Amygdala Sensitivity to Race Is Not Present in - Am MIT spezialisierte KI-Forschung

Amygdala Sensitivity to Race Is Not Present in
Childhood but Emerges over Adolescence

Eva H. Telzer1, Kathryn L. Humphreys2, Mor Shapiro2,
and Nim Tottenham2

Abstrakt

■ Neuroimaging research in adults has consistently found
that differential perception of race is associated with in-
creased amygdala activity. We hypothesized that such neural
biases unlikely reflect innate processes but instead emerge
over development. In the current study, we used fMRI to ex-
amine the neurodevelopmental trajectory of the amygdala in
response to race across childhood and adolescence ranging
aus 4 Zu 16 Jahre. Thirty-two youths viewed African Ameri-

can and European American faces during a functional brain
scan. Results suggest that differential amygdala response to
African American faces does not emerge until adolescence,
reflecting the increasing salience of race across development.
Zusätzlich, greater peer diversity was associated with attenu-
ated amygdala response to African American faces, vorschlagen-
ing that intergroup racial contact may reduce the salience
of race. ■

EINFÜHRUNG

Although explicit cultural norms in the United States may
endorse egalitarian values and nonprejudiced attitudes,
African Americans (AAs) continue to be evaluated differ-
ently from other racial/ethnic groups (Rosette, Leonardelli,
& Phillips, 2008; Plant & Devine, 1998; Dovidio, Kawakami,
Johnson, Johnson, & Howard, 1997). Zum Beispiel, AA faces
are detected more quickly in visual search tasks (Levin,
2000) and produce an attentional bias during a dot-probe
Aufgabe (Richeson & Trawalter, 2008; Trawalter, Todd, Baird, &
Richeson, 2008), suggesting that AA faces hold increased
saliency in adulthood. Neuroimaging research in adults
has consistently found that this differential perception
Ist, in part, associated with increased amygdala activity.
European American (EA) adults show increased amygdala
Aktivität, even in the absence of conscious awareness, In
response to AA relative to EA faces (Cunningham et al.,
2004). Darüber hinaus, EA adults who harbor implicit negative
attitudes toward AAs show greater amygdala activation
while viewing AA relative to EA faces (Phelps et al.,
2000). Interessant, heightened amygdala response to AA
faces is found for both EA and AA adults (Lieberman, Hariri,
Jarcho, Eisenberger, & Bookheimer, 2005). This height-
ened amygdala response is thought to be involved in auto-
matic, subconscious responses to race, reflecting the
learned cultural knowledge that AAs are treated differently,
and such cultural knowledge is shared across individuals
from diverse backgrounds (Lieberman et al., 2005; Phelps
et al., 2000). Given that the value placed on racial groups is
socially constructed (Eberhardt, 2005), we hypothesized that

1University of Illinois, 2Universität von Kalifornien, Los Angeles

such biases unlikely reflect innate processes but instead
emerge over developmental time through learning. Im
current study, we used fMRI to examine the neurodevelop-
mental trajectory of the amygdala response to race across
Kindheit und Jugend.

Cultural norms and biases about race develop over the
course of childhood and adolescence. When social groups
are treated or labeled differently in childrenʼs environ-
ment, children learn that certain categories are salient
(z.B., Wettrennen), whereas others are not (z.B., handedness;
Bigler & Liben, 2007). At a very young age, children learn
that individuals can be sorted into social categories, solch
as race. Zum Beispiel, infants as young as 3–6 months can
perceptually discriminate between racial groups (Bar-
Haim, Ziv, Lamy, & Hodes, 2006), and preschool-aged
children can accurately identify othersʼ racial group mem-
bership (Aboud, 2003). Von 6 Jahre, some children dem-
onstrate implicit biases about race (Baron & Banaji,
2006), und von 10 Jahre, children internalize the social
and moral norms of their culture, demonstrating in-
creased knowledge regarding racial stereotypes and cul-
tural norms (Apfelbaum, Pauker, Ambady, Sommers, &
Norton, 2008).

The amygdala is involved in processing of stimuli that
have an acquired emotional significance based on previous
experience and plays a role in sensitivity to the salience of
environmental cues (Cunningham & Brosch, 2012; Santos,
Mier, Kirsch, & Meyer-Lindenberg, 2011; Fitzgerald, Angstadt,
Jelsone, Nathan, & Phan, 2006; Fudge & Emiliano, 2003;
Whalen et al., 2001). Whereas brain regions such as the
cerebellum respond to visual and perceptual differences in
ones environment, such as shades of color (Claeys et al.,
2003), the amygdala responds to emotionally salient stimuli

Zeitschrift für kognitive Neurowissenschaften 25:2, S. 234–244

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(Cunningham & Brosch, 2012; Whalen et al., 2001). Der
amygdala responds to both negatively and positively
valenced stimuli (Hennenlotter et al., 2005; Breiter et al.,
1996), highlighting its role in learning about the emotional
significance of the environment in general. daher, Die
amygdala is well positioned to acquire affective associations
learned in the social environment, such as those associated
with race. In addition to responding to emotionally salient
stimuli based on experience, the amygdala is involved in
fear-related learning, detecting and responding to threats,
and encoding the hedonic value of learned and unlearned
Reize (Fanselow & Sturm, 2003; LeDoux, 2003).

Both human and animal work shows that the amygdala
is an early developing brain structure (Payne, Machado,
Bliwise, & Bachevalier, 2010). Structurally, the amygdala
undergoes rapid development early in life (Tottenham,
Hare, & Casey, 2009). Tatsächlich, the basic neuroanatomical
architecture of the human amygdala is present by birth
(Ulfig, Setzer, & Bohl, 2003; Humphrey, 1968). Obwohl
structurally mature by early childhood, the amygdala
undergoes massive changes in functional processing dur-
ing adolescence, increasing in responsiveness to social
Reize (Moore et al., 2012; Guyer et al., 2008; Nelson,
Leibenluft, McClure, & Pine, 2005). Zum Beispiel, the onset
of puberty is associated with enhanced amygdala activa-
tion to facial stimuli (Moore et al., 2012). Adolescence is
also a time when race becomes increasingly salient. Für
Beispiel, adolescents enter high school where ethnic
clubs and coalitions form, and youth begin to explore the
meaning and importance of ethnicity and race (Roberts
et al., 1999). Darüber hinaus, the transition to adolescence is
marked by a greater awareness of racial stereotypes and
norms (Apfelbaum et al., 2008). The social reorientation of
the amygdala (Nelson et al., 2005), coupled with more
mature cognitive skills (Bigler & Liben, 2007), as well as an
increasing salience of race, renders the early adolescent
Jahre, particularly amenable to enhanced amygdala response
to race.

In the current study, we sought to understand how
experience alters race-related processing in the amyg-
dala. Erste, we examined age-related differences in amyg-
dala response to race to test whether the pattern of
amygdala response to AA and EA faces that is observed
in adulthood is present in early childhood or whether it
emerges across development. We examined amygdala
sensitivity to race across a wide developmental age range,
spanning 4–16 years. Zusätzlich, we examined how neural
responses to race may differ across ethnically diverse
youth. Children from diverse ethnic and racial backgrounds
living in similar geographical areas are exposed to similar
messages about race throughout their environment (Averhart
& Bigler, 1997). daher, we expected that children from
both EA and AA backgrounds will show a similar neuro-
developmental increase to AA faces, similar to the findings
of Lieberman and colleagues (2005), who found that both
AA and EA adults showed heightened amygdala response
to AA faces.

Zweite, we examined whether childrenʼs social envi-
ronment modulates the amygdala response to race. Prior
work has highlighted the importance of diverse social
environments, such as neighborhood and school diver-
Stadt, in shaping perceptions of race. Zum Beispiel, 3-month-
old infants exhibit a preference for faces from their own
racial group (d.h., in-group bias), but this bias is only pres-
ent for infants living in racially homogeneous neighbor-
hoods; infants living in a heterogeneous environment do
not exhibit an in-group bias (Bar-Haim et al., 2006). In
addition, children from racially mixed schools are less
likely to develop race-related favorable in-group biases
and negative out-group biases (Rutland, Cameron, Milne,
& McGeorge, 2005). Contact between individuals from
diverse backgrounds may reduce the salience of inter-
group boundaries, producing more individuated and
personalized relationships (Dovidio & Gaertner, 1999).
In the current study, we examined the independent con-
tribution of childrenʼs neighborhood and peer diversity
on their amygdala response to race to examine whether
more racially heterogeneous contexts would decrease
the amygdalaʼs response to AA faces. We expected that
heightened amygdala response to AA faces would only
be present among children and adolescents in racially
homogenous contexts, because of the increased salience
of race for these youth.

METHODEN

Teilnehmer

Participants included 32 healthy children and adolescents
(20 Männer), ages 4–16.5 years (Durchschnittsalter = 11.3 Jahre,
SD = 3.95 Jahre). Age was evenly distributed among male
and female participants (weiblich, 5–16 years; männlich, 4–
16 Jahre). Participants were predominantly from AA
(n = 11) and EA (n = 11) backgrounds, with the remain-
ing from Asian American (n = 6) and Latin American
(n = 4) backgrounds. Participants from AA and EA back-
grounds were similar in age (AA, Durchschnittsalter = 12.18 Jahre,
SD = 3.69, age range = 4.6–16.5 years; EA, Durchschnittsalter =
11.75, SD = 4.08, age range = 5.3–16.5). All children
were physically and psychiatrically healthy, which was
confirmed by a telephone screening. Childrenʼs IQs were
within the normal range (mean = 110.7, SD =16.8) als
estimated via two subtests from the Wechsler Abbre-
viated Scale of Intelligence (Wechsler, 1999). All partici-
pants were right-handed.

Procedures

Individual Difference Measures

Parents completed several measures about their child.

Alter. Parents indicated their childʼs date of birth. Chil-
drenʼs age at the time of the scan was measured by taking

Telzer et al.

235

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the difference in months between the childʼs birth date
and the date of the scan.

Peer and neighborhood diversity. Parents indicated the
racial diversity of their childʼs peers by answering two
Fragen, “Are your childʼs friends…” and “Are the other
children in your childʼs current school…”: 1 = all his or
her race, 2 = mostly his or her race, 3 = mixed, 4 =
some his or her race, 5 = not at all his or her race.
These two items were averaged to create one index of
peer diversity where lower scores indicated greater
homogeneity of peers. Using the same 5-point scale, Par-
ents indicated their childʼs neighborhood diversity with
the following item: “Is the neighborhood your child
grows up in….” Five parents did not provide Peer and
Neighborhood Diversity scores, including three EA, eins
AA, and one Latin American participant.

fMRT-Aufgabe

During the fMRI scan, participants completed two func-
tional runs of the Emotional Matching Task, adapted
from Hariri et al. (2002) and Lieberman and colleagues
(2005). During each run, two blocks of emotional faces
were interleaved with two blocks of a sensorimotor con-
trol task (shapes). For the face blocks, children were pres-
ented with a trio of faces and were instructed to make a
button response to indicate which of the two faces at the
bottom was expressing the same emotion or felt the same
as the face on top. The faces were displaying one of three
emotions: Angry, Happy, or Neutral, and all were taken
from the NimStim Set of Facial Expressions (Tottenham,
Tanaka, et al., 2009). For the shapes blocks, children were
presented with a trio of shapes and selected one of the two
shapes at the bottom that was identical to the shape on
top. Each block consisted of six faces or shapes, welche
were each presented for 5 Sek. Participants completed
two runs of the Emotional Matching Task. Similar to the
paradigm used by Lieberman and colleagues (2005), Par-
ticipants played one run in which all the faces were EA
and one run in which all the faces were AA. Run order
was counterbalanced across participants. Participants were
never instructed to attend to race.

fMRT-Datenerfassung

Participants were scanned on a Siemens Trio 3.0-T MRI
scanner. For each participant, an initial 2-D spin echo
Bild (repetition time = 4000 ms, Echozeit = 40 ms,
matrix size = 256 × 256, 4 mm thick, 0 mm gap) im
oblique plane was acquired to enable prescription of slices
obtained in the structural and functional scans. A whole-
Gehirn, high-resolution, T1*-weighted anatomical scan
(MPRAGE; 192 × 192 in-plane resolution, 250 mm field
of view; 176 mm × 1 mm sagittal slices) was acquired for
each subject for registration and localization of functional

data into Talairach space. The Emotional Matching Task was
presented on a computer screen through MR-compatible
goggles. The task was completed during two functional
scannt. Ninety-nine T2*-weighted EPIs were collected (repe-
tition time = 2000, Echozeit = 30 ms, flip angle = 90°,
matrix size = 64 × 64, 34 Scheiben, 4 mm voxel, skip 0 mm) bei
an oblique angle of approximately 30°.

fMRT-Datenanalyse

Functional imaging data were preprocessed and analyzed
with the Analysis of Functional Neuroimaging (AFNI) weich-
ware package (Cox, 2006). All data were free of movement
greater than 2.5 mm in any direction. Preprocessing for
each participantʼs images included slice time correction
to adjust for temporal differences in slice acquisition within
each volume, spatial realignment to correct for head motion,
registration to the first volume of each run, spatial smooth-
ing using anisotropic 6 mm Gaussian kernel, FWHM to
increase the signal to noise ratio, and transformation into
the standard coordinate space of Talairach and Tournoux
(Talairach & Tournoux, 1988) with parameters obtained
from the transformation of each subjectʼs high-resolution
anatomical scan. Talairach transformed images had a re-
sampled resolution of 3 mm3. Time series were normalized
to percent signal change to allow comparisons across runs
and individuals by dividing signal intensity at each time
point by the mean intensity for that voxel and multiplying
the result by 100.

The functional runs were concatenated before creating
each participantʼs individual-level model, which included
three regressors for each of the stimulus types (AA faces,
EA faces, and shapes) by convolving the stimulus timing
files with canonical hemodynamic response function. Six
motion parameters were included as separate regressors
for a total of nine regressors. General linear modeling
was performed to fit the percent signal change time
courses to each regressor. Linear and quadratic trends
were modeled in each voxel time course to control for
correlated drift.

Nächste, the individual level regression coefficients were
submitted to random effects, group level analyses. Wir
conducted regression analyses using the 3dRegAna pro-
gram within AFNI to explore how neural responses to AA
and EA faces changed as a function of age and diversity. Alter
and diversity scores were each entered as regressors. Cor-
rection for multiple comparisons was applied at the cluster
level following Monte Carlo simulations conducted in the
AlphaSim program within AFNI. This method controls for
type I errors, offering a reasonable correction for multiple
tests during group level analyses in ROIs. Results of the
AlphaSim indicated a voxel-wise threshold of p < .05 com- bined with a minimum cluster size of eight voxels for the bilateral amygdala (Phan, Fitzgerald, Nathan, & Tancer, 2006), corresponding to p < .05, false discovery rate cor- rected. Non a priori regions outside the amygdala were corrected for multiple comparisons within the whole brain 236 Journal of Cognitive Neuroscience Volume 25, Number 2 D o w n l o a d e d f r o m l l / / / / j f / t t i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e d o u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 5 / 2 2 5 2 / 3 2 4 / 1 2 9 3 4 4 4 / 9 1 4 7 3 7 o 8 c 8 n 2 _ 1 a / _ j 0 o 0 c 3 n 1 1 _ a p _ d 0 0 b 3 y 1 g 1 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 i 2 3 e s / j / f t . u s e r o n 1 7 M a y 2 0 2 1 Table 1. Behavioral Responses on the Emotional Matching Task Condition AA Faces Children (4–9 years) Early adolescents (10–13 years) Late adolescents (14–16.5 years) EA Faces Children (4–9 years) Early adolescents (10–13 years) Late adolescents (14–16.5 years) Shapes Children (4–9 years) Early adolescents (10–13 years) Late adolescents (14–16.5 years) Mean Reaction Time (SD) Mean Accuracy (%) (SD) 1850.85 (536.62) 1606.69 (474.40) 1359.89 (333.85) 1876.21 (515.04) 1597.56 (410.75) 1506.06 (408.01) 1284.60 (534.54) 1040.78 (337.94) 919.09 (202.71) 91.7 (7.8) 92.5 (6.1) 95.8 (5.6) 93.4 (7.7) 93.4 (10.2) 93.7 (7.2) 89.8 (16.4) 86.7 (18.0) 96.3 (3.8) For descriptive purposes only, participants were broken up into three age groups: children (n = 10), early adolescence (n = 10), and late adoles- cence (n = 12). Statistical analyses treated age as a continuous variable. at p < .01 with a minimum cluster size of 56 voxels. All analyses controlled for participantsʼ own race. RESULTS Behavioral Performance on the Emotional Matching Task Separate repeated-measures ANOVAs were performed using the within subjects factor of Condition (AA faces, EA faces, shapes) and the between-subject factor of Age on the dependent measures of mean RT and percentage accuracy. We found a significant main effect for Condition on RT (F = 81.03, p < .001), such that participants were faster at matching shapes than either face condition (see Table 1). There was no main effect of Age or interaction of Condition × Age. There was also a main effect for Con- dition on accuracy. Participants made more errors when matching shapes than either face condition (F = 8.68, p < .05). There was no main effect of Age or interaction of Condition × Age. These findings show that younger children and older adolescentsʼ performance is similar on the task, with high performance levels across age, sug- gesting that it is a developmentally appropriate paradigm. The behavioral data suggest that the shapes condition was experienced quantitatively differently than the face conditions, and therefore, we used the implicit baseline (crosshair fixation) rather than shapes to contrast with the faces in the fMRI analyses. Amygdala Response to Race across Development Our first analyses examined whether the amygdala in our child and adolescent sample coincides with the adult template used for registration. We created an average anatomical from all participants in the study. As shown in Figure 1, the anatomical average from our developmental population shows that the amygdala region coincides with the adult template. Figure 1. The amygdala in the adult template (left) corresponds to the amygdala in the average anatomical template from the developmental population in the current study (right). xyz coordinates are 18 −3 −9. Telzer et al. 237 D o w n l o a d e d f r o m l l / / / / j t t f / i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e d o u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 5 / 2 2 5 2 / 3 2 4 / 1 2 9 3 4 4 4 / 9 1 4 7 3 7 o 8 c 8 n 2 _ 1 a / _ j 0 o 0 c 3 n 1 1 _ a p _ d 0 0 b 3 y 1 g 1 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 i 2 3 e s / j t / . f u s e r o n 1 7 M a y 2 0 2 1 D o w n l o a d e d f r o m l l / / / / j t t f / i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e d o u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 5 / 2 2 5 2 / 3 2 4 / 1 2 9 3 4 4 4 / 9 1 4 7 3 7 o 8 c 8 n 2 _ 1 a / _ j 0 o 0 c 3 n 1 1 _ a p _ d 0 0 b 3 y 1 g 1 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 i 2 3 e s / j f . t / Figure 2. (A) The bilateral amygdala to AA–EA faces correlated positively with age. This neurodevelopmental amygdala increase is specific to AA faces such that (B) the right amygdala response to AA faces relative to baseline correlated positively with age, whereas (C) the amygdala does not show a developmental increase in response to EA faces. (D) The age effect with the 95% confidence interval. Where the confidence interval does not include 0 on the y-axis (depicted with an arrow), participants are showing a significant differential response to AA faces. Our first primary goal was to examine whether there were neurodevelopmental changes to AA faces relative to EA faces. In whole-brain regression analyses, we corre- lated age with neural activation to AA–EA faces. As shown in Figure 2A, with age, children showed increased bilat- eral amygdala activation to AA–EA faces (right: xyz = 16 −2 −8, t(30) = 3.67, p < .05, corrected; left: xyz = −14 −2 −7, t(30) = 2.37, p < .05, corrected). Next, we examined whether this neurodevelopmental increase in amygdala response to AA–EA faces is specific to AA faces, EA faces, or both. We correlated age with neural activation in the contrast of AA faces-baseline and EA faces-baseline separately in whole-brain analyses. Developmental increases in the amygdala were specific to AA faces. Whereas activation in the right amygdala significantly increased to AA faces across development 238 Journal of Cognitive Neuroscience Volume 25, Number 2 u s e r o n 1 7 M a y 2 0 2 1 Table 2. Whole-brain Significant Activations for AA and EA Faces that Correlated Positively with Age and Peer Diversity Anatomical Region BA (a) AA > EA Faces and Age

VLPFC

(B) AA Faces (Relative to Baseline) and Age

VLPFC

Middle occipital gyrus

18/19

Culmen

VLPFC

Anterior cingulate

Insula

−34

−29

−45

−10

−33

−61

−85

−88

−64

−10

−19

−1

−7

5.78

3.82

4.58

3.34

4.40

4.91

280

−3.37

−3.04

−3.68

1879

1161

1723

129

1020

127

BA refers to putative Brodmannʼs area; L and R refer to left and right hemispheres; X, j, and z refer to Talairach coordinates; t refers to the t score at
those coordinates (local maxima); k refers to the number of voxels in each significant cluster. The following abbreviations are used for the names of
specific regions: DLPFC = dorsolateral; pFC, VLPFC = ventrolateral pFC.

Non a priori regions outside the amygdala were corrected for multiple comparisons within the whole brain at p < .05 with a minimum cluster size of 146 voxels. (t(31) = 3.41, p < .05, corrected; Figure 2B), age did not correlate with amygdala activation to EA faces (Fig- ure 2C). A repeated-measures ANOVA using the within- subject factor Race (AA and EA) and the between subject factor of Age on the dependent measure of percent BOLD signal change in the amygdala, revealed a signifi- cant Race × Age interaction, F(1, 30) = 14.6, p < .001. Given this developmental increase that is specific to AA faces, we explored at what age the amygdala responds differentially to AA faces. We ran follow-up analyses using the margins function in STATA11 (StataCorp, College Station, TX). Figure 2D displays the age effect with the 95% confidence interval. Where the confidence interval does not include 0 on the y axis, the participants are showing a significant differential response to Black faces. The margin becomes significant around age 14 (z = 2.51, p = .01, 95% CI [0.32, 2.66]). Together, these findings indicate that there are age-related changes in the process- ing of AA but not EA faces, such that amygdala sensitivity to AAs is not present in early childhood but emerges during adolescence. For other significant regions that correlated with age to AA and EA faces, see Table 2A–C. Our next goal was to examine whether AA and EA par- ticipants showed similar neurodevelopmental trajectories to AA and EA faces. We extracted parameter estimates from the right amygdala to EA faces and AA faces and ran sepa- rate regression analyses in SPSS for each ethnic group, ex- amining how age related to amygdala response to EA and AA faces separately. Both EA (B = 0.79, SE = 0.20, β = .80, p < .005) and AA (B = 0.42, SE = 0.11, β = .80, p < .005) participants showed increased right amygdala activation to AA faces with age, but neither group showed increased amygdala response to EA faces with age. These findings sug- gest that the amygdala becomes increasingly sensitive to AA faces with development, and this neurodevelopmental tra- jectory is similar for individuals from AA and EA backgrounds. Finally, as a control to ensure that it is possible to get amygdala response in our younger children, we examined whether all age groups show differential amygdala response to emotional faces (angry). Because we found evidence of a developmental increase in the amygdala to AA faces, we examined the contrast of EA angry faces > baseline, als
we anticipated that angry faces would produce a stable
signal across all age groups. For descriptive purposes, Wir
divided our sample into three age groups, Kinder (Alter
4–9, n = 10), early adolescents (ages 10–13, n = 10), Und
adolescents (ages 14–16.5, n = 12). We observed differen-
tial amygdala response to EA angry faces > baseline in each
age group in the amygdala (Kinder: right amygdala: xyz =
−25 −1 −20, T(9) = 3.88, P < .05, left amygdala: xyz = 22 10 −3, t(9) = 2.95, p < .05; early adolescents: right amyg- dala: xyz = 22 1 −18, t(9) = 3.34, p < .05; adolescents: right amygdala: xyz = −28 2 −13, t(11) = 3.77, p < .05, left amygdala: xyz = 20 9 −10, t(11) = 3.28, p < .05). Moreover, in a whole-brain regression analysis, correlating age with brain activation to EA angry faces, we do not find an age-related Telzer et al. 239 D o w n l o a d e d f r o m l l / / / / j t t f / i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e d o u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 5 / 2 2 5 2 / 3 2 4 / 1 2 9 3 4 4 4 / 9 1 4 7 3 7 o 8 c 8 n 2 _ 1 a / _ j 0 o 0 c 3 n 1 1 _ a p _ d 0 0 b 3 y 1 g 1 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 i 2 3 e s / j f / . t u s e r o n 1 7 M a y 2 0 2 1 Figure 3. Children with more diverse peers show dampened amygdala activation to AA faces. increase or decrease in the amygdala. Therefore, across the ages tested, we obtained a stable amygdala response. Amygdala Response to Race as a Function of Neighborhood and Peer Diversity Next, we tested whether racially diverse contexts would modulate the amygdala response to race. Prior work has highlighted the importance of diverse social environ- ments, such as neighborhood and school diversity, in re- ducing racial in- and out-group biases (Bar-Haim et al., 2006; Rutland et al., 2005). Given the specificity of the amygdala to AA faces, we examined whether racial diver- sity of childrenʼs neighborhood and peers would modu- late this amygdala response. In separate whole-brain analyses, we correlated neighborhood and peer diversity with neural activation to AA faces (relative to baseline), controlling for participantsʼ own race. Whereas neighbor- hood diversity was not related to amygdala response to AA faces, greater peer diversity was associated with atten- uated right amygdala response to AA faces (xyz = 16 −2 −8, t(25) = −3.27, p < .05, corrected; Figure 3),1 sug- gesting that more racially homogenous peer groups (re- gardless of racial composition) relate to greater amygdala response to AA faces. These findings suggest that chil- Figure 4. The left amygdala to AA relative to EA faces correlated negatively with mean RT to AA relative to EA faces. Adolescents who matched AA faces more quickly than EA faces showed enhanced amygdala activation to AA relative to EA faces. drenʼs peer environment can shape how race is pro- cessed in the brain. No other brain regions correlated with racial diversity. Finally, given that the amygdala cluster found for peer diversity was in the same region as that found for age, we conducted regression analyses in which we simultaneously entered peer diversity and age to predict amygdala response to AA faces, controlling for participantsʼ own race. Results show that age and peer diversity each independently predicted amygdala activation to AA faces (age: B = 0.29, SE = 0.11, β = .42, p < .05; peer diversity: B = −1.38, SE = 0.55, β = −.41, p < .05). Age accounted for 35.9% of the variance, and peer diversity accounted for an addi- tional 11.3%. Together, age and peer diversity explained nearly half (47.2%) of the amygdala response to AA faces. Neural and Behavioral Response to Race To examine whether the amygdala response to race was related to childrenʼs behavior, we conducted multiple regression analyses in which we examined how the amyg- dala response to AA relative to EA faces predicted partici- pantsʼ mean RT when matching the emotion of AA relative to EA faces. The behavioral bias was calculated by subtracting the standardized mean RT to EA faces from the standardized mean RT to AA faces. Negative scores indicate faster RTs to AA faces and positive scores indicate faster RTs to EA faces. We controlled for age and partici- pantsʼ race. As shown in Figure 4, participants who showed greater activation to AA relative to EA faces in the left amyg- dala were also faster at matching AA relative to EA faces. These behavioral data suggest that amygdala response to AA faces was associated with a decrease in speed in behav- ioral responding to AA faces. DISCUSSION The social environment plays a large role in shaping affec- tive perceptions of race (Bar-Haim et al., 2006). The amyg- dala is involved in nonconscious processing of stimuli that have an acquired emotional significance based on previous experience and plays a role in sensitivity to the salience of environmental cues (Cunningham & Brosch, 2012; Santos et al., 2011; Fitzgerald et al., 2006; Fudge & Emiliano, 2003; 240 Journal of Cognitive Neuroscience Volume 25, Number 2 D o w n l o a d e d f r o m l l / / / / j f / t t i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e d o u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 5 / 2 2 5 2 / 3 2 4 / 1 2 9 3 4 4 4 / 9 1 4 7 3 7 o 8 c 8 n 2 _ 1 a / _ j 0 o 0 c 3 n 1 1 _ a p _ d 0 0 b 3 y 1 g 1 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 i 2 3 e s / j . f / t u s e r o n 1 7 M a y 2 0 2 1 Whalen et al., 2001). Thus, the amygdala is particularly amenable to learning about socially constructed values placed on social groups, such as those about race. We find that the amygdala becomes increasingly sensitive to AA faces across development, with activation to AA faces only becoming significant around 14 years. The heightened amygdala activity to AA faces previously reported in adults (Lieberman et al., 2005; Cunningham et al., 2004; Phelps et al., 2000) is not present during early childhood and only becomes evident during adolescence. Thus, amygdala responsivity to race is likely the result of a developmental process in which the amygdala acquires emotional knowl- edge learned over development, becoming more sensitive to AA faces. This heightened amygdala response to AA faces may reflect learned cultural knowledge, such as implicit and explicit stereotypes. Across development, youth inter- nalize cultural biases and norms in their environment (Apfelbaum et al., 2008). Additionally, this response may reflect the increasing salience of race that occurs during adolescence that is not associated with bias, such as adoles- centsʼ ethnic identity explorations. For example, adoles- cents enter high school where ethnic clubs and coalitions may form and youth begin to explore their ethnic identity (Roberts et al., 1999). Therefore, the amygdala response may reflect increased learning, exploration, and awareness of race. Future research should explore whether cultural biases, awareness or endorsement of stereotypes, or ethnic identity exploration explain the age effect to race found in the current study. Alternatively, the increasing amygdala response to race may be driven by intrinsic factors of the child, such as puberty, rather than exposure to cultural messages. Indeed, prior research has found that puberty is associated with increased amygdala response to emo- tional stimuli (Moore et al., 2012), and pubertal hormones may partly drive the social reorientation of the amygdala during adolescence (Nelson et al., 2005). Future research should examine how pubertal hormones relate to the neural processing of race. Children from both EA and AA backgrounds showed a similar neurodevelopmental increase in the amygdala to AA faces, consistent with behavioral research showing that AA youth internalize socially constructed views held by the dominant culture (Averhart & Bigler, 1997; Spencer & Markstrom-Adams, 1990) and neuroimaging research among AA adults showing heightened amygdala response to AA faces (Lieberman et al., 2005). Individuals from diverse ethnic and racial groups are exposed to similar cul- tural messages, and with age youth may internalize these messages, attaining the cultural knowledge that AA indi- viduals are treated differently. Alternatively, the amygdala response in our AA and EA samples may be tapping differ- ent processes. For the AA participants, the heightened amygdala response may be following a developmental path parallel to AA youthsʼ explorations of their ethnic identity, which increases during high school more so than for EA youth (Phinney, 1996). For the EA participants, the height- ened amygdala response may reflect the development of cultural biases. Thus, race may be salient for each ethnic group but for different reasons, and so the same neuro- developmental activations may be reflecting different underlying processes. Future research should attempt to understand the mechanisms driving the amygdala response in different ethnic populations. The amygdala is involved in the detection of motiva- tionally relevant and salient aspects of oneʼs environment (Cunningham & Brosch, 2012; Santos et al., 2011; Fitzgerald et al., 2006; Fudge & Emiliano, 2003; Whalen et al., 2001). When the amygdala detects salience, via substantial pro- jections to primary and high-order sensory and motor areas of the brain, it guides further neural processing to appropriately respond, potentially impacting behavior (Cunningham & Brosch, 2012; Davis & Whalen, 2001). We observed that children who showed a stronger amyg- dala response to AA faces relative to EA faces were also faster at matching AA faces, suggesting that the heightened amygdala response to AA faces resulted in faster RTs. This finding provides support that the amygdala is involved in detecting salience of the stimulus, which may be part of the process whereby learning affective properties of social stimuli occurs. If the amygdala were responding to negativ- ity, one might expect this to influence childrenʼs behavior through avoidance (i.e., slower RTs to matching AA faces). Thus, within this experimental context, the behavioral data suggest that the amygdala response may be signaling the increasing saliency of AA that accompanies age. Indeed, on its own, heightened salience of social groups increases negative out-group behaviors and positive in-group behav- iors (Patterson & Bigler, 2006; Claeys et al., 2003; Bigler, Brown, & Markell, 2001; Tajfel & Turner, 1979; Tajfel, 1978) and may be the basis for the high correlation be- tween negative appraisals of AAs and amygdala re- sponse that have been observed in adulthood (Phelps et al., 2000). In addition to the amygdala, the fusiform gyrus (FG) and ventrolateral pFC ( VLPFC) were specifically recruited to AA faces as children got older. The FG is a brain region involved in face perception (Haxby, Hoffman, & Gobbini, 1999) and visual expertise (Kanwisher, McDermott, & Chun, 1997). Therefore, as children get older, they may have more experience and exposure to AA individuals, thus developing greater expertise and recruiting the FG to AA faces. The VLPFC is thought to mediate evaluative and reg- ulatory processes and may modulate amygdala reactivity (Passarotti, Sweeny, & Pavuluri, 2009). Moreover, with age individuals are better able to regulate affective responses (Yurgelun-Todd, 2007). Therefore, as children get older, AA individuals may become more emotionally salient, as evidenced by the amygdala response, and the VLPFC may come on-line. Our second goal was to understand how childrenʼs social environment may alter the amygdala response to race by examining childrenʼs peer and neighborhood contexts. The salience of social categories, such as race, varies according to social contexts (Turner, Hogg, Oakes, Reicher, Telzer et al. 241 D o w n l o a d e d f r o m l l / / / / j f / t t i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e d o u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 5 / 2 2 5 2 / 3 2 4 / 1 2 9 3 4 4 4 / 9 1 4 7 3 7 o 8 c 8 n 2 _ 1 a / _ j 0 o 0 c 3 n 1 1 _ a p _ d 0 0 b 3 y 1 g 1 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 i 2 3 e s / j f / . t u s e r o n 1 7 M a y 2 0 2 1 & Wetherell, 1987). Increased racial diversity may reduce the salience of AA faces. Our results revealed that when children had more cross-race friends and schoolmates, they were less likely to exhibit a neural bias to AA faces, consis- tent with a body of work highlighting the benefit of racially diverse schools for decreasing in-group biases ( Juvonen, Nishina, & Graham, 2006; Rutland et al., 2005). This atten- uation of amygdala response suggests that intergroup racial contact may reduce the salience of race. Contact between members of different racial groups may expose children to more diverse views, producing more individuated and per- sonalized relationships across racial groups (Dovidio & Gaertner, 1999). Even for AAs themselves, contact between individuals from diverse backgrounds may reduce the sa- lience of intergroup boundaries (Dovidio & Gaertner, 1999). Thus, interventions designed to reduce the develop- ment of racial biases could focus on providing children with opportunities to interact with individuals from diverse back- grounds, thereby potentially decreasing the salience of race. Interestingly, childrenʼs neighborhood diversity was not related to their neural processing of race. Perhaps neighborhood diversity results in fewer opportunities to interact with individuals of different racial backgrounds compared with diversity in schools, which provide hourly interactions with oneʼs peers. Because our participants spanned a broad age range from 4 to 16 years, it was important to demonstrate that warping to the adult template did not bias the results toward less amygdala activation in younger individuals, thereby driving our race-related developmental effects. We addressed this issue in two ways. First, we created an anatomical average of our developmental participants and overlay it on the adult template. The anatomical average from our developmental population shows that the amygdala region coincides with the adult template. Second, we examined neural activation in the amygdala to angry faces across age and show that we get differen- tial amygdala response in the youngest participants in response to emotional stimuli. In fact, there are no age- related changes in amygdala response to emotional faces; children, young adolescents, and older adolescents all show enhanced activation to angry faces. Moreover, results from our primary analysis show that children across our entire age range evidence stable amygdala activation to EA faces. Together, this suggests that warping the child brains to the adult template did not bias the results toward less amygdala activation in younger children. Recent advances in developmental neuroscience have shown that pediatric and adult neuroimaging data can be analyzed in the same strerotactic space. For instance, Kang, Burgund, Lugar, Petersen, and Schlaggar (2003) and Burgund and colleagues (2002) found that atlas-transformed brain mor- phology, BOLD responses, and locations of functional acti- vation foci are consistent between 7- and 8-year-old children and adults. In conclusion, the findings in the current study demon- strate the continuous functional maturation of the amyg- dala in response to social groups across development spanning a large age range of children from 4 to 16 years. The differential response of the amygdala to AA faces does not emerge until adolescence, suggesting that the in- creasing salience of race across development may shape the functional architecture of the amygdala. Importantly, these findings suggest that neural biases to race are not innate and that race is a social construction, learned over time. Acknowledgments This work was supported by NIMH R01MH091864 (NT). Reprint requests should be sent to Eva H. Telzer, Department of Psychology, University of Illinois, 603 E. Daniel St, Champaign, IL 61820, or via e-mail: ehtelzer@illinois.edu. Note 1. The n in each racial group is too small to warrant a formal separate analysis. However, for descriptive purposes, we pre- sent the findings for EA (n = 8) and AA (n = 10) participants. Although the relationship between peer diversity and amygdala response is not significant for either group alone, both EA (B = −1.12, SE = 1.15, β = −.37) and AA (B = −1.98, SE = 1.04, β = −.56) participants show similar decreases in amygdala response to AA faces with more diverse peers. REFERENCES Aboud, F. E. (2003). The formation of ingroup favoritism and outgroup prejudice in young children. Developmental Psychology, 39, 48–60. Apfelbaum, E. P., Pauker, K., Ambady, N., Sommers, S. R., & Norton, M. I. (2008). Learning (not) to talk about race: When older children underperform in social categorization. Developmental Psychology, 44, 1513–1518. Averhart, C. 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