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Threat Facilitates Subsequent Executive Control During Anxious Mood PDF

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Emotion ©2011AmericanPsychologicalAssociation 2011,Vol.11,No.6,1291–1304 1528-3542/11/$12.00 DOI:10.1037/a0026152 Threat Facilitates Subsequent Executive Control During Anxious Mood Jeffrey L. Birk Tracy A. Dennis TuftsUniversity CityUniversityofNewYork Lisa M. Shin Heather L. Urry TuftsUniversityandMassachusettsGeneralHospital/Harvard TuftsUniversity MedicalSchool Dualcompetitionframework(DCF)positsthatlow-levelthreatmayfacilitatebehavioralperformanceby influencingexecutivecontrolfunctions.Anxietyisthoughttostrengthenthiseffectbyenhancingthreat’s affective significance. To test these ideas directly, we examined the effects of low-level threat and experimentallyinducedanxietyononeexecutivecontrolfunction,theefficiencyofresponseinhibition. In Study 1, briefly presented stimuli that were mildly threatening (i.e., fearful faces) relative to nonthreatening(i.e.,neutralfaces)ledtofacilitatedexecutivecontrolefficiencyduringexperimentally inducedanxiety.Nosucheffectwasobservedduringanequallyarousing,experimentallyinducedhappy moodstate.InStudy2,weassessedtheeffectsoflow-levelthreat,experimentallyinducedanxiety,and individualdifferencesintraitanxietyonexecutivecontrolefficiency.ConsistentwithStudy1,fearful relative to neutral faces led to facilitated executive control efficiency during experimentally induced anxiety.Nosucheffectwasobservedduringanexperimentallyinducedneutralmoodstate.Moreover, individual differences in trait anxiety did not moderate the effects of threat and anxiety on executive control efficiency. The findings are partially consistent with the predictions of DCF in that low-level threatimprovedexecutivecontrol,atleastduringastateofanxiety. Keywords:low-levelthreat,executivecontrol,stateanxiety,traitanxiety,dualcompetitionframework Thehumanattentionalsystemisbiasedtowardthedetectionof scribed(LoBue,2010),thespecificattentionalmechanismsunder- threatening information. Indeed, the presence of threat-related lyingthreat’seffectonbehaviorremainlargelyunexplained. information (e.g., fearful faces) leads to enhancement at an early One candidate attentional mechanism is executive control, stage of visual processing (Phelps, Ling, & Carrasco, 2006) and which refers to the ability to organize cognitive and sensory capturesattentionsoefficientlythatthreatscanbedetectedevenin processing in a goal-directed way in order to guide appropriate the absence of conscious awareness (Dijksterhuis & Aarts, 2003; action (Miller & Cohen, 2001). An essential component of exec- Marcos&Redondo,2005).Althoughtheevolutionaryrootsofthis utive control is response inhibition, namely the ability to inhibit remarkable prioritization of threat detection are fairly well de- dominant or prepotent responses as needed (Miyake, Friedman, Emerson,Witzki,&Howerter,2000).Responseinhibitionmaybe critical for promoting adaptive functions following threat experi- ThisarticlewaspublishedOnlineFirstNovember7,2011. ences. Imagine, for example, that you are having a conversation JeffreyL.BirkandHeatherL.Urry,DepartmentofPsychology,Tufts with a friend during a hike in the woods when suddenly you University;TracyA.Dennis,DepartmentofPsychology,HunterCollege, encounterapoisonoussnake.Themostadvantageousresponsein CityUniversityofNewYork;LisaM.Shin,DepartmentofPsychology, thisscenariowouldinvolverapidlydetectingthesnakewhilealso Tufts University and Psychiatric Neuroimaging Group, Massachusetts inhibiting previously planned actions related to the conversation. GeneralHospital/HarvardMedicalSchool. To explain how the executive control of attention influences the WethankTuftsUniversityfortheirassistanceinfundingthisworkvia effectofthreatonbehavior,Pessoa(2009)proposeddualcompe- two Graduate Student Research Awards to J. L. Birk. We also thank S. Bansil,A.Bellet,P.Bene,K.Brethel,E.Brown,C.Callahan,R.Citron,E. titionframework(DCF). Davidowitz,M.DeMatteo,S.DelDonno,R.Gavrielov,J.Laks,B.Meller, AccordingtoDCF,threat-relatedstimulicarryaffectivesignif- D.Millstein,M.Patlingrao,V.Peisch,P.Pensuwan,P.Pop,C.Rucinski, icance,whichaltersperformancebystrengtheningsensoryrepre- M. Santarsieri, S. Sloley, R. Trumball, V. Tran, A. Wei, and J. Yih for sentationsattheperceptuallevelandbyprioritizingattentionatthe assistancewithdatacollectionandprocessing.Finally,wethankP.Opitz, executive level. The model predicts that, although threat consis- S. Cavanagh, and J. DiCorcia for valuable feedback and technical assis- tently leads to prioritized perceptual processing, its effect on tance,andF.WilhelmandP.PeykformakingANSLAB,asuiteofopen executivecontroldependsdramaticallyonthelevelofthreat(e.g., source Matlab routines used to process physiological data, available as high/extreme,low/mild).Specifically,DCFleadstotheprediction freeware in the Society for Psychophysiological Research Software Re- thatahigh-threatstimulusplacesapremiumonprocessingofthe pository(http://www.sprweb.org/repository). CorrespondenceconcerningthisarticleshouldbeaddressedtoHeather threatand,thus,reducestheexecutiveresourcesthatareavailable L.Urry,DepartmentofPsychology,TuftsUniversity,490BostonAvenue, to manage responses to stimuli other than the threat itself (i.e., Medford,MA02155.E-mail:[email protected] “hard prioritization”). A low-threat stimulus, on the other hand, 1291 1292 BIRK,DENNIS,SHIN,ANDURRY placesasmallerpremiumonprocessingofthethreatand,thus,has during experimentally manipulated happiness. In Study 2, we minoreffectsontheavailabilityofexecutiveresourcestomanage compare the effect of threat during experimentally manipulated responses to stimuli other than the threat (i.e., “soft prioritiza- anxiety to the effect of threat during an experimentally manipu- tion”). In fact, according to DCF, low-level threats prioritize lated neutral mood state. Because there is some indication that subsequent processing at the location of the threat “as part of experimental demonstrations of threat-related attentional capture additionalinformationgathering”(p.161)andthusshouldenhance may reflect an interaction between state and trait anxiety (for a subsequenttaskperformanceatthatsamelocation. review, see Williams, Mathews, & MacLeod, 1996), we also Several studies have provided evidence for the conclusion that examinewhethereffectsoflow-levelthreatandmanipulatedanx- threat can improve executive control function (Blanchette, 2006; ietydependonindividualdifferencesintraitanxiety. Cohen, Henik, & Mor, 2011; Öhman, Flykt, & Esteves, 2001; In both studies, consistent with the research on optimizing Schimmack,2005;butseeKrysko&Rutherford,2009).However, mood-induction procedures (for a review, see Gilet, 2008), we missing from these studies is a consideration of state-dependent usedanovel,multimodalmood-inductionprocedurethatincorpo- influences (e.g., mood state). Indeed, DCF proposes that the ex- ratednarrativetext,mood-congruentmusic,emotionalimages,and ecutiveenhancementseeninresponsetolow-levelthreatshouldbe imagined self-involvement. To measure the executive control of especially pronounced for people in an elevated state of anxiety attention,participantscompletedamodifiedversionoftheAtten- becauseelevatedanxietyheightenstheperceivedaffectivesignif- tion Network Test (ANT; Fan, McCandliss, Sommer, Raz, & icanceofthethreat.Consistentwiththisidea,thepresentationof Posner, 2002) following the mood-induction procedure. In this low-threat stimuli (i.e., fearful faces) relative to neutral stimuli variantoftheANT(Dennisetal.,2008),eachtrialwaspreceded (i.e., neutral faces) improved subsequent executive control effi- byathreat-related(fearfulface)ornonthreat-related(neutralface) ciency for people with high levels of naturally occurring state stimulus.Fearfulfacesconstitutelow-levelthreatstimulibecause anxiety but not for people with low levels of anxiety (Dennis, they do not directly jeopardize one’s survival, but they neverthe- Chen,&McCandliss,2008).Similarly,asecondstudyfoundthat lessclearlysignifythepotentialpresenceofdanger.Foreachcell higher levels of executive control efficiency following presenta- ofthedesign,wecalculatedaconflictscore—abehavioralindexof tion of fear-evoking stimuli (e.g., pictures of sharks) but not theefficiencywithwhichapersonusesexecutivecontroltoinhibit neutralstimuli(e.g.,picturesoffish)werecorrelatedwithhigher adominantresponse.Asatestofspecificity,wealsocalculatedan levels of naturally occurring state anxiety (Finucane & Power, alertingscore,abehavioralindexoftheefficiencywithwhichone 2010). However, because naturally occurring state anxiety might achieves a state of readiness to act. To be certain that our novel covarysystematicallywithunmeasured/unknownextraneousvari- mood-induction procedures were effective, we measured self- ables (e.g., arousal, intelligence, personality traits), it is possible reportedaffectandfacialelectromyography(EMG)overthecor- thatathirdvariablemightexplainthemoderatingeffectsofstate rugator supercilii muscle region, which is active when frowning anxietythatwereobservedinthesetwostudies. (Cacioppo,Martzke,Petty,&Tassinary,1988). Finally,asharedlimitationofexistingstudiesistheirfocuson one, unpleasant arousing mood state, namely anxiety. It may be Study 1 that threat affects subsequent executive control efficiency during anxiousmoodstatesbecauseanxietyrepresentsastateofheight- In Study 1, we tested the hypothesis that the presentation of a ened arousal. In that case, any aroused state, even one that is fearful face relative to a neutral face increases executive control pleasant rather than unpleasant, might similarly influence execu- efficiency in response to target stimuli presented subsequently at tivecontrolefficiency.Indeed,thereissomeevidencethatastate thesamespatiallocation.Giventhewell-establishedlinkbetween ofhappiness,whichisanaroused,positivelyvalencedmoodstate, anxiety and threat, it was possible that the facilitative effect of impairsperformanceefficiencyontasksrequiringexecutiveatten- low-levelthreatonexecutivecontrolefficiencywouldoccuronly tion, such as the Flanker task (Rowe, Hirsh, & Anderson, 2007) forpeopleinananxiousmoodstateandnotforpeopleinahappy and the alternating Stroop task (Phillips, Bull, Adams, & Fraser, mood state. However, if the effect of an anxious mood state is 2002),ascomparedwithaneutralstate(butseeFinucane,White- explained by arousal, then a similar effect would be observed man, & Power, 2009). Finucane (2011) demonstrated that exper- during the equally arousing happy mood state. Finally, although imentally induced fearful and angry mood states improved exec- our primary focus was to determine whether low-level threat utivecontrolefficiencyrelativetoaneutralmoodstate,buttoour would facilitate subsequent executive control, we also examined knowledge, only one study has directly compared effects of ex- whether threat would influence attention more broadly, in which perimentallyinducedanxiousandpositive(“nonanxious”)moods casewewouldalsoobserveeffectsoflow-levelthreatonalerting. onexecutivecontrolefficiency(Pacheco–Unguetti,Acosta,Calle- jas, & Lupia´n˜ez, 2010). In that study, there was no effect of Method inducedmoodstateonexecutivecontrolefficiency.Itisimportant that neither Finucane (2011) nor Pacheco–Unguetti et al. (2010) Participants evaluatedtheeffectofthreat. Inwhatfollows,wepresenttheresultsoftwostudies,foreach Sixty-one undergraduate students from Tufts University (35 of which the goal was to determine whether low-level threat female;mean[M] (cid:1)20.10years,standarddeviation[SD] (cid:1) age age stimuli would, as predicted by DCF, facilitate subsequent execu- 1.76) participated for course credit or monetary compensation. tivecontrolefficiencyinthecontextofanexperimentallymanip- Participantswere67.2%Caucasian,19.7%AsianorAsianAmer- ulatedstateofanxiety.InStudy1,wecomparetheeffectofthreat ican, and 8.2% Black or African American; 4.9% declined to during experimentally manipulated anxiety to the effect of threat providethisinformation.Ofthetotalsample,9.8%endorsedbeing THREATFACILITATESSUBSEQUENTEXECUTIVECONTROL 1293 of Hispanic origin. All study procedures were approved by the &Cacioppo,2003).CorrugatorEMGactivitywasmeasuredusing InstitutionalReviewBoardatTuftsUniversity,andallparticipants shielded 4-mm Ag/AgCl electrodes. Following site preparation providedinformedconsentpriortoparticipatinginthestudy. with an electrode preparation pad, a pair of sensors was attached directlyabovetheeyebrowandagroundelectrodewasattachedto themiddleoftheforeheadconsistentwithguidelinesbyFridlund Materials andCacioppo(1986). Moodmanipulation. Moodwasmanipulatedonabetween- EMG data were collected using MP 150 hardware and Acq- subjectsbasis.Participantswererandomlyassignedtooneoftwo Knowledge3.8.2software(Biopac,Goleta,CA).Dataweresam- groups.Thirtyparticipantscompletedananxiousmoodinduction, pled at 1,000 Hz and filtered from 5 Hz to 3 kHz (60-Hz notch and 31 participants completed an equally arousing happy mood filteron)online.Offline,datawereresampledto400Hz,rectified, induction. All participants also completed a nonaroused neutral andsmoothedwitha16-Hzlow-passfilter,decimatedto4Hz,and moodinduction,whichservedasabasisforcomparisoninmood- further smoothed with a 1-s prior moving average filter. These manipulationchecks.Intheanxiousmoodinduction,participants steps were completed in part with Matlab software (Mathworks, imagined being a passenger in a drunk-driving car accident and Natick, MA) using ANSLAB routines (Wilhelm & Peyk, 2005). helpinginjuredpeopleinitsaftermath.Inthehappymoodinduc- Corrugatoractivitywasaveragedacrossthe4-mindurationofeach tion, participants imagined walking with a friend on a warm day mood-inductionconditionandlog-transformedtoachievenormal- amid picnicking families, playing children, and running dogs. In ity. the neutral mood induction, participants imagined a sequence of AttentionNetworkTest. Aftereachofthetwomoodinduc- mundane activities such as shopping for groceries, doing small tions,participantscompletedamodifiedversionoftheANT(Fan tasksathome,andcallingafamilymember. etal.,2002)similartothatusedbyDennisandcolleagues(2008). Mood-congruent instrumental music, selected based on pilot The modified ANT allowed us to measure the efficiency of the testing of effects on self-reported affect, was played during each executive control and alerting attention networks. The task of scene(selectionsavailableuponrequest).Inaddition,picturesthat participantswastorespondviamousebuttonswiththethumbsof accompanied the narrative were shown. Pictures with established both hands to indicate whether a centrally located arrow (i.e., positive, negative, and neutral valence were selected from the target) pointed to the left or right. Each trial lasted for 4,050 ms InternationalAffectivePictureSeries1(Lang,Bradley,&Cuthbert, (seeFigure1fortrialstructure).Sincetheeffectsofinducedmood 2005); supplementary pictures were obtained from Shutterstock could be time limited, we excluded trials from the original ANT (http://www.shutterstock.com) and Wellcome Images (http:// thatmeasuretheefficiencyoftheorientingnetwork.Wereasoned images.wellcome.ac.uk). that this would minimize the time it would take to complete the The mood-induction procedure was programmed in E-Prime taskwhileatthesametimemaximizingourabilitytotesthypoth- 1.2.1 software (Psychology Software Tools, Sharpsburg, PA). esized effects on executive control efficiency and to examine Eachmoodinductioncontained12pictureswithassociatedstory- specificitybyretainingtrialsthatassessalertingefficiency. linetext,andlasted4min.Eachpicturewaspresentedwithtextat To assess executive control efficiency, the target was sur- thetopofthescreenfor12sandthenwithouttextforanother8s. roundedbyfourstimuli(i.e.,flankers),twoeachontheleftandthe Participantswereinstructedtoreadthetextfirstandthenviewthe right. There were three varieties of flankers that differed across imagewhileimaginingthatthedepictedeventswereoccurringin trialtypes:congruent(i.e.,arrowspointinginthesamedirectionas their real lives. The order of the neutral and anxious (or happy) the target), incongruent (i.e., arrows pointing in the opposite di- moodinductionswascounterbalancedacrossparticipants. rection of the target), and neutral (i.e., horizontal lines with no Moodmeasures. Participantscompletedacomputerizedver- directional information). Greater conflict is present on trials in sionofthePositiveandNegativeAffectSchedule(PANAS;Wat- which the flanking arrows are incongruent with the target arrow, son,Clark,&Tellegen,1988)beforeandaftereachmoodinduc- relative to trials in which the flanking arrows are congruent or tion. Participants reported how much they felt each of 20 neutral. As is standard practice with the ANT (Fan et al., 2002), adjectives“rightnow,atthisverymoment”ona5-pointscalefrom conflict scores were generated by subtracting congruent-flanker 1, very slightly or not at all, to 5, extremely. We computed a reaction time from incongruent-flanker reaction time for correct positive affect (PA) score by summing scores for enthusiastic, trials. Smaller conflict scores indicate greater efficiency of exec- proud,inspired,anddetermined,andanegativeaffect(NA)score utivecontrol,specificallyresponseinhibition. by summing scores for distressed, upset, guilty, scared, hostile, To assess alerting, on each trial the target was preceded by a irritable,ashamed,nervous,jittery,andafraid.Wealsocomputed double warning cue (i.e., asterisks above and below the fixation anarousalscorebysummingscoresforinterested,excited,alert, cross)ortherewasnocueprecedingthetarget(i.e.,fixationcross attentive, strong, and active. These latter six items are typically only). Greater alerting is present on trials in which the target is includedasitemsonthePAscale(Watsonetal.,1988),butPatrick precededbythedoubleasterisks,relativetotrialsinwhichthereis and Lavoro (1997) demonstrated that they index arousal, since nocue.Again,followingstandardpracticewiththeANT(Fanet both positive and negative pictures elicit higher ratings on these al., 2002), alerting scores were generated by subtracting double- items,relativetoneutralpictures. cue reaction time from no-cue reaction time for correct trials. In addition, for a more objective assessment of emotional va- lence, we measured EMG activity over the corrugator supercilii muscle region. Relative to a neutral baseline, greater corrugator 1Identification numbers for IAPS pictures: 2570, 2751, 2810, 5760, activity is observed for unpleasant emotions, whereas lower cor- 5800,7022,7035,7041,7060,7090,7100,7175,7211,7217,7233,7235, rugatoractivityisobservedforpleasantemotions(Larsen,Norris, and7920. 1294 BIRK,DENNIS,SHIN,ANDURRY Figure1. TrialstructureforthemodifiedAttentionNetworkTest.Inthisexampletrial,theparticipantsawa fearfulfacewhichwasfollowedbyafixationcrossandthenadoublecue(whichwilltendtoincreasealerting efficiencyrelativetotrialswithnocue).Thedoublecuewasfollowedbyanotherfixationcrossandthenthe targetwaspresented.Thetargetinthiscasewasaright-facingarrowwhichwassurroundedbyleft-facingarrow flankers.Thus,thiswasanincongruenttrial(whichwilltendtoincreaseconflict,i.e.,decreaseexecutivecontrol efficiency,relativetotrialswithcongruentarrows).Thetrialendswithavariable-intervalfixationcrossthat makeseachtriallastatotalof4,050ms. Higher alerting scores indicate greater efficiency in achieving a presented in four experimental blocks of 48 trials each. Partici- stateofactionreadiness. pants were instructed to take a break between each block. At the Toexaminetheinfluenceofthreatonsubsequentefficiencyof endofthesession,participantsprovideddemographicinformation the executive control and alerting attentional networks, each trial andweredebriefedaboutthenatureofthestudy. beganwiththepresentationofoneoftwotypesoffaces—fearful (i.e.,threat-related)orneutral.FollowingDennisetal.(2008),the Results delay between the offset of the face and the onset of the target varied randomly between 900 ms and 1800 ms (see Figure 1). Data Retention and Analysis Faces from 12 actors were selected from the NimStim collection (Tottenham et al., 2009). Greater threat is present on trials pre- For each reaction time measure (conflict score and alerting cededbyfearfulfacesthanbyneutralfaces. score), Mahalanobis distance was computed for each participant Conflictandalertingscoreswerecalculatedseparatelyforfear- acrosslevelsofthewithin-subjectsindependentvariablestoiden- fulandneutralfacetrialsforeachmoodgroup.Accuracydatawill tify multivariate outliers. One participant was excluded from the not be reported for Study 1 or Study 2 because the ANT was analysis of conflict scores and two participants were excluded designed to measure efficiency, which is our primary dependent fromtheanalysisofalertingscoresonthebasisofthesetests(p(cid:2) variable of interest. Furthermore, the accuracy data are not nor- .001). In the general linear model (GLM) analyses below, we mallydistributed. reportresultsofmultivariateFstatistics((cid:3)(cid:1).05)andfollow-up Fisher’s least significant difference tests as needed to interpret Procedure significantresults. Participantscompletedthetasksinthefollowingorder:baseline Manipulation Checks PANAS, mood induction 1, PANAS, ANT 1, PANAS, mood induction 2, PANAS, ANT 2, PANAS. Each ANT began with a Todeterminewhetherthemoodmanipulationsachievedapos- practiceblockof12trialsthatwereaccompaniedbyreactiontime itively valenced mood state in the happy group and a negatively andaccuracyfeedback.Atotalof192no-feedbacktrialswerethen valenced mood state in the anxious group, repeated-measures THREATFACILITATESSUBSEQUENTEXECUTIVECONTROL 1295 GLMswereperformedwithineachgrouptoexaminetheeffectsof buttherewasnosuchdifferenceinpreinductionarousalineither arousal (arousing, nonarousing) and period (preinduction, postin- thehappy(p(cid:1).954)oranxiousgroup(p(cid:1).890). duction) on self-reported PA and NA from the PANAS. In addi- Finally, to determine whether the anxious and happy mood tion, paired samples t tests were performed within each group to stateswereequallyarousing,wedirectlycomparedarousalscores compare the arousing (anxious, happy) conditions to the nonar- for the happy and anxious groups using independent samples t ousing (neutral) condition on mean corrugator activity, a second tests. The postinduction arousal score was higher in the anxious indexofmoodvalence. groupthaninthehappygroup,t(59)(cid:1)2.11,p(cid:1).039.However, Inthehappygroup,asignificantinteractionofArousal(cid:4)Period the preinduction arousal score was also marginally higher in the emergedforPAscores,F(1,30)(cid:1)18.61,p(cid:2).001,(cid:5)2(cid:1).383,but anxious group than in the happy group, t(59) (cid:1) 1.80, p (cid:1) .077. p not for NA scores, F(1, 30) (cid:1) 1.10, p (cid:1) .303, (cid:5)2 (cid:1) .035. As Takingthispreinductiondifferenceintoaccount(i.e.,bysubtract- p shown in Table 1, postinduction PA was higher for the arousing ing preinduction levels of arousal from postinduction levels), the (happy) than the nonarousing (neutral) condition (p (cid:2) .001), but groupdifferencewasnolongersignificant,t(59)(cid:1).086,p(cid:1).932. there was no such difference in preinduction PA (p (cid:1) .291). In Collectively, these arousal results suggest that the happy and addition, mean corrugator activity was significantly lower (i.e., anxious mood states were achieved with an equivalent degree of less“frowning”)duringthearousing(happyM(cid:1)0.60,SD(cid:1)0.33) arousalinthetwogroups. than during the nonarousing (neutral M (cid:1) 0.74, SD (cid:1) 0.35) condition, t(30) (cid:1) (cid:6)2.42, p (cid:1) .022. Collectively, these PA and Hypothesis Testing corrugator results indicate that the happy induction elicited the intendedpositivemoodstate. Executive control. GLMs were conducted for the arousing In the anxious group, there was a significant interaction of moodconditionstoassesstheeffectsofthreat(fearfulface,neutral Arousal(cid:4)PeriodforNAscores,F(1,29)(cid:1)29.26,p(cid:2).001,(cid:5)2(cid:1) face),moodgroup(happy,anxious),andtime(early[blocks1–2], p .502,butnotforPAscores,F(1,29)(cid:1)1.82,p(cid:1).187,(cid:5)2(cid:1).059. late[blocks3–4])onconflictscores.Therewerenomaineffects p AsshowninTable1,postinductionNAwashigherforthearous- ofthreat,F(1,58)(cid:1)2.75,p(cid:1).103,(cid:5)2(cid:1).045,ormoodgroup, p ing(anxious)thanthenonarousing(neutral)condition(p(cid:2).001), F(1, 58) (cid:1) 0.01, p (cid:1) .938, (cid:5)2 (cid:2) .001. There was a significant p buttherewasnosuchdifferenceinpreinductionNA(p(cid:1).562). interaction of Time (cid:4) Threat, F(1, 58) (cid:1) 6.07, p (cid:1) .017, (cid:5)2 (cid:1) p Inaddition,meancorrugatoractivitywassignificantlygreater(i.e., .095.Forearlytrials,whichoccurredsoonafterthemoodinduc- more“frowning”)duringthearousing(anxiousM(cid:1)0.69,SD(cid:1) tion,conflictscoresweresignificantlylowerforthefearful(M(cid:1) 0.34)thanduringthenonarousing(neutralM(cid:1)0.60,SD(cid:1)0.32) 88.33, SD (cid:1) 37.03) relative to the neutral (M (cid:1) 107.09, SD (cid:1) condition, t(29) (cid:1) 2.25, p (cid:1) .032. Collectively, these NA and 43.94)condition,p(cid:1).002.Forlatetrials,however,conflictscores corrugatorresultsindicatethattheanxiousmoodinductionelicited didnotdifferforthefearful(M(cid:1)106.60,SD(cid:1)42.56)relativeto theintendednegativemoodstate. the neutral (M (cid:1) 104.70, SD (cid:1) 53.03) condition, p (cid:1) .723. Next,todeterminewhetherthemoodmanipulationsachieveda Critically,therewasasignificantThreat(cid:4)MoodGroupinterac- state of arousal in both groups, repeated-measures GLMs were tion,F(1,58)(cid:1)4.37,p(cid:1).041,(cid:5)2(cid:1).070.Intheanxiousgroup, p performed within each group to examine the effects of arousal conflictscoresweresignificantlydiminishedfortrialsprecededby (arousing, nonarousing) and period (preinduction, postinduction) fearful faces relative to neutral faces, p (cid:1) .012, but no such onself-reportedarousalfromthePANAS.Asexpected,therewere differencewasobservedinthehappygroup,p(cid:1).757(seeFigure significantArousal(cid:4)Periodinteractionsforthehappy,F(1,30)(cid:1) 2a). There was no significant interaction of Time (cid:4) Threat (cid:4) 7.34, p (cid:1) .011, (cid:5)2 (cid:1) .197, and anxious, F(1, 29) (cid:1) 15.78, p (cid:2) MoodGroup,F(1,58)(cid:2)0.01,p(cid:1).957,(cid:5)2(cid:2).001. p p .001, (cid:5)2 (cid:1) .352, groups. As shown in Table 1, postinduction To examine the nature of the conflict effect in the anxious p arousal was higher for the arousing (anxious or happy) than the group,afollow-upGLMforonlytheseparticipantswasconducted nonarousing (neutral) condition in both groups (both ps (cid:2) .001), toexaminetheeffectsofthreatandflanker(incongruent,neutral, Table1 Self-ReportedAffectBeforeandAftertheMoodInductionsinStudy1 Preinduction Postinduction Arousingmood Nonarousingmood Arousingmood Nonarousingmood Anxiousgroup PA 8.37(3.66) 8.23(3.36) 6.77(2.80) 7.57(3.20) NA 13.33(3.46) 12.87(4.02) 18.83(6.16)a 11.40(1.48)a Arousal 15.60(4.98) 15.43(4.66) 17.77(4.22)a 13.70(4.19)a Happygroup PA 7.13(3.55) 7.77(3.79) 9.90(3.18)a 7.03(3.39)a NA 14.00(3.24) 13.71(3.55) 11.84(3.28) 12.45(3.19) Arousal 13.42(4.48) 13.48(5.09) 15.48(4.23)a 12.23(4.18)a Note. Mean(SD)self-reportedaffectscoresfromthePositiveandNegativeAffectSchedule(PANAS;Watson, Clark,&Tellegen,1988).PAandNAdenote,respectively,positiveaffectscoresandnegativeaffectscores. aMeanssharingthissamesuperscriptwithinarowaresignificantlydifferentatp(cid:2).05. 1296 BIRK,DENNIS,SHIN,ANDURRY did not differ between the arousing (anxious M (cid:1) 101.34, SD (cid:1) 33.86) and nonarousing (neutral M (cid:1) 104.83, SD (cid:1) 38.34) con- ditions,t(28)(cid:1)(cid:6)0.56,p(cid:1).577.Similarly,forthehappygroup, conflict scores did not differ between the arousing (happy M (cid:1) 102.01,SD(cid:1)32.12)andnonarousing(neutralM(cid:1)99.55,SD(cid:1) 29.16)conditions,t(30)(cid:1)0.61,p(cid:1).545.Theseresultsshowthat arousing mood alone was not sufficient to influence executive controlefficiencyforeithergroup. Alerting. GLMswereconductedforthearousingmoodcon- ditions to assess the effects of threat (fearful face, neutral face), mood group (happy, anxious), and time (early, late) on alerting scores.Themaineffectoftimewastheonlysignificantfindingin this analysis, F(1, 57) (cid:1) 10.31, p (cid:1) .002, (cid:5)2 (cid:1) .153, indicating p thatalertingefficiencywashigheroverallinthelaterelativetothe earlytrials,allotherFs(cid:2)1.89.Thus,therewasnothreat-related facilitation of alerting efficiency in either group (see Figure 2b), which suggests specificity in the threat-related facilitation of ex- ecutivecontrolefficiencyintheanxiousgroup. Toexaminetheeffectofarousingmoodalone,separatepaired t tests were conducted for each mood group to compare alerting scores for the arousing (anxious or happy) condition to the non- arousing(neutral)condition.Fortheanxiousgroup,alertingscores did not differ between the arousing (anxious M (cid:1) 18.66, SD (cid:1) 21.82)andnonarousing(neutralM(cid:1)23.11,SD(cid:1)18.80)condi- tions, t(28) (cid:1) (cid:6)1.03, p (cid:1) .310. Similarly, for the happy group, alerting scores did not differ between the arousing (happy M (cid:1) 25.93, SD (cid:1) 18.68) and nonarousing (neutral M (cid:1) 28.17, SD (cid:1) Figure2. Thesetwopanelsdepictattentionnetworkefficiencyscoresin 20.60) conditions, t(29) (cid:1) (cid:6)0.65, p (cid:1) .521. These results show Study1forreactiontime(RT)differencesasafunctionofhappy(HAP) that arousing mood alone was not sufficient to influence alerting andanxious(ANX)moodstatesfortrialsprecededbyfearfulfaces(black efficiencyforeithergroup. bars)andneutralfaces(graybars).Thetopfigureshowsconflictscores (incongruent—congruent)andthebottomfigureshowsalertingscores(no cue—doublecue).Notethatconflictscoresareinverselyrelatedtoexec- Discussion utivecontrolefficiency.Errorbarsrepresent(cid:7)1standarderror. Usinganovel,multimodalmood-inductionprocedure,wefound thatthreat-relatedstimulifacilitatedsubsequentexecutivecontrol congruent) on raw reaction times. An interaction of Threat (cid:4) efficiencyduringexperimentallyinducedstateanxiety.Therewas Flanker,F(2,27)(cid:1)7.46,p(cid:1).003,(cid:5)2(cid:1).356,indicatedthatRT no parallel effect of threat on alerting efficiency, which suggests p werefastertoincongruent-flankertrialsprecededbyfearfulfaces that threat had a specific effect on executive control. Finally, (M (cid:1) 597.71, SD (cid:1) 61.69) versus neutral faces (M (cid:1) 608.67, threat-related facilitation of executive control efficiency occurred SD(cid:1)75.74),p(cid:1).023.Inaddition,RTwereslowerforcongruent- only during the anxious mood state and not during the equally flankertrialsprecededbyfearfulfaces(M(cid:1)505.90,SD(cid:1)55.22) arousing,happymoodstate,whichsuggeststhattheeffectofthreat versus neutral faces (M (cid:1) 497.79, SD (cid:1) 58.64), but this was a on executive control efficiency during experimentally induced marginallysignificanteffect,p(cid:1).072.RTwerealsosignificantly stateanxietycannotbeexplainedbyarousal.Thesefindingsindi- slower for neutral-flanker trials preceded by fearful faces (M (cid:1) cate that state anxiety may potentiate the facilitative effect of 494.15 SD (cid:1) 47.88) versus neutral faces (M (cid:1) 486.11, SD (cid:1) low-levelthreatonsubsequentexecutivecontrolefficiency. 49.74),p(cid:1).044.Thispatternofresultsindicatesthatthethreat- TherewerefourimportantlimitationstoStudy1.First,among related conflict effect described above for participants in an anx- peoplewithhightraitanxiety,whichisthedispositionaltendency iousmoodwasexplainedbyathreat-relatedspeedingofRTinthe tobeanxious,threathasbeenshowntohaveaninhibitingeffecton incongruentconditionandamarginalthreat-relatedslowinginthe executive control (Derakshan & Eysenck, 2009; Derryberry & congruent condition. Thus, the attentional effect of threat among Reed,2002;Eysenck,Derakshan,Santos,&Calvo,2007;Wood, peopleinananxiousmoodisnotsimplyathreat-relatedenhance- Mathews, & Dalgleish, 2001). It may be that the threat-related mentofattentiontothecentrallocationorathreat-relatedincrease facilitation of executive control efficiency observed during state in general effort, which would be manifested as faster responses anxiety in Study 1 would be absent or perhaps even reversed in for trials preceded by fearful faces relative to neutral faces irre- participants with high trait anxiety. Second, although unlikely spectiveofflankertype. given random assignment of participants to conditions, it is pos- Toexaminetheeffectofarousingmoodalone,separatepaired sible that the observed effect of threat on executive efficiency t tests were conducted for each mood group to compare conflict mighthaveexistedintheseparticipantspriortoanyexperimental scores for the arousing (anxious or happy) condition to the non- manipulation of mood. Third, although the anxious and happy arousing(neutral)condition.Fortheanxiousgroup,conflictscores moods were purposefully matched for arousal, the mood states THREATFACILITATESSUBSEQUENTEXECUTIVECONTROL 1297 might have differed in other undesirable ways. For example, in the high trait-anxiety group (n (cid:1) 59; M (cid:1) 49.39, SD (cid:1) 6.38, happiness might be uniquely associated with alterations in self- range:40–65). focused attention (Green, Sedikes, Saltzberg, Wood, & Forzano, Moodmanipulation. Participantswererandomlyassignedto 2003)oragenerallyreducedcognitivecapacity(Mackie&Worth, oneoftwogroups.Sixtyparticipantscompletedtheanxiousmood 1989). Fourth, although we confirmed that the mood manipula- induction,and61participantscompletedtheneutralmoodinduc- tions induced arousing negative and positive moods using mea- tion. sures that were sensitive to valence and arousal, we could not Moodmeasures. Inadditiontorecordingcorrugatoractivity, specificallyconfirmthatparticipantswereanxiousinresponseto participants completed a computerized version of the State–Trait theanxiousmoodmanipulation.Toaddressthesefourlimitations Inventory of Cognitive and Somatic Anxiety (STICSA; Grös, whilereplicatingtheprimaryfinding,weconductedStudy2. Antony, Simms, & McCabe, 2007) before and after each task. Participants reported how much they felt each of 20 descriptions “rightnow,atthisverymoment”ona4-pointscalefrom1,notat Study 2 all,to4,verymuchso.Wecomputedasomaticanxietyscoreand AsinStudy1,weexaminedtheeffectsoflow-levelthreatand acognitiveanxietyscoreforeachtimepointinthestudy. experimentally induced state anxiety on executive control effi- Participants also reported how they felt before and after each ciencyasmeasuredusingourmodifiedversionoftheANT.Each task using mood ratings that were modified from the adjective trialoftheANTwasprecededbyathreateningornonthreatening tripletsusedbyTamir,John,Srivastava,andGross(2007).These stimulus(fearfulorneutralface,respectively).Forthisfollow-up ratingscapturedanxiety(e.g.,anxious,worried,fearful)andgen- study, we added a measure of trait anxiety and a premood- eral arousal (e.g., active, alert, keyed up) (see Nitschke, Heller, inductionANT.Thisenabledustotestwhethertheeffectsofthreat Palmieri, & Miller, 1999). They also included filler items corre- andstateanxietydependonleveloftraitanxiety,andtoruleout spondingtoothernegativeandpositiveemotionalstatessoasnot the possibility that our two mood groups differed in attentional toemphasizenegativemoodcategories(andanxietyspecifically) efficiencypriortothemoodinduction.Moreover,ratherthanusing and thereby reduce demand characteristics (data not reported). a happy mood induction as our comparison group, participants Participantsreportedhowstronglytheyfelteachsetofadjectives were randomly assigned to an anxious or neutral mood group. “rightnow,atthisverymoment”ona7-pointscalefrom0,notat Finally, we replaced the valence and arousal mood-manipulation all,to6,verymuch. measuresfromStudy1withspecificself-reportmeasuresofanx- ANT. Beforeandafterthemoodinduction,participantscom- iety before and after every mood induction. Consistent with the pletedatwo-blockversionoftheANTadministeredinStudy1to rationaleforandfindingsfromStudy1,wetestedthehypothesis assess the efficiency of executive control and alerting attention that the presentation of a fearful face relative to a neutral face networks. Since the threat-related effects were strongest early in wouldincreaseexecutivecontrolefficiencyinanxiouspeople. the task during Study 1, the overall length of the ANT was shortenedinStudy2tofocusonattentionaleffectsoccurringclose to the end of the mood induction. Based on tests of internal Method consistency reliability of the ANT data from Study 1, we deter- minedthatthetasklengthcouldbehalvedwhilestillmaintaining ThemethodsusedforStudy2wereidenticaltothosedescribed adequate reliability (Cronbach’s alpha (cid:1) .70 for all cells of the forStudy1,exceptasnotedbelow. studydesign). Participants Procedure One hundred twenty-one undergraduate students from Tufts Participants completed the tasks in the following order: a dot University (69 female; M (cid:1) 20.55 years, SD (cid:1) 4.45 years) age age probe task (data not presented as they are not the focus of this participated for course credit or monetary compensation. Partici- report), demographic questionnaire and STAI, STICSA/triplets, pants were 71.1% Caucasian, 20.7% Asian or Asian American, baselineANT,STICSA/triplets,moodinduction(ANXorNEU), 4.1% Black or African American, and 0.8% American Indian or STICSA/triplets,experimentalANT,STICSA/triplets. AlaskaNative;3.3%declinedtoprovidethisinformation.Ofthe totalsample,14.9%endorsedbeingofHispanicorigin. Results Materials Data Retention and Analysis Trait Anxiety Groups Using Mahalanobis distance, three participants were excluded from the analysis of conflict scores and four participants were Participants completed the trait form (Y-2) of the State–Trait excludedfromtheanalysisofalertingscoresonthebasisofthese AnxietyInventoryforAdults(STAI;Spielberger,1983).Thetotal tests (p (cid:2) .001). Additionally, two participants were excluded scoresfortheSTAIcoveredawidespectrum(M(cid:1)40.79,SD(cid:1) because they represented age outliers within their respective 10.21, median (cid:1) 39, min (cid:1) 21, max (cid:1) 65). Participants with groups(i.e.,greaterthan35yearsold).Finally,oneparticipantwas scores less than or equal to the median were included in the low automaticallyexcludedfromtheanalysisofconflictscoresdueto traitanxietygroup(n(cid:1)62;M(cid:1)32.50,SD(cid:1)4.79,range:21–39), missingdata.Onehundredfifteenparticipantswereincludedinthe andparticipantswithscoresgreaterthanthemedianwereincluded manipulation checks that follow (34 anxious mood, low trait 1298 BIRK,DENNIS,SHIN,ANDURRY anxiety;24anxiousmood,hightraitanxiety;26neutralmood,low anxious and neutral mood groups for low TA, p (cid:1) .223, or high trait anxiety; 31 neutral mood, high trait anxiety). There was no TA,p(cid:1).496,participants.Therewasalsonomaineffectoftrait significant difference in level of trait anxiety for participants anxiety, F(1, 112) (cid:1) 0.810, p (cid:1) .370, (cid:5)2 (cid:1) .007. Therefore, p assigned to the anxious (M (cid:1) 39.84, SD (cid:1) 10.25) and neutral despitetheevidenceoftheself-reportmeasuresandunlikeStudy (M (cid:1) 41.47, SD (cid:1) 10.08) mood conditions, t(114) (cid:1) 0.86, p (cid:1) 1, participants in the anxious mood induction did not exhibit .392,suggestingthatstateandtraitanxietywerenotconfounded. greaterexpressive“frowning”behavior,ascomparedwithpartic- ipantsintheneutralinduction. Manipulation Checks Accounting for Individual Differences in the Todeterminewhetherthemoodmanipulationsweresuccessful, Effectiveness of Induced Mood GLMswereperformedtoassesstheeffectsofmoodgroup(anx- ious, neutral), trait anxiety group (low TA, high TA), and period Given the great importance of the corrugator measure as an (preinduction,postinduction)ontheSTICSAandontheanxiety- index of the mood manipulation success that is relatively free of relevant adjective triplets from the mood ratings. In addition, a demandcharacteristics,theabsenceofamoodeffectoncorrugator GLM was also performed to examine the effects of mood group activity suggests that anxious mood may not have been success- (anxious, neutral) and trait anxiety group (low TA, high TA) on fullyinducedinallparticipants.Wethereforeretainedonlythose meancorrugatoractivityduringthemoodmanipulations. participants for whom the target mood was manipulated to a There was a trend toward a main effect of mood group, F(1, sufficientdegreeinsubsequentanalyses.Specifically,participants 112)(cid:1)2.99,p(cid:1).087,(cid:5)2(cid:1).026,andasignificantmaineffectof intheanxiousmoodgroupwereretainedonlyiftheydisplayedat p periodonSTICSAsomaticanxietyscores,F(1,112)(cid:1)15.96,p(cid:2) least a 50% increase in corrugator activity. Participants in the .001, (cid:5)2 (cid:1) .125. These effects were qualified by a significant neutralmoodgroupwereretainedonlyiftheydisplayedlessthan p MoodGroup(cid:4)Periodinteraction,F(1,112)(cid:1)34.82,p(cid:2).001, a 50% increase in corrugator activity. Proportional change in (cid:5)2 (cid:1) .237, which revealed that preinduction scores were not corrugator activity was computed by subtracting the average ac- p different for the anxious (M (cid:1) 14.52, SD (cid:1) 3.41) relative to the tivityduringthe5-speriodatthestartoftheinduction(baseline) neutral(M(cid:1)14.85,SD(cid:1)3.41)group,p(cid:1).620,butpostinduc- from the average activity during the 5-s period at the end of the tionscoresweregreaterfortheanxious(M(cid:1)16.94,SD(cid:1)4.18) inductionanddividingthisdifferencebythebaseline. relativetotheneutral(M(cid:1)14.30,SD(cid:1)4.07)group,p(cid:1).001.As Intotal,70participantswereretainedforthefinalanalyses(17 anticipated,therewasamaineffectoftraitanxietysuchthathigh anxious mood, low trait anxiety; 11 anxious mood, high trait TAparticipantshadoverallhigherscores(M(cid:1)16.06,SD(cid:1)3.52) anxiety;19neutralmood,lowtraitanxiety;23neutralmood,high than low TA participants (M (cid:1) 14.19, SD (cid:1) 3.63), F(1, 112) (cid:1) trait anxiety). As intended, the STAI scores for retained partici- 8.34,p(cid:1).005,(cid:5)2(cid:1).069.Therewerenoothersignificanteffects pantsweresimilartoscoresinthefullsample(lowTAgroup:n(cid:1) involvingtraitanxpiety,however,indicatingthattheimpactofthe 36;M(cid:1)32.56,SD(cid:1)4.78,range:22–39;highTAgroup:n(cid:1)34; moodinductionsonsomaticanxietydidnotdifferasafunctionof M(cid:1)49.65,SD(cid:1)6.46,range:40–65).Inaddition,therewasno traitanxiety. significant difference in TA level for participants assigned to the TheanalysisofSTICSAcognitiveanxietyscoresrevealedonly anxious(M(cid:1)39.68,SD(cid:1)10.29)andneutral(M(cid:1)41.64,SD(cid:1) theexpectedmaineffectoftraitanxietygroupsuchthat,asabove, 10.31)moodconditions,t(68)(cid:1)0.781,p(cid:1).437,suggestingagain highTAparticipantshadoverallhigherscores(M(cid:1)17.00,SD(cid:1) thatstateandtraitanxietywerenotconfounded.Participantswho 4.50) than low TA participants (M (cid:1) 12.70, SD (cid:1) 4.50), F(1, were retained based on the criteria above did not differ from 112) (cid:1) p (cid:2) .001, (cid:5)2 (cid:1) .186. There were no other significant participantswhowerenotretainedinTA,overallconflictscores, effectsorinteractionsp,allotherFs(cid:2)0.91. oroverallalertingscores,allps(cid:1).220. Theparallelanalysisonscoresfortheanxious,worried,fearful and active, alert, keyed up triplets revealed significant Mood Hypothesis Testing Group(cid:4)Periodinteractions,F(1,112)(cid:1)39.21,p(cid:2).001,(cid:5)2(cid:1) p Executivecontrol. AGLMtestedtheeffectsofthreat(fear- .259 and F(1, 112) (cid:1) 28.59, p (cid:2) .001, (cid:5)2 (cid:1) .203, respectively. p fulface,neutralface),moodgroup(neutral,anxious),traitanxiety Preinduction scores were not different for the anxious relative to group(lowTA,highTA),andperiod(preinduction,postinduction) theneutralgroup,ps(cid:8).05butpostinductionscoresweregreater onconflictscores.2AsinStudy1,therewasamaineffectofthreat, fortheanxiousrelativetotheneutralgroup,ps(cid:2).05.Therewere F(1,66)(cid:1)19.91,p(cid:2).001,(cid:5)2(cid:1).204,suchthatconflictscores no interactive effects involving trait anxiety. However, high TA p fortrialsprecededbyfearfulfaces(M(cid:1)88.02,SD(cid:1)43.09)were participantsendorsedfeelingmoreanxious,worried,fearful(M(cid:1) lowerthanconflictscoresfortrialsprecededbyneutralfaces(M(cid:1) 2.16,SD(cid:1)0.91)thanthelowTAparticipants(M(cid:1)1.56,SD(cid:1) 106.94,SD(cid:1)45.68).Therewasalsoamaineffectofperiod,F(1, 0.91), F(1, 112) (cid:1) 12.47, p (cid:1) .001, (cid:5)2 (cid:1) .100. High TA partic- p 66) (cid:1) 7.44, p (cid:1) .008, (cid:5)2 (cid:1) .101, such that conflict scores were ipantsalsoendorsedfeelinglessactive,alert,keyedup(M(cid:1)3.11, p lowerduringthepostinductionANTthanduringthepreinduction SD(cid:1)1.37)thanthelowTAparticipants(M(cid:1)3.92,SD(cid:1)1.36), ANT. No other main effects or interactions were significant, all F(1,112)(cid:1)10.05,p(cid:1).002,(cid:5)2(cid:1).082. p In the analysis of mean corrugator activity during the mood manipulations, contrary to expectations, there was no effect of 2Unlike in Study 1, a factor of Time (i.e., early vs. late trials) is not mood group, F(1, 112) (cid:1) 0.13, p (cid:1) .719, (cid:5)p2 (cid:1) .001, and no includedintheanalysesbecauseeachANTconsistsonlyofblocks1and interactionofmoodgroupandtraitanxiety,F(1,112)(cid:1)1.80,p(cid:1) 2,whicharetogetherequivalenttotheearlytrialsforwhichthefacilitation .182, (cid:5)2 (cid:1) .016. No significant difference existed between the effectwasdiscoveredinStudy1. p THREATFACILITATESSUBSEQUENTEXECUTIVECONTROL 1299 Fs (cid:2) 1.78, all ps (cid:1) .187. However, the critical Threat (cid:4) Mood threatandflanker(incongruent,neutral,congruent)onrawRT.An Group(cid:4)Periodinteractionwassignificant,F(1,66)(cid:1)8.61,p(cid:1) interaction of Threat (cid:4) Flanker emerged, F(2, 26) (cid:1) 12.64, p (cid:2) .005,(cid:5)2(cid:1).115. .001, (cid:5)2 (cid:1) .493. As in Study 1, RT were faster to incongruent- p p Tointerpretthisinteraction,weranseparateGLMstestingthe flanker trials preceded by fearful faces (M (cid:1) 561.12, SD (cid:1) effectsofthreat,moodgroup,andtraitanxietygrouponpreinduc- 105.20)versusneutralfaces(M(cid:1)585.79,SD(cid:1)96.20),p(cid:1).007. tionandpostinductionconflictscores.Asexpected,thepreinduc- RT were marginally slower for congruent-flanker trials preceded tion analysis revealed no main effects or interactions, all Fs (cid:2) by fearful faces (M (cid:1) 490.30, SD (cid:1) 92.81) versus neutral faces 1.67,allps(cid:1).200.Incontrast,thepostinductionanalysisrevealed (M(cid:1)473.29,SD(cid:1)83.92),p(cid:1).077.RTwerealsosignificantly amaineffectofthreat,F(1,66)(cid:1)14.02,p(cid:2).001,(cid:5)2(cid:1).175,and, slower for neutral-flanker trials preceded by fearful faces (M (cid:1) p most importantly, a significant interaction of Threat (cid:4) Mood 485.53, SD (cid:1) 81.59) versus neutral faces (M (cid:1) 466.71, SD (cid:1) Group, F(1, 66) (cid:1) 9.37, p (cid:1) .003, (cid:5)2 (cid:1) .124. For the anxious 78.68),p(cid:1).019.AsinStudy1,thispatternofresultssuggeststhat p mood group, conflict scores for trials preceded by fearful faces theattentionaleffectofthreatamongpeopleinananxiousmoodis (M(cid:1)68.34,SD(cid:1)41.96)werelowerthanconflictscorespreceded notsimplyathreat-relatedenhancementofattentiontothecentral byneutralfaces(M(cid:1)111.90,SD(cid:1)53.44),p(cid:2).001.However, locationorathreat-relatedincreaseingeneraleffort,whichwould fortheneutralmoodgroup,conflictscoresfortrialsprecededby manifest as faster responses for trials preceded by fearful faces fearful faces (M (cid:1) 90.52, SD (cid:1) 41.22) were not significantly relativetoneutralfacesforallthreeflankertypes. different from conflict scores for trials preceded by neutral faces Alerting. A GLM was conducted to examine the effects of (M (cid:1) 94.89, SD (cid:1) 52.43), p (cid:1) .587. Again, there were no threat (fearful face, neutral face), mood group (neutral, anxious), significanteffectsinvolvingtraitanxietygroup(seeFigure3a). trait anxiety group (low TA, high TA), and period (preinduction, Toexaminethenatureoftheconflicteffectintheanxiousmood postinduction)onalertingscores.Therewerenosignificantmain group, we computed a follow-up GLM to examine the effect of effectsorinteractions,allFs(cid:2)2.27,allps(cid:1).137.Consistentwith Study1,thissuggestsspecificityinthethreat-relatedfacilitationof executive control efficiency in the anxious group. See Figure 3b for a presentation of the alerting scores for all levels of threat, moodgroup,andtraitanxietygroupforthepostinductionANT.3 Discussion Consistentwithourhypothesis,participantseffectivelyinduced into an anxious mood state demonstrated facilitated efficiency of executive control following stimuli that were mildly threatening relative to nonthreatening. No such effect was present for partic- ipantseffectivelyinducedintoaneutralmoodstateandtherewere no threat-related or overall effects involving state anxiety on alertingefficiency.Moreover,therewerenoeffectsoftraitanxiety onexecutivecontrolefficiency.Infact,notonlydidthefacilitative effect of threat as a function of experimentally manipulated state anxietynotdependontraitanxiety,traitanxietyalsodidnotaffect executive control as a main effect. In sum, these results suggest that an attentional boost follows threat during an anxious state, regardless of one’s level of anxious disposition (at least within a normal range); this attentional benefit specifically takes the form ofimprovedefficiencyofexecutivecontrol. General Discussion Taken together, and consistent with DCF, the present studies demonstrate a replicable, threat-driven improvement in executive 3In order to test for possible effects of trait anxiety as a continuous measure, the five analyses for the manipulation check were conducted Figure3. Thesetwopanelsdepictattentionnetworkefficiencyscoresin againasGLMswithSTAIscoreasacovariateinsteadofTAGroupasa Study2forreactiontime(RT)differencesasafunctionofneutral(NEU) two-level factor. Similarly, for the final set of retained participants, the andanxious(ANX)moodstatesandlowandhightraitanxiety(TA)for three GLMs for conflict reaction time scores and the GLM for alerting trialsprecededbyfearfulfaces(blackbars)andneutralfaces(graybars). reactiontimescoreswererepeatedwithSTAIscoreasacovariate.Apart Theseresultspertaintoparticipantsforwhomthemoodinductionswere from the same four main effects of trait anxiety reported in the original effective.Thetopfigureshowsconflictscores(incongruent—congruent) analyses (i.e., STICSA somatic; STICSA cognitive, anxious, worried, andthebottomfigureshowsalertingscores(nocue—doublecue).Note fearfultriplet;active,alert,keyeduptriplet),therewerenoothersignificant that conflict scores are inversely related to executive control efficiency. effectsinvolvingtraitanxietyinanyofthesenineGLMs,allFs(cid:2)1.64,all Errorbarsrepresent(cid:7)1standarderror. ps(cid:1).204. 1300 BIRK,DENNIS,SHIN,ANDURRY control efficiency as a function of low-level threat during state sponds to one or more brain regions (Pessoa, 2009). Substantial anxiety. Study 1 reveals that this effect is not simply due to the evidence shows that the registration of affective significance is arousingnatureofanxiety;Study2revealsthatthiseffectdoesnot carriedoutbytheamygdala(Vuilleumier,2005).Perceptualcom- depend on the level of trait anxiety. Both studies suggest that petition in the visual domain is purported to occur in extrastriate threat-related facilitation of attention during an anxious state is regions(Pinsk,Doniger,&Kastner,2004).Executivecompetition specific to executive control efficiency as no such effect was ispurportedtooccurinfrontalregionssuchasanteriorcingulate observedforalertingefficiency. cortex (ACC; Westlye, Grydeland, Walhovd, & Fjell, 2011), Althoughpreviousstudieshaveshownthatlow-levelthreatcan which is known to operate in conjunction with dorsolateral pre- causeprioritizedattentionalprocessinginanxiouspeople,presum- frontal cortex (DLPFC; Silton et al., 2010). We cannot confirm ablyduetothreat’seffectsonexecutiveattention(e.g.,Fox,Russo, these neural mechanisms in this behavioral study; future studies Bowles, & Dutton, 2001), few have addressed the combined ef- mustinvestigatewhetherthefacilitatedbehavioraleffectisaccom- fects of threat and state anxiety on the efficiency of executive panied by an amygdalar response to threat stimuli that correlates attentionspecifically,andtodatethosestudieshaveonlyexamined withactivationinDLPFCorACC.Ifso,theaccumulatedevidence individual differences in anxious mood (Dennis et al., 2008; Fi- wouldsuggestthatfunctioninginaprefrontal–amygdalacircuitis nucane & Power, 2010). In the present studies we randomly themechanismunderlyingtheimprovedexecutivecontrolfollow- assignedparticipantstomoodconditionsinordertoeliminatethe ingtheperceptionofthreatduringanxiety. influence of potentially confounding extraneous variables (e.g., Ithasbeensuggestedthatfearfulfaces(unlikeangryfaces)are enduringpersonalitycharacteristicsthatsystematicallypredispose ambiguous,inthesensethattheysignalthepresenceofthreatbut particular people to experience state anxiety). This point is espe- notthesource(Whalenetal.,2001).Itmaybethatambiguityas cially important when attempting, as we did, to disentangle the to source of threat in the context of anxiety was important in effectsofstateandtraitanxiety. increasing perceived affective significance, which then signaled It is interesting that experimentally manipulated anxiety failed the need for executive control to resolve the ambiguity. Whalen toindependentlyfacilitateexecutivecontrolinthesestudies,which (1998)hassuggestedthatthistypeofambiguityismediatedbythe is consistent with findings of Pacheco–Unguetti and colleagues amygdala, which is consistent with DCF’s suggestion that the (2010). This null result for anxiety seems at odds with studies amygdalatracksaffectivesignificance.Ifthataccountisaccurate, suggesting that other negative emotions, specifically fear and onemightaskwhywedidnotobserveacorrespondingincreasein anger,canfacilitateexecutivecontrolevenintheabsenceofthreat alerting efficiency in conjunction with the increase in executive (Finucane, 2011). However, Gable and Harmon–Jones (2010a; controlefficiency.Oneexplanationisthattheasteriskcuesusedin 2010b) have suggested that any emotion that engenders suffi- the modified ANT may have been too subtle. Pacheco–Unguetti ciently high levels of motivational intensity—that is, “impetus to andcolleagues(2010)usedanauditorytonetoinducealertingin act”—should enhance attentional focusing (Gable & Harmon– their version of the ANT, and observed greater alerting in the Jones, 2010a; Gable & Harmon–Jones, 2010b). Although atten- anxious compared to the positive group. A stronger alerting ma- tionalfocusingisnotisomorphicwithexecutivecontrol,itmaybe nipulation may have been necessary to prompt a threat-related that our experimental manipulation of anxiety only engendered boostinalertinginthepresentstudy. sufficient motivational intensity to enhance executive control Notably, we did not observe facilitated executive control effi- when coupled with threat, a notion that is reminiscent of DCF’s ciencyinresponsetothreatinthehappyandneutralmoodstates. considerationofaffectivesignificance. This result is not fully consistent with the predictions of DCF or with previous studies (e.g., Cohen et al., 2011). Although the modelpositsthattheeffectoflow-levelthreatonexecutivecontrol Implications for Dual Competition Framework should be greatest during anxiety, it also suggests that threat AccordingtoDCF,threat-relatedstimulicarryaffectivesignif- shouldinfluenceexecutivecontrol,thoughtoalesserdegree,even icance, which alters behavior by strengthening sensory represen- forpeoplenotinananxiousstate.Anxiety-relatedvariationsinthe tations at the perceptual level and by prioritizing attention at the evaluationofaffectivesignificancemayhaveplayedthekeyrole executive level. The present findings are entirely consistent with in modulating executive control in the present studies; affective this formulation, at least with regard to manipulated anxiety. We significance may have simply been too low for people in happy speculate that the mechanism for threat-related facilitation of ex- andneutralmoodstoproduceaneffectonexecutiveattention. ecutive control during state anxiety was a soft prioritization of perceptual processing, specifically processing the spatial location The Role of Conflict Monitoring and Adaptation ofthreat.Inotherwords,experimentallymanipulatedstateanxiety led participants to pay greater attention to the spatial location of In line with the conflict-monitoring hypothesis of the ACC fearfulfacesthanneutralfaces,whichfacilitatedexecutivecontrol (Botvinick, Cohen, & Carter, 2004), we propose that reduced efficiency.However,wehavenodirectmeasureofattentiontothreat conflict scores in the present studies reflect the outcome of en- in the ANT. In future work, it would be useful to manipulate the hancedconflictmonitoringandresolutionprocesses.Wespeculate validitywithwhichthefacespredictthespatiallocationofthetarget. thatthisenhancementisinitiatedbytheamygdala,whichsignals Iffearfulfaceswereinvalidcuesforthespatiallocationofthetarget, the high affective significance of threat stimuli in the context of DCF would predict threat-related hampering (certainly not threat- anxiousmood.Next,theamygdalacommunicateswiththeACC, relatedfacilitation)ofexecutivecontrol. which signals the need to recruit executive control such that DCF is founded largely on neuroanatomical and functional DLPFC is engaged to facilitate subsequent conflict resolution. neuroimaging findings such that each aspect of the model corre- When the target finally appears, conflict is reduced, response

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Threat Facilitates Subsequent Executive Control During Anxious Mood Jeffrey L. Birk subsequent processing at the location of the threat as part of
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