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Influence of Repressive Coping Style on Cortical Activation during Encoding of Angry Faces PDF

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RESEARCHARTICLE Influence of Repressive Coping Style on Cortical Activation during Encoding of Angry Faces AstridVeronikaRauch1*,LenaterHorst1,VictoriaGabrielePaul1,JochenBauer1, Udo Dannlowski2, Carsten Konrad2, Patricia Ohrmann1, Harald Kugel3, Boris Egloff4, Volker Arolt1, Thomas Suslow5 1.DepartmentofPsychiatry,UniversityofMuenster,Muenster,Germany,2.DepartmentofPsychiatry, UniversityofMarburg,Marburg,Germany,3.DepartmentofClinicalRadiology,UniversityofMuenster, Muenster,Germany,4.DepartmentofPsychology,UniversityofMainz,Mainz,Germany,5.Departmentof PsychosomaticMedicine,UniversityofLeipzig,Leipzig,Germany *[email protected] Abstract Background: Coping plays an important role for emotion regulation in threatening situations. The model of coping modes designates repression and sensitization as two independent coping styles. Repression consists of strategies that shield the individual from arousal. Sensitization indicates increased analysis of the OPENACCESS environment in order to reduce uncertainty. According to the discontinuity Citation:RauchAV,terHorstL,PaulVG,BauerJ, hypothesis, repressors are sensitive to threat in the early stages of information DannlowskiU,etal.(2014)InfluenceofRepressive CopingStyleonCorticalActivationduring processing. While repressors do not exhibit memory disturbances early on, they EncodingofAngryFaces.PLoSONE9(12): manifest weak memory for these stimuli later. This study investigates the e112398.doi:10.1371/journal.pone.0112398 discontinuity hypothesis using functional magnetic resonance imaging (fMRI). Editor:LindaChao,UniversityofCalifornia,San Francisco,UnitedStatesofAmerica Methods: Healthy volunteers (20 repressors and 20 sensitizers) were selected Received:September7,2012 from a sample of 150 students on the basis of the Mainz Coping Inventory. During Accepted:October16,2014 the fMRI experiment, subjects evaluated and memorized emotional and neutral Published:December11,2014 faces. Subjects performed two sessions of face recognition: immediately after the Copyright:(cid:2)2014Rauchetal.Thisisanopen- fMRI session and three days later. accessarticledistributedunderthetermsofthe Results: Repressors exhibited greater activation of frontal, parietal and temporal CreativeCommonsAttributionLicense,which permitsunrestricteduse,distribution,andrepro- areas during encoding of angry faces compared to sensitizers. There were no ductioninanymedium,providedtheoriginalauthor differences in recognition of facial emotions between groups neither immediately andsourcearecredited. after exposure nor after three days. Funding:TheDeutscheForschungsgemeinschaft (DFG)supportedpartofthisresearch(RA2009/1- Conclusions: The fMRI findings suggest that repressors manifest an enhanced 2).Thefundershadnoroleinstudydesign,data collectionandanalysis,decisiontopublish,or neural processing of directly threatening facial expression which confirms the preparationofthemanuscript.Noadditional assumption of hyper-responsivity to threatening information in repression in an externalfundingwasreceivedforthisstudy. early processing stage. A discrepancy was observed between high neural CompetingInterests:Theauthorshavedeclared thatnocompetinginterestsexist. PLOSONE | DOI:10.1371/journal.pone.0112398 December11,2014 1/21 CopingStyleandEncodingofAngryFaces activationinencoding-relevantbrainareasinresponsetoangryfacesinrepressors and no advantage in subsequentmemory for these faces compared to sensitizers. Introduction Coping is the process of attempting to manage the demands created by stressful events [1]. Coping strategies can be adaptive or maladaptive. The development of psychiatric diseases and the maintenance of physical health is influenced by coping, which can also influence interventions by aiming to reduce the health risks of stress [2]. Therefore, knowledge about coping is important for developing a better understanding of psychiatric diseases. The model of coping modes [3,4] describes habitual behavior tendencies in reaction to anxiety-provoking and threatening situations. Referring to this model, the terms ‘repression’ and ‘sensitization’ are introduced as independent coping styles. Repression ischaracterized bycognitive avoidance ofthreatening stimuli in order to shield the organism from distressing stimuli and to prevent an uncontrollable increase in arousal. The behavioral aim of repressors is to divert attention from threat-relevant cues. In contrast, sensitization is associated with vigilance and the ability to tolerate arousal but not uncertainty. Although repressors describe themselves as low-anxious, they exhibit higher than average levels of physiological arousal [5–7] and are at an increased risk of stress related health problems [8]. The use of avoidant coping strategies is associated with a worse outcome in depression and several somatic diseases (for a review, see [2]). Previous coping studies have revealed several differences in memory processing for emotional stimuli. Generally, repressors tend to avoid threats in order to prevent arousal, but the repressive discontinuity hypothesis postulates that, when exposed, repressors are not generally insensitive to threat at an early stage of memory processing. Instead, repressors showed a decline in memory for threateningstimuliafter3days,butnotduringimmediaterecall(30 minutesafter stimulus exposure) [9,10]. However, the discontinuity hypothesis has not been tested in the context of neuroimaging. Thus, the cerebral correlates of encoding remain unclear in repression. Only a few imaging studies have investigated emotion processing in repression. Sander et al. [11] found an increased activation in repressors in left temporoparietal regions during the presentation of sad intonations. Cortical activation was also found to be larger for repressors in prefrontal areas. In a previous study [12] we demonstrated that repressors showed heightened activation in frontal and temporo-occipital areas during the perception of fearful and happy faces. Repressors exhibited an increased connectivity between medial prefrontal areas and the amygdala during the presentation of fearful faces. PLOSONE | DOI:10.1371/journal.pone.0112398 December11,2014 2/21 CopingStyleandEncodingofAngryFaces Facial expressions play an important role in social signaling and for determiningenvironmentalconditions[13,14].Correctlyinterpretingthreatening faces is crucial for detecting environmental danger. So far, emotional faces have not been applied to test the discontinuity hypothesis. Hence, we employed emotional faces in this context. In the past, several functional magnetic resonance imaging (fMRI) studies investigated emotional face processing. Increased activation during the processing of emotional faces was found in several regions, including prefrontal, temporoparietal, limbic, visual areas, amygdala, putamen and cerebellum ((for reviews see [15–17]). A recent study [18] investigated memory for emotional stimuli and revealed several areas that were involved in recognizingemotionalfaces.Theseincludedprefrontalandfrontalregions,aswell as temporal and parietal areas. Encoding of emotional faces was associated with activity in the dorsolateral prefrontal cortex (DLPFC), frontal areas, amygdala, temporo-occipital regions and parietal lobule [19,20]. Taken together,the neurobiological mechanisms ofthe repressive discontinuity hypothesis in the context of processing emotional faces remain unknown. Therefore,theaimofthisfMRIstudyistotestthishypothesisataneurobiological level. Repressors are expected to evaluate emotional faces as less threatening than sensitizers. Based on previous studies it is hypothesized that repressors will show anincreased response tothreateningfaces (i.e.angryandfearfulfacialexpression) in cortical regions which are known to play a role in emotion regulation and memory encoding, such as the prefrontal and temporo-parietal areas. Therefore, repressors’ memory for threatening faces should be better directly after encoding compared to that of sensitizers. Furthermore, it was hypothesized that due to cognitive avoidance repressors will show a decreased memory for threatening faces after three days compared to sensitizers. Methods Participants Forty healthy female, right-handed university students participated in this fMRI study. All subjects had no history of psychiatric, neurological, or other relevant medical diseases, were free of psychotropic medication and had a normal or corrected-to-normal (by contact lenses) vision. Handedness was assessed with the HandednessQuestionnaire[21].VisualacuitywascheckedpriorthefMRIsession by reading a miniature Snellen eye chart. Study participants were selected from a sample of 150 students on the basis of their scores on the German version of the Mainz Coping Inventory, MCI [22]. Twenty persons with high scores on the cognitive avoidance scale (.66th percentile of the screening sample) and low scores on the vigilance scale (,33th percentile) were included as consistent repressors in the present study, whereas 20 persons with high scores on the vigilance scale (.66th percentile of the screening sample) and low scores on the cognitive avoidance scale (,33th percentile) were included as consistent sensitizers. Cronbach’s a was 0.84 for the MCI avoidance scale and 0.89 for the PLOSONE | DOI:10.1371/journal.pone.0112398 December11,2014 3/21 CopingStyleandEncodingofAngryFaces Table1.Descriptivecharacteristicsofstudyparticipants. Sensitizers Repressors Mean SD Mean SD MCIvigilance 28.2 3.3 14.0 2.7 MCIcognitiveavoidance 16.5 4.1 27.5 2.4 Age 22.9 2.3 22.4 2.7 VerbalIQ(MWT-B) 115.8 10.9 117.0 13.0 PictureCompletion(WAIS-R) 14.5 1.3 14.2 1.9 STAI-State 36.0 6.7 31.6 4.7 STAI-Trait 36.1 8.7 29.2 4.2 BDI 3.9 4.7 2.0 2.0 Note.MCI:MainzCopingInventory;WAIS-R:WechslerAdultIntelligenceScale–revisedform;STAI:State-TraitAnxietyInventory,stateandtraitversion; BDI:BeckDepressionInventory. doi:10.1371/journal.pone.0112398.t001 MCI vigilance scale. Repressors differed significantly (Ps,0.001) from sensitizers on both MCI scales (see table 1). Subjects were screened for imaging safety concerns, and gave written informed consent, which was approved by the ethics committee of the Medical Faculty at the University of Muenster. All subjects received a compensation of 35 EUR after their participation. The same subjects participated also in a second experiment which was published previously [23]. In thisexperimentadetection task wasapplied tomeasure automaticresponsivityto facial emotions as a function of coping style. The detection task was always conductedbeforethepresentmemoryexperimentanduseddifferentfacialstimuli [24]. In the detection task participants were shown happy, angry, fearful, and neutral facial expressions masked by neutral faces and they had to guess which of these facial expressions has been briefly presented [23]. Questionnaires and Neuropsychological Measures TheMCI[25]isastimulus-response inventorythatassessesvigilantandcognitive avoidant coping strategies in 4 ego-threatening (e.g., public speaking) and 4 physically threatening situations (e.g., riding with an inexperienced driver). For each situation, 5 vigilant or sensitizing items (e.g., information search, anticipation of negative events) and 5 cognitively avoidant or repressive items (e.g., denial, attention diversion) are administered in a true—false response format. Scored answers are summed for vigilant and avoidant items across all 8 situations. The State-Trait Anxiety Inventory, STAI, [26] was administered in its German version[27],aswellastheBeckDepressionInventory,BDI,[28,29].TheMultiple choice vocabulary test, MWT-B, [30] was applied to assess the verbal intelligence of study participants. Picture Completion is a subtest of the Wechsler Adult Intelligence Scale, WAIS-R, [31] which measures visual perception and recognitionofessentialdetailsofobjects.Statisticalanalyseswereperformedusing SPSS Statistics 17.0.0 (Statistical Package for the Social Sciences, 2008). PLOSONE | DOI:10.1371/journal.pone.0112398 December11,2014 4/21 CopingStyleandEncodingofAngryFaces fMRI Stimulus Material and Procedure The study protocol included an event-related fMRI experiment and two sessions of face recognition tasks outside the scanner after the fMRI session. The first was 30 minutes after, and the second 3 days after. All three tasks included colored facial stimuli (angry, happy, fearful and neutral) [32] of 15 male and 15 female subjects. Facial stimuli were presented pseudo-randomly. Only during the fMRI experiment was a no face condition (a grey rectangle) added to optimize the paradigmwithnullevents[33].InthefMRIexperiment,150trialswerepresented. Each emotional face condition and the no face condition consisted of 30 trials. Thetotaldurationoftheexperimentwas17.5 minutes.Afixationcrossof500 ms preceded the emotion face, which was presented for 3000 ms followed immediately by a time interval (3500 ms) for rating the facial emotion as rather threatening (20.5, 21.5) or rather non-threatening (+0.5, +1.5) by pressing a button. Thus, both negative numbers were accompanied by the adjective ‘‘threatening’’, whereas both positive numbers were accompanied by the adjective ‘‘non-threatening’’. One-half of the samples gave positive responses with the left hand and the other half used the right hand for positive responses. Subjects were instructed to evaluate and memorize the faces. The experiment was programmed usingthesoftwarepackageInquisit[34].Twofixedrandomorderswerepresented which were counterbalanced across subjects. The type of evaluative response and reaction latencies were registered. Face stimuli were presented via a projector (Sharp XG-PC10XE). The head position was stabilized with a vacuum head cushion. The two recognition tasks consisted of 120 original faces shown during the fMRI session and 120 distractors (30 angry, 30 happy, 30 fearful and 30 neutral facesof other individuals [32]).Twofixedrandomordersof faceswerepresented. The order of faces was counterbalanced across subjects. Subjects were instructed to rate whether they have seen the faces during the fMRI experiment or not. Participants rated the facial emotion as original seen during the fMRI session (‘‘Yes’’) or as distractors (‘‘No’’) by pressing a button. Each stimulus was presented until the subject responded. fMRI Data Acquisition and fMRI Data Analyses T * functional data were acquired at a 3-Tesla scanner (Gyroscan Intera 3T, 2 Philips Medical Systems, Best, The Netherlands) using a single shot echoplanar sequence with parameters selected to minimize distortion while retaining adequate signal to noise ratio and T * sensitivity, according to suggestions made 2 by Robinson et al. [35]. Volumes consisting of 36 axial slices were acquired (matrix 642, resolution 3.5 mm * 3.5 mm * 3.5 mm; time repetition 52.5 s, time echo 535 ms, flip angle 590˚) 420 times in an event-related design. T -weighted 1 inversion recovery and a high-resolution T -weighted 3D sequence (isotropic 1 pixel, 0.53 mm) were also acquired. Functional imaging data were motion corrected using a set of 6 rigid body transformations determined for each image. We applied slice timing and images were spatially normalized to standard MNI PLOSONE | DOI:10.1371/journal.pone.0112398 December11,2014 5/21 CopingStyleandEncodingofAngryFaces space (Montreal Neurological Institute) as well as smoothed (Gaussian kernel, 6- mm FWHM) using Statistical Parametric Mapping 5 (SPM5; Wellcome Department of Cognitive Neurology, London, UK, http://www.fil.ion.ucl.ac.uk/ spm)implementedinMatlab7.1.Checksonmotionartifactswereperformedand analyses used the estimated motion parameters as covariates of no interest in the event related design matrix. Statistical analysis was performed by modeling the different conditions (angry, fearful, happy, and neutral) as variables within the context of the general linear model. A whole-brain analysis was conducted to determine which brain regions were differentially activated as a function of coping style. For each emotion condition (fearful, angry, and happy), activation data (t maps) were calculated for subjects relative to the neutral control face condition. Furthermore, activation data of the neutral versus angry contrast were calculated for each subject. Random effects analysis (t-tests for independent samples) was performed to examine brain activation differences between groups (on the contrasts: fearful vs. neutral, angry vs. neutral, happy vs. neutral and neutral vs. angry). Coordinates of significant activations (p,0.05, k550, False Discovery Rate) were converted into Talairach and Tournoux [36] space using the Talairach Daemon [37]. Results Questionnaires and Neuropsychological Measures Repressors and sensitizers did not differ in mean age, verbal intelligence 29 or visual perception and organization ((WAIS-R, (Ps.0.53)). Cronbach’s a for the STAI state version was 0.86. Sensitizers were more anxious than repressors (t (38) 522.42, P,0.05). Cronbach’s a for the STAI trait was 0.92. As expected, sensitizers had higher trait anxiety scores than repressors (t (38) 523.16, P,0.005). Cronbach’s a for the BDI was 0.83. Sensitizers and repressors did not differ on the BDI (P.0.10) (see table 1). Behavioral Performance We conducted t-tests for sum scores of evaluation ratings and latencies for all facial conditions (fearful, angry, happy, and neutral). Only ratings for the neutral faceconditionshowedasignificantdifferencebetweenrepressorsandsensitizers(t (38) 52.10, p50.042), but not for latency (t (38) 51.69, p50.20). Repressors rated neutral faces as less threatening than sensitizers. Repressors tended to rate angry (t (38) 51.59, p50.12) and fearful (t (38) 51.46, p50.15) faces as less threatening than sensitizers, but these results, as for happy faces (t (38) 50.11, p50.92),didnotreachstatisticalsignificance.Inallconditions,sensitizersshowed longer latencies than repressors, but differences were not significant (see table 2). To examine the influence of coping on recognition memory, we carried out 26264ANOVAswithgroupasbetween-groupindependentvariableandtimeof recall(immediatevs.delayedrecall)andfacecondition(fearful,angry,happy,and PLOSONE | DOI:10.1371/journal.pone.0112398 December11,2014 6/21 CopingStyleandEncodingofAngryFaces Table2.Faceevaluationtask(fMRIexperiment):evaluativeratingsandlatenciesasafunctionoffacialexpressionandcopingstyle(lowvaluesindicate moreperceivedthreat). Sensitizers Repressors Mean SD Mean SD Angryfaces Rating 20.9 0.3 20.6 0.6 Latency 639.3 221.4 603.3 158.7 Happyfaces Rating 1.3 0.3 1.3 0.4 Latency 583.8 205.3 526.3 143.3 Fearfulfaces Rating 20.2 0.4 0.0 0.7 Latency 617.9 230.4 544.1 147.1 Neutralfaces Rating 0.3 0.3 0.5 0.5 Latency 587.64 205.6 514.4 145.1 doi:10.1371/journal.pone.0112398.t002 neutral) as repeated measure independent variables. First, we analyzed total numbers of correctly remembered original faces (hits). There were main effects of time (F (1,38) 516.87, p,0.001) and emotion (F (3,36) 54.10, p50.013) for correctly recognized original faces. Subjects showed significantly more hits in the immediate than in the delayed recognition task and the most hits for the happy condition. No effects involving groups were found in the hit condition. Second, analysisoffalsepositivelyidentifieddistractorsdidnotrevealamaineffectfortime or group, buta main effect for emotion was observed(F (3,36)510.29,p,0.001). According to the results of paired t-tests, subjects produced more false alarms for happyandangryfacescomparedtoneutralfaces(p,0.05)(seetable 3fordetails). Additionally,weassessedparticipants’discriminationperformancebycalculatinga discriminationindexH-FA(hitrate(numberofhits/30)–falsealarmrate(number offalsealarms/210))(seetable 4).TheANOVAresultsbasedonthediscrimination index showed significant main effects of time (F (1,38) 518.88, p,0.001) and emotion(F(3,36)53.82,p50.018).Participantsmanifestedabetterdiscrimination performanceimmediatelyafterthefMRIexperimentcomparedtothedelayedrecall conditionandrecognizedhappyfacesbetterthanneutralfaces.Noeffectsinvolving groups were observed for the discrimination index. Whole Brain FMRI Results: Between Group Comparison Brain response to angry faces compared to neutral faces Repressors showed greater activation in the superior, medial, middle and inferior frontalgyrus,superiortemporalgyrus,precuneus,cerebellum,andinferiorparietal lobe relative to sensitizing individuals. No significantly increased brain activation wasfoundinsensitizingindividuals.Alldataaresignificantatp,0.05,k550,False Discovery Rate corrected (FDR). Please see table 5 and Fig. 1 for details. PLOSONE | DOI:10.1371/journal.pone.0112398 December11,2014 7/21 CopingStyleandEncodingofAngryFaces Table3.Immediate(time1)anddelayed(time2)recognitionperformanceasafunctionoffacialexpressionandcopingstyle(numberofcorrectrecognitions (hits)andfalsealarms). Sensitizers Repressors Mean SD Mean SD Hits Angrytime1 15.9 5.9 14.9 6.1 Angrytime2 13.0 6.7 12.1 4.2 Happytime1 16.7 6.6 16.0 6.4 Happytime2 14.4 7.3 11.9 5.8 Fearfultime1 15.7 7.1 12.2 5.5 Fearfultime2 13.7 6.9 10.9 4.8 Neutraltime1 13.9 6.9 13.2 5.0 Neutraltime2 12.7 7.6 10.0 4.9 Falsealarms Angrytime1 7.7 6.8 7.4 4.8 Angrytime2 7.9 7.1 6.9 4.6 Happytime1 7.7 7.7 7.3 6.2 Happytime2 7.7 7.4 6.9 6.3 Fearfultime1 6.0 6.0 5.2 4.8 Fearfultime2 7.3 6.6 5.1 5.0 Neutraltime1 6.2 6.9 5.7 5.3 Neutraltime2 7.0 7.1 5.4 4.2 Note:time15after30minutes;time25after3days. doi:10.1371/journal.pone.0112398.t003 Table4.Discriminationperformance(H(hitrate(numberofhits/30))–FA(falsealarmrate(numberoffalsealarms/210)))asafunctionoffacialexpression andcopingstyle. Sensitizers Repressors Mean SD Mean SD H-FAAngrytime1 0.49 0.17 0.46 0.19 H-FAAngrytime2 0.40 0.20 0.37 0.12 H-FAHappytime1 0.52 0.20 0.50 0.20 H-FAHappytime2 0.44 0.22 0.36 0.17 H-FAFearfultime1 0.49 0.22 0.38 0.17 H-FAFearfultime2 0.42 0.21 0.34 0.15 H-FANeutraltime1 0.43 0.21 0.41 0.15 H-FANeutraltime2 0.39 0.23 0.31 0.15 Note:time15after30minutes;time25after3days. doi:10.1371/journal.pone.0112398.t004 PLOSONE | DOI:10.1371/journal.pone.0112398 December11,2014 8/21 CopingStyleandEncodingofAngryFaces Table5.Brainregionswithheightenedactivationofrepressorscomparedtosensitizersintheangryversusneutralfacecontrast. BrainRegion BAa Hemisphereb MNICoordinates Size Z-scorec x y z Inferiorfrontalgyrus 9 R 60 12 28 633 5.06 Medialfrontalgyrus 6 R 10 222 56 153 3.99 Middlefrontalgyrus 46 L 250 32 20 213 3.98 Superiorfrontalgyrus 10 R 30 60 20 64 3.72 Superiortemporalgyrus 39 R 36 260 30 64 3.87 Inferiorparietallobule 40 L 264 230 28 97 3.61 Precuneus 7 L 26 270 48 760 3.94 Cerebellum Declive L 230 260 230 534 3.96 aBrodmannareas. bL5left,R5right. cCoordinatesofthemaximalpointofactivationandtheassociatedZ-valuesareshown.Theactivationsaresignificantatp,0.05,k550(FDRcorrected). doi:10.1371/journal.pone.0112398.t005 Brain response to fearful faces compared to neutral faces Repressive and sensitizing individuals showed no significant differences in brain activation (p,0.05, k550, FDR). Brain response to happy faces compared to neutral faces No significant differences were found in brain activation between repressive and sensitizing individuals (p,0.05, k550, FDR). Brain response to neutral faces compared to angry faces Sensitizers showed greater activation in the superior, medial, middle and inferior frontal gyrus, superior, middle and inferior temporal gyrus, fusiform gyrus, middle occipital gyrus, precentral gyrus, substantia nigra, thalamus, Fig.1.Between-groupdifferencesinbrainresponsetoangryfacialexpressioncomparedtoneutral faces.Enhancedbrainactivationsofrepressorscomparedtosensitizers(MNIcoordinates).Theactivations aresignificantatp,0.05,k550(FDRcorrected).Reader’srightissubjects’right. doi:10.1371/journal.pone.0112398.g001 PLOSONE | DOI:10.1371/journal.pone.0112398 December11,2014 9/21 CopingStyleandEncodingofAngryFaces hippocampus, precuneus, cerebellum, and inferior parietal lobule relative to repressive individuals. No significantly increased brain activation was found in repressive individuals. All data are significant at p,0.05, k550, False Discovery Rate corrected (FDR). Please see table 6 for details. Correlations between brain response to angry faces with memory for angry faces To explore the relationship between activation in those brain areas where repressorsmanifestedoveractivationcomparedwithsensitizersandrecognitionof angry faces we calculated for each participant mean activation scores averaged across all voxels for each of the eight clusters where group differences were observed. Then Product–moment correlations were calculated between mean cluster activation scores and memory for angry faces at time 1 and time 2 were calculated for sensitizers and repressors separately (see Table 7). Even though no significant correlations betweenbrainresponseandrecognitionperformancewere found interesting differential correlation patterns were observed. Almost all correlationsofbrainactivationwithimmediaterecognitionhadapositivesignfor both study groups. Sensitizers showed also exclusively positive correlations between brain response and delayed recognition butrepressors manifestedin four cases negative correlations with delayed recognition for angry faces. Finally,anindexforlossofdiscriminationaccuracyovertimewascomputedby subtracting discrimination accuracy at time 2 from discrimination accuracy at time1(H-FAtime1–H-FAtime2).Sensitizersshowedfivenegativecorrelations between brain response to angry faces and loss of discrimination accuracy over time. This means that higher activation in response to angry faces at encoding tended to be associated with less memory loss for angry faces over time. Repressors manifested instead exclusively positive correlations between brain response to angry faces and loss of discrimination accuracy. Thus, for repressors higher activation in response to angry faces at encoding tended to be related to more memory loss for angry faces. To compare strength of correlation of brain response with memory loss between groups Fisher’s Z was applied. For the correlation betweenmiddle frontal gyrus activation and memory loss a significant difference between groups was obtained ((z51.64, p50.05, one-tailed). For the correlations of inferior frontal gyrus and superior temporal gyrus activation with memory loss marginally significant differences between groups in height of correlation were observed ((z51.47 and z51.34 p,0.10, one-tailed). Discussion The aim of this study was to test the discontinuity hypothesis using fMRI. We investigated the influence of coping style on brain activation during memory encoding of emotional faces and the relationship between coping style and the recall of threatening and non-threatening faces. To date, the discontinuity hypothesis has been investigated by behavioral studies only. PLOSONE | DOI:10.1371/journal.pone.0112398 December11,2014 10/21

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Department of Clinical Radiology, University of Muenster, aCoordinates of the maximal point of activation within clusters, k5 cluster size. Implicit learning tasks seem to be more appropriate to reveal repressive Rohrmann S, Rohrmann S, Hennig J, Netter P (2002) Manipulation of physiological
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