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Negative stereotype activation alters interaction between neural correlates of arousal, inhibition and cognitive control

Negative stereotype activation alters interaction between neural correlates of arousal,... doi:10.1093/scan/nsr052 SCAN (2012) 7, 771^781 Negative stereotype activation alters interaction between neural correlates of arousal, inhibition and cognitive control 1 1 2 1 Chad E. Forbes, Christine L. Cox, Toni Schmader, and Lee Ryan 1 2 Department of Psychology, University of Arizona, Tucson, AZ, USA and Department of Psychology, University of British Columbia, Vancouver, B.C., Canada Priming negative stereotypes of African Americans can bias perceptions toward novel Black targets, but less is known about how these perceptions ultimately arise. Examining how neural regions involved in arousal, inhibition and control covary when negative stereotypes are activated can provide insight into whether individuals attempt to downregulate biases. Using fMRI, White egalitarian-motivated participants were shown Black and White faces at fast (32 ms) or slow (525 ms) presentation speeds. To create a racially negative stereotypic context, participants listened to violent and misogynistic rap (VMR) in the background. No music (NM) and death metal (DM) were used as control conditions in separate blocks. Fast exposure of Black faces elicited amygdala activation in the NM and VMR conditions (but not DM), that also negatively covaried with activation in prefrontal regions. Only in VMR, however, did amygdala activation for Black faces persist during slow exposure and positively covary with activation in dorsolateral prefrontal cortex while negatively covarying with activation in orbitofrontal cortex. Findings suggest that contexts that prime negative racial stereotypes seem to hinder the downregulation of amygdala activation that typically occurs when egalitarian perceivers are exposed to Black faces. Keywords: stereotypes; stereotype inhibition; implicit and explicit processing; social neuroscience; amygdala; prefrontal cortex INTRODUCTION Using functional magnetic resonance imaging (fMRI) and functional connectivity analyses, the present study tested the Cultural stereotypes seem as ubiquitous in our society as hypothesis that in contexts that prime negative racial stereo- synapses are in our brains. Although our perceptions of types, exposure to black faces will engender prolonged, others can often be biased by these stereotypes, their effects rather than reduced, activation in brain regions implicated can be regulated with time, cognitive resources and motiv- in arousal that alters the interaction between regions impli- ation (Devine et al., 2002; Payne, 2005). Social neuroscience cated in inhibition and control. Such findings would provide research suggests that these processes are subserved by neural support from a cognitive neuroscience perspective for the regions associated with arousal, inhibition and control parameters under which people control their negative (Amodio et al., 2003; Cunningham et al., 2004). Prior re- biases toward racial out-groups. search, however, has only examined these regulatory pro- cesses in neutral contextswhere negative stereotypes are not already activated by other features of the context. But Neural underpinnings of stereotype activation, what happens when a White individual encounters a Black inhibition and control male while negative Black stereotypes are cued by the context Encountering a stereotyped out-group member engenders a (e.g. by racially stereotypic music)? Is the individual moti- fast activation of positive and negative cognitive associations vated and able to regulate activated stereotypes despite the and valenced feelings associated with one’s representation of context justifying that stereotype, or does this situation pro- the group (Devine, 1989; Lepore and Brown, 1997). vide a ‘license to stereotype’ (Crandall and Eshleman, 2003)? Activation of these attitudes and stereotypes can be coun- tered with a slower, more controlled response reflecting an individual’s desire to evaluate the out-group member in a non-biased manner (Wilson and Brekke, 1994; Wegener and Received 22 September 2010; Accepted 4 August 2011 Advance Access publication 27 September 2011 Petty, 1997; Payne, 2005). Recent advances in social neuro- The authors would like to thank Cindy Woolverton, Shawn Marks, and Tanya Nemec for their assistance science suggest that fast and slow cognitive processes with data analysis, and all participants for their time, cooperation, and ability to resist head banging while in (e.g. stereotype activation and subsequent downregulation of the scanner. Financial support for this project was provided by a grant from the Arizona Department of Health Services to the Arizona Alzheimer’s Research Center, Cognition and Neuroimaging Laboratories at the that activation) are neurally, as well as behaviorally, identi- University of Arizona (HB2354). fiable and may interact at multiple speeds of cognitive Correspondence should be addressed to Chad E. Forbes, PhD, Department of Psychology, University of Delaware, 222 Wolf Hall, Newark, DE 19716, USA. E-mail: cforbes@psych.udel.edu processing (Forbes and Grafman, 2010). The Author (2011). Published by Oxford University Press. For Permissions, please email: journals.permissions@oup.com 772 SCAN (2012) C.E. Forbes et al. In particular, activation in the amygdala differentiates re- to be cognitively taxing and dependent on finite metabolic actions to White and Black faces (Hart et al., 2000; Phelps resources (Richeson et al., 2003). The goal of the present study was to test the hypothesis et al., 2000; Lieberman et al., 2005; Ronquillo et al., 2007). Amygdala activity has also been positively correlated with that the amygdala activation cued by novel Black faces would Whites’ implicit associations between ‘Black’ and ‘bad’ not be downregulated when those faces are encountered in a (Phelps et al., 2000) and habituates more slowly to previously context that primes and reinforces violent or hostile negative seen Black vs White faces (Hart et al., 2000). The amyg- racial stereotypes. Past research suggests that individuals’ dala may reflect the immediate and often negative perceptions of out-group members are predicated on the arousal induced by exposure to Black faces that are stereo- situation (Wittenbrink et al., 2001; Blair, 2002). Even manip- typically associated with hostility and aggression (Phelps ulating music played in the background can have robust et al., 2000). effects on stereotype activation and overt perceptions of The amygdala likely relays this arousing information to out-group members (Johnson et al., 2000; Rudman and ventromedial regions of the brain. Medial regions of pre- Lee, 2002). frontal cortex (PFC), including orbitofrontal cortex (OFC), may be particularly important for regulating perceivers’ Hypotheses evaluations of Black targets (Forbes and Grafman, 2010). The OFC is highly interconnected with the amygdala and These findings suggest that contextual cues that sustain negative stereotype activation might either, (i) legitimate lateral PFC (according to non-human primate studies and the application of a negative stereotype to a target out-group human diffusion tensor imaging studies; Amaral and Price, 1984; Ghashghaei and Barbas, 2002; Ghashghaei et al., 2007; individual (i.e. reduce motivation; Crandall and Eshleman, Bracht et al., 2009), and may integrate information pertinent 2003), or (ii) make it more cognitively demanding to down- to negative affect received from the amygdala (Banks et al., regulate what is now an enhanced or prolonged negative 2007) with information relevant to current context and past arousing reaction (i.e. reduce ability). Both routes to the biased response could result in enhanced amygdala activity experiences from the lateral PFC (Rolls, 2008). Specific to to Black (vs White) faces that persist even when the faces are stereotype inhibition, OFC may play a critical role in mod- ulating amygdala activity elicited by exposure to Black faces presented at speeds that normally allow for cognitive control. This primary hypothesis was addressed using fMRI. if that initial bias conflicts with one’s explicit, overarching egalitarian motives represented in lateral PFC (Beer et al., Black and White male faces were presented at fast (32 ms) 2003; Blair, 2004; Elliott and Deakin, 2005; Cunningham and or slow (525 ms) presentation rates to self-reportedly Zelazo, 2007). non-biased White participants while (NM), violent and Downregulation of the amygdala can be achieved by lat- misogynistic rap (VMR) or death metal (DM) played in eral PFC indirectly via the OFC and thalamus, as well as the background. Past research has demonstrated that rap music is stereotypically associated with Blacks more than directly via a dorsolateral pathway in the external capsule (Fuster, 1997; Nolte 2002; Bracht et al., 2009). Whites, whereas rock music is stereotypically associated Cunningham et al. (2004) demonstrated how lateral PFC with Whites more than Blacks (Rentfrow et al., 2009). We regions might downregulate amygdala activation in a social hypothesized that when subjects heard no music in the back- context by manipulating the speed at which Black and White ground findings would replicate Cunningham et al. (2004); faces were presented to self-reportedly non-biased White whereas fast exposure to Black (vs White) faces should elicit increased amygdala activity, slow exposure to Black faces participants, controlling whether faster or slower cognitive processing would be involved. When participants were pre- should elicit increased DLPFC activity in lieu of amygdala sented with a Black face at faster exposure speeds (30 ms), activity. and thus did not have the time to initiate controlled pro- Since contextual support for negative stereotypes of cesses, they exhibited an enhanced amygdala response com- Blacks should elicit a prolonged arousal response to novel pared to implicitly presented White faces. When Black faces out-group members, we further hypothesized increased were presented at slower exposure rates (525 ms), however, amygdala activity in response to both fast and slow exposure to Black faces when subjects are also listening to no music this difference in amygdala activity was not evident. Instead, enhanced activity was seen in the dorsolateral prefrontal (VMR). If this prolonged amygdala response prompts cortex (DLPFC), among other regions, that was correlated (rather than reduces) a motivation to downregulate stereo- with decreases in amygdala activity in response to slower type activation in attempts to perceive out-group members exposure to Black faces. Thus while exposure to Black faces in a non-biased manner, we might expect increases in OFC may elicit an initial, arousing reaction as indexed by and DLPFC activity in response to Black faces (vs White increased amygdala activity, with time and motivation, faces). Functional connectivity analyses will provide a more increased DLPFC activity at slower processing speeds sug- sensitive, direct assessment of the degree to which amygdala gests individuals are able to downregulate this response. activity modulates OFC and DLPFC activity at both fast and Furthermore, the downregulation of this response is thought slow presentation speeds. Stereotypes and neural control SCAN (2012) 773 We included a DM condition to prime negative affect but and to press a mouse button (left or right) corresponding not negative Black stereotypes. Thus, in this context, we did to which side of the crosshair an image appeared. Stimuli not expect differences in amygdala, OFC or DLPFC activa- were randomized and counterbalanced according to stimu- tion when participants were exposed to Black vs White faces lus type and left/right presentation. Each face was presented (regardless of presentation speed). once and masked with its corresponding scrambled, Fourier-transformed image. For the fast presentation condi- tion, each trial began with the presentation of a cross hair in MATERIALS AND METHODS the middle of the screen, followed by the presentation of a Participants Black or White face for 32 ms, and then the presentation of Twenty-three White undergraduates (10 males, 13 females; its Fourier-transform for 525 ms (Morris et al., 1998; Whalen aged 18–21 years) participated for course credit. Participants et al., 1998). For the slow presentation condition, a Black were screened to exclude drug and/or alcohol abuse, neuro- face or White face was presented for 525 ms, followed by its logical disorder, head injury with sequellae, psychiatric ill- Fourier-transform for 32 ms. Trial lengths were held con- ness and contraindications to MRI. The data from two stant at 557 ms. Each block consisted of 96 total trials that participants were excluded due to aberrant artifacts incurred contained 12 each of Black and White faces presented at both during data acquisition. presentation speeds, and 24 fast and slow square trials. All sessions began with the NM block so that baseline reactions Materials to fast and slow Black faces could be obtained. The two In an initial pilot study, White participants (n¼ 20) listened remaining music blocks were counterbalanced. to five songs from the VMR and DM genres and rated each Once participants completed the task, they were asked to song on its valenced feeling, stereotypicality of Black or complete a series of questionnaires outside the scanner, White Americans, tempo and association to violence. Out including questions specific to what they thought the pur- of this analysis, ‘Straight Outta Compton’ by N.W.A. (VMR) pose of the study was and whether they saw fast face primes. and ‘Only One’ by Slipknot (DM) were selected because they Participants were then extensively probed for knowledge or were equivalent in negative affect, perceived violence and suspicion specific to both impressions of true study purposes tempo, but the VMR song was rated more stereotypic of and awareness of fast faces, before being debriefed. No par- Black Americans than the DM song. ticipants reported any connections between the music and race of the faces, nor did anyone report seeing any fast dur- Task and procedure ation faces. As part of a larger study, the experiment was a within-subjects design that used a task similar to that of Image acquisition and analysis Cunningham et al. (2004). Participants believed they were Functional images were acquired on a 3.0T scanner with an serving as a control group for a study investigating neural eight-channel phased-array coil (HD Signa Excite, General relationships between music and spatial processing in pro- Electric, Milwaukee, WI, USA). Using a 2D single shot spiral sopagnosic patients. Participants were exposed to either fast in/spiral out sequence (Glover and Law, 2001; TR¼ 2300 ms, (32 ms) or slow (525 ms) presentations of emotionally neu- TE¼ 30 ms, FOV¼ 240 mm, matrix¼ 64 64), 35 sections tral Black and White male faces. Three fMRI scans were were collected (3 mm, 0.5 mm skip) aligned obliquely in the completed while either NM, VMR or DM played in the coronal plane, tilted 308 anterior of the anterior commis- background over headphones. sure/posterior commissure axis. Audiovisual stimuli were During the task, participants were instructed to focus on a projected through VisuaStim digital goggles and headphones crosshair in the middle of the screen (ISI jittered (Resonance Technologies, Inc.). High-resolution anatomical 1945–2445 ms) presented through goggles in the scanner, 3D SPGR as well as T1-weighted anatomical images were obtained for localization of functional activity and for regis- tration of fMRI data sets to stereotactic space according to the Montreal Neurological Institute (MNI) template. Although rock music is stereotypically associated with Whites, our participants were White and there is no empirical precedent for expecting amygdala activation in response to in-group faces. Thus, we did not Images were reconstructed offline and preprocessed using hypothesize (nor did we find) a symmetrical effect indicating activation of biases to white faces in this Statistical Parametric Mapping software (SPM2; Wellcome condition. 2 Department of Cognitive Neurology, London, UK). All Participants were specifically told that we were interested in facial and object recognition, audio perception and how these processes are related to each other in the brain. They were informed that prosopagnosics have volumes were realigned to the third volume to correct for trouble identifying faces but may also have acute sensitivities to different kinds of music and certain musical movement. The data in each section were interpolated in patterns and that many researchers argue that these deficiencies and/or sensitivities may be due to the audio time to match the acquisition timing of the middle section. perception region of the brain experiencing chronic increased stimulation which in turn causes a decrease in activity in the facial and spatial recognition regions of the brain. Participants thought then that they were part In order to minimize normalization errors due to partial of a control group for prosopagnosics being run in the same study and that their efforts would provide insight brain coverage, spatial normalization parameters were esti- in to how and why certain audio stimuli may affect one’s attempts to accurately identify faces and objects mated by first co-registering each participant’s mean func- that are located in different areas of one’s visual field. In light of this cover story, no participants accurately identified the true purpose of the study. tional image to their T1 anatomical image, warping the T1 774 SCAN (2012) C.E. Forbes et al. image to the standard MNI T1 template, and then applying conditions, and planned contrasts were conducted to assess the resulting normalization parameters to each functional for differences between the VMR condition compared to the image (Ashburner and Friston, 1999). The normalized other two conditions specifically. images were resliced to 3 3 3 mm voxels and smoothed Psychophysiological interaction (PPI) analyses were car- with an isotropic 6 mm FWHM Gaussian kernel. The time ried out in order to determine how activity in the amygdala series in each voxel was high-pass filtered to 1/128 Hz and modulated activity in other brain regions specifically for averaged over all voxels and scans within a session. Black > White faces (Friston et al., 1997; as described in Functional images were analyzed using SPM2 with statis- Das et al., 2005). In those contrasts displaying amygdala ac- tical analyses performed by first modeling neural activity tivation at the group level, time series from individuals’ vol- with a delta function at stimulus onset; stimulus duration umes of amygdala activation (NM fast: n¼ 14; VMR fast: was defined according to length of presentation (for faces, n¼ 12; VMR slow: n¼ 9) were extracted, and PPI models either 32 or 525 ms; Williams et al., 2006). The ensuing were created modeling the interaction with amygdala activity BOLD response was modeled by convolving these delta func- for Black > White faces. Resulting contrast maps were taken tions with a canonical hemodynamic response function to a random effects analysis (one-sample t-test), the results (HRF; Friston et al., 1995). The resulting time courses of which are presented, set at a threshold of P < 0.005 with were downsampled to form covariates in a General Linear a small volume correction and cluster size threshold of three Model. Covariates were modeled for the canonical HRFs of voxels (Friston et al., 1997; Das et al., 2005). the Black and White faces separately for each presentation Stereotactic coordinates were generated in the standard speed in each condition previously described. MNI brain by SPM, and are reported here in MNI space. For manipulation checks, comparisons between the music and NM conditions were carried out, and confirmed Behavioral measures that activation in the auditory cortex was observed within Post-scanning, participants were asked to rate how each song the context of music primes. In addition, comparisons made them feel (1¼ very negative, 7¼ very positive), how between the face and no face (squares) conditions (col- stereotypic of Black Americans each song was (1¼ not lapsed across music) confirmed activation in the fusiform stereotypic, 7¼ very stereotypic), and whether they owned face area. a copy of the song. None of the participants owned a copy of the songs used. To assess explicit prejudice levels, partici- Region of interest and connectivity analyses pants were asked to complete the Modern Racism Scale Contrasts of parameter estimates were used in a second-level (MRS, 4-point scale; McConahay, 1986) and the analysis, which treated participants as a random effect Motivation to Respond Without Prejudice Scale (MRWPS, (one-sample t-test); contrasts of interest taken to the 9-point scale; Plant and Devine, 1998). second-level included fast Black > fast White faces and slow Black > slow White faces in each condition. Results from these analyses are presented for a priori regions of interest RESULTS (ROI) based on previous research as outlined in the intro- Behavioral measures duction (amygdala, OFC and DLPFC) thresholded at Participants reported explicit prejudice levels (M¼ 2.34, P < 0.01 with a small volume correction and cluster size s.d.¼ 0.31) that were significantly below the scale midpoint threshold of three voxels (following methods consistent (3), t(20)¼9.57, P < 0.001. Participants also reported with previous studies on amygdala activity and face process- being more internally motivated to respond without preju- ing but with a more conservative threshold, e.g. Killgore and dice (M¼ 7.82, s.d.¼ 1.32) compared to the scale midpoint Yurgelun-Todd, 2004; Das et al., 2005; Liddell et al., 2005). (5), t(20)¼ 9.77, P < 0.001, and marginally more externally For completeness, contrasts comparing fast White > fast motivated (M¼ 5.81, s.d.¼ 1.90), t(20)¼ 1.95, P¼ 0.06. Black and slow White > slow Black were also carried out. Thus, our sample reported being explicitly non-prejudiced Results are presented in Supplementary Table S1 (significant and motivated to regulate bias. results were only observed in the fast but not slow condition Repeated measures analyses revealed that participants for these comparisons). Anatomical regional masks rated the VMR (M¼ 3.86, s.d.¼ 1.39) and DM (M¼ 3.19, were based on standardized neuroanatomical divisions s.d.¼ 1.69) songs as comparable in the negative affect they (Lancaster et al., 2000; Tzourio-Mazoyer et al., 2002). engendered, F(1, 20)¼ 1.89, P¼ 0.18. Participants also rated We also conducted analyses comparing activation levels the VMR song (M¼ 5.81, s.d.¼ 0.81) as more stereotypic of between the three conditions. Effect size estimates were ex- Blacks than Whites (M¼ 4.00, s.d.¼ 1.82), F(1, 20)¼ 14.14, tracted from the aforementioned regions of interest for each P < 0.01, and the DM song as more stereotypic of Whites participant, focusing on the fast Black face > fast White face and slow Black face > slow White face contrasts. Repeated Caution is stressed in interpretation of these comparisons because the no music condition was always first measures Analyses of Variance (ANOVA) were conducted and there is a tendency for the amygdala response to habituate to stimuli (albeit at a slower rate) over time to assess differences in neural activity between the three (Hart et al., 2000; Olsson et al., 2005). Stereotypes and neural control SCAN (2012) 775 Table 1 Activation for Black compared to White faces at fast (32 ms) presentation speeds in a priori regions of interest Condition L/R T-value MNI No. of voxels Brain Region xy z No music Amygdala L 4.36 20 4 12 22 R 3.24 32 2 18 15 Orbital frontal L 4.83 34 54 279 L 4.28 32 54 0 87 L 4.29 32 54 2 155 R 2.87 26 50 27 R 3.02 28 48 421 R 3.72 28 60 14 112 Dorsolateral PFC L 3.28 48 34 22 20 L 2.77 44 36 22 6 Violent rap Amygdala R 2.71 36 2 24 3 Orbital frontal L 2.69 46 48 65 R 2.94 38 56 0 10 Death metal N/A All activations significant at P < 0.01, small volume corrected with a cluster threshold of three voxels. L, left; R, right; PFC, prefrontal cortex; N/A, not applicable. Fig. 1 Amygdala activation for Black > White faces as a function of fast (32 ms) or slow (525 ms) presentation speeds. (M¼ 4.81, s.d.¼ 1.47) than Blacks (M¼ 2.81, s.d.¼ 1.66), F(1, 20)¼ 17.14, P < 0.01. ROI and connectivity analyses: NM condition As predicted, when participants heard NM in the back- ground, fast exposure to Black faces, compared to White faces, elicited bilateral amygdala activation, replicating Cunningham et al.’s (2004) findings (Figure 1). Greater ac- tivity was also found in bilateral OFC and left DLPFC in response to Black > White faces at fast exposure (Table 1). Furthermore, PPI analyses revealed that amygdala activity negatively covaried with activity in left OFC and bilateral DLPFC (Figure 2 and Table 3). Therefore, even though we observed greater activation for Black compared to White faces in both amygdala and prefrontal regions, further ana- Fig. 2 PPI analyses demonstrating regions of the DLPFC that negatively covaried lyses suggested a more subtle inverse relationship between with amygdala activity in response to fast exposure to Black faces in the (A) No Music these regions. condition and (B) VMR condition. Slower exposure to faces revealed no differential amygdala activity for Black > White faces (Figure 1). Consistent with the interpretation that individuals were able to successfully ROI and connectivity analyses: death metal condition downregulate initial amygdala activity at slower presentation speeds, exposure to Black (vs White) faces for 525 ms elicited When participants received fast exposure to faces during greater activity in right DLPFC (Table 2). Given that there DM, no differential amygdala, OFC or DLPFC activity was was no differential amygdala activation present at slow pres- observed in response to Black > White faces. DM also did not entation speeds, PPI analyses were not conducted in this elicit any differential amygdala activity in response to condition. Black > White faces presented at slower exposure rates 776 SCAN (2012) C.E. Forbes et al. Table 2 Activation for Black compared to White faces at slow (525 ms) Table 3. Regions of activation that positively or negatively covaried with presentation speeds in a priori regions of interest amygdala activation for Black compared to White faces at fast (32 ms) and slow (525 ms) presentation speeds Condition L/R T-value MNI No. of voxels Brain Region Condition L/R T-value MNI No. of voxels xy z Brain Region xy z No Music Dorsolateral PFC R 2.88 28 256 14 32 ms Violent Rap Positive: No Music fast Amygdala R 3.06 22 4 24 3 N/A Orbital frontal L 2.83 38 30 49 Negative: No Music fast Dorsolateral PFC L 2.84 42 2 46 6 Orbital frontal L 3.25 40 32 819 Death metal Dorsolateral PFC L 3.51 18 40 26 59 Orbital frontal R 2.94 42 52 16 11 L 4.62 48 18 42 45 R 3.03 26 56 14 11 L 3.20 32 30 40 21 Dorsolateral PFC R 2.77 38 36 34 5 L 3.71 12 38 44 67 R 2.93 22 44 28 3 All activations significant at P < 0.01, small volume corrected with a cluster threshold R 3.88 22 32 34 84 of three voxels. R 3.71 18 46 28 32 L, left; R, right; PFC, prefrontal cortex. Positive: Violent Rap fast Orbital frontal L 3.36 42 52 268 L 3.07 28 48 46 Dorsolateral PFC L 2.96 22 10 60 5 (Figure 1), but greater activation to Black > White faces was Negative: Violent Rap fast Orbital frontal R 3.46 46 38 14 41 observed in right OFC and right DLPFC (Table 2). In light of Dorsolateral PFC R 4.64 28 26 44 241 there being no differential amygdala activation at either pres- 525 ms entation speed, PPI analyses were not conducted within the Positive: Violent Rap slow DM condition. Thus, exposure to DM appeared to create a Dorsolateral PFC R 3.41 22 24 54 8 Negative: Violent Rap slow context where Black and White faces were perceived in a Orbital frontal L 4.12 30 34 222 similar manner at fast presentation speeds. At slower pres- entation speeds, greater OFC and DLPFC activation to Black All activations significant at P < 0.005, small volume corrected with a cluster thresh- faces is consistent with an interpretation that exposure to old of three voxels. Black faces still cued individuals to engage in more con- L, left; R, right; PFC, prefrontal cortex; N/A, not applicable; Positive, positive covari- ation; Negative, negative covariation. trolled processing. ROI and connectivity analyses: VMR condition NM (Figure 2 and Table 3). At slow exposure rates, these Our key hypotheses concerned patterns of activation in the relationships reversed: amygdala activity to Black > White context of racially stereotypic (VMR) music. Consistent with faces positively covaried with right DLPFC activity, but the NM condition, fast exposure to Black (vs White) faces in negatively covaried with left OFC activity (Figure 3 and the context of VMR elicited greater activity in the right Table 3). We consider possible interpretations of these rela- amygdala (Figure 1) and bilateral OFC (Table 1). tionships below. Examining overall neural responses to Black faces with slow exposure in a negative stereotypic context also revealed ROI comparisons between music conditions a pattern consistent with predictions. Although participants Repeated measures ANOVAs and planned contrasts in the NM condition seemed to have downregulated their (VMR¼ 2, NM¼1, DM¼1) were conducted on effect amygdala response at slow presentation speeds, when listen- size estimates extracted from the ROIs to assess the unique ing to VMR, participants exhibited greater right amygdala effects of negative stereotypic primes on participants’ neural activity during slow exposure to Black compared to White response to the exposure of fast and slow Black faces. We faces (Figure 1). In addition, consistent with the interpret- focus on right lateralized comparisons. ation that participants might still have been motivated to try Amygdala Activity. With respect to fast exposure to Black to control stereotype activation, exposure to VMR also re- faces, we found a main effect for right amygdala activity, sulted in increased left OFC and DLPFC activity for F(1, 20)¼ 5.35, P < 0.01, that resulted from greater amygdala Black > White faces (Table 2). PPI analyses revealed other intriguing points of contrast to 4 Participants in the fMRI experiment also rated each song on how positive or negative it made them feel, how the NM Condition. At fast exposure rates, amygdala activity stereotypic it was of Black or White Americans, tempo, how violent it was and how much they liked each song and each music genre post-experiment. We extracted effect size estimates from the amygdala ROI and for Black > White faces positively covaried with activity in left entered all of the aforementioned variables in to a simultaneous regression analysis predicting amygdala OFC and left DLPFC but negatively covaried with activation activity in response to black faces presented at either fast or slow presentation times. These analyses yielded in a more extensive region of right DLPFC than that seen in no significant effects for any of the variables, all P’s > 0.10. Stereotypes and neural control SCAN (2012) 777 Behavioral follow-up A follow-up behavioral study tested our assumption that prolonged exposure to VMR primes engenders more nega- tively biased perceptions of novel Black faces. After complet- ing a between-subjects version of the same face priming task that our primary sample completed in the scanner, 41 White participants with high motivation to be non- prejudiced (M ¼ 6.81, MD ¼ 6.40; M ¼ 7.73, EMNP EMNP IMNP MD ¼ 7.80) were asked to estimate the number of IMNP Black and White faces they saw during the task that were angry and happy (in actuality, all faces had neutral expres- sions). We hypothesized that participants would estimate having seen a greater number of angry Black faces when making this judgment if they had completed the face prim- ing task while listening to VMR. Given prior evidence that individuals are more likely to perceive anger in out-group faces (Hugenberg and Bodenhausen, 2003; Dunham, 2011), we reasoned that if VMR makes negative racial stereotypes salient (i.e. Rudman and Lee, 2002), this evaluative bias Fig. 3 PPI analyses conducted on amygdala activation in the VMR condition in would be accentuated. response to slow exposure to Black faces revealed (A) negative covariation between A 3 (Condition: NM, VMR and DM) 4 (Face type: the amygdala and OFC (red voxels), but (B) positive covariation between the amyg- Angry Black, Angry White, Happy Black and Happy dala and DLPFC (green voxels). White) mixed factors ANOVA with repeated measures on the latter variable yielded a main effect for condition (P < 0.01) and face type (P < 0.001), which was qualified by activity in the VMR condition compared to the NM and DM a significant interaction, F (1,38)¼ 3.90, P¼ 0.001. conditions, F(1, 20)¼ 7.08, P < 0.02. As predicted, this pat- Participants in the VMR condition reported seeing more tern persisted in response to slow exposure to Black faces, angry black faces (M¼ 67.88, s.d.¼ 31.34) compared to F(1, 20)¼ 5.01, P < 0.02; planned contrasts revealed greater angry white faces (M¼ 53.76, s.d.¼ 33.20; P < 0.001) and amygdala activity in VMR compared to the NM and DM more angry black faces compared to individuals listening conditions, F(1, 20)¼ 5.01, P < 0.04. to either NM (M¼ 21.86, s.d.¼ 27.54; P < 0.001) or DM OFC Activity. Analyses conducted on OFC activity in re- (M¼ 35.00, s.d.¼ 31.62; P < 0.01). There were no differences sponse to fast exposure of Black faces yielded a main effect in percentage of angry black faces reported between partici- for condition, F(1, 20)¼ 3.71, P < 0.04; participants elicited pants in the DM and NM conditions (P’s¼ 0.30). These somewhat more OFC activity in response to Black faces patterns provide further evidence that exposure to VMR when primed with VMR compared to NM and DM, elicits biased perceptions toward novel out-group members F(1, 20)¼ 4.13, P¼ 0.056. Although there was also a main that are consistent with the negative stereotypic prime, des- effect for response to slower exposure of Black faces, pite chronic egalitarian goals. F(1, 20)¼ 12.01, P < 0.001, planned contrasts revealed greater OFC activity in the NM and DM conditions com- DISCUSSION pared to the VMR condition F(1, 20)¼ 12.02, P < 0.01. This study examined whether people attempt to and succeed DLPFC Activity. Finally, analyses conducted on DLPFC at downregulating a biased response to an out-group indi- activity revealed condition main effects in response to both vidual even when the context primes a negative stereotype. fast exposure, F(1, 20)¼ 7.34, P < 0.01, and slow exposure, The findings were consistent with the interpretation that, F(1, 20)¼ 3.20, P¼ 0.05, of Black faces. Planned contrasts although White individuals are successful at controlling an indicated that greater DLPFC activity was elicited in the initial arousal reaction to a Black target in a neutral (NM) NM and DM conditions compared to the VMR condition context, this arousal response is not downregulated in the (F’s > 4.83, P’s < 0.05). presence of negative stereotypical cues (while listening to VMR). Specifically, findings in a neutral context replicated earlier work by Cunningham et al. (2004) by showing that It should be noted that we also found decreased right DLPFC activity in slow White > Black face contrasts in fast exposure to Black faces engendered an enhanced amyg- the VMR condition compared to the NM and DM conditions, suggesting that VMR is generally reducing right DLPFC activity compared to other regions for reasons that are unclear. Conversely, we found greater increases dala response. Functional connectivity analyses extended in left DLPFC activity in slow Black > White face contrasts in the VMR condition compared to the NM and DM past work by indicating that increased amygdala activity at conditions, which is consistent with within condition effects. Nevertheless, it is ultimately difficult to interpret fast presentation speeds was negatively associated with acti- these between condition comparisons because activity in the PFC could stem from different kinds of psychological processing varying by context. vation of the OFC and DLPFC. 778 SCAN (2012) C.E. Forbes et al. In contrast, when faces were encountered in a racially An alternative explanation for the pattern of findings is stereotypic context (via VMR exposure), we observed that VMR reinforces the initial negative response to Black increased amygdala activity in response to fast exposure faces and elicits biased elaboration of it. Crandall and to Black faces as well as increased activation in the OFC Eshleman’s (2003) suppress–justification account of preju- (but not in the DLPFC) that was greater than that found dice suggests that certain stereotyped contexts could lead in Black stereotype-neutral contexts (i.e. NM and DM). Whites to feel justified in viewing Blacks in a negative Furthermore, this increased amygdala activity in response light. Thus, when listening to VMR, controlled processes to Black faces was evident even at slower exposure rates, (as evidenced by increases in the DLPFC) could be recruited suggesting that in this context, the initial arousal elicited to justify rather than suppress biased appraisals of the faces. Other research has found evidence of increased activation in by exposure to a Black face was not downregulated. a large portion of lateral PFC when perceivers apply stereo- Participants in the VMR condition also showed increased types to targets (Mitchell et al., 2009). Further evidence is activation in the left OFC and DLPFC to Black vs White faces. Importantly, these results were unique to situations needed to disambiguate these two accounts; however, the that primed negative stereotypes of Blacks, as these patterns coordinates within DLPFC that covaried with amygdala ac- were not evident when Black faces were presented to partici- tivity in this study were much more consistent with those found in studies involving cognitive control of emotional pants in a negative affective context alone (i.e. the DM con- distractors (e.g. Warren et al., 2010), than they were with dition). In conjunction with the behavioral findings, findings from studies such as Mitchell et al. (2009). these patterns suggest that in negative Black stereotypical A seemingly surprising finding in this study was the evi- contexts, Whites who on average are motivated to be dence for OFC and DLPFC activity in response to fast ex- non-prejudiced, might still react with bias to members of a posure to Black faces. Prior research on the neural substrates racial out-group. involved in prejudice control has suggested that the OFC and Functional connectivity analyses provided further insight DLPFC regulate biases through more explicit or slow pro- into the potential neural dynamics underlying these findings. cessing. We not only observed evidence of overall activation The standard account of down-regulating a negative affective in these regions after fast exposure to black faces, but OFC response suggests a negative relationship between neural re- and DLPFC were differentially, yet reliably, associated with gions associated with control (e.g. DLPFC) and those asso- amygdala activity at both fast and slow presentation speeds. ciated with arousal (e.g. amygdala). When Black faces were Such findings imply that processes involved in monitoring presented at fast exposure rates, there was a negative associ- and control of racial bias might be initiated quite quickly if ation between activation in the amygdala and right DLPFC not automatically, and indeed, research utilizing in both the NM and VMR conditions. However, only in the event-related brain potential (ERP) methodology provides VMR condition did we observe a positive association be- direct evidence that conflict induced by racial primes can tween amygdala activation and activation in both the left initiate neural control mechanisms as early as 50 ms after a OFC and DLPFC to fast presented Black faces. In contrast, given prime (Amodio et al., 2004; Amodio et al., 2008; for a at slow exposure rates amygdala activation to Black recent review, see Bartholow, 2010). (vs White) faces negatively covaried with activation in the To better understand these effects, it could be useful to OFC and positively covaried with activation in the right conceive of automatic/implicit/fast and controlled/explicit/ DLPFC. slow cognitive processes as lying on a temporal continuum There are two possible interpretations for the lack of of control as opposed to representing two orthogonal con- down-regulation observed in a stereotypic context. On the structs (e.g. Cunningham and Johnson 2007; Devine and one hand, exposure to an ongoing stereotypic context could Sharp, 2009; Forbes and Grafman, 2010). The OFC may elicit a prolonged negative response that taxes processing aid in selecting and actively inhibiting neural circuits asso- demands in regions implicated in cognitive control. As a ciated with arousing responses anywhere along this con- result, efforts to control bias are initiated but unsuccessful tinuum, i.e. either quickly or more slowly, possibly at down-regulating affect. This interpretation is consistent through a dynamic interaction between regions such as the with other evidence that reduced executive functioning amygdala and DLPFC (Rule et al., 2002). In conjunction predicts greater biases in response to exposure to out- with known neuroanatomical feedforward and feedback group members (Richeson et al., 2003; Payne, 2005). Our loops, functional and anatomical connectivity between participants were highly motivated to respond without cortical and subcortical regions and neural transmission prejudice and thus likely to engage in bias control speeds on the order of 0.5–50 m/s within the cortex in gen- (Amodio et al., 2003). At the same time, our task did not eral (Fuster, 1997; Buzsaki, 2006), it is difficult to imagine require participants to overtly control stereotypic responses, how the OFC and DLPFC could only be involved in the and we have no direct evidence that attempts to control bias were initiated in the VMR condition. Thus, we cannot Although note that Cunningham et al. (2004) also found increased activity in left superior frontal sulcus conclusively interpret activity in the DLPFC as control in (x¼21, y¼ 30, z¼ 39), a region located in Brodmann’s area 9 and considered to be part of the DLPFC, in particular. the B > W face contrast in the fast condition. Stereotypes and neural control SCAN (2012) 779 dynamic modulation of behavior later in the information that they are activated independently and process different processing stream (e.g. 525 ms or slower). Consistent with aspects of the task. The differential co-activation observed in the different task conditions argues against this interpret- this reasoning, evidence suggests that ventromedial PFC re- gions are equally involved at fast and slow speeds of process- ation, but the possibility cannot be ruled out with current ing of self-relevant schematic material (Rameson et al., methods. 2010), and egalitarian minded individuals activate goals to Using a social neuroscience approach, this study provides regulate their racial biases after subliminal exposure to Black insight into how non-prejudiced individuals sometimes per- face primes (Moskowitz et al., 2000); goals that are likely ceive Black Americans in a negative stereotypic way when represented in medial and lateral PFC regions specifically something as subtle as a rap song is played in the back- ground. Our results suggest that non-prejudiced Whites (Barbey et al., 2009). Thus, our evidence of OFC and might not control a negative response to Black targets en- DLPFC activation after only brief exposure to affectively countered in a negative stereotypic context due to a complex charged primes adds to a growing literature supporting a interaction between neural systems involved in arousal, in- more continuum-based view of fast and slow social cognitive hibition and control. processes. We acknowledge the limitations of drawing strong tem- poral conclusions from these data given the sub-optimal SUPPLEMENTARY DATA temporal resolution of fMRI methodology. Although our Supplementary data are available at SCAN online. scanning parameters were optimized for our rapid event-related design, including a jittered inter-stimulus REFERENCES interval that allowed us to deconvolve the signal, the nature of the hemodynamic response limits the ability to Amaral, D.G., Price, J.L. (1984). Amygdalo-cortical projections in the monkey (Macaca fascicularis). 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Negative stereotype activation alters interaction between neural correlates of arousal, inhibition and cognitive control

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Abstract

doi:10.1093/scan/nsr052 SCAN (2012) 7, 771^781 Negative stereotype activation alters interaction between neural correlates of arousal, inhibition and cognitive control 1 1 2 1 Chad E. Forbes, Christine L. Cox, Toni Schmader, and Lee Ryan 1 2 Department of Psychology, University of Arizona, Tucson, AZ, USA and Department of Psychology, University of British Columbia, Vancouver, B.C., Canada Priming negative stereotypes of African Americans can bias perceptions toward novel Black targets, but less is known about how these perceptions ultimately arise. Examining how neural regions involved in arousal, inhibition and control covary when negative stereotypes are activated can provide insight into whether individuals attempt to downregulate biases. Using fMRI, White egalitarian-motivated participants were shown Black and White faces at fast (32 ms) or slow (525 ms) presentation speeds. To create a racially negative stereotypic context, participants listened to violent and misogynistic rap (VMR) in the background. No music (NM) and death metal (DM) were used as control conditions in separate blocks. Fast exposure of Black faces elicited amygdala activation in the NM and VMR conditions (but not DM), that also negatively covaried with activation in prefrontal regions. Only in VMR, however, did amygdala activation for Black faces persist during slow exposure and positively covary with activation in dorsolateral prefrontal cortex while negatively covarying with activation in orbitofrontal cortex. Findings suggest that contexts that prime negative racial stereotypes seem to hinder the downregulation of amygdala activation that typically occurs when egalitarian perceivers are exposed to Black faces. Keywords: stereotypes; stereotype inhibition; implicit and explicit processing; social neuroscience; amygdala; prefrontal cortex INTRODUCTION Using functional magnetic resonance imaging (fMRI) and functional connectivity analyses, the present study tested the Cultural stereotypes seem as ubiquitous in our society as hypothesis that in contexts that prime negative racial stereo- synapses are in our brains. Although our perceptions of types, exposure to black faces will engender prolonged, others can often be biased by these stereotypes, their effects rather than reduced, activation in brain regions implicated can be regulated with time, cognitive resources and motiv- in arousal that alters the interaction between regions impli- ation (Devine et al., 2002; Payne, 2005). Social neuroscience cated in inhibition and control. Such findings would provide research suggests that these processes are subserved by neural support from a cognitive neuroscience perspective for the regions associated with arousal, inhibition and control parameters under which people control their negative (Amodio et al., 2003; Cunningham et al., 2004). Prior re- biases toward racial out-groups. search, however, has only examined these regulatory pro- cesses in neutral contextswhere negative stereotypes are not already activated by other features of the context. But Neural underpinnings of stereotype activation, what happens when a White individual encounters a Black inhibition and control male while negative Black stereotypes are cued by the context Encountering a stereotyped out-group member engenders a (e.g. by racially stereotypic music)? Is the individual moti- fast activation of positive and negative cognitive associations vated and able to regulate activated stereotypes despite the and valenced feelings associated with one’s representation of context justifying that stereotype, or does this situation pro- the group (Devine, 1989; Lepore and Brown, 1997). vide a ‘license to stereotype’ (Crandall and Eshleman, 2003)? Activation of these attitudes and stereotypes can be coun- tered with a slower, more controlled response reflecting an individual’s desire to evaluate the out-group member in a non-biased manner (Wilson and Brekke, 1994; Wegener and Received 22 September 2010; Accepted 4 August 2011 Advance Access publication 27 September 2011 Petty, 1997; Payne, 2005). Recent advances in social neuro- The authors would like to thank Cindy Woolverton, Shawn Marks, and Tanya Nemec for their assistance science suggest that fast and slow cognitive processes with data analysis, and all participants for their time, cooperation, and ability to resist head banging while in (e.g. stereotype activation and subsequent downregulation of the scanner. Financial support for this project was provided by a grant from the Arizona Department of Health Services to the Arizona Alzheimer’s Research Center, Cognition and Neuroimaging Laboratories at the that activation) are neurally, as well as behaviorally, identi- University of Arizona (HB2354). fiable and may interact at multiple speeds of cognitive Correspondence should be addressed to Chad E. Forbes, PhD, Department of Psychology, University of Delaware, 222 Wolf Hall, Newark, DE 19716, USA. E-mail: cforbes@psych.udel.edu processing (Forbes and Grafman, 2010). The Author (2011). Published by Oxford University Press. For Permissions, please email: journals.permissions@oup.com 772 SCAN (2012) C.E. Forbes et al. In particular, activation in the amygdala differentiates re- to be cognitively taxing and dependent on finite metabolic actions to White and Black faces (Hart et al., 2000; Phelps resources (Richeson et al., 2003). The goal of the present study was to test the hypothesis et al., 2000; Lieberman et al., 2005; Ronquillo et al., 2007). Amygdala activity has also been positively correlated with that the amygdala activation cued by novel Black faces would Whites’ implicit associations between ‘Black’ and ‘bad’ not be downregulated when those faces are encountered in a (Phelps et al., 2000) and habituates more slowly to previously context that primes and reinforces violent or hostile negative seen Black vs White faces (Hart et al., 2000). The amyg- racial stereotypes. Past research suggests that individuals’ dala may reflect the immediate and often negative perceptions of out-group members are predicated on the arousal induced by exposure to Black faces that are stereo- situation (Wittenbrink et al., 2001; Blair, 2002). Even manip- typically associated with hostility and aggression (Phelps ulating music played in the background can have robust et al., 2000). effects on stereotype activation and overt perceptions of The amygdala likely relays this arousing information to out-group members (Johnson et al., 2000; Rudman and ventromedial regions of the brain. Medial regions of pre- Lee, 2002). frontal cortex (PFC), including orbitofrontal cortex (OFC), may be particularly important for regulating perceivers’ Hypotheses evaluations of Black targets (Forbes and Grafman, 2010). The OFC is highly interconnected with the amygdala and These findings suggest that contextual cues that sustain negative stereotype activation might either, (i) legitimate lateral PFC (according to non-human primate studies and the application of a negative stereotype to a target out-group human diffusion tensor imaging studies; Amaral and Price, 1984; Ghashghaei and Barbas, 2002; Ghashghaei et al., 2007; individual (i.e. reduce motivation; Crandall and Eshleman, Bracht et al., 2009), and may integrate information pertinent 2003), or (ii) make it more cognitively demanding to down- to negative affect received from the amygdala (Banks et al., regulate what is now an enhanced or prolonged negative 2007) with information relevant to current context and past arousing reaction (i.e. reduce ability). Both routes to the biased response could result in enhanced amygdala activity experiences from the lateral PFC (Rolls, 2008). Specific to to Black (vs White) faces that persist even when the faces are stereotype inhibition, OFC may play a critical role in mod- ulating amygdala activity elicited by exposure to Black faces presented at speeds that normally allow for cognitive control. This primary hypothesis was addressed using fMRI. if that initial bias conflicts with one’s explicit, overarching egalitarian motives represented in lateral PFC (Beer et al., Black and White male faces were presented at fast (32 ms) 2003; Blair, 2004; Elliott and Deakin, 2005; Cunningham and or slow (525 ms) presentation rates to self-reportedly Zelazo, 2007). non-biased White participants while (NM), violent and Downregulation of the amygdala can be achieved by lat- misogynistic rap (VMR) or death metal (DM) played in eral PFC indirectly via the OFC and thalamus, as well as the background. Past research has demonstrated that rap music is stereotypically associated with Blacks more than directly via a dorsolateral pathway in the external capsule (Fuster, 1997; Nolte 2002; Bracht et al., 2009). Whites, whereas rock music is stereotypically associated Cunningham et al. (2004) demonstrated how lateral PFC with Whites more than Blacks (Rentfrow et al., 2009). We regions might downregulate amygdala activation in a social hypothesized that when subjects heard no music in the back- context by manipulating the speed at which Black and White ground findings would replicate Cunningham et al. (2004); faces were presented to self-reportedly non-biased White whereas fast exposure to Black (vs White) faces should elicit increased amygdala activity, slow exposure to Black faces participants, controlling whether faster or slower cognitive processing would be involved. When participants were pre- should elicit increased DLPFC activity in lieu of amygdala sented with a Black face at faster exposure speeds (30 ms), activity. and thus did not have the time to initiate controlled pro- Since contextual support for negative stereotypes of cesses, they exhibited an enhanced amygdala response com- Blacks should elicit a prolonged arousal response to novel pared to implicitly presented White faces. When Black faces out-group members, we further hypothesized increased were presented at slower exposure rates (525 ms), however, amygdala activity in response to both fast and slow exposure to Black faces when subjects are also listening to no music this difference in amygdala activity was not evident. Instead, enhanced activity was seen in the dorsolateral prefrontal (VMR). If this prolonged amygdala response prompts cortex (DLPFC), among other regions, that was correlated (rather than reduces) a motivation to downregulate stereo- with decreases in amygdala activity in response to slower type activation in attempts to perceive out-group members exposure to Black faces. Thus while exposure to Black faces in a non-biased manner, we might expect increases in OFC may elicit an initial, arousing reaction as indexed by and DLPFC activity in response to Black faces (vs White increased amygdala activity, with time and motivation, faces). Functional connectivity analyses will provide a more increased DLPFC activity at slower processing speeds sug- sensitive, direct assessment of the degree to which amygdala gests individuals are able to downregulate this response. activity modulates OFC and DLPFC activity at both fast and Furthermore, the downregulation of this response is thought slow presentation speeds. Stereotypes and neural control SCAN (2012) 773 We included a DM condition to prime negative affect but and to press a mouse button (left or right) corresponding not negative Black stereotypes. Thus, in this context, we did to which side of the crosshair an image appeared. Stimuli not expect differences in amygdala, OFC or DLPFC activa- were randomized and counterbalanced according to stimu- tion when participants were exposed to Black vs White faces lus type and left/right presentation. Each face was presented (regardless of presentation speed). once and masked with its corresponding scrambled, Fourier-transformed image. For the fast presentation condi- tion, each trial began with the presentation of a cross hair in MATERIALS AND METHODS the middle of the screen, followed by the presentation of a Participants Black or White face for 32 ms, and then the presentation of Twenty-three White undergraduates (10 males, 13 females; its Fourier-transform for 525 ms (Morris et al., 1998; Whalen aged 18–21 years) participated for course credit. Participants et al., 1998). For the slow presentation condition, a Black were screened to exclude drug and/or alcohol abuse, neuro- face or White face was presented for 525 ms, followed by its logical disorder, head injury with sequellae, psychiatric ill- Fourier-transform for 32 ms. Trial lengths were held con- ness and contraindications to MRI. The data from two stant at 557 ms. Each block consisted of 96 total trials that participants were excluded due to aberrant artifacts incurred contained 12 each of Black and White faces presented at both during data acquisition. presentation speeds, and 24 fast and slow square trials. All sessions began with the NM block so that baseline reactions Materials to fast and slow Black faces could be obtained. The two In an initial pilot study, White participants (n¼ 20) listened remaining music blocks were counterbalanced. to five songs from the VMR and DM genres and rated each Once participants completed the task, they were asked to song on its valenced feeling, stereotypicality of Black or complete a series of questionnaires outside the scanner, White Americans, tempo and association to violence. Out including questions specific to what they thought the pur- of this analysis, ‘Straight Outta Compton’ by N.W.A. (VMR) pose of the study was and whether they saw fast face primes. and ‘Only One’ by Slipknot (DM) were selected because they Participants were then extensively probed for knowledge or were equivalent in negative affect, perceived violence and suspicion specific to both impressions of true study purposes tempo, but the VMR song was rated more stereotypic of and awareness of fast faces, before being debriefed. No par- Black Americans than the DM song. ticipants reported any connections between the music and race of the faces, nor did anyone report seeing any fast dur- Task and procedure ation faces. As part of a larger study, the experiment was a within-subjects design that used a task similar to that of Image acquisition and analysis Cunningham et al. (2004). Participants believed they were Functional images were acquired on a 3.0T scanner with an serving as a control group for a study investigating neural eight-channel phased-array coil (HD Signa Excite, General relationships between music and spatial processing in pro- Electric, Milwaukee, WI, USA). Using a 2D single shot spiral sopagnosic patients. Participants were exposed to either fast in/spiral out sequence (Glover and Law, 2001; TR¼ 2300 ms, (32 ms) or slow (525 ms) presentations of emotionally neu- TE¼ 30 ms, FOV¼ 240 mm, matrix¼ 64 64), 35 sections tral Black and White male faces. Three fMRI scans were were collected (3 mm, 0.5 mm skip) aligned obliquely in the completed while either NM, VMR or DM played in the coronal plane, tilted 308 anterior of the anterior commis- background over headphones. sure/posterior commissure axis. Audiovisual stimuli were During the task, participants were instructed to focus on a projected through VisuaStim digital goggles and headphones crosshair in the middle of the screen (ISI jittered (Resonance Technologies, Inc.). High-resolution anatomical 1945–2445 ms) presented through goggles in the scanner, 3D SPGR as well as T1-weighted anatomical images were obtained for localization of functional activity and for regis- tration of fMRI data sets to stereotactic space according to the Montreal Neurological Institute (MNI) template. Although rock music is stereotypically associated with Whites, our participants were White and there is no empirical precedent for expecting amygdala activation in response to in-group faces. Thus, we did not Images were reconstructed offline and preprocessed using hypothesize (nor did we find) a symmetrical effect indicating activation of biases to white faces in this Statistical Parametric Mapping software (SPM2; Wellcome condition. 2 Department of Cognitive Neurology, London, UK). All Participants were specifically told that we were interested in facial and object recognition, audio perception and how these processes are related to each other in the brain. They were informed that prosopagnosics have volumes were realigned to the third volume to correct for trouble identifying faces but may also have acute sensitivities to different kinds of music and certain musical movement. The data in each section were interpolated in patterns and that many researchers argue that these deficiencies and/or sensitivities may be due to the audio time to match the acquisition timing of the middle section. perception region of the brain experiencing chronic increased stimulation which in turn causes a decrease in activity in the facial and spatial recognition regions of the brain. Participants thought then that they were part In order to minimize normalization errors due to partial of a control group for prosopagnosics being run in the same study and that their efforts would provide insight brain coverage, spatial normalization parameters were esti- in to how and why certain audio stimuli may affect one’s attempts to accurately identify faces and objects mated by first co-registering each participant’s mean func- that are located in different areas of one’s visual field. In light of this cover story, no participants accurately identified the true purpose of the study. tional image to their T1 anatomical image, warping the T1 774 SCAN (2012) C.E. Forbes et al. image to the standard MNI T1 template, and then applying conditions, and planned contrasts were conducted to assess the resulting normalization parameters to each functional for differences between the VMR condition compared to the image (Ashburner and Friston, 1999). The normalized other two conditions specifically. images were resliced to 3 3 3 mm voxels and smoothed Psychophysiological interaction (PPI) analyses were car- with an isotropic 6 mm FWHM Gaussian kernel. The time ried out in order to determine how activity in the amygdala series in each voxel was high-pass filtered to 1/128 Hz and modulated activity in other brain regions specifically for averaged over all voxels and scans within a session. Black > White faces (Friston et al., 1997; as described in Functional images were analyzed using SPM2 with statis- Das et al., 2005). In those contrasts displaying amygdala ac- tical analyses performed by first modeling neural activity tivation at the group level, time series from individuals’ vol- with a delta function at stimulus onset; stimulus duration umes of amygdala activation (NM fast: n¼ 14; VMR fast: was defined according to length of presentation (for faces, n¼ 12; VMR slow: n¼ 9) were extracted, and PPI models either 32 or 525 ms; Williams et al., 2006). The ensuing were created modeling the interaction with amygdala activity BOLD response was modeled by convolving these delta func- for Black > White faces. Resulting contrast maps were taken tions with a canonical hemodynamic response function to a random effects analysis (one-sample t-test), the results (HRF; Friston et al., 1995). The resulting time courses of which are presented, set at a threshold of P < 0.005 with were downsampled to form covariates in a General Linear a small volume correction and cluster size threshold of three Model. Covariates were modeled for the canonical HRFs of voxels (Friston et al., 1997; Das et al., 2005). the Black and White faces separately for each presentation Stereotactic coordinates were generated in the standard speed in each condition previously described. MNI brain by SPM, and are reported here in MNI space. For manipulation checks, comparisons between the music and NM conditions were carried out, and confirmed Behavioral measures that activation in the auditory cortex was observed within Post-scanning, participants were asked to rate how each song the context of music primes. In addition, comparisons made them feel (1¼ very negative, 7¼ very positive), how between the face and no face (squares) conditions (col- stereotypic of Black Americans each song was (1¼ not lapsed across music) confirmed activation in the fusiform stereotypic, 7¼ very stereotypic), and whether they owned face area. a copy of the song. None of the participants owned a copy of the songs used. To assess explicit prejudice levels, partici- Region of interest and connectivity analyses pants were asked to complete the Modern Racism Scale Contrasts of parameter estimates were used in a second-level (MRS, 4-point scale; McConahay, 1986) and the analysis, which treated participants as a random effect Motivation to Respond Without Prejudice Scale (MRWPS, (one-sample t-test); contrasts of interest taken to the 9-point scale; Plant and Devine, 1998). second-level included fast Black > fast White faces and slow Black > slow White faces in each condition. Results from these analyses are presented for a priori regions of interest RESULTS (ROI) based on previous research as outlined in the intro- Behavioral measures duction (amygdala, OFC and DLPFC) thresholded at Participants reported explicit prejudice levels (M¼ 2.34, P < 0.01 with a small volume correction and cluster size s.d.¼ 0.31) that were significantly below the scale midpoint threshold of three voxels (following methods consistent (3), t(20)¼9.57, P < 0.001. Participants also reported with previous studies on amygdala activity and face process- being more internally motivated to respond without preju- ing but with a more conservative threshold, e.g. Killgore and dice (M¼ 7.82, s.d.¼ 1.32) compared to the scale midpoint Yurgelun-Todd, 2004; Das et al., 2005; Liddell et al., 2005). (5), t(20)¼ 9.77, P < 0.001, and marginally more externally For completeness, contrasts comparing fast White > fast motivated (M¼ 5.81, s.d.¼ 1.90), t(20)¼ 1.95, P¼ 0.06. Black and slow White > slow Black were also carried out. Thus, our sample reported being explicitly non-prejudiced Results are presented in Supplementary Table S1 (significant and motivated to regulate bias. results were only observed in the fast but not slow condition Repeated measures analyses revealed that participants for these comparisons). Anatomical regional masks rated the VMR (M¼ 3.86, s.d.¼ 1.39) and DM (M¼ 3.19, were based on standardized neuroanatomical divisions s.d.¼ 1.69) songs as comparable in the negative affect they (Lancaster et al., 2000; Tzourio-Mazoyer et al., 2002). engendered, F(1, 20)¼ 1.89, P¼ 0.18. Participants also rated We also conducted analyses comparing activation levels the VMR song (M¼ 5.81, s.d.¼ 0.81) as more stereotypic of between the three conditions. Effect size estimates were ex- Blacks than Whites (M¼ 4.00, s.d.¼ 1.82), F(1, 20)¼ 14.14, tracted from the aforementioned regions of interest for each P < 0.01, and the DM song as more stereotypic of Whites participant, focusing on the fast Black face > fast White face and slow Black face > slow White face contrasts. Repeated Caution is stressed in interpretation of these comparisons because the no music condition was always first measures Analyses of Variance (ANOVA) were conducted and there is a tendency for the amygdala response to habituate to stimuli (albeit at a slower rate) over time to assess differences in neural activity between the three (Hart et al., 2000; Olsson et al., 2005). Stereotypes and neural control SCAN (2012) 775 Table 1 Activation for Black compared to White faces at fast (32 ms) presentation speeds in a priori regions of interest Condition L/R T-value MNI No. of voxels Brain Region xy z No music Amygdala L 4.36 20 4 12 22 R 3.24 32 2 18 15 Orbital frontal L 4.83 34 54 279 L 4.28 32 54 0 87 L 4.29 32 54 2 155 R 2.87 26 50 27 R 3.02 28 48 421 R 3.72 28 60 14 112 Dorsolateral PFC L 3.28 48 34 22 20 L 2.77 44 36 22 6 Violent rap Amygdala R 2.71 36 2 24 3 Orbital frontal L 2.69 46 48 65 R 2.94 38 56 0 10 Death metal N/A All activations significant at P < 0.01, small volume corrected with a cluster threshold of three voxels. L, left; R, right; PFC, prefrontal cortex; N/A, not applicable. Fig. 1 Amygdala activation for Black > White faces as a function of fast (32 ms) or slow (525 ms) presentation speeds. (M¼ 4.81, s.d.¼ 1.47) than Blacks (M¼ 2.81, s.d.¼ 1.66), F(1, 20)¼ 17.14, P < 0.01. ROI and connectivity analyses: NM condition As predicted, when participants heard NM in the back- ground, fast exposure to Black faces, compared to White faces, elicited bilateral amygdala activation, replicating Cunningham et al.’s (2004) findings (Figure 1). Greater ac- tivity was also found in bilateral OFC and left DLPFC in response to Black > White faces at fast exposure (Table 1). Furthermore, PPI analyses revealed that amygdala activity negatively covaried with activity in left OFC and bilateral DLPFC (Figure 2 and Table 3). Therefore, even though we observed greater activation for Black compared to White faces in both amygdala and prefrontal regions, further ana- Fig. 2 PPI analyses demonstrating regions of the DLPFC that negatively covaried lyses suggested a more subtle inverse relationship between with amygdala activity in response to fast exposure to Black faces in the (A) No Music these regions. condition and (B) VMR condition. Slower exposure to faces revealed no differential amygdala activity for Black > White faces (Figure 1). Consistent with the interpretation that individuals were able to successfully ROI and connectivity analyses: death metal condition downregulate initial amygdala activity at slower presentation speeds, exposure to Black (vs White) faces for 525 ms elicited When participants received fast exposure to faces during greater activity in right DLPFC (Table 2). Given that there DM, no differential amygdala, OFC or DLPFC activity was was no differential amygdala activation present at slow pres- observed in response to Black > White faces. DM also did not entation speeds, PPI analyses were not conducted in this elicit any differential amygdala activity in response to condition. Black > White faces presented at slower exposure rates 776 SCAN (2012) C.E. Forbes et al. Table 2 Activation for Black compared to White faces at slow (525 ms) Table 3. Regions of activation that positively or negatively covaried with presentation speeds in a priori regions of interest amygdala activation for Black compared to White faces at fast (32 ms) and slow (525 ms) presentation speeds Condition L/R T-value MNI No. of voxels Brain Region Condition L/R T-value MNI No. of voxels xy z Brain Region xy z No Music Dorsolateral PFC R 2.88 28 256 14 32 ms Violent Rap Positive: No Music fast Amygdala R 3.06 22 4 24 3 N/A Orbital frontal L 2.83 38 30 49 Negative: No Music fast Dorsolateral PFC L 2.84 42 2 46 6 Orbital frontal L 3.25 40 32 819 Death metal Dorsolateral PFC L 3.51 18 40 26 59 Orbital frontal R 2.94 42 52 16 11 L 4.62 48 18 42 45 R 3.03 26 56 14 11 L 3.20 32 30 40 21 Dorsolateral PFC R 2.77 38 36 34 5 L 3.71 12 38 44 67 R 2.93 22 44 28 3 All activations significant at P < 0.01, small volume corrected with a cluster threshold R 3.88 22 32 34 84 of three voxels. R 3.71 18 46 28 32 L, left; R, right; PFC, prefrontal cortex. Positive: Violent Rap fast Orbital frontal L 3.36 42 52 268 L 3.07 28 48 46 Dorsolateral PFC L 2.96 22 10 60 5 (Figure 1), but greater activation to Black > White faces was Negative: Violent Rap fast Orbital frontal R 3.46 46 38 14 41 observed in right OFC and right DLPFC (Table 2). In light of Dorsolateral PFC R 4.64 28 26 44 241 there being no differential amygdala activation at either pres- 525 ms entation speed, PPI analyses were not conducted within the Positive: Violent Rap slow DM condition. Thus, exposure to DM appeared to create a Dorsolateral PFC R 3.41 22 24 54 8 Negative: Violent Rap slow context where Black and White faces were perceived in a Orbital frontal L 4.12 30 34 222 similar manner at fast presentation speeds. At slower pres- entation speeds, greater OFC and DLPFC activation to Black All activations significant at P < 0.005, small volume corrected with a cluster thresh- faces is consistent with an interpretation that exposure to old of three voxels. Black faces still cued individuals to engage in more con- L, left; R, right; PFC, prefrontal cortex; N/A, not applicable; Positive, positive covari- ation; Negative, negative covariation. trolled processing. ROI and connectivity analyses: VMR condition NM (Figure 2 and Table 3). At slow exposure rates, these Our key hypotheses concerned patterns of activation in the relationships reversed: amygdala activity to Black > White context of racially stereotypic (VMR) music. Consistent with faces positively covaried with right DLPFC activity, but the NM condition, fast exposure to Black (vs White) faces in negatively covaried with left OFC activity (Figure 3 and the context of VMR elicited greater activity in the right Table 3). We consider possible interpretations of these rela- amygdala (Figure 1) and bilateral OFC (Table 1). tionships below. Examining overall neural responses to Black faces with slow exposure in a negative stereotypic context also revealed ROI comparisons between music conditions a pattern consistent with predictions. Although participants Repeated measures ANOVAs and planned contrasts in the NM condition seemed to have downregulated their (VMR¼ 2, NM¼1, DM¼1) were conducted on effect amygdala response at slow presentation speeds, when listen- size estimates extracted from the ROIs to assess the unique ing to VMR, participants exhibited greater right amygdala effects of negative stereotypic primes on participants’ neural activity during slow exposure to Black compared to White response to the exposure of fast and slow Black faces. We faces (Figure 1). In addition, consistent with the interpret- focus on right lateralized comparisons. ation that participants might still have been motivated to try Amygdala Activity. With respect to fast exposure to Black to control stereotype activation, exposure to VMR also re- faces, we found a main effect for right amygdala activity, sulted in increased left OFC and DLPFC activity for F(1, 20)¼ 5.35, P < 0.01, that resulted from greater amygdala Black > White faces (Table 2). PPI analyses revealed other intriguing points of contrast to 4 Participants in the fMRI experiment also rated each song on how positive or negative it made them feel, how the NM Condition. At fast exposure rates, amygdala activity stereotypic it was of Black or White Americans, tempo, how violent it was and how much they liked each song and each music genre post-experiment. We extracted effect size estimates from the amygdala ROI and for Black > White faces positively covaried with activity in left entered all of the aforementioned variables in to a simultaneous regression analysis predicting amygdala OFC and left DLPFC but negatively covaried with activation activity in response to black faces presented at either fast or slow presentation times. These analyses yielded in a more extensive region of right DLPFC than that seen in no significant effects for any of the variables, all P’s > 0.10. Stereotypes and neural control SCAN (2012) 777 Behavioral follow-up A follow-up behavioral study tested our assumption that prolonged exposure to VMR primes engenders more nega- tively biased perceptions of novel Black faces. After complet- ing a between-subjects version of the same face priming task that our primary sample completed in the scanner, 41 White participants with high motivation to be non- prejudiced (M ¼ 6.81, MD ¼ 6.40; M ¼ 7.73, EMNP EMNP IMNP MD ¼ 7.80) were asked to estimate the number of IMNP Black and White faces they saw during the task that were angry and happy (in actuality, all faces had neutral expres- sions). We hypothesized that participants would estimate having seen a greater number of angry Black faces when making this judgment if they had completed the face prim- ing task while listening to VMR. Given prior evidence that individuals are more likely to perceive anger in out-group faces (Hugenberg and Bodenhausen, 2003; Dunham, 2011), we reasoned that if VMR makes negative racial stereotypes salient (i.e. Rudman and Lee, 2002), this evaluative bias Fig. 3 PPI analyses conducted on amygdala activation in the VMR condition in would be accentuated. response to slow exposure to Black faces revealed (A) negative covariation between A 3 (Condition: NM, VMR and DM) 4 (Face type: the amygdala and OFC (red voxels), but (B) positive covariation between the amyg- Angry Black, Angry White, Happy Black and Happy dala and DLPFC (green voxels). White) mixed factors ANOVA with repeated measures on the latter variable yielded a main effect for condition (P < 0.01) and face type (P < 0.001), which was qualified by activity in the VMR condition compared to the NM and DM a significant interaction, F (1,38)¼ 3.90, P¼ 0.001. conditions, F(1, 20)¼ 7.08, P < 0.02. As predicted, this pat- Participants in the VMR condition reported seeing more tern persisted in response to slow exposure to Black faces, angry black faces (M¼ 67.88, s.d.¼ 31.34) compared to F(1, 20)¼ 5.01, P < 0.02; planned contrasts revealed greater angry white faces (M¼ 53.76, s.d.¼ 33.20; P < 0.001) and amygdala activity in VMR compared to the NM and DM more angry black faces compared to individuals listening conditions, F(1, 20)¼ 5.01, P < 0.04. to either NM (M¼ 21.86, s.d.¼ 27.54; P < 0.001) or DM OFC Activity. Analyses conducted on OFC activity in re- (M¼ 35.00, s.d.¼ 31.62; P < 0.01). There were no differences sponse to fast exposure of Black faces yielded a main effect in percentage of angry black faces reported between partici- for condition, F(1, 20)¼ 3.71, P < 0.04; participants elicited pants in the DM and NM conditions (P’s¼ 0.30). These somewhat more OFC activity in response to Black faces patterns provide further evidence that exposure to VMR when primed with VMR compared to NM and DM, elicits biased perceptions toward novel out-group members F(1, 20)¼ 4.13, P¼ 0.056. Although there was also a main that are consistent with the negative stereotypic prime, des- effect for response to slower exposure of Black faces, pite chronic egalitarian goals. F(1, 20)¼ 12.01, P < 0.001, planned contrasts revealed greater OFC activity in the NM and DM conditions com- DISCUSSION pared to the VMR condition F(1, 20)¼ 12.02, P < 0.01. This study examined whether people attempt to and succeed DLPFC Activity. Finally, analyses conducted on DLPFC at downregulating a biased response to an out-group indi- activity revealed condition main effects in response to both vidual even when the context primes a negative stereotype. fast exposure, F(1, 20)¼ 7.34, P < 0.01, and slow exposure, The findings were consistent with the interpretation that, F(1, 20)¼ 3.20, P¼ 0.05, of Black faces. Planned contrasts although White individuals are successful at controlling an indicated that greater DLPFC activity was elicited in the initial arousal reaction to a Black target in a neutral (NM) NM and DM conditions compared to the VMR condition context, this arousal response is not downregulated in the (F’s > 4.83, P’s < 0.05). presence of negative stereotypical cues (while listening to VMR). Specifically, findings in a neutral context replicated earlier work by Cunningham et al. (2004) by showing that It should be noted that we also found decreased right DLPFC activity in slow White > Black face contrasts in fast exposure to Black faces engendered an enhanced amyg- the VMR condition compared to the NM and DM conditions, suggesting that VMR is generally reducing right DLPFC activity compared to other regions for reasons that are unclear. Conversely, we found greater increases dala response. Functional connectivity analyses extended in left DLPFC activity in slow Black > White face contrasts in the VMR condition compared to the NM and DM past work by indicating that increased amygdala activity at conditions, which is consistent with within condition effects. Nevertheless, it is ultimately difficult to interpret fast presentation speeds was negatively associated with acti- these between condition comparisons because activity in the PFC could stem from different kinds of psychological processing varying by context. vation of the OFC and DLPFC. 778 SCAN (2012) C.E. Forbes et al. In contrast, when faces were encountered in a racially An alternative explanation for the pattern of findings is stereotypic context (via VMR exposure), we observed that VMR reinforces the initial negative response to Black increased amygdala activity in response to fast exposure faces and elicits biased elaboration of it. Crandall and to Black faces as well as increased activation in the OFC Eshleman’s (2003) suppress–justification account of preju- (but not in the DLPFC) that was greater than that found dice suggests that certain stereotyped contexts could lead in Black stereotype-neutral contexts (i.e. NM and DM). Whites to feel justified in viewing Blacks in a negative Furthermore, this increased amygdala activity in response light. Thus, when listening to VMR, controlled processes to Black faces was evident even at slower exposure rates, (as evidenced by increases in the DLPFC) could be recruited suggesting that in this context, the initial arousal elicited to justify rather than suppress biased appraisals of the faces. Other research has found evidence of increased activation in by exposure to a Black face was not downregulated. a large portion of lateral PFC when perceivers apply stereo- Participants in the VMR condition also showed increased types to targets (Mitchell et al., 2009). Further evidence is activation in the left OFC and DLPFC to Black vs White faces. Importantly, these results were unique to situations needed to disambiguate these two accounts; however, the that primed negative stereotypes of Blacks, as these patterns coordinates within DLPFC that covaried with amygdala ac- were not evident when Black faces were presented to partici- tivity in this study were much more consistent with those found in studies involving cognitive control of emotional pants in a negative affective context alone (i.e. the DM con- distractors (e.g. Warren et al., 2010), than they were with dition). In conjunction with the behavioral findings, findings from studies such as Mitchell et al. (2009). these patterns suggest that in negative Black stereotypical A seemingly surprising finding in this study was the evi- contexts, Whites who on average are motivated to be dence for OFC and DLPFC activity in response to fast ex- non-prejudiced, might still react with bias to members of a posure to Black faces. Prior research on the neural substrates racial out-group. involved in prejudice control has suggested that the OFC and Functional connectivity analyses provided further insight DLPFC regulate biases through more explicit or slow pro- into the potential neural dynamics underlying these findings. cessing. We not only observed evidence of overall activation The standard account of down-regulating a negative affective in these regions after fast exposure to black faces, but OFC response suggests a negative relationship between neural re- and DLPFC were differentially, yet reliably, associated with gions associated with control (e.g. DLPFC) and those asso- amygdala activity at both fast and slow presentation speeds. ciated with arousal (e.g. amygdala). When Black faces were Such findings imply that processes involved in monitoring presented at fast exposure rates, there was a negative associ- and control of racial bias might be initiated quite quickly if ation between activation in the amygdala and right DLPFC not automatically, and indeed, research utilizing in both the NM and VMR conditions. However, only in the event-related brain potential (ERP) methodology provides VMR condition did we observe a positive association be- direct evidence that conflict induced by racial primes can tween amygdala activation and activation in both the left initiate neural control mechanisms as early as 50 ms after a OFC and DLPFC to fast presented Black faces. In contrast, given prime (Amodio et al., 2004; Amodio et al., 2008; for a at slow exposure rates amygdala activation to Black recent review, see Bartholow, 2010). (vs White) faces negatively covaried with activation in the To better understand these effects, it could be useful to OFC and positively covaried with activation in the right conceive of automatic/implicit/fast and controlled/explicit/ DLPFC. slow cognitive processes as lying on a temporal continuum There are two possible interpretations for the lack of of control as opposed to representing two orthogonal con- down-regulation observed in a stereotypic context. On the structs (e.g. Cunningham and Johnson 2007; Devine and one hand, exposure to an ongoing stereotypic context could Sharp, 2009; Forbes and Grafman, 2010). The OFC may elicit a prolonged negative response that taxes processing aid in selecting and actively inhibiting neural circuits asso- demands in regions implicated in cognitive control. As a ciated with arousing responses anywhere along this con- result, efforts to control bias are initiated but unsuccessful tinuum, i.e. either quickly or more slowly, possibly at down-regulating affect. This interpretation is consistent through a dynamic interaction between regions such as the with other evidence that reduced executive functioning amygdala and DLPFC (Rule et al., 2002). In conjunction predicts greater biases in response to exposure to out- with known neuroanatomical feedforward and feedback group members (Richeson et al., 2003; Payne, 2005). Our loops, functional and anatomical connectivity between participants were highly motivated to respond without cortical and subcortical regions and neural transmission prejudice and thus likely to engage in bias control speeds on the order of 0.5–50 m/s within the cortex in gen- (Amodio et al., 2003). At the same time, our task did not eral (Fuster, 1997; Buzsaki, 2006), it is difficult to imagine require participants to overtly control stereotypic responses, how the OFC and DLPFC could only be involved in the and we have no direct evidence that attempts to control bias were initiated in the VMR condition. Thus, we cannot Although note that Cunningham et al. (2004) also found increased activity in left superior frontal sulcus conclusively interpret activity in the DLPFC as control in (x¼21, y¼ 30, z¼ 39), a region located in Brodmann’s area 9 and considered to be part of the DLPFC, in particular. the B > W face contrast in the fast condition. Stereotypes and neural control SCAN (2012) 779 dynamic modulation of behavior later in the information that they are activated independently and process different processing stream (e.g. 525 ms or slower). Consistent with aspects of the task. The differential co-activation observed in the different task conditions argues against this interpret- this reasoning, evidence suggests that ventromedial PFC re- gions are equally involved at fast and slow speeds of process- ation, but the possibility cannot be ruled out with current ing of self-relevant schematic material (Rameson et al., methods. 2010), and egalitarian minded individuals activate goals to Using a social neuroscience approach, this study provides regulate their racial biases after subliminal exposure to Black insight into how non-prejudiced individuals sometimes per- face primes (Moskowitz et al., 2000); goals that are likely ceive Black Americans in a negative stereotypic way when represented in medial and lateral PFC regions specifically something as subtle as a rap song is played in the back- ground. Our results suggest that non-prejudiced Whites (Barbey et al., 2009). Thus, our evidence of OFC and might not control a negative response to Black targets en- DLPFC activation after only brief exposure to affectively countered in a negative stereotypic context due to a complex charged primes adds to a growing literature supporting a interaction between neural systems involved in arousal, in- more continuum-based view of fast and slow social cognitive hibition and control. processes. We acknowledge the limitations of drawing strong tem- poral conclusions from these data given the sub-optimal SUPPLEMENTARY DATA temporal resolution of fMRI methodology. Although our Supplementary data are available at SCAN online. scanning parameters were optimized for our rapid event-related design, including a jittered inter-stimulus REFERENCES interval that allowed us to deconvolve the signal, the nature of the hemodynamic response limits the ability to Amaral, D.G., Price, J.L. (1984). Amygdalo-cortical projections in the monkey (Macaca fascicularis). 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Journal

Social Cognitive and Affective NeuroscienceOxford University Press

Published: Oct 27, 2012

Keywords: stereotypes stereotype inhibition implicit and explicit processing social neuroscience amygdala prefrontal cortex

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