
one notable exception — a study in which fMRI was
recorded during performance on a racial attitude IAT,
which requires cognitive control — ACC activity was
associated with the ability to identify the correct (that
is, non-biased) task response64. This finding is concep-
tually consistent with evidence from ERP studies that
link ACC activity to conflict processing and the engage-
ment of top-down control128. Similar patterns have been
observed when people are confronted with explicit
feedback about their bias. In two fMRI studies in which
subjects completed IAT measures of racial attitudes, false
IAT feedback to the subjects indicating that he or she
showed racial prejudice elicited heightened ACC activity,
and this degree of activity was associated with feelings
of guilt — a self-regulatory emotion that promotes pro-
social behaviours129,130. Together, these findings suggest
that the ACC supports the detection of one’s unwanted
social biases and the engagement of cognitive control in
order to avoid the expression ofbias.
Lateral prefrontal cortex. As noted above, the PFC is
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
in the left lateral PFC is linked to the implementation of
action, whereas activity in the right lateral PFC is linked
to action inhibition (in right-handed individuals)133,134.
On the basis of research in cognitive neuroscience, lateral
PFC regions have been proposed as primary substrates of
cognitive control of prejudice27,37,64.
Until recently, fMRI studies of responses towards
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
outgroup members were viewed passively by subjects.
Interestingly, these studies consistently revealed activ-
ity in regions of the PFC, most often the right IFG, in
response to explicit presentations of black faces com-
pared with white faces37,45,48. Although this finding is
difficult to interpret in terms of control, the established
role of the right IFG in response inhibition suggests that
exposure to black faces in these tasks may have spon-
taneously elicited a form of inhibitory control, perhaps
owing to subjects’ concern about appearing prejudiced.
In addition, right IFG activity has been reported to cor-
relate negatively with amygdala activity in response to
viewing black faces. This could suggest a potential regu-
latory circuit for prejudice control37,45; however, as these
data are merely correlational and as there is little direct
connectivity between the IFG and amygdala135, it is more
likely that right IFG activity reflects a form of response
inhibition rather than the direct downregulation of
amygdala activity.
The role of lateral PFC activity in the cognitive con-
trol of race biased behaviour was examined directly
in an electroencephalography (EEG) study, in which
subjects completed a task that assessed the behavioural
inhibition of stereotypes (the weapons identification
task)136. Greater dlPFC activity was found to be associ-
ated with better behavioural control (as modelled using
the process dissociation procedure), indicating a direct
link between the dlPFC and control of stereotyping.
Furthermore, the relation between dlPFC activity and
stereotype control was mediated by greater attentional
orienting to black faces than white faces, as indexed
by the P2 component of the ERP. This pattern sug-
gested that PFC activity tuned perceptual attention to
relevant stimuli, which in turn facilitated behavioural
Figure 5 | Regulation network. Neural structures supporting the regulation of
intergroup responses. Conflicts between a biased tendency and either internal goals or
external cues (for example, social norms) are processed in the dorsal anterior cingulate
cortex (dACC) and rostral ACC (rACC), respectively. The medial prefrontal cortex (mPFC)
is involved in perspective taking and mentalizing, and activation in this region provides
further representation of interpersonal cues to guide regulatory processing. Intergroup
response goals are represented in the lateral PFC and implemented in behaviour in
coordination with the striatum and motor cortex. dlPFC, dorsolateral PFC; IFG, inferior
frontal gyrus.
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control (BOX2). Findings consistent with this idea have
been reported in studies using EEG, fMRI and brain
lesion approaches in combination with behavioural
tasks designed to assess elements of prejudice con-
trol51,64,126,137,138. Together, these studies have begun to
identify the specific pathways through which the PFC
guides the control of intergroup responses.
Medial prefrontal cortex. As noted in previous sections,
the mPFC contributes to aspects of both stereotyping
and prejudice. However, this region is large, heterogene-
ous and widely interconnected, and emerging theories
and research suggest that its function may be closely
tied to regulatory processes as well. Amodio and Frith62
proposed that, given its role in mentalizing, the mPFC
supports the regulation of behavioural responses accord-
ing to social cues. Early evidence from ERP data sug-
gested that activity in the mPFC and/or rostral ACC
was uniquely associated with behavioural control that
is guided by external social cues, whereas activity in
the dorsal ACC was associated with internally con-
trolled behaviour124. In addition, ventral portions of the
mPFC are interconnected with the OFC and amygdala,
and through these connections it may support the top-
down modulation of emotional responses139. Hence, the
mPFC may support the regulation of intergroup affect,
such as threat or contempt, although this hypothesis
remains to be tested. Considered broadly, emerging evi-
dence regarding mPFC function suggests that it has a
larger role in cognitive control than previously thought,
particularly in the context of regulating complex social
responses.
A network for the regulation of prejudice and stereotyping.
Self-regulation is critical for the adaptive expression
of social behaviour. This is especially true with regard
to stereotyping and prejudice given the potential
(unwanted) influence of implicit biases on behaviour.
The putative network of neural regions involved in the
regulation of intergroup responses includes ACC and
PFC regions that have been implicated in existing mod-
els of cognitive control, as well as additional regions that
facilitate control in social contexts (FIG.5). Specifically,
although the dorsal ACC and lateral PFC may carry out
the domain-general functions of detecting conflict and
implementing top-down control, the mPFC and rostral
ACC are important for guiding control that is based on
social cues, such as norms against expressing prejudice,
and intergroup emotions.
From brain to society
Research on the neural basis of prejudice occupies a
special position at the interface of the natural and social
sciences, and as such it is uniquely situated to bring
neuroscientific advances to bear on real-world social
issues. To date, the neuroscientific analysis of prejudice
has advanced theories of how prejudices are formed,
expressed and potentially controlled, and these can be
used to inform interventions aimed at reducing discrimi-
nation. For example, research linking implicit prejudice
and stereotyping to different neural substrates suggests
that these two forms of bias are subserved by different
learning and memory systems — a clue that interventions
to reduce prejudice and stereotyping may require differ-
ent approaches10,28. Implicit prejudice has been linked to
fear conditioning involving the amygdala, whereas ste-
reotype associations appear to reflect conceptual learn-
ing systems involving the temporal cortex and PFC.
Importantly, learning and expression differ consider-
ably between these systems: fear conditioning may be
acquired in a single trial and expressed primarily through
behavioural freezing, anxiety and heightened vigilance18,
whereas conceptual associations require many exposures
for acquisition and are expressed through high-level rep-
resentations of impressions and goal-directed actions140.
Social cognition studies have begun to adopt interven-
tion strategies that are consistent with this analysis, using
tasks in which images of racial outgroup members are
repeatedly paired with positive images and appetitive
responses or with counter-stereotypical concepts, in an
effort to selectively target the affective or semantic mem-
ory systems underlying implicit prejudices and stereo-
types, respectively29,30,141. By considering the operations
of these different neural systems, researchers are gaining
a better understanding of how and under what condi-
tions different forms of bias are activated, expressed and
potentially extinguished.
Despite some success in reducing behavioural and
physiological expressions of implicit bias in the labo-
ratory29,30,141,142, most forms of implicit learning are
resistant to extinction140,143. Implicit racial biases are
particularly difficult to change in a cultural milieu that
constantly reinforces racial prejudices and stereotypes
(for example, in mainstream media). Thus, although
attempts to undo learned intergroup associations are
laudable, such strategies may be ineffective for reducing
the expression of bias in behaviour outside the labora-
tory. Instead, interventions that enhance the cognitive
control of behaviour should be more effective. Such
control-based strategies may not reduce prejudice in
the mind, but they can prevent its effect on potential
victims. Over time, control-driven changes in behaviour
may become habitual, and prejudiced and stereotypical
associations in the mind may weaken7,144. Neuroscience
models suggest that control-based interventions should
focus on (at least) two separate processes: those for mon-
itoring unwanted racially biased tendencies and those
involved in the top-down control of behaviour. Strategies
to enhance ACC-mediated conflict-monitoring processes
include interventions that increase people’s awareness of
the potential for bias, increase attention to specific cues
indicating that control may be needed (for example, the
appearance of an outgroup member in an interaction)
and increase the sensitivity of conflict monitoring systems
(for example, by activating cognitive conflict prior to an
intergroup response)145,146. For the effective control of
behaviour, conflict monitoring processes must be paired
with top-down response plans. To this end, psychological
research has shown that goal strategies that link a spe-
cific cue (for example, ‘if I meet a black person’) with a
pre-planned response (for example, ‘I will ignore his or
her race’ or ‘I will respond more carefully’) are especially
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effective at facilitating the control of implicit stereotypes
in behaviour147. By helping to inspire and explicate strate-
gies such as these, the neuroscience of prejudice is already
beginning to inform policy and interventions aimed at
reducing prejudice in society.
Beyond its implications for social issues regarding
intergroup relations, research on the neural basis of
prejudice provides a context for understanding neural
function as it relates to the real-world lives of human
beings. Many areas of social neuroscience consider the
effects of social factors on neural function, but the neu-
roscience of prejudice is a particularly rich topic as it
considers the roles of personal attitudes and motivations,
social norms and social emotions as they relate to com-
plex interpersonal behaviours. If the human brain has
evolved to support survival and prosperity in a complex
social environment, then a research approach that con-
siders this range of factors will be needed to truly under-
stand neural function. Research on the neural basis of
prejudice is an important step in this direction.
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Acknowledgements
Work on this article was supported by a National Science
Foundation grant (BCS 0847350). The author thanks mem-
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K. Ratner and the three anonymous reviewers for their helpful
feedback on earlier versions of this article.
Competing interests statement
The author declares no competing interests.
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... In this review, we first build on gender schema theories (GSTs; Bem, 1981;Martin and Halverson, 1981;Bem, 1983) and neural models of stereotypes (Amodio, 2014) to identify several neural and cognitive processes that may explain why some parents are more likely to apply gender socialization practices than other parents. Subsequently, empirical evidence for direct associations between cognitive and neural processes and gender socialization is discussed. ...
... In addition, neuroscientists have developed a neural model of implicit stereotypes (Stanley et al., 2008;Amodio, 2014) reflecting several neural processes that could underlie parental gender socialization. In this neural model, the temporal pole functions as a hub for social (stereotype) knowledge (Olson et al., 2013). ...
... In this neural model, the temporal pole functions as a hub for social (stereotype) knowledge (Olson et al., 2013). Based on this stereotype knowledge, the amygdala automatically evaluates socially salient (both negative and positive) stimuli and facilitates the allocation of the appropriate attentional processes to respond (Amodio, 2014). However, relying solely on automatic evaluations to drive our behaviors is not an optimal strategy in our complex social environment, and a certain level of control over the influence of stereotypes on behavior would be necessary. ...
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Parental gender socialization refers to ways in which parents teach their children social expectations associated with gender. Relatively little is known about the mechanisms underlying gender socialization. An overview of cognitive and neural processes underlying parental gender socialization is provided. Regarding cognitive processes, evidence exists that parents’ implicit and explicit gender stereotypes, attitudes, and gendered attributions are implicated in gender socialization. Other cognitive factors, such as intergroup attitudes, gender essentialism, internal motivation for parenting without gender stereotypes, gender identity, and conflict resolution are theoretically relevant mechanisms underlying gender socialization, but need further investigation. Regarding neural processes, studies demonstrated that attentional processing, conflict monitoring, behavior regulation, and reward processing might underlie stereotypes and biased behavior. However, more research is necessary to test whether these neural processes are also related to parental gender socialization. Based on this overview, a framework is presented of neural and cognitive factors that were theoretically or empirically related to gender socialization.
... Ethnic inter-group bias refers to biased mental processing of ethnic in-group and out-group members. It can encompass in-group favoritism or out-group derogation, being either unconscious or conscious (Hewstone et al., 2002;Amodio, 2014). In this way, inter-group bias is an adjacent concept to prejudices that are negative evaluations or emotional reactions toward an out-group member on the basis of preconceptions (Amodio, 2014). ...
... It can encompass in-group favoritism or out-group derogation, being either unconscious or conscious (Hewstone et al., 2002;Amodio, 2014). In this way, inter-group bias is an adjacent concept to prejudices that are negative evaluations or emotional reactions toward an out-group member on the basis of preconceptions (Amodio, 2014). Inter-group bias has enormous societal significance by providing a psychological foundation for ethnic discrimination (Greenwald and Pettigrew, 2014). ...
... This evidence has remained ignored, however, in the neuroscientific models of ethnic inter-group bias and prejudices. That is, the neuroscientific models on the topic have postulated a single model of neural responses toward ethnic in-and outgroups, without paying attention to the specific ethnic group in question, or ethnic minority or majority status (Amodio, 2014;Molenberghs and Louis, 2018). Thus, there is an urgent need to gain evidence whether distinct or overlapping brain regions are involved in inter-group bias among ethnic minority vs. majority members, and whether similar or different brain regions are involved in response to different ethnic groups. ...
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Introduction This meta-analysis investigated (1) whether ethnic minority and majority members have a neural inter-group bias toward each other, and (2) whether various ethnic groups (i.e., White, Black, and Asian) are processed in the brain differently by the other respective ethnicities. Methods A systematic coordinate-based meta-analysis of functional magnetic resonance imaging (fMRI) studies was conducted using Web of Science, PubMed, and PsycINFO (altogether 50 datasets, n = 1211, 50.1% female). Results We found that ethnic minority members did not show any signs of neural inter-group bias (e.g., no majority-group derogation). Ethnic majority members, in turn, expressed biased responses toward minority (vs. majority) members in frontal, parietal, temporal, and occipital regions that are known to be involved in e.g., facial processing, attention, and perspective-taking. We also found differences in neural response patterns toward different ethnic groups (White, Black, and Asian); broadest biases in neural response patterns were evident toward Black individuals (in non-Black individuals). Heterogeneity was mostly minor or low. Discussion Overall, the findings increase understanding of neural processes involved in ethnicity perception and cognition as well as ethnic prejudices and discrimination. This meta-analysis provides explanations for previous behavioral reports on ethnic discrimination toward minority groups.
... Prejudice is the attitude toward others based on their group membership, and it is intrinsically related to affective and cognitive processes, such as social categorization and stereotyping (Amodio, 2014). Negative beliefs about the outgroup influence choices, judgments, and behaviors (Sellaro et al., 2015) and can give rise to discrimination and prejudice to outgroup members (Amodio, 2014). ...
... Prejudice is the attitude toward others based on their group membership, and it is intrinsically related to affective and cognitive processes, such as social categorization and stereotyping (Amodio, 2014). Negative beliefs about the outgroup influence choices, judgments, and behaviors (Sellaro et al., 2015) and can give rise to discrimination and prejudice to outgroup members (Amodio, 2014). In contrast, individuals judge more positively members of the same group when compared to another racial group, a phenomenon called ingroup favoritism (Taylor & Doria, 1981). ...
... Different cerebral cortical regions are involved in prejudice, mainly associated with social perception and evaluation . One of the primary brain areas associated with prejudice is the medial prefrontal cortex (MPFC), an area involved in several cognitive activities, such as social perception, categorization, stereotyping, and regulation/control of behavioral responses in social contexts (Amodio & Frith, 2006;Amodio, 2014;Sellaro et al., 2015). Sellaro et al. (2015) investigated the causal role of MPFC in stereotype neutralization using tDCS. ...
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Machine learning (ML) is a subarea of artificial intelligence which uses the induction approach to learn based on previous experiences and make conclusions about new inputs (Mitchell, Machine learning. McGraw Hill, 1997). In the last decades, the use of ML approaches to analyze neuroimaging data has attracted widening attention (Pereira et al., Neuroimage 45(1):S199–S209, 2009; Lemm et al., Neuroimage 56(2):387–399, 2011). Particularly interesting recent applications to affective and social neuroscience include affective state decoding, exploring potential biomarkers of neurological and psychiatric disorders, predicting treatment response, and developing real-time neurofeedback and brain-computer interface protocols. In this chapter, we review the bases of the most common neuroimaging techniques, the basic concepts of ML, and how it can be applied to neuroimaging data. We also describe some recent examples of applications of ML-based analysis of neuroimaging data to social and affective neuroscience issues. Finally, we discuss the main ethical aspects and future perspectives for these emerging approaches.
... We cannot right wrongs by adding bias, prejudice, stigma and discrimination in any direction. There are recognized brain structures in the limbic system such as the amygdala, insula, hippocampus as well as the anterior cingulate cortex and associated pathways addressing automatic and controlled output of negative stimuli of bias, stereotyping, stigma, prejudice thoughts and fear of the "racially other" [3,4]. These feelings have been serving evolutionary existential purposes but are problematic and unwelcomed for an orderly function of a heterogeneous and civilized society. ...
... Ainsi, ce phénomène qui conduit les individus à préférer leurs propres groupes sociaux s'inscrit directement dans la façon que notre cerveau a d'orchestrer nos comportements. C'est, en effet, ce qu'ont mis en avant les neurosciences sociales depuis une vingtaine d'années, éclaircissant sous un angle nouveau le fonctionnement des préjugés (Amodio, 2014 ;Kubota et al., 2012). Visant la compréhension des processus psychologiques à travers le prisme des connaissances et outils de la fonction neuronale, les neurosciences sociales ont permis des découvertes cruciales sur la façon dont les humains perçoivent les groupes, forment et expriment des préjugés, ainsi que sur leurs conséquences sur les comportements intergroupes . ...
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Ce travail de thèse est pensé comme une contribution originale à la recherche sur le handicap, en adoptant une approche neuro-socio-cognitive de la perception sociale. L’objectif général est de poursuivre l’analyse des facteurs contribuant à comprendre les barrières à l’inclusion sociale auxquelles font face les personnes en situation de handicap. Dans ces recherches, nous discutons l’opérationnalisation du handicap comme une catégorie univoque. Nous abordons également la question centrale de la mesure en psychologie sociale et comment celle-ci peut approcher au mieux la réalité sociale. A partir de ces réflexions, ce travail de thèse suit une ambition double. D’une part, élargir l’objet d’étude du handicap en prenant en compte l’(in)visibilité de la déficience. D’autre part, diversifier les outils d’investigation concernant notre objet d’étude. Pour répondre à ceci, la thèse s’organise autour de deux volets principaux. Le premier volet, centré sur la perception de Soi des individus en situation de handicap, propose quatre études dont deux font l’objet de publications. Globalement, les résultats mettent en évidence le rôle déterminant de facteurs subjectifs, tels que les jugements de soi en terme de compétence. Le deuxième volet de cette thèse se focalise sur l’étude de la perception sociale à l’égard des personnes en situation de handicap. Il s’organise autour de sept études dont quatre s’inscrivent dans deux publications. Les résultats mettent en évidence l’impact du handicap sur la perception d’autrui, modulé par la visibilité de celui-ci. Pour mettre en évidence l’ensemble de ces résultats, les études se sont appuyées sur des outils relevant de la psychologie cognitive, de la cognition sociale et des neurosciences sociales. Cette approche neuro-socio-cognitive permet d’aborder les questions soulevées par le handicap de façon transversale tel que le préconise actuellement la recherche dans ce domaine.
... A very important aspect of intergroup empathy bias is that such bias appears to depend highly on the social motivation of the perceiver. Indeed, various studies have shown that self-categorization along an in-group/out-group distinction is flexible and that recategorization with an arbitrarily defined group may be sufficient to overcome automatic response biases [168]. This is of special importance in the context of radicalization since young people recategorize their group of membership by establishing new kinship relations as an effect of propaganda [103]. ...
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Background: The sympathy-empathy (SE) system is commonly considered a key faculty implied in prosocial behaviors, and SE deficits (also called callous-unemotional traits, CUTs) are associated with nonprosocial and even violent behaviors. Thus, the first intuitive considerations considered a lack of SE among young people who undergo radicalization. Yet, their identification with a cause, their underlying feelings of injustice and grievance, and the other ways in which they may help communities, suggest that they may actually have a lot of empathy, even an excess of it. As a consequence, the links between SE and radicalization remain to be specified. This critical review aims to discuss whether and how SE is associated with developmental trajectories that lead young people to radicalization. Method: We first recall the most recent findings about SE development, based on an interdisciplinary perspective informed by social neuroscience. Then, we review sociological and psychological studies that address radicalization. We will critically examine the intersections between SE and radicalization, including neuroscientific bases and anthropologic modulation of SE by social factors involved in radicalization. Results: This critical review indicates that the SE model should clearly distinguish between sympathy and empathy within the SE system. Using this model, we identified three possible trajectories in young radicalized individuals. In individuals with SE deficit, the legitimization of violence is enough to engage in radicalization. Concerning individuals with normal SE, we hypothesize two trajectories. First, based on SE inhibition/desensitization, individuals can temporarily join youths who lack empathy. Second, based on an SE dissociation, combining emotional sympathy increases for the in-group and cognitive empathy decreases toward the out-group. Conclusions: While confirming that a lack of empathy can favor radicalization, the counterintuitive hypothesis of a favorable SE development trajectory also needs to be considered to better specify the cognitive and affective aspects of this complex phenomenon.
... These psychological responses could result, in part, from a variety of factors, including: less intergroup contact in high-prejudice communities (Tropp & Pettigrew, 2006); stronger norms around anti-Black prejudice as socially acceptable (Crandall et al., 2002), because individuals in high-prejudice communities are repeatedly exposed to environmental cues that differentiate and marginalize people on the basis of skin tone (Vuletich & Payne, 2019); because high-prejudice communities are places where racialization is a salient axis of intergroup conflict (Cikara, 2021;Pietraszewski, 2021); or even concerns about being exposed as prejudiced in the context of the study (Amodio 2014;Chekroud et al. 2014). We are unable to test these and other competing (though not mutually exclusive) psychological, intergroup, and contextual explanations because, like all meta-analyses, we are limited by the data that could be reliably coded across the individual studies that we have included. ...
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We evaluated the hypothesis that neural responses to racial out-group members vary systematically based on the level of racial prejudice in the surrounding community. To do so, we conducted a spatial meta-analysis, which included a comprehensive set of studies (k=22; N=481). Specifically, we tested whether community-level racial prejudice moderated neural activation to Black (vs. White) faces in primarily White participants. Racial attitudes, obtained from Project Implicit, were aggregated to the county (k=17; N=10,743) in which each study was conducted. Multi-level kernel density analysis demonstrated that significant differences in neural activation to Black (vs. White) faces in right amygdala, dorsal anterior cingulate cortex, and dorsolateral prefrontal cortex were detected more often in communities with higher (vs. lower) levels of explicit (but not implicit) racial prejudice. These findings advance social-cognitive neuroscience by identifying aspects of macro-social contexts that may alter neural responses to out-group members.
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Previous research has suggested that race-specific features are automatically processed during face perception, often with out-group faces treated categorically. Functional imaging has illuminated the hemodynamic correlates of this process, with fewer studies examining single-neuron responses. In the present experiment, epilepsy patients undergoing microwire recordings in preparation for surgical treatment were shown realistic computer-generated human faces, which they classified according to the emotional expression shown. Racial categories of the stimulus faces varied independently of the emotion shown, being irrelevant to the patients’ primary task. Nevertheless, we observed race-driven changes in neural firing rates in the amygdala, anterior cingulate cortex, and hippocampus. These responses were broadly distributed, with the firing rates of 28% of recorded neurons in the amygdala and 45% in the anterior cingulate cortex predicting one or more racial categories. Nearly equal proportions of neurons responded to White and Black faces (24% vs. 22% in the amygdala and 26% vs. 28% in the anterior cingulate cortex). A smaller fraction (12%) of race-responsive neurons in the hippocampus predicted only White faces. Our results imply a distributed representation of race in brain areas involved in affective judgments, decision making, and memory. They also support the hypothesis that race-specific cues are perceptually coded even when those cues are task-irrelevant.
Article
While social neuroscience has already provided evidence for a deficit of affective empathy in racial prejudice, little is known about other less visible social categories when considered as an outgroup. We studied the process of empathy through event-related brain potentials (ERPs). We focused on the group "people with disabilities" as they are the target of a large amount of prejudice. Twenty-six participants performed a pain decision task. The mean amplitudes of N1, P2, N2–N3 and P3 components were recorded. Our results are consistent with previous work on prejudice, showing that the pain detection is modulated by group membership (with disabilities vs. without disabilities) on N2–N3, suggesting a better neural decoding of pain vs. non-pain in the without-disability condition. Critically, no effect of early sensory components (N1, P2) was found, and P3 was not moderated by disability. These findings indicate a different time course of empathic responses depending on the condition, suggesting that people with disabilities trigger specific empathic responses. Our results contribute to disentangling perceptual processes from affective empathy reactions.
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Previous studies have demonstrated that the activation of stereotype conflict is similar to the N400 congruency effect shown by the activation of semantic violation. In order to distinguish the differences between the two, the first experiment used gender stereotype trait words as target stimuli, and used “male/female” and “synonym of trait words/antonym of trait words” as priming stimuli respectively, so that the subjects completed the consistency determination task. In experiment 2, gender stereotyped behavior pictures were used as target stimuli, and “male/female” was used as priming stimuli, so that the subjects completed the task of consistency determination. The results showed that both gender stereotype conflict and semantic violation could induce N400 a congruency effect. Importantly, the N400 amplitude and response latency induced by gender stereotype activation are both smaller than those induced by semantic activation. These results show that stereotype activation is distinct from semantic activation, further demonstrating that the brain preferentially processes information related to gender stereotypes, and gender stereotype cognitive processing is more likely to happen than semantic knowledge processing.
Article
Three studies examined the moderating role of motivations to respond without prejudice (e.g., internal and external) in expressions of explicit and implicit race bias. In all studies, participants reported their explicit attitudes toward Blacks. Implicit measures consisted of a sequential priming task (Study 1) and the Implicit Association Test (Studies 2 and 3). Study 3 used a cognitive busyness manipulation to preclude effects of controlled processing on implicit responses. In each study, explicit race bias was moderated by internal motivation to respond without prejudice, whereas implicit race bias was moderated by the interaction of internal and external motivation to respond without prejudice. Specifically, high internal, low external participants exhibited lower levels of implicit race bias than did all other participants. Implications for the development of effective self-regulation of race bias are discussed.
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A neglected question regarding cognitive control is how control processes might detect situations calling for their involvement. The authors propose here that the demand for control may be evaluated in part by monitoring for conflicts in information processing. This hypothesis is supported by data concerning the anterior cingulate cortex, a brain area involved in cognitive control, which also appears to respond to the occurrence of conflict. The present article reports two computational modeling studies, serving to articulate the conflict monitoring hypothesis and examine its implications. The first study tests the sufficiency of the hypothesis to account for brain activation data, applying a measure of conflict to existing models of tasks shown to engage the anterior cingulate. The second study implements a feedback loop connecting conflict monitoring to cognitive control, using this to simulate a number of important behavioral phenomena.
Article
Anterior cingulate cortex (ACC) is a part of the brain's limbic system. Classically, this region has been related to affect, on the basis of lesion studies in humans and in animals. In the late 1980s, neuroimaging research indicated that ACC was active in many studies of cognition. The findings from EEG studies of a focal area of negativity in scalp electrodes following an error response led to the idea that ACC might be the brain's error detection and correction device. In this article, these various findings are reviewed in relation to the idea that ACC is a part of a circuit involved in a form of attention that serves to regulate both cognitive and emotional processing. Neuroimaging studies showing that separate areas of ACC are involved in cognition and emotion are discussed and related to results showing that the error negativity is influenced by affect and motivation. In addition, the development of the emotional and cognitive roles of ACC are discussed, and how the success of this regulation in controlling responses might be correlated with cingulate size. Finally, some theories are considered about how the different subdivisions of ACC might interact with other cortical structures as a part of the circuits involved in the regulation of mental and emotional activity.
Article
In the intergroup relations literature, theories of control concern the interplay of basic cognitive mechanisms of self-regulation with intrapersonal and societal-level goals and motivations. By integrating multiple levels of analysis, this literature has been uniquely positioned to advance our understanding of control as it operates in the complex social world that the human self-regulatory system has evolved to negotiate. In this chapter, we review research and theoretical models of control that have emerged from intergroup approaches. The first section of this chapter describes four theoretical models of control that have been central to research in prejudice, stereotyping, and intergroup relations. The next section highlights some of the recent exciting advances in the study of control in the intergroup domain and notes how they are refining and, in some cases, redefining basic conceptions of control as a self-regulatory process. The final section outlines what we see as some major challenges faced by current theories of control, both in the intergroup domain and in the broader psychological literature.
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Social neuroscience is a young and thriving area of research in psychology that integrates diverse literatures and methodologies to address broad questions about the brain and behavior. But despite the excitement and activity generated by this approach, its contribution to ideas in social psychology is sometimes questioned. This article discusses the ways in which social neuroscience research may or may not contribute to theoretical issues in social psychology. Still a young field, much research in this area has focused on issues of brain mapping and methodological development, with less emphasis on generating and testing novel social psychological hypotheses. The challenges to theoretical advancement, including psychometric and methodological issues, are considered, and a set of guidelines for conducting theoretically-informative social neuroscience is offered. In the final analysis, it is argued that neuroscience has much to offer to social psychology, both theoretically and methodologically, but that like any new approach, these contributions will take time to realize.
Article
Previous research on the neural underpinnings of empathy has been limited to affective situations experienced in a similar way by an observer and a target individual. In daily life we also interact with people whose responses to affective stimuli can be very different from our own. How do we understand the affective states of these individuals? We used functional magnetic resonance imaging to assess how participants empathize with the feelings of patients who reacted with no pain to surgical procedures but with pain to a soft touch. Empathy for pain of these patients activated the same areas (insula, medial/anterior cingulate cortex) as empathy for persons who responded to painful stimuli in the same way as the observer. Empathy in a situation that was aversive only for the observer but neutral for the patient recruited areas involved in self-other distinction (dorsomedial prefrontal cortex) and cognitive control (right inferior frontal cortex). In addition, effective connectivity between the latter and areas implicated in affective processing was enhanced. This suggests that inferring the affective state of someone who is not like us can rely upon the same neural structures as empathy for someone who is similar to us. When strong emotional response tendencies exist though, these tendencies have to be overcome by executive functions. Our results demonstrate that the fronto-cortical attention network is crucially involved in this process, corroborating that empathy is a flexible phenomenon which involves both automatic and controlled cognitive mechanisms. Our findings have important implications for the understanding and promotion of empathy, demonstrating that regulation of one's egocentric perspective is crucial for understanding others.
Article
Three studies tested basic assumptions derived from a theoretical model based on the dissociation of automatic and controlled processes involved in prejudice. Study 1 supported the model's assumption that high- and low-prejudice persons are equally knowledgeable of the cultural stereotype. The model suggests that the stereotype is automatically activated in the presence of a member (or some symbolic equivalent) of the stereotyped group and that low-prejudice responses require controlled inhibition of the automatically activated stereotype. Study 2, which examined the effects of automatic stereotype activation on the evaluation of ambiguous stereotype-relevant behaviors performed by a race-unspecified person, suggested that when subjects' ability to consciously monitor stereotype activation is precluded, both high- and low-prejudice subjects produce stereotype-congruent evaluations of ambiguous behaviors. Study 3 examined high- and low-prejudice subjects' responses in a consciously directed thought-listing task. Consistent with the model, only low-prejudice subjects inhibited the automatically activated stereotype-congruent thoughts and replaced them with thoughts reflecting equality and negations of the stereotype. The relation between stereotypes and prejudice and implications for prejudice reduction are discussed.