cortex xxx (2013) 1e15 Availableonlineatwww.sciencedirect.com Journalhomepage:www.elsevier.com/locate/cortex Research report Exaggerated object affordance and absent automatic inhibition in alien hand syndrome Jennifer McBridea,b,*, Petroc Sumnerc, Stephen R. Jacksond, Nin Bajaje and Masud Husainf,g aInstituteofCognitiveNeuroscience,UniversityCollegeLondon,AlexandraHouse,London,UK bInstituteofNeurology,UniversityCollegeLondon,AlexandraHouse,London,UK cSchoolofPsychology,CardiffUniversity,TowerBuilding,ParkPlace,Cardiff,UK dSchoolofPsychology,UniversityofNottingham,Nottingham,UK eDepartmentofClinicalNeurology,NottinghamUniversityHospitalsNHSTrust,Queen’sMedicalCentre,Nottingham,UK fNuffieldDepartmentofClinicalNeurosciences,UniversityofOxford,UK gDepartmentofExperimentalPsychology,UniversityofOxford,UK a r t i c l e i n f o a b s t r a c t Articlehistory: Patients with alien hand syndrome (AHS) experience making apparently deliberate and Received13September2012 purposeful movements with their hand against their will. However, the mechanisms Reviewed17October2012 contributingtotheseinvoluntaryactionsremainpoorlyunderstood.Here,wedescribetwo Revised6December2012 experimentalinvestigationsinapatientwithcorticobasalsyndrome(CBS)withalienhand Accepted11January2013 behaviourinherrighthand.First,weshowthatresponseswiththealienhandaremade ActioneditorSergioDellaSala significantly more quickly to images of objects which afford an action with that hand Publishedonlinexxx comparedtoobjectswhichaffordanactionwiththeunaffectedhand.Thisfindingsug- geststhatinvoluntarygraspingbehavioursinAHSmightbeduetoexaggerated,automatic Keywords: motor activation evoked by objects which afford actions with that limb. Second, using Alienlimb abackwards masked primingtask, we found normal automatic inhibition ofprimed re- Objectaffordance sponsesinthepatient’sunaffectedhand,butimportantlytherewasnoevidenceofsuch Automaticinhibition suppression in the alien limb. Taken together, these findings suggest that grasping be- Maskedpriming haviours in AHS may result from exaggerated object affordance effects, which might potentiallyarisefromdisruptedinhibitionofautomaticallyevokedresponses. ª2013ElsevierLtd.Allrightsreserved. 1. Introduction releasingobjectsoncegrasped(seee.g.,BiranandChatterjee, 2004; Della Sala et al., 1991). Despite the fact that such in- Althoughmosthealthyadultsfeelthattheyhaveagreatdeal dividuals make seemingly deliberate and purposeful move- of control over their actions, some neurological patients do mentswiththeir“alien”hand,thereiscleardisparitybetween not.Patientswithalienhandsyndrome(AHS)mayinvoluntarily actions performed by the alien limb and the intentions of graspobjectsplacedwithintheirreach,experiencingdifficulty patients,whosubjectivelyreportthatthehandisnotunder * Corresponding author. Institute of Cognitive Neuroscience, University College London, Alexandra House, 17 Queen Square, London WC1N3AR,UK. E-mailaddresses:[email protected],[email protected](J.McBride). 0010-9452/$eseefrontmatterª2013ElsevierLtd.Allrightsreserved. http://dx.doi.org/10.1016/j.cortex.2013.01.004 Pleasecitethisarticleinpressas:McBrideJ,etal.,Exaggeratedobjectaffordanceandabsentautomaticinhibitioninalienhand syndrome,Cortex(2013),http://dx.doi.org/10.1016/j.cortex.2013.01.004 2 cortex xxx (2013) 1e15 theircontrol.Instead,theyreportthatthealienlimbbehaves irrelevanttotheobserver’stask.Ofcourse,inhealthypeople, asthoughithasamindofitsownorisbeingcontrolledbyan objectsdonotalwayselicitactionstowardsthem;thatwould external agent (e.g., Assal et al., 2007; Biran and Chatterjee, makepeopleentirelystimulus-bound.Hencethereisaneed 2004; Fitzgerald et al., 2007). Although these remarkable tosuppresssuchautomaticallyevokedaffordances.Indeedin grasping behaviours in AHS are now well-documented, we healthyobservers,thereisnowcompellingevidencethatre- understandverylittleaboutthemechanismsthatmightun- sponsesautomaticallyprimedbytheenvironmentcanalsobe derliesuchbehaviour. automatically suppressed (for reviews see Eimer and AHSisarelativelyraresyndrome(forareview,seeFisher, Schlaghecken,2003;McBrideetal.,2012a;Sumner,2007). 2000), so detailed investigation has been correspondingly Usingabackwardsmaskedprimingparadigm,Eimerand sparse.Someofthemostdetailedexperimentalworkcomes Schlaghecken (1998) showed that participants’ responses to from Riddoch and her colleagues (e.g., Humphreys and targets were typically speeded if targets were preceded by Riddoch, 2000; Riddoch et al., 1998). They instructed a pa- a compatible prime (a prime associated with the same tient with bilateral AHS to reach out and grasp a cup with responseasthetarget)comparedtowhentargetsfollowedan ahand.Thepatientwasabletodothiscorrectlyaslongasthe incompatible prime (a prime associated with the opposite cup’s handle was on the same side as the hand they were response to the target). Thus, a target directing a left hand instructedtousetograspthecup.However,ifthehandlewas response is faster if preceded by a (backward-masked) left on the opposite side, “interference” errors were generated prime relative to a right prime. However, when the interval withthepatientreachingwithwhicheverhandmatchedthe between masked-prime and target is extended beyond side the cup’s handle was on. For example, if instructed to w150 msec this usual positive compatibility effect (PCE) graspacupwiththerighthandwhenthecup’shandlewasto actually reverses to produce a negative compatibility effect the left, the patient would often erroneously grasp the cup (NCE;EimerandSchlaghecken,1998).Now,atargetdirecting withthelefthand.Theseeffectsareunlikelytobeperceptual a left hand response is actually slower if it is preceded by or attentional because fewer interference errors were made a(backward-masked)leftprimerelativetoarightprime. whenthetaskwastopointratherthantograsp,orwhenthe Aslongasappropriatestimuliareused(seeSchlaghecken patientrespondedtoLEDsinsteadoftocups.Thesefindings etal.,2007;LlerasandEnns,2004;Sumner,2008),thisNCEcan suggestthat,forthispatient,simpleobservationofagrasp- be interpreted as reflecting automatic suppression of the ableobjectmightbesufficienttoelicittheassociatedmotor primed response (see e.g., Eimer and Schlaghecken, 2003; plan for interacting with that object, even when the plan Ja´skowski,2007,2008,2009;Sumner,2007).Accordingtothese conflictswithcurrentgoals(seealsoBlakemoreetal.,2002). sensorimotor accounts of the NCE, initial motor activation Indeed,suchinvoluntarygraspingbehaviourinAHSmay evoked by the prime is subsequently suppressed when the be related to the longstanding view that, even in healthy primeisremovedoranovelstimulus(themask)isaddedto adults,viewingvisualobjectscanautomaticallyprimeactions the scene (e.g.,Boy et al., 2008; Ja´skowski, 2007, 2008, 2009). intheobserver.AHSmightrepresentanexaggeratedformof This suppression means that it takes longer to initiate the such automatic priming. Gibson (1979) described “affordan- suppressed response relative to a response which has not ces”aspropertiesofobjectsintheenvironmentwhichprime beeninhibited,therebyproducingtheNCE. an observer to act. For example, seeing a teapot with the SumnerandHusain(2008)suggestedthatsuchautomatic handletotherightmightautomaticallyprimetheobserverto suppression of automatically evoked responses might be reach out with the right hand to grasp the handle. Object crucialforgoal-directedbehaviourbecauseitfreesanorgan- affordanceeffectssuchasthesehavebeenextensivelystud- ism from stimulus-bound responses, and provides a level ied in healthy adults using stimulus-response compatibility playingfieldforalternativeactionstooccuraccordingtothe paradigms(e.g.,ChoandProctor,2010;Derbyshireetal.,2006; current goals of an animal. Consistent with this proposal, Iani et al., 2011; McBride et al., 2012b; Pellicano et al., 2010; Vainioandcolleagueshavereportedthatautomaticinhibition Phillips and Ward, 2002; Tucker and Ellis, 1998, 2001). For isnotrestrictedtomasked-primeparadigms,butalsooccurs example,TuckerandEllis(1998)presentedpicturesofobjects whenresponsesareaffordedbygraspablestimuli(e.g.,Vainio, which healthy observers classified as upright or inverted as 2009;Vainioetal.,2011;VainioandMustonen,2011). quickly and accurately as possible using a manual button Such considerations naturally raise the possibility of press.Crucially, the objects could be presentedso that they grasping behaviour in AHS arising from disruption of auto- maximallyaffordedaresponsewitheithertheleftortheright matic inhibitory mechanisms which, in healthy observers, hand.Althoughthisleft/rightorientationwasirrelevanttothe halt inappropriate activation of responses afforded by the participants’ task, responses were significantly faster and environment (see also Blakemore et al., 2002; Giovannetti moreaccuratewhenparticipantsrespondedwithahandthat et al., 2005). At present, however, there is very little direct wascongruentwiththe(task-irrelevant)responseaffordedby evidencetosupportthishypothesis,althoughtherearesome theobject. suggestivepiecesofevidence.Inhealthyadults,thesupple- Thesefindings, and themanyothers likethem (e.g.,Cho mentary motor area (SMA) in the medial frontal lobes is and Proctor, 2010; Derbyshire et al., 2006; Iani et al., 2011; associatedbothwithsimplyviewinggraspableobjectswith- McBrideetal.,2012b;Pellicanoetal.,2010;PhillipsandWard, outreachingforthem(e.g.,Gre`zesandDecety,2002)aswellas 2002; Tucker and Ellis, 1998, 2001), suggest that through with successful automatic inhibition of primed responses experience observers associate objects with particular ac- indexedbytheNCE(e.g.,Boyetal.,2010a,2011;Sumneretal., tions, and that these actions can be (partially) evoked by 2007).Intriguingly,AHShaslongbeenassociatedwithdamage perceptual processing of the object even when they are tothesesamemedialfrontalregions(e.g.,Bakheitetal.,2013; Pleasecitethisarticleinpressas:McBrideJ,etal.,Exaggeratedobjectaffordanceandabsentautomaticinhibitioninalienhand syndrome,Cortex(2013),http://dx.doi.org/10.1016/j.cortex.2013.01.004 cortex xxx (2013) 1e15 3 MarchettiandDellaSala,1998).AHSisincreasinglyrecognised in the alien hand compared to the non-alien hand (and in corticobasal syndrome (CBS, to distinguish it from the relative to healthy controls); and (ii) automatic inhibition pathologicentity,corticobasaldegeneration, CBD; seeBoeve of automatically evoked responses is reduced in the alien etal.,2003).CBSisarare(annualincidencerateshavebeen limb. estimatedataround.02per100,000individuals;Winteretal., 2010),slowlyprogressive,neurodegenerativeconditionwhich affects cortical regions as well as the basal ganglia (e.g., 2. Case report Fitzgeraldetal.,2007;Murrayetal.,2007;Riddochetal.,1998; TiwariandAmar,2008).Interestingly,CBSisalsoassociated PatientSAwasa72-year-old,right-handedwomanwhofirst with metabolic impairment in the SMA (e.g., Garraux et al., reportednoticinghersymptoms3yearspreviouslywhenshe 2000). hadafall.Atthattime,itwasobservedthatherspeechhad Tothebestofourknowledge,patientswithAHShavenot a telegraphic quality. She developed progressive difficulty previously been tested on object affordance “compatibility” speaking and writing, swallowing, and controlling her right tasks, or paradigms designed to investigate automatic inhi- hand.Shebegantouseherrightarmlessfrequently.Although bitionofprimedactions(e.g.,maskedpriming).Wemetwith shecouldvoluntarilymoveitifnecessary,therewasalackof fourpatientswithCBS(seeTable1forasummaryofpatients’ spontaneous use. Soon, she began to experience difficulty details),butunfortunatelythemotorsymptomsexperienced chopping vegetables using the right hand. She encountered bythreeofthesepatientsweresoseverethattheywerenot problems with her right hand grip, but at that time had no able to complete basic motor tasks. However, one patient, difficultylettingobjectsgo.Priortotesting,shenotedthather PatientSA,wasabletomakespeededmanualresponseswith walking had slowed. She began to experience difficulties either hand according to stimuli presented. Patient SA had standingfromaseatedposition.Therewasnofamilyhistory AHS which affected her right hand (involuntary grasping ofneurodegenerativedisease. movements to objects placedwithinherreach),and no evi- On examination, she had a profound expressive aphasia denceofalienbehaviourinherlefthand(seeTable1). and impaired articulation. However, she was able to com- Here we report results from two experiments conducted prehend 3-stage commands well. Visual fields were full to with Patient SA. Experiment 1 was designed to investigate confrontation. There was no evidence of visual or tactile whetherobjectaffordanceeffectswerestrongerinthealien extinction. Eye movements were full, but she was slow to hand relative to the unaffected hand. Our second study initiate saccades, particularly towards the left compared to comparedautomaticinhibitionofactioninthetwohands.If therightandtherewassomeevidenceofgazeimpersistence. grasping behaviour in AHS arises because of disruption of SuchoculomotordeficitsarenotuncommoninCBSpatients. normal automatic suppression of afforded responses, one Therewas no facial weaknessand palatal movements were mightpredictthat(i)objectaffordanceeffectsareexaggerated normal.Therewasnopoutreflex. Table1ePatientdetails.DetailsofthefourpatientswithCBS.PatientsFC,DH,andEFhadmotordeficitsthatwereso severethattheywerenotabletoperformsimplemotortasks.Theexperimentsreportedherewereconductedonlywith PatientSA.DIndicatesthatasymptomwaspresent,eindicatesthatthesymptomwasnotdetected. PatientSA PatientFC PatientDH PatientEF Demographics Age(years) 72 70 61 59 Sex(M/F) F F M F Manualsymptoms Involuntarygrasping þRighthand þBothhands e þLefthand Difficultyreleasing e þBothhands e e Intermanualconflict e e e e Armlevitation e þBothhands e e Mirrormovements e e e e Dystonia e þBothhands e e Dyspraxia e þBothhands þSomeinlefthand þ Tremor e e e e Rigidity þRightarm þBothhands þEspeciallyleftarm þ Impairedeye-movements Voluntarysaccades Horizontal Slowedtoleft Impaired e e Vertical e Impaired Impaired e Reflexivesaccades Horizontal e Impaired e Impaired Vertical e Impaired e e Othersymptoms Telegraphicspeech þ þ þ þ Visualextinction e þ e e Tactileextinction e þ þ e Pleasecitethisarticleinpressas:McBrideJ,etal.,Exaggeratedobjectaffordanceandabsentautomaticinhibitioninalienhand syndrome,Cortex(2013),http://dx.doi.org/10.1016/j.cortex.2013.01.004 4 cortex xxx (2013) 1e15 There was rigidity of the right arm and poor fine finger caudateheadbilaterally.Thesefindingswouldbeconsistent movements, but good strength throughout. The right hand with the clinical diagnosis of CBS. Selected images in Fig. 1 showed evidence of mild alien hand behaviour, with invol- demonstrate loss of volume of the left medial frontal and untarygraspingofanyobjectthatwasbroughtclosetoit.The parietalcortexwithapathologicallywidenedcingulatesulcus patientwasadamantthatshewasnotwillingthehandtodo (whitearrowhead);lossofcorticalvolumeadjacenttoawid- this, and she could not stop this behaviour even when she ened intraparietal sulcus particularly involving the superior madeanefforttodoso.Therewasnoevidenceofalienhand parietal lobe, most prominently on the left (yellow arrow- behaviourinthelefthand. head); widened sulci over superior parietal and frontal re- Examinationdidnotrevealanydystoniaorlimbapraxia, gions, including the left central sulcus (red arrowhead); and aboveandbeyondtheproblemsassociatedwithfinecontrolof reductionincaudateheadvolumebilaterally(leftsidemarked therighthandmovements.Therewasnoamorphosynthesis withgreenarrowhead). in the left hand. When she walked, there was reduced arm SA completed the two different experiments on two dif- swing,moreprominentlyontherightthanontheleft,butshe ferentdays,approximately4weeksapart.Theaffordancetask had a good stride length and postural reflexes were intact. was performed first. This study was approved by the local Therewasnoevidenceofsomeoftheotherbehaviourswhich humansubjectsethicscommitteeandthepatientgavewrit- are common in AHS: no levitation of either arm, no mirror teninformedconsentpriortotesting. movements,andnointermanualconflictbetweenthehands. Overall, the clinical presentation was considered to be con- sistentwithCBS. 3. Experiment 1jObject affordance task Magnetic resonance imaging (MRI; Fig. 1) demonstrated cortical atrophy,slightly more prominent over parietalthan Stimuli, task, response measurement and analysis follow frontal regions and in the left hemisphere compared to the closely from those reported in McBride et al. (2012b) which right. In addition, there was reduction in volume of the reporteddatafromyounghealthyindividuals. Fig.1eMRIbrainscansofPatientSA.(A)Sagittalimagedemonstratinglossofvolumeoftheleftmedialfrontalandparietal cortexwithapathologicallywidenedcingulatesulcus.(B)Coronalimagesshowinglossofcorticalvolumeparticularlyofthe superiorparietallobeadjacenttoawidenedintraparietalsulcus,mostprominentlyontheleft,togetherwithreductionin caudateheadvolumebilaterally.(C)Axialimagesdemonstratingwidenedsulcioversuperiorparietalandfrontalregions, includingtheleftcentralsulcus.Whitearrowhead[cingulatesulcus;yellowarrowhead[leftintraparietalsulcus;red arrowhead[leftcentralsulcus;greenarrowhead[caudatehead. Pleasecitethisarticleinpressas:McBrideJ,etal.,Exaggeratedobjectaffordanceandabsentautomaticinhibitioninalienhand syndrome,Cortex(2013),http://dx.doi.org/10.1016/j.cortex.2013.01.004 cortex xxx (2013) 1e15 5 3.1. Stimuliandtask experimenterifshehaddifficultyrecognisinganyoftheob- jects(shedidnotreportanydifficulty).Therewereanequal Eachtrialbeganwithpresentationofablackfixationcrosson numberoftrialscontainingstimuliofeachcategory(kitchen awhitebackgroundonaCRTmonitor(seeFig.2).Thiscross or toolbox), and an equal number of congruent and incon- subtended1degree(cid:2)1degreeofvisualangle,andwaspre- gruenttrialswithtargetsofeachcategory(kitchenortoolbox) sented in the centre of the screen for 1500 msec. Following in each block. Order of presentation was shuffled randomly a blank interval (200 msec), an imageof a target object was and independently for each block, and which image of the presented at screen centre for 2000 msec. Stimuli were pic- targetcategorywaspresentedwasdeterminedrandomlyand tures of ten household objects taken from the Object independentlyonatrial-by-trialbasis. Databank (courtesy of Michael J. Tarr, Brown University, http://www.tarrlab.org/)andVerfailleandBousten’s3Dobject 3.2. Apparatus database (see Verfaillie and Boutsen, 1995; Boutsen et al., 1998). Objects were matched for orientation. Five objects Stimuliweredisplayedona21inchCRTmonitor(1024(cid:2)768) belonged in a kitchen (fork, frying pan, knife, saucepan, whichtheparticipantviewedbinocularlyfromadistanceof spoon),andfiveinatoolbox(chisel,pliers,saw,screwdriver, 60 cm. Stimulus timing and presentation was locked to the spanner).Imagessubtended10.6e17.3degreesofvisualangle screen refresh rate of 100 Hz. Stimuli were presented using horizontally, and 2.8e5.3 degrees of visual angle vertically. aPCrunningPresentationsoftware(version13.1;http://www. Objectswereorientedwiththeirhandlesaffordinganaction neurobs.com). withtheleftorrighthand. Responsesweremeasuredusingtwospeciallydesignedde- The participant was instructed to respond by making vices,constructedfromarolledaneroidsphygmomanometer ashort,sharpsqueezeofagripforcemeasuringdevice(details cuff(Boso-clinicusI,ref:030-0-111),inflatedto20mmHg,con- below) with the left hand for kitchen objects, and with the nectedtoapressuretransducer.Onedevicewasheldineach right hand for toolbox objects. Therefore, depending on the hand, and the participant was instructed to make their re- orientationoftheobjectpresented,theobjectcouldaffordan sponsesbymakingashort,sharpsqueezeoftherolledcuffand actionthatwaseither“congruent”or“incongruent”withthe then release their grip. Grip force was converted to voltage requiredresponse.Thenexttrialbeganfollowingablankin- which was digitised and stored using a LabJack U3 HV data terval(1000msec). acquisitiondevicewithDAQFactorysoftware.Dataweresam- Before the experiment began, the participant practiced pledat1000Hz.Theparticipantwasencouragedtorespondas makingresponseswhileobservingtheoutputfromthepres- quicklyaspossiblewhilemaintainingahighlevelofaccuracy, sure transducers on a computer screen. Following a short butnoresponsefeedbackwasgivenduringtheexperiment. practiceblock(12trials)PatientSAcompletedtwosessionson the same day, each containing 4 blocks of 64 trials each, 3.3. Dataanalysis totalling512trialsafterpractice.Therewasanopportunityto restbetweenblocks.Allobjectswerepresentedatleastonce Continuous force recordings were locked to stimulus onset during practice, and Patient SA was instructed to tell the andepochedintoperiodsof2500msec,beginning500msec Fig.2eSequenceofeventsinatypicalaffordanceexperimenttrial.PatientSAwasinstructedtomakeasqueezeresponse withthelefthandiftheobjectpresentedbelongedinakitchen,andtomakeasqueezeresponsewiththerighthandifthe objectbelongedinatoolbox(thelatterisdepictedhere).Eachobjectcouldbepresentedintwopossibleorientations,sothat itaffordedanactionwiththeleftorrighthand(alltrialtypeswereequiprobable).Objectorientationwasirrelevanttothe task. Pleasecitethisarticleinpressas:McBrideJ,etal.,Exaggeratedobjectaffordanceandabsentautomaticinhibitioninalienhand syndrome,Cortex(2013),http://dx.doi.org/10.1016/j.cortex.2013.01.004 6 cortex xxx (2013) 1e15 before target onset. Data were smoothed using a simple 5- than3SDsfromeachparticipants’meanRTforthathandfor point moving average to reduce high-frequency noise. The thatcondition).Theelderlyhealthycontrolshadfasteroverall resulting waveforms were baseline corrected on a trial-by- RTs (mean ¼ 609 msec) and showed a smaller congruency trialbasisaccordingtotheaveragebaselineactivityforeach effect [mean ¼ 14 msec; congruency effect was reliable in responsedeviceduringthe200msecpre-stimulusperiodon elderly controls: t(24) ¼ 3.15, p ¼ .004] than for Patient SA’s eachtrial. alienhand.1TodirectlycomparetheperformanceofPatient A response (either correct or incorrect) was said to have SA’s alien hand to that of healthy elderly controls, we con- occurred in a trial if at any point after the target stimulus vertedtheoverallmeanRTandaffordanceeffectforthealien onsetuntiltheendofthetrial,twocriteriaweresatisfied:(i) handtoz-scores,calculatedaccordingtotheelderlycontrols’ the force measured was greater than 3 SDs from the mean samplemeansandSDs.Thez-scoresfortheaffordanceeffect forcemeasuredduringthepre-stimulusbaselineperiodand and overall RT shown for Patient SA’s alien hand were 2.82 that was followedby at least 18/20pointsthat also reached and4.24,respectively.Asthesearebothbeyondthe95%limits thisthreshold;and(ii)therewasanincreaseinresponsebyat (two-tailed) of the controls’ distributions (95% limits are least .01 V over the following 100 points or less. Response indicatedbyaz-scoreof1.96),itisunlikelythatPatientSA’s onset time (RT) was defined as the first point that satisfied effectsaresimplyanextremecaseinthenormalelderlydis- these criteria. Peak response was determined as the max- tribution,andthattheseeffectsareduetoage.2 imum amplitude of the response made in a trial that was To investigate how often differences like those exhibited surroundedbypointsoneithersidewiththesameorlower bySA’salienlimbexistinhealthycontrols,weanalysedthe amplitude. individual affordance effects for left and right hands in the Outliersweredefinedasanyresponsesgreaterthanthree younghealthycontrolspreviouslyreportedbyMcBrideetal. standarddeviations(SDs)awayfromthemeanresponsetime (2012a), plus the previously unpublished data from elderly forthathand,inthatcondition(congruentorincongruent)in healthy controls, mentioned above. None of these healthy thattestingsession.Remainingcorrectresponsetimeswere adultsshowedthesamepatternofeffectsshownbySA,with enteredintoa2(hand)(cid:2)2(congruency)(cid:2)2(session)factorial a significant interaction between the effects of hand and analysisofvariance(ANOVA).Therewasnosignificanteffect congruency,andasignificantasymmetryinoverallRT. of session (morning or afternoon) on RTs, and the effect of However,overallRTsinSA’salienhandwerelongerthan session did not interact with any of the other factors (all those recorded in the non-alien hand, as well as those F’s < 1). Therefore, subsequent analyses collapse across reported in young and elderly controls. Therefore, we per- session. formedfurtheranalysestoinvestigatethepossibilitythatthe differenceincongruencyeffectacrossPatientSA’shandswas simply attributable to the difference in baseline RT. We re- 4. Results and discussion plotted the congruency effect as a function of RT in a delta plot(seevandenWildenbergetal.,2010,forareviewofthis The key motivation in conducting Experiment 1 was to techniqueanditsadvantages).Foreachhandseparately,un- examinewhetherresponseswithPatientSA’salienhandwere trimmed(includingthose trialsconsidered“outliers”forthe moresusceptibletoprimingbyobjectaffordancesrelativeto ANOVAanalysis)correctRTsweredividedaccordingtotrial responseswithhernon-alienhand.Herresponsesweregen- congruency (congruent or incongruent), rank-ordered, and erally slower than those reported for healthy adults on this then divided into eight bins of equal size. On two trials, no task(seeMcBrideetal.,2012b).Moreover,SA’sleft(non-alien) correct response was detected. Data for these trials were hand responses were significantly faster than right (alien) replacedwiththemeancorrectRTforthathandandcondition hand responses [see Fig. 3; left mean ¼ 836 msec vs (thisisameanstokeepthetotalnumberoftrialsthesamein right¼1090msec,F(1,497)¼307.47,p<.001].Furthermore, each condition and dividable by 8, to avoid problems asso- stimuliwhichaffordedacongruentresponseproducedfaster ciatedwithunequalbinsizes).ThemeanRTineachbinfor reactions than stimuli which afforded an incongruent each condition was then calculated and the difference be- response [incongruent mean ¼ 983 msec; congruent tweenincongruentandcongruenttrialsisplottedagainstthe mean¼944msec;F(1,497)¼7.13,p¼.008].Importantly,the mean RT for that bin (see Fig. 3B), giving the size of the congruencyeffectwasmuchlargerforthealienthanforthe non-alien hand [significant congruency (cid:2) hand interaction: F(1,497)¼6.62,p¼.010].ThisinteractionisshowninFig.3A. 1We have also examined the median affordance effects from untrimmed RTs. For Patient SA: median affordance effect non- Thecongruencyeffectshowninthealienhand(76msec) alien hand ¼ 7 msec; median affordance effect alien wasseveraltimeslargerthanwehavefoundusingidentical hand ¼ 67 msec; for healthy elderly controls: mean of median apparatus in healthy young controls (mean of median affordanceeffect¼13msec. RTs¼16msec,seeMcBrideetal.,2012b).Wealsohaveasyet 2Recently,CrawfordandGarthwaite(2012)havesuggestedan unpublished data on this task from elderly healthy controls alternativemethodfortestingwhetheraneffectshownbyasingle (N¼26;aged54e75years;meanage¼64years;onepartic- caseislikelytohavebeendrawnfromthecontroldistributionby ipant,whoshowedanaverageaffordanceeffectof(cid:3)111msec, conducting a t-test following Crawford and Howell’s (1998) method. For completeness, we have also conducted this test, wasremovedasanoutlier).AsforPatientSA,wecalculated whichalsoindicatesthattheaffordanceeffect[t(24)¼2.76,p<.01] each elderly control’s mean RT for each condition, after andoverallRT[t(24)¼4.16,p<.01]forPatientSA’salienhandare removingoutliers(followingthesamecriteriaforoutlierse- reliablydifferentfromthoseshowninelderlycontrols(testsare lectionasforPatientSA,weremovedanyRTthatwasgreater two-tailed). Pleasecitethisarticleinpressas:McBrideJ,etal.,Exaggeratedobjectaffordanceandabsentautomaticinhibitioninalienhand syndrome,Cortex(2013),http://dx.doi.org/10.1016/j.cortex.2013.01.004 cortex xxx (2013) 1e15 7 A B Fig.3eResultsfromaffordancetask.(A)MeanRTs(trimmedatD/L3SDsfromthemeanforeachhandineachcondition) forresponsesmadewitheachhandtotargetstimulithatwereincongruent(greybars)orcongruent(whitebars)withthe responseaffordedbytheobject.ErrorbarsdenoteD/L1SEM.(B)MeanoctilecongruencyeffectsasafunctionofmeanRT forthatoctile,calculatedseparatelyfortheleft(greyline)andright(blackline)hands.Datafromthelongestandmost variableRTbinareincludedforcompleteness,andareshownhereasdottedlinesandopensymbols.*DenotesBonferroni- correctedt-testp<.05. congruency effect for that part of the RT distribution. Un- t(24)¼.55,p¼.59;elderlycontrols’meanaffordanceeffectfor trimmed RTs such as these typically have long “tails” pro- kitchen stimuli ¼ 12 msec; for toolbox stimuli ¼ 16 msec; duced from a number of slow outlier responses (see also t(24) ¼ .570, p ¼ .574]. Therefore, there is no indication that Maylor et al., 2011). The increased variability and reduced thereisanyreliabledifferenceintheaffordanceselicitedby reliabilityoflongRTsmeanthatitisdifficulttodrawmean- differentstimulustypes. ingfulconclusionsfromthelastbin,butitisincludedinthe As noted in the methods, the particular object presented figureforcompleteness(dottedlines). was determined randomly and independently for each trial If the difference in congruency effects across the two (while the number of trials in each condition was held handswassimplyinlinewithdifferencesinbaselineRT,then constant). Therefore, perhaps the very large affordance wemightexpectsimilarcongruencyeffectsinthosepartsof effectshowninPatientSA’sright(alien)handisduetoasubset theRTdistributionwhichoverlapacrossthetwohands(i.e., of toolbox stimuli which by chance appeared more (or less) for responses which onset between approximately often than the others. To investigate this possibility, we cal- 800e1000msecafterthestimulusappeared).However,there culated how often each particular toolbox object was pre- isclearseparationbetweenthecongruencyeffectsshownby sented. Four out of the five objects appeared 54 or 55 times theleftandrighthandsinthispartoftheRTdistribution,soit each,andoneitem(thechisel)waspresented39times.Reas- seems unlikely that the interaction between the effects of suringly,therewasnoreliableinteractionbetweentheafford- handandcongruencyisbeingdrivenbydifferencesinbase- ance effect and the particular toolbox exemplar presented lineRT.Thus,PatientSAshowsasignificantlylargerafford- [congruency (cid:2) object interaction: F(4, 239) ¼ 1.20, p ¼ .31]. ancecongruencyeffectwhenmakingresponseswithheralien Furthermore, we repeated the analysis of correct RTs after (right)hand,comparedtohernon-alien(left)hand,suggesting removingthosetrialswhichcontainedtherelativelyinfrequent thatanobject’saffordancehadanexaggeratedeffectonher exemplar (the chisel). The affordance effect shown for the alienlimbcomparedtotheunaffectedhand. remaining toolbox items remains very large and statistically The stimulus-response mappings Patient SA used in significant (incongruent mean ¼ 1122 msec; congruent Experiment 1 were held constant over the course of the mean¼1064msec;congruencyeffect¼58msec,p¼.03). experiment.ThiswastopreventanypossibledifficultiesPa- Errorswereveryinfrequent(anabove-thresholdresponse tient SA might have experienced with task-switching if we was made by the erroneous hand on only 10/512 trials e had changed the stimulus-response mapping part-way approximately2%ofalltrials).Thiserrorrateissimilartothat through the experiment (see Alvarez and Emory, 2006, for whichweobservedinyoung(approximately5%),andelderly discussion).Toexaminewhetherthereisanydifferenceinthe (approximately3%)healthycontrols.Oftheseerrorsmadeby affordance effects normally produced by different stimulus PatientSA,8/10weremadebytheright(alien)handwhenthe types, we analysed affordance effectsto these same stimuli taskrequiredaresponsewiththelefthand.Sixerrorswere fromyoung(previouslyreportedinMcBrideetal.,2012b)and detected by the alien limb in response to affordance incon- elderly(previouslyunpublished)healthycontrolparticipants, gruent trials (in other words, when the object presented where stimulus-response mapping was counterbalanced requiredalefthandresponse,butwasorientedsuchthatit across participants. Young and elderly healthy controls affordedaright-handresponse),and2errorsinresponseto showed comparable affordance effectsfor kitchen and tool- affordancecongruenttrials.Errorswerenotconfinedtoone box stimuli [young controls’ mean affordance effect for particularstimulus,andinsteadwerespreadacross7differ- kitchenstimuli¼18msec;fortoolboxstimuli¼15msec;no ent exemplars. As errors were so infrequent, they were not reliable difference of stimulus type on affordance effect: analysedanyfurther. Pleasecitethisarticleinpressas:McBrideJ,etal.,Exaggeratedobjectaffordanceandabsentautomaticinhibitioninalienhand syndrome,Cortex(2013),http://dx.doi.org/10.1016/j.cortex.2013.01.004 8 cortex xxx (2013) 1e15 SAcompleted8blocks(4ofeachSOAcondition)of84trials 5. Experiment 2jMasked priming task each,makingatotalof672trials. Aschematicofthestimuliandtimingsforthistaskcanbe InExperiment2,weusedabackwardsmaskedprimingtask seen in Fig. 4. Note that the total presentation time of each (adapted from Sumner et al., 2007) to investigate automatic stimulus (prime, mask, target) was the same in both SOA inhibitionofresponsesthathadbeenautomaticallyprimedin conditions. thealienandnon-alienhands. Thetargetstimulusappearedafterthemaskhadonset,and In order to be sure of producing automatic priming and waseitheraleft-,orright-pointingdoublearrowhead(sothatit inhibitionofresponses,itwasnecessarytochangetheinter- waseithercompatibleorincompatiblewiththeprimestimu- val between masked-prime and target. There are several lus). The target appeared in one of three possible locations, methodsreportedin theliteratureto achievethis.Onepos- centred 5 degrees of visual angle to the left, to the right, or sibilitywouldbetopresentthetargetstimulusoncethemask abovethecentreofthescreen.Theparticipantwasinstructed had offset, and change the duration of the mask. However, toignorethetarget’sposition,andtorespondtothedirection shortermaskswouldbeexpectedtomasktheprimestimulus ofthisarrowheadbysqueezingwitheitherthelefthand(for lesswell,whichcouldhavestrongeffectsontheprimingof left-pointingtargets)ortherighthand(forright-pointingtar- responses. Alternatively, some researchers have usedmeta- gets) as quickly and accurately as possible. In each block of contrast masking e that is, to use a stimulus which masks trials there were an equal number of trials with each target theprimebysurroundingit.However,suchmasksareprob- type(left-,andright-pointing)ineachpossibleposition(left-, lematicbecausemaskscanactasprimestimuliintheirown right-, above-centre), with each prime type (compatible and right e as masks of this type typically contain elements of incompatible), presented in randomly shuffled order deter- both possible primes, any NCE obtained using such a mask mined independently for each block. The target stimulus may not be produced by response inhibition, but by mask- remained on the screen for 200 msec. There was a blank inducedprimingoftheresponseoppositethatevokedbythe intertrialinterval(ITI)of2500msecbeforethenexttrialbegan prime (see “object updating” e.g., Lleras and Enns, 2004; withthefixationcross.Datarecordingandanalysisprocedures Sumner,2008).Aswewereinterestedintheeffectsofauto- werethesameasthoseusedintheaffordanceexperiment. maticresponseinhibition,wesoughttoavoidthispossibility. Left-andright-pointingdoublearrowheads(e.g.,“<<”and Thus, we followed a well-established, standard method “>>”)servedasprimesandtargets.Thelinesmakingupthese reportedpreviouslyintheliteraturewhichisknowntoreliably stimuliwereeach1degreeofvisualanglelong,andthelines producePCEsandNCEsandwhichkeepsthedurationsofeach ineacharrowheadhadanangularseparationof60(cid:4)(30(cid:4)above stimulus (prime, mask, and target) constant. We presented and below the horizontal). Masks were constructed of 30 masks and primes simultaneously in short stimulus onset pseudo-randomlyorientatedlinesarrangedintoa6(cid:2)5grid asynchrony(SOA)conditions,andintroducedablankscreen centred over the centre of the screen. To prevent any per- betweenmaskandtargetinlongSOAconditions(seee.g.,Boy ceptual interactions between prime and mask modulating etal.,2010a;2010b;BoyandSumner,2010;Schlagheckenetal., priming effects (see “object updating” accounts of the NCE 2006,2003;SchlagheckenandEimer,2002;Schlagheckenand e.g., Lleras and Enns, 2004) lines in the mask avoided any Maylor,2005).Itispossiblethatdifferencesintheshort-and orientationwithin5degreesofthelinesmakinguptheprime long-SOAtrialsequencemayaffectglobalRTseforexample andtarget.Thelinesinthemaskwerebetween1.5and3de- theoffsetofthemaskinthelongSOAconditionmayserveas grees of visual angle long. Line length and orientation were awarningsignalthatthetargetisabouttoappearandthus determinedrandomlywithintheselimitsandindependently speedresponsesinthelongSOAcondition.However,assuch for each line in the mask. Thus, the mask was between effectsareexpectedtohaveaglobalinfluenceonRTs,andnot 3.5 (cid:2) 3.5e5.5 (cid:2) 5.5 degrees of visual angle, centred on the affect one condition (compatible or incompatible) or hand centre of the screen. A new mask was constructed on each (alienornon-alien)morethantheother,theyshouldnotbe trialtopreventperceptuallearningofthemaskwhichcouldin able to account for any differences in compatibility effect turnleadtoincreasedprimeidentification(e.g.,Schlaghecken showninthedifferenthands. etal.,2008).SuchmaskshavebeenshownnottoinvokeNCEs by object updating (Sumner, 2008) or by perceptual in- 5.1. Stimuliandtask teractions (Boy and Sumner, 2010), and thus any NCEs observedcanbeattributedtomotorinhibition. Eachtrialbeganwithpresentationofawhitefixationcrosson amid-greybackground.Thiscrosssubtended1degree(cid:2)1de- 5.2. Proceduretodeterminethresholdforprime greeof visual angle,and waspresentedin the centreofthe perception screenfor500msec.Followingablankintervalof200msec, theprimeappearedinthecentreofthescreenandremained Priortothemainexperiment,thedurationoftheprimewas for50msec(seebelowforhowthisdurationwasdetermined). settobelowthethresholdforconsciousperception(50msec Theprimewasthenreplacedwiththemaskwhichremained duration) using a psychophysical staircase procedure. Here, onthescreenfor100msec.Twomask-targetSOAswereused oneachtrialaprimeandmaskwerepresentedwithnotarget, inthisexperiment;20msec(shortSOA,whichwasexpected and the participant was instructed to make a 2-alternative to produce a PCE) and 150 msec (long SOA, which was forced-choice button-press according to the direction of the expectedtoproduceanNCE).SOAconditionswerepresented primestimulus.Theparticipantwasinstructedtomaketheir inalternatingblocks,startingwithalongSOAblock.Patient best guess if they were unsure of prime direction, to Pleasecitethisarticleinpressas:McBrideJ,etal.,Exaggeratedobjectaffordanceandabsentautomaticinhibitioninalienhand syndrome,Cortex(2013),http://dx.doi.org/10.1016/j.cortex.2013.01.004 cortex xxx (2013) 1e15 9 A B Fig.4eStimulussequenceandrelevanttimingsinthemasked-primetask.Inthisexample,asprimeandtargetare associatedwiththesameresponse,bothtrialsshownarecompatibletrials.Thepatientmadespeededsqueezeresponses accordingtothedirectionofthetargetpresentedoneachtrial(alltrialtypeswereequiprobable).(A)Showsthetrial sequenceforshortSOAblocks,and(B)showsthetrialsequenceforlongSOAblocks.Thetargetarrowheadsappeared 5degreesofvisualangleeithertotheleft,right,orabovethecentreofthescreen.Alltrialtypeswereequiprobable.Aleft handsqueezeresponsetoleft-pointingtarget,primecompatibletrialsareshownhere.Notethatthemaskandtargetwere eachpresentedforthesametotaltimeinbothshortandlongSOAconditions. concentrateonbeingaccurate,andthatspeedwasunimpor- This method of analysis is essentially like the delta plot tant for this part of the task. The prime duration began at method used to analyse the data in Experiment 1. Trials in 120msec,andthenwasvariedaccordingtoafixed-step,1-up/ whichnocorrectresponsewasdetectedwerereplacedwith 2-downprocedure:Aftertwoconsecutivecorrectresponsesto themeancorrectresponsetimeforthathand,condition,and primespresentedatthesameduration,primedurationwas mask-targetSOA(thisisameanstokeepthetotalnumberof reduced by 10 msec on the next trial; after an incorrect trialsthesameineachconditionanddividableby8,toavoid response it was increased by 10 msec, within a range of problemsassociatedwithunequalbinsizes).Then,response 10e200 msec. This staircase procedure terminated after 10 timeswerere-calculatedrelativetotheprimeonset(weadded “reversals”. The fastest prime duration was 60 msec (which theprime-targetSOAforeachtrialtotheRTforthattrial),and waspresentedtwice,andtheprimewasincorrectlyidentified rank-orderedforeachhand(leftorright)foreachcondition onthesecondpresentation),andthemeanprimedurationat (incompatible or compatible) across SOA conditions. This the reversals was 84 msec. Thus, for the remainder of the meantthattherewassomere-shufflingofresponsesacross experimenttheprimedurationwassetto50msec,whichwas SOAconditions(becauseaslowresponseonashortSOAtrial the faster than the fastest prime duration measured during may have a longer prime-locked response time than a fast thestaircase(andwasnotreliablyidentified),andfasterthan response on a long SOA trial). These prime-locked response theaveragedurationofthereversals. times were divided into 8 bins of equal size. The average compatibility effect (average incompatible RT (cid:3) average 5.3. Prime-lockedanalysisofRTs compatibleRT)foreachbinforeachhandwascalculated,and plottedrelativetothemeanRTforthatbinandhand.Lastly, We followed the method described in Schlaghecken et al. thestatisticalsignificanceofthecompatibilityeffectineach (2011) to analyse RTs in a masked priming task relative to bin was determined by conducting a Bonferroni-corrected prime onset rather than to target onset. Schlaghecken and unpairedt-testontheresponsetimesineachbin. colleaguesnotedthattheincreasedtrial-to-trialvariabilityin olderadults’RTsmayobscuretheprimingeffectsthatwould be revealed by traditional analyses which separate target- 6. Results and discussion locked RTs according to prime-target compatibility and mask-target SOA on each trial. In fact, Schlaghecken et al. The results for the masked-prime experiment are shown in (2011) showed that calculating RTs relative to prime onset Fig.5.TheunaffectedlefthandshowedthepatternofRTef- could reveal a reliable NCE in older participants’ RTs when fects that would be expected from healthy individuals in thesewerenotshownbytraditionalanalyses. amaskedprimingtask(e.g.,SchlagheckenandEimer,2002). Pleasecitethisarticleinpressas:McBrideJ,etal.,Exaggeratedobjectaffordanceandabsentautomaticinhibitioninalienhand syndrome,Cortex(2013),http://dx.doi.org/10.1016/j.cortex.2013.01.004 10 cortex xxx (2013) 1e15 Schlaghecken et al. (2012) have recently suggested that prime-locked distributional analyses like those performed herecanproduce‘significant’effectsinsomelatencybinsby chance.Herewedonotrelyonsearchingforsignificantbins, butrathercomparethewholepatternbetweenthealienand non-alienhands.Nevertheless,wehavealsotestedthepos- sibilitythatthepatternshownbythealienhandcouldariseby chance from a ‘healthy’ distribution of data. We pooled the prime-locked RT data from the non-alien hand across com- patibleandincompatibleconditionsandrandomlyre-labelled trials as incompatible and compatible. We then re-ran the distributional analyses described here. After repeating this process100times,noneofthe100randomlyre-sampleddata Fig.5eResultsfrommaskedprimingtask.Meanoctile setsshowedthesamereliablePCEin6/8RTbinsasshownby compatibilityeffectsasafunctionofmeanRT(relativeto thealienhand(andonly3outof100showedareliablePCEin primeonset)forthatoctile,calculatedseparatelyfortheleft anyof5/8bins).Thus,wesuggestthatitisveryunlikelythat (greyline)andright(blackline)handsfollowingthe responsesfromPatientSA’salienhandactuallybelongtothe proceduredescribedinSchlagheckenetal.(2011).Data same distribution as that of her non-alien hand, and only fromthelongestandmostvariableRTbinareincludedfor showedaconsistentlysignificantPCEduetochance. completeness,andareshownhereasdottedlinesand Thus,thereisnoevidenceofautomaticmotorinhibitionof opensymbols.Theleft(non-alien)handshowsapatternof primed responses, indexed by the NCE, for responses made compatibilityeffectswhichissimilartothatreported withthealienhand.Itisunlikelythatthisdisruptedinhibition elsewhereinhealthycontrols(PCEfollowedbyNCE).The ismerelyduetoageornon-specificeffectsofdisease,because right(alien)handshowsastrongPCE,andthereisno reliableinhibitionisshownforresponsesmadewiththeleft evidencethatthisturnsnegativeastheprime-response (non-alien)hand. intervalincreases.*DenotesBonferroni-correctedt-test p<.05. 6.1. Spatialcongruencyeffects The design of the masked priming experiment required the targetstimulustobepresentedinadifferentlocationtothe For fast responses (which occurred most quickly after the primeandmask(toavoidspatialandtemporaloverlappingof prime was presented), RTs were faster for compatible trials stimuli in the short SOA condition). Thus, on each trial the relative to incompatible trials (a PCE). For responses that targetwaspresentedtotheleft,totheright,orabovecentral occurred later, responses were faster on incompatible trials fixation.Thisspatialaspectofthetargetstimulusmighthave relativetocompatibletrials(aNCE).Thereissomeevidence affectedperformanceinamannersimilartotheSimoneffect thattheprimingeffectmayhavereturnedtopositiveagainat (e.g.,LuandProctor,1995,forareview)andthespatialStroop thetailendofthedistributioninbin8,whichisalsoconsistent effect (e.g., Banich et al., 2000). Thus, the design of this withpreviousstudies(seee.g.,SumnerandBrandwood,2008), experimentprovidesanopportunitytoinvestigateanyeffect butthis effectmay have beenskewed by outliersin thetail ofspatialcongruencyinPatientSA. end of the distribution, and did not reach statistical signifi- Afterremovalofanyresponsewhichoccurredþ/(cid:3)3SDs cance(Bonferroni-correctedp>.1). away from the mean of that condition, we calculated mean A very different pattern emerged in the RTs for the re- RTs for each hand for spatially incongruent, neutral, and sponses made with the alien hand. Here, responses were congruenttrials.ThesemeanRTsare showninFig.6A.The consistentlyfasterforcompatibletrialsrelativetoincompat- expected location congruency effects were observed: re- ible trials (a PCE), and there was no evidence of this effect sponseswerefastestwhenthetargetappearedinalocation turningnegative(NCE),evenatlaterpointsinthedistribution. thatwascongruentwiththerequiredresponse,andslowest WhilethereisasmalldifferenceintheoverallRTsfortheleft when the target appeared in a location that was congruent and the right hands, responses made with the left hand with the response opposite that required to the target showedasignificantNCEbyaround850msecaftertheprime {incongruent condition; [F(2, 627) ¼ 7.37, p ¼ .001]}. Also, as hadonset,whereasright-handresponsesstillshowedasig- expected,responsesmadewiththeleft(non-alien)handwere nificant PCE at 1050 msec.Thus, these distributions suggest significantlyfasterthanresponsesmadewiththeright(alien) thatthis difference incompatibilityeffectisnotlikelyto be hand[F(1,627)¼51.12,p<.001].Importantly,theinteraction duetotheslightlylongerrightthanlefthandresponses. betweentheeffectsofhandandcongruencydidnotapproach Furthermore,we suggestthat it is unlikelythat the inhi- statisticalsignificance[F(2,627)<1]. bition has simply been delayed in right-hand responses. Asnotedabove,adeltaplotcanbeamoresensitivewayof Mayloretal.(2011)reportedreliableNCEsforelderlypartici- examiningRTeffectsthancomparingaverageRTs.Therefore, pantsforresponseswhichoccurredby500msecafterprime we have plotted the spatial congruency effect (incongruent onset,whereasPatientSAhereshowedaprimingeffectthat RT(cid:3)congruentRT)over8RTbins(seeFig.6B)accordingtothe wasstillpositiveforresponseswhichwererecordedaround procedures described above. The pattern of spatial congru- 1050msecaftertheprimehadonset. encyeffectswassimilarforbothhands,andtheeffectdidnot Pleasecitethisarticleinpressas:McBrideJ,etal.,Exaggeratedobjectaffordanceandabsentautomaticinhibitioninalienhand syndrome,Cortex(2013),http://dx.doi.org/10.1016/j.cortex.2013.01.004
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