Neural mechanisms mediating optimism bias. Positive Negative t-test (P) Correlation with optimism

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1 oi:.8/nture68 Neurl mehnisms meiting optimism is Tli Shrot,, Alison M. Riri, Cne M. Rio & Elizeth A. Phelps Humns expet positive events in the future even when there is no eviene to support suh expettions. For exmple, people expet to live longer n e helthier thn verge, they unerestimte their likelihoo of getting ivore, n overestimte their prospets for suess on the jo mrket. We exmine how the rin genertes this pervsive optimism is. Here we report tht this teneny ws relte speifilly to enhne tivtion in the mygl n in the rostrl nterior ingulte ortex when imgining positive future events reltive to negtive ones, suggesting key role for res involve in monitoring emotionl sliene in meiting the optimism is. These re the sme regions tht show irregulrities in epression, whih hs een relte to pessimism. Aross iniviuls, tivity in the rostrl nterior ingulte ortex ws orrelte with trit optimism. The urrent stuy highlights how the rin my generte the teneny to engge in the projetion of positive future events, suggesting tht the effetive integrtion n regultion of emotionl n utoiogrphil informtion supports the projetion of positive future events in helthy iniviuls, n is relte to optimism. People ten to mke overly onfient, positive preitions out the future, whih re often inurte,. The teneny to expet goo things in the future is known s optimism. Extreme optimism n e hrmful s it n promote n unerestimtion of risk n poor plnning 5. In ontrst, pessimisti view is orrelte with severity of epression symptoms. A moerte optimisti illusion, however, n motivte ptive ehviour in the present towrs future gol, n hs een relte to mentl 6 n physil 7 helth. How oes the helthy rin generte the teneny to rete imges of positive future events? It hs een suggeste tht imgining future events emns system tht lls on the pst to retrieve piees of informtion tht re then reonstrute to form representtions of possile future senrios 8,9. In orne with this proposl, it hs een shown tht imgining the future epens on the sme neurl networks tht re tive when relling the pst 8,,. One possiility is tht in optimists this network is more tive when imgining future positive events thn negtive ones. It is lso possile tht other rin res moulte this network in wy tht ises it to engge in positive future projetions. To exmine the neuroiologil sis of optimism we ollete funtionl mgneti resonne imging (fmri) t while prtiipnts thought of utoiogrphil events relte to esription of life episoe (for exmple, winning n wr or the en of romnti reltionship ). The wor pst or future inite if they shoul think of n event tht ourre in the pst or one tht might our in the future. Trils were lssifie into positive, negtive n neutrl oring to prtiipnts rtings. The men numer of trils rte s neutrl ws negligile, n thus ll ontrsts of interest were onute etween positive n negtive trils only. After snning, prtiipnts rte their memories n projetions on six ftors relte to their sujetive experiene. The men sores for these rtings n sttistil nlysis re presente in Tle n Supplementry Tle. Finlly, prtiipnts omplete the LOT-R (Life Orienttion Test-Revise) sle tht mesures trit optimism (see Methos for etils). As represente in Tle n Supplementry Tle, future positive events were rte s more positive thn pst positive events, n were imgine to e loser in temporl proximity then future negtive events n ll pst events (Fig. ). Negtive future events were experiene with weker sujetive sense of pre-experiening, n were more likely to e imgine from n outsier viewing in, thn positive future events n ll pst events (Fig. ). The more optimisti prtiipnts were, s inite y the LOT-R sores, the more likely they were to expet positive events to hppen loser in the future thn negtive events, n to experiene them with greter sense of pre-experiening (Fig., ). Next we exmine the fmri t to ientify the unerlying neurl mehnisms. Funtionl regions of interest (ROIs) were ientifie y ontrsting ll trils with fixtion (P,.5, unorrete). The four regions tht survive this threshol (Fig., ) inlue the rostrl nterior ingulte ortex (racc, Fig. ), extening into the ventrl meil prefrontl ortex t more lenient threshol (see Fig., ), the posterior ingulte ortex (, Fig. ), n the orsl meil prefrontl ortex (Fig. ), ll of whih hve een reporte to hve key role in retrievl of utoiogrphil memory n in imgining future events 8. The fourth region ws the right mygl, whih hs een shown to e importnt for emotion s Tle Mens n sttistis for sujetive rtings oring to tril type Sle Pst Negtive t-test (P) Correltion with optimism Negtive t-test (P) Correltion with optimism Vlene..,. NS.77.7,. NS Arousl NS.5.6 NS NS Time of event.7.8,.5,.5.. NS NS Sujetive temporl istne..7 NS NS.. NS NS Perspetive.96.7,.5 NS.9.9 NS NS Reliving or pre-experiening.59.7,.5, NS NS Viviness.66.8 NS NS.9.9 NS NS Vlene (, positive; 6, negtive), rousl (, low; 6, high), time of event (yers from present, 5), sujetive temporl istne (, ner; 6, fr), perspetive (, involve; 6, oserver), reliving or preexperiening (, low; 6, high) n viviness (, low; 6, high). NS, not signifint. Deprtment of Psyhology, New York University, New York, USA. Wellome Deprtment of Imging Neurosiene, Institute of Neurology, University College Lonon, Lonon WCN BG, UK.

2 Time from present (yers) 5 Time in future from present, positive negtive (yers) Negtive Pst Sujetive intensity 5 Pre-experiening, positive negtive Negtive Pst Figure relte to expete time of event n sense of preexperiening future events., events re pereive to e loser in time to the present thn negtive future events n ll pst events., Negtive future events re experiene with weker sense of preexperiening thn positive future events n ll pst events., The ifferene in expete time of event for future positive n negtive events orrelte with the optimism level efine y rnking prtiipnts oring to their LOT-R sores (r 5.7, P,.5)., The ifferene in pre-experiening for positive n negtive events orrelte with trit optimism (r 5 8, P,.5). N 5 5; error rs, 6s.e.m.; P,.5. influene on utoiogrphil memory (Fig. ; the left mygl ws lso oserve t more lenient threshol, Fig., ). Among these ROIs, the loo-oxygention-level-epenent (BOLD) signl in the mygl n racc ws reue when imgining negtive future events reltive to positive future events n to ll pst events (Fig., ). The ifferenes in BOLD signl etween positive n negtive future events remine signifint when ontrolling for ifferenes in the sense of pre-experiening (mygl, P,.; racc, P,.5). For struturl ROI nlysis (Supplementry Fig. ) n explortory whole-rin nlysis, see Supplementry Informtion. To exmine if these ifferenes were relte to optimism ross iniviuls, we ientifie voxels within the four funtionl ROIs in whih hnges in BOLD signl uring future positive trils reltive to future negtive trils were orrelte with prtiipnts optimism sore on the LOT-R sle (see Supplementry Informtion for itionl nlysis n isussion). We foun positive orreltion in the racc (Fig. e). No signifint orreltion with optimism ws oserve in the other ROIs. Finlly, funtionl onnetivity nlysis revele strong orreltion etween tivity in the racc n tivity in the mygl ilterlly while imgining future positive events; this orreltion ws weker n less extensive when imgining future negtive events (Fig. f). Our ehviourl results suggest tht, wheres the pst is onstrine, the future is open to interprettion llowing sujets to istne themselves from possile negtive events n move loser towr positive ones. Aross iniviuls, this teneny ws ssoite with trit optimism. The rin imging finings provie possile mehnism meiting the ehviourl oservtions. Signl hnge (%) Signl hnge (%) e Signl hnge in the racc (%) future positive future negtive Negtive Negtive Pst Pst f Funtionl onnetivity with racc R R Amygl y = 9 DMPFC racc x = positive negtive Amygl, y = 9 L L Figure Ativity in the mygl n racc n its reltion to optimism. ROIs efine y ontrsting ll trils with fixtion (shown t P,.5, unorrete) revel tivity of, the mygl (Tlrih oorintes of pek voxel:, 9, ) n, the racc (Bromn Are (BA) :,, ), the (BAs n :, 7, 9) n the orsl meil prefrontl ortex (DMPFC; BA 6: 8,, 5). BOLD signls in oth, the mygl n, the racc reue while imgining negtive future events. e, BOLD signl in the racc uring future positive events versus future negtive events orrelte with trit optimism (9, 9, ; r 5, P,.5). f, Funtionl onnetivity mp with racc s the see revels extensive orreltion with the mygl signl uring future positive trils, n restrite orreltion uring future negtive trils. The re squre in inites the region shown in f. N 5 5; error rs, 6s.e.m.; P,.5.

3 VMPFC DMPFC Amygl, R Pst events Amygl, L VMPFC Amygl, R events Amygl, L Figure Brin tivity uring pst n future trils. Regions tht emerge when ontrsting,, pst trils with fixtion n,, future trils with fixtion t threshol of P,.5, unorrete, revel enggement of regions shown previously 8 to e involve in utoiogrphil memory n future projetion inluing ventrl meil prefrontl ortex (VMPFC) n the (, ), s well s the mygl ilterlly (, ). Reue BOLD signl ws oserve in the mygl n racc uring imgintion of negtive future events reltive to positive future events n ll pst events, suggesting tht the optimism is my e relte to reution in negtive future thought. The mygl hs oumente role in the moultion y emotion of ognitive proesses inluing memory n eision mking (for review, see ref. 5). Our t exten the role of the mygl to inlue simultion of future emotionl events. Previous stuies exmining the neurl mehnisms unerlying imgintion of nonemotionl future events i not oserve mygl involvement 8,. Thus, s in the se of memory, we suggest tht the mygl my e seletively engge in imgining future emotionl events, rther thn serving generl funtion in imgining the future. The racc hs strong reiprol onnetion with the mygl n other regions tht onvey emotionl n motivtionl informtion 6. It hs een implite in tsks involving self-refletion, suh s mking positive self-referentil jugments 7, refleting on hopes n rems 8, initing preferenes 9 n juging the trustworthiness of others. This hs use some reserhers to suggest tht when informtion is juge to e self-relevnt, tivity in the racc will onvey the vlene 7. However, others hve suggeste more generl role for the racc in ssessing the sliene of emotionl n motivtionl informtion n regulting emotionl responses oringly. Consistent with this notion it hs een shown tht the racc s response to positive n negtive stimuli reflets situtionl speifiity. For exmple, oth the racc n mygl were more sensitive to positive stimuli in sujets tht fouse on otining gols (promotionl ontext) n to negtive stimuli in sujets tht fouse on voiing filure (prevention ontext). Furthermore, hnges in racc tivity hve een relte to extintion of fer onitioning, n to reue nxiety uring shok expetny ; this possily inites iminishe sliene of negtive stimuli tht my onsequently lower mygl tivity. Consistent with these previous stuies, we suggest tht in the urrent stuy the racc is trking the sujetive sliene of the stimuli y ssessing emotionl, motivtionl n utoiogrphil informtion, n possily y regulting suh signls. This suggestion is supporte y the strong funtionl onnetivity oserve etween the racc n mygl uring imgintion of positive future events. In ition, iniviul ifferenes in trit optimism were relte to the reltive level of racc enggement when imgining positive future senrios versus negtive ones. We speulte tht racc tivity uring imgintion of future events reflets self-regultory fous tht unerlies is in ttention n vigilne towrs positive future events n wy from negtive ones, moulting enggement of other regions tht provie emotionl n utoiogrphil informtion tht is proesse n reomine to onstrut future senrios. These finings my provie insight to the mehnisms unerlying epression. Depressive symptoms re ssoite with pessimism n with iffiulties in reting etile imges of future events 5. It hs een suggeste tht mlfuntion of neurl pthwy inorporting the racc n the mygl my use epression y leing to erese regultory ffets of the racc over the mygl n other regions involve in emotionl proessing. stuies re neee to etermine whether these normlities re relte iretly to the rekown of optimism in epression, speifilly euse it reltes to the wy in whih epresse ptients imgine future events. One shoul note tht when ompring pst n future trils we re lso ompring imgining n rememering. Thus, our t nnot inite whether the positivity is is funtion of time (tht is, it will emerge only when thinking out the future) or whether it reflets teneny to engge in positive thought when not onstrine y relity. We speulte tht the ifferene is not etween pst n future, lthough in prtie the optimism is mnifests itself in thoughts of the future euse the sene of ftul onstrints leves plenty of room to lter expettions. Expeting positive events, n generting ompelling mentl imges of suh events, my serve n ptive funtion y motivting ehviour in the present towrs future gol. We o not suggest tht simulting negtive events is not importnt for survivl. However, ponering on suh events my interfere with ily tivities y promoting negtive effets suh s nxiety n epression. The urrent stuy highlights how the rin my generte the teneny to engge in the projetion of positive future events, suggesting tht the racc moultes tivity in rin res tht re involve in emotionl proessing n utoiogrphil retrievl to rete positive imges of the future. METHODS SUMMARY Funtionl MRI imges were ollete from prtiipnts while they thought of utoiogrphil events relte to esription of life episoe tht ppere on sreen for s. Either the wor pst or future inite if they shoul think of n event tht ourre in the pst or tht might our in the future. The prtiipnts were instrute to press utton one the memory or projetion ws eginning to form in their min, n gin when they finishe elorting on it. They then h s to rte the memory or projetion for emotionl rousl n s to rte it for vlene. There were 8 trils tht were lssifie into positive, negtive or neutrl oring to the prtiipnts rtings. After snning, prtiipnts rte their memories or projetions on six ftors relte to their sujetive experiene, n omplete the LOT-R sle tht mesures trit optimism. Prtiipnts were rnke oring to their optimism sore on the LOT-R sle. These optimism rnkings were orrelte with the ifferene in sores (future positive minus future negtive) on the ehviourl sles. For eh prtiipnt time series ws rete initing the temporl position of the ifferent tril types triggere from the prtiipnt s first utton press. Funtionl ROIs were ientifie y ontrsting ll trils with fixtion (P,.5, unorrete) using rnom effets GLM (Generl Liner Moel) on the t of ll prtiipnts. The time ourses of tivtion were extrte from these regions for eh tril type n prtiipnt for sttistil nlysis. Next, we ientifie voxels within the ROIs in whih hnges in BOLD signl uring future positive trils reltive to future negtive trils were orrelte with the LOT-R sores. Aitionlly, funtionl onnetivity mps 6 were rete for the ifferent tril types (Flse Disovery Rte (FDR) orretion threshol t t 5 5.5).

4 Full Methos n ny ssoite referenes re ville in the online version of the pper t Reeive 7 July; epte 9 Septemer 7. Pulishe online XX 7.. Weinstein, N. D. Unrelisti optimism out future life events. J. Pers. So. Psyhol. 9, 86 8 (98).. Hoh, S. Conterftul resoning n ury in preiting personl events. J. Exp. Psyhol., 79 7 (98).. Drevets, W. C. et l. Sugenul prefrontl ortex normlities in moo isorers. Nture 86, 8 87 (997).. Strunk, D. R., Lopez, H. & DeRueis, R. J. Depressive symptoms re ssoite with unrelisti negtive preitions of future life events. Behv. Res. Ther., (6). 5. Lovllo, D. & Khnemn, D. Delusions of suess: how optimism unermines exeutives eisions. Hrv. Bus. Rev. 56, 56 6 (). 6. Tylor, S. E. & Brown, J. D. Illusion n well-eing: soil psyhologil perspetive on mentl helth. Psyhol. Bull., 9 (988). 7. Sheier, M. F. & Crver, C. S. Dispositionl optimism n physil well-eing: the influene of generlize outome expetnies on helth. J. Pers. 55, 69 (987). 8. Ais, D. R., Wong, A. T. & Shter, D. L. Rememering the pst n imgining the future: ommon n istint neurl sustrtes uring event onstrution n elortion. Neuropsyhologi 5, 6 77 (7). 9. Tulving, E. in The Missing Link in Cognition (es Terre, H. S. & Metlfe, J.) 56 (Oxfor Univ. Press, New York, 5).. Oku, J. et l. Thinking of the pst n pst: the roles of the frontl pole n the meil temporl loe. Neuroimge 9, 69 8 ().. Hssis, D., Kumrn, D., Vnn, S. D. & Mrguire, E. A. Ptients with hippompl mnesi nnot imgine new experienes. Pro. Ntl A. Si. USA, 76 7 (7).. D Argemeu, A. & Vn er Linen, M. Influene of ffetive mening on memory for ontextul informtion. Emotion, 7 88 ().. Sheier, M. F., Crver, C. S. & Briges, M. W. Distinguishing optimism from neurotiism (n trit nxiety, self-mstery, n self-esteem): reevlution of the Life Orienttion Test. J. Pers. So. Psyhol. 67, 6 78 (99).. Shrot, T., Mrtorell, E. A., Delgo, M. R. & Phelps, E. A. How personl experiene moultes the neurl iruitry of memories of Septemer. Pro. Ntl A. Si. USA, 89 9 (7). 5. Phelps, E. A. Emotion n ognition: insights from stuies of the humn mygl. Annu. Rev. Psyhol. 57, 7 5 (6). 6. Vogt, B. A. & Pny, D. N. Cingulte ortex of the rhesus monkey: II. ortil fferents. J. Comp. Neurol. 6, 7 89 (987). 7. Morn, J. M., Mre, C. N., Hetherton, T. F., Wyln, C. L. & Kelley, W. M. Neurontomil eviene for istint ognitive n ffetive omponents of self. J. Cogn. Neurosi. 8, (6). 8. Johnson, S. C., Bxter, L. C., Wiler, L. S., Pipe, J. G., Heisermn, J. E. & Prigtno, G. P. Neurl orreltes of self-refletion. Brin 5, 88 8 (). 9. Pulus, M. P. & Frnk, L. R. Ventromeil prefrontl ortex tivtion is ritil for preferene jugments. Neuroreport, 5 ().. Winston, J. S., Strnge, B. A., O Doherty, J. & Doln, R. J. Automti n intentionl rin responses uring evlution of trustworthiness of fes. Nture Neurosi. 5, 77 8 ().. Bush, G., Luu, P. & Posner, M. I. Cognitive n emotionl influenes in nterior ingulte ortex. Trens Cogn. Si., 5 ().. Cunninghm, W. A., Rye, C. L. & Johnson, M. K. Neurl orreltes of evlution ssoite with promotion n prevention regultory fous. Cogn. Affet. Behv. Neurosi. 5, (5).. Phelps, E. A., Delgo, M. R., Nering, K. I. & LeDoux, J. E. Extintion lerning in humns: role of the mygl n vmpfc. Neuron, ().. Simpson, J. R., Snyer, A. Z. Jr, Gusnr, D. A. & Rihle, M. E. Emotion-inue hnges in humn meil prefrontl ortex: I. uring ognitive tsk performne. Pro. Ntl A. Si. USA 98, (). 5. Willims, J. M., Ellis, N. C., Tyers, C., Hely, H., Rose, G. & MLeo, A. K. The speifiity of utoiogrphil memory n imgeility of the future. Mem. Cognit., 6 5 (996). 6. Roeroek, A., Formisno, E. & Goeel, R. Mpping irete influene over the rin using Grnger uslity n fmri. Neuroimge 5, (5). 7. Pruessner, J. C. et l. Volumetry of hippompus n mygl with highresolution MRI n three-imensionl nlysis softwre: minimizing the isrepnies etween lortories. Cere. Cortex, (). Supplementry Informtion is linke to the online version of the pper t Aknowlegements This stuy ws supporte y the NIMH (E.A.P.), the Sever Fountion (grnt to NYU s Center for Brin Imging), n Mrgret n Hermn Sokol Postotorl Fellowship (T.S.). We thnk S. H. Mulhern, D. C. Johnson n J. K. Szry for help in t nlysis, n D. Shiller, J. H. MDermott n Y. Trope for isussion. Author Contriutions T.S. esigne the stuy. T.S. n E.A.P. interprete the t n wrote the pper. T.S. n A.M.R. evelope stimuli, gthere ehviourl pilot t, n onute ehviourl t nlysis. T.S. gthere fmri t. T.S. onute neuroimging nlyses with the help of A.M.R. n C.M.R., n with vie of E.A.P. Whole-rin explortory nlysis ws onute y C.M.R. Author Informtion Reprints n permissions informtion is ville t Corresponene n requests for mterils shoul e resse to E.A.P. (liz.phelps@nyu.eu).

5 oi:.8/nture68 METHODS Prtiipnts. Prtiipnts (8) were reruite through poste vertisements. Three were eliminte from the nlysis euse of either exessive he movement or spikes ientifie in the fmri t. The remining 5 prtiipnts were inlue in the nlysis (7 mles, 8 femles; men ge,. yr; rnge, 8 6 yr). Before snning, prtiipnts omplete sreening form for signifint meil onitions. All prtiipnts gve informe onsent n were pi for their prtiiption. Stimuli. Stimuli onsiste of 8 short esriptions of possile life episoes (Supplementry Informtion). The orer n onition in whih episoes were presente were rnom. Beuse prtiipnts h teneny to generte positive events, we neee to provie more negtive n neutrl life episoes to otin n equl numer of trils rte s positive n negtive. Proeure. Before the snning session, prtiipnts went through four prtie trils. The session egn with short struturl sn, followe y four funtionl sns ( min eh), eh onsisting of trils of s. Finlly, n itionl longer struturl sn ws performe. Stimuli were presente y mens of mirror mounte on the he oil. In eh tril, esription of possile life episoe n the wor initing the onition of the tril (either pst or future ) ppere for s. The prtiipnts were instrute to press utton one mentl memory or projetion ws eginning to form in their min, n gin if they finishe elorting on it efore the en of the tril. The prtiipnt then h s to rte the memory or projetion for emotionl rousl (, not rousing;, somewht rousing;, highly rousing) n s to rte it for vlene (, positive;, neutrl;, negtive). Finlly, fixtion ross ws presente for s. If the prtiipnt i not press the utton to inite tht they were thinking of speifi memory or projetion, or i not inite rousl n/or vlene y pressing the utton ox, tht tril ws exlue from the finl t nlysis. The numer of trils rte s positive n negtive i not iffer for future or pst onitions. Retion times, initing when mentl memory or projetion ws eginning to form in the prtiipnts min, i not iffer for the ifferent tril types (rnge of mens:.9.6 s). After the snning session, prtiipnts were presente with the sme trils gin on omputer sreen, n were ske to rte their memories or projetions on six sles pte from previous stuy : viviness, sense of reliving or preexperiening, estimtion of the time of the event, sujetive temporl istne from the event, perspetive (involve or oserver), rousl n vlene. Finlly, prtiipnts omplete the LOT-R sle tht mesures trit optimism pessimism on sle from (pessimisti) to (optimisti). Prtiipnts were generlly optimisti; the men optimism sore ws Behviourl nlysis. Prtiipnts were rnke oring to their optimism sore on the LOT-R sle from the most optimisti to the lest optimisti. These optimism rnkings were orrelte with the ifferene sores (future positive minus future negtive) on the ehviourl sles. MRI snning n t nlysis. The stuy ws onute t the NYU Center for Brin Imging using T Siemens Allegr snner n Siemens he oil. Antomil imges were quire using MPRge sns, followe y -mm-thik xil slies (prllel to the nterior ingulte posterior ingulte plne). Positioning of the slies ws se on n AAsout sequene. Funtionl sns use grient eho sequene: TR (time of repetition) 5 s, TE (time of eho) 5 5 ms, FA (flip ngle) 5 9, mtrix 5 6 6, FOV (fiel of view) 5 9 mm, slie thikness 5 mm. A totl of 9 xil slies prllel to the AC PC plne were smple for whole-rin overge. The in-plne resolution ws mm mm. Imging t were nlyse using Brin Voyger softwre (QX, version.7). Dt were temporlly n sptilly smoothe ( mm full-with t hlfmximum) n motion-orrete. Iniviul t were trnsforme into Tlirh spe for group nlysis. For eh prtiipnt, time series ws rete initing the temporl position of the ifferent tril types triggere from the prtiipnt s first utton press to the onset of the rting sle. Tril types were lssifie into six groups oring to the prtiipnts rousl n vlene rtings uring the fmri snning: future positive, future negtive, future neutrl, pst positive, pst negtive n pst neutrl. Negtive events were those reeiving high () or meium () rousl rting n negtive vlene (). events were those reeiving high () or meium () rousl rting n positive vlene (). Neutrl events were those reeiving low rousl rting () n neutrl vlene (). Trils reeiving ny other omintion of responses oul not e hrterize in either group n were not inlue in the nlysis. The seventh tril type ws fixtion. Funtionl ROIs were ientifie y ontrsting ll trils with fixtion (P,.5, unorrete) using rnom effets GLM on the t of ll prtiipnts. We extrte the time ourse of tivtion from these regions for the ifferent tril types for eh prtiipnt. The verge perentge signl hnge from 6 s to 8 s fter the prtiipnt s first utton press in these ROIs were lulte for eh tril type n prtiipnt. Sttistil tests were performe on these vlues. Within these funtionl ROIs, we onute voxel-wise nlysis of ovrine ontrsting future positive trils with future negtive trils using the LOT-R optimism sore s ovrite (P,.5). In ition, priori regions of interest the right n left mygl were ientifie oring to lnmrks esrie previously 7. Extrtion of time ourses n sttistil nlysis ws one in the sme mnner s for the funtionl ROIs. Funtionl onnetivity nlysis 6 ws onute using the racc ROI s see. Mps were rete to ientify voxels where BOLD signl ws temporlly orrelte with the see uring future positive trils n uring future negtive trils (FDR orretion threshol t t 5 5.5). To ientify other voxels in the rin tht showe stronger BOLD responses uring one tril type thn the other, we onute whole-rin explortory nlysis on group t using rnom effets GLM (t P,., unorrete;. ontiguous voxels mm ).

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