Facial Expressions, Their Communicatory Functions and Neuro-Cognitive Substrates Author(s): R. J. R.

Blair Source: Philosophical Transactions: Biological Sciences, Vol. 358, No. 1431, Decoding, Imitating and Influencing the Actions of Others: The Mechanisms of Social Interaction (Mar. 29, 2003), pp. 561-572 Published by: The Royal Society Stable URL: http://www.jstor.org/stable/3558134 . Accessed: 13/07/2013 02:05
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Ekman 1997). although the expression may to observers. To do this.Instead. 206. Health. they are important signals to modulate currentbehavioural responding. and that they impart specific informationto the observer. The argument is simply that ? 2003 The Royal Society Phil.1220 561 This content downloaded from 14. Department of of Bethesda.THE ROYAL c-m RJP SOCIETY Publishedonline5 February 2003 Facial expressions. Displays of anger or embarrassment. are appetitive unconditioned stimuli which increase the probabilityof actions to which they appear causally related (Matthews & Wells 1999). Instead social context predicts probability of emotional expression in humans as it does probabilityof non-verbal displays in non-human species (Cheney & Seyfarth1980. J. B (2003) 358. Izard & Malatesta 1987. Services. Similarly. referenceto the neuro-psychiatric be made. Instead.2002.the neuro-cognitive systems necessary for their production. USA (blairj@intra. the empirical literature indicate that individuals display emotional expressions automatically as a function of the degree to which they feel a particularemotion (Fridlund 1991.3 on Sat. Camras 1994). autism 1. psychopathyand acquired sociopathy will Finally.235.in reparatory contrast.it has been suggested that sad facial expressions also act as aversive unconditioned stimuli discouraging actions that caused the display of sadness in anotherindividual and motivating behaviours (Blair 1995). Trans. Keltner & Anderson 2000).139. 561-572 DOI 10.infantsmilingfromthe age of 10 months is almost entirelydependent on visual contact with the caregiver: without such contact the infantis very unlikelyto smile (Jones & Raag 1989. amygdala. therehave been suggestionsthatemotional expressionsare automatic displays that occur as a function of the emotional experience of the individual (Darwin 1872. Thus. Keywords: facial expressions. INTRODUCTION Facial expressions are a crucial component of human emotional and social behaviour and are believed to represent innate and automatic behaviour patterns (Darwin 1872). Buck 1984. In autism.1098/rstb. Lond. and happiness are reinforcersthat modulate the probin the abilitythat a particularbehaviour will be performed future.MD 20892-2670. 1991). In contrastto the communicatoryfunctionassumption.gov) Human emotional expressionsserve a crucial communicatoryrole allowing the rapid transmissionof valence informationfrom one individual to another. Thus. In acquired sociopathy. 13 Jul 2013 02:05:56 AM All use subject to JSTOR Terms and Conditions . the expression is an automatic consequence of the individual's experience does not (Ekman 1997).the response to abilityto representthe referent with socialization resultingin an individual fearfuland sad expressions is attenuated and this interferes who fails to learn to avoid actions that result in harm to others. Fridlund 1991).National Institute Cognitive ofMental Health. do not act as unconditioned stimuliforaversiveconditioning or instrumentallearning. Program. I will make one fundamental assumption: that facial expressions of emotion do indeed have a communicatory function.the appropriateprocessing of emotional expressions is impaired. imitating and influencingthe actions of others: the mechanisms of social interaction'. Hinde 1985).R. 2670. 1993). the argumenthere is not thatthe display of Importantly. The purpose of this paper is to consider facial expressions: the stimuli that elicit their presentation. psychopath. Happy expressions. Indeed. In psychopathy. According to these authors. the expressions the neuro-cognitivesystemsthat interpret produced by others and the conditions under which the interpreter may respond to the emoter thus closing the communicatoryloop. theircommunicatory functions and neuro-cognitive substrates R. Blair Mood and AnxietyDisorders Unit on Affective Neuroscience. Jones et al.the transmissionof inforimpartinformation mation is not theirfunction. fearfulfaces have been seen as aversive to unconditioned stimulithat rapidlyconvey information othersthat a novel stimulusis aversiveand should be avoided (Mineka & Cook 1993). In these conditions. the sadness suggestionwill be that expressions of fearfulness. Similarly. Disgusted but are used most freexpressions are also reinforcers One contribution of 15 to a Theme Issue 'Decoding.R. However. communication. This paper will review the literatureon the neural mechanisms necessary for this communication: both the mechanisms involved in the production of of the emotional expressions of others. quently to provide informationabout foods (Rozin et al.particularly in situations involving hierarchy interactions (Blair & Cipolotti 2000. it is argued that the basic response to emotional expressions remains intact but that there is impaired of the individual displayingthe emotion. Soc. emotional expressions and those involved in the interpretation disorders of autism.nimh.it is argued.nih. an emotional expressionimplies intentto convey a specific message to the observer. participantssmile more at a humourous video or show greaterdistressto the sound of an individual in distressif they are togetherwith another rather than iftheyare alone (Chovil 1991. the response to angryexpressions in particularis attenuated resultingin reduced regulation of social behaviour. Health Human Room MSC and 15K National Institute NorthDrive.

this strict dichotomy overstates the empirical picture. investigationsof patients with Parkinson's disease and otherpatients with damage to the basal ganglia report marked reductions in the production of spontaneous emotional expressions. such patients also show some impairment in the production of posed emotional displays. 1996). Weddell 1994. of valencesfornovel objectscan be Indeed. But emotional expressions are not automatically elicited under these conditions. The childwillthenform the direction theappropriate association between theinformation comand the objectto municated by thecaregiver's expression had been attending which the caregiver (Moses et al. Basal ganglia and frontalcortex are representedas reciprocally interconnected such that damage to either 2. and associatethe new wordwiththisnovel attending durobject(Baldwinetal.in particular basal ganglia. willeither A particularly clear illustration of the communicatory can be seen afteran of emotionalexpressions function of a novel object. A simpleconditioning to is. 1990.3 on Sat. 1990. Pairing theinfant theCS to elicita conditioned response.This maternalfearful the US. other studies find significantimpairment in the production of both posed and spontaneous emotional expressions in patients with frontal cortex lesions (Weddell et al. responds expression. 2001). if to the child is social attending one referencing. of the US with the the novel object. 1990. Soc. however. and is object.calculatethe object that theyare to. monkey.theymay follow 'display rules'. However. such patients show reduced displays of emotional expressions when watching emotionally arousing videos relative to comparison individuals (Borod et al.562 communication and neuralsubstrates R. Similarly. A schematic of regions known to be involved is presentedin figure 1. Smith et al.If the caregiver displays the child will approachthe novel a happy expression.If the determined by the caregiver's of fearor disgust. serve a communicatory function emotionalexpressions have evolvedso thatinformation on thevalence thatthey can be transmitted between of objects/situations rapidly foran emotional Thus. one major task faced by the child in middle childhood is to learn the culture's display rules governingthe conditionsthat are appropriate for the display of specific emotions. Weddell 1994). 13 Jul 2013 02:05:56 AM All use subject to JSTOR Terms and Conditions . 1990. 1996).139. Thus. display not occuror be considerably muted. conditions when thereis an emotionalstimulusin the affectthe production of spontaneous expressions (Borod and thereis an audience to perceivethe et al. the communication of names to objects. 1997) and a verysimilar processhas and labelledobservational been shownin othermonkeys fear(Mineka& Cook 1993). age-related changes were demonstratedin the abilityof childrento cover theirdisappointment at the discovery that their giftfor helping out an than the gifttheyhad been adult was much less interesting expecting. controlled expressions (Weddell et al. Inter(Klinnert socialreferencing is seenin chimpancomparable estingly. It has been argued that the neuropsychological data about the production of emotional expressions echo this dichotomy (Rinn 1984. However. the processof observational fear Mineka characterizes framework withinan aversiveconditioning (Mineka & Cook 1993). EXPRESSIONS The greateroutput fromthe frontalcortex representsthe The suggestiondeveloped above is that emotional fact that while frontal cortical lesions cause significant are communicatory to expressions signalsthatfunction impairment to both the production of spontaneous and convey valence information rapidly to conspecifics.Whenhearing associatethisword withwhatever novel object matically is in theirimmediatefield of view. monkey comes to showa fearful reaction to thenovelobject. Similarly. Smith et al. whereas cortical regionsare necessaryforcontrolled emotional displays but not automatic emotional displays (Rinn 1984). Lond. involves like the communication oftheaffective with a CS thatcorassociation information to the communicator's referent.This processis knownas social referencing seen in childrenfromthe age of eightto ten months etal. thenovelobject.allows CS.In humans. Weddell 1994. zees (Russellet al. lesions to the basal ganglia disproportionately they likely Specifically. of valence to objects. J. response. THE PRODUCTION OF EMOTIONAL structure impairsthe production of emotional expressions. In a classic study of the development of display rules and control over emotional expressions.there have been reportsthat lesions of frontalcortex impairthe ability of the patient to pose emotional expressions but spare the production of spontaneous emotional expressions (Hopf et al. 1983. 1988. R. Hopf et al. The data therefore suggest that sub-cortical regions. 1988. 1990. important triggers conspecifics.the displaysan expression caregiver childwillavoid the novelobject. Blair Facial expressions. Walker-Andrews 1998). in theinfant a fearful ditioned reaction. Instead. of theirattention. and cortical regions. There is thus a suggestion of spontaneous or overlearned emotional expressions to emotional stimuliin the presence of observers as well as controlled or posed emotional expressions as a functionof display rules.children do not autoobjects. 1992). particularly frontal cortex. Indeed. societal proscriptions as to what emotion should be displayed in given circumstances and how intensely it should be displayed (Ekman & Friesen 1969). are involved in both the production of spontaneous and controlled emotional displays. Pitcairn et al.though to a lesser degree (Borod et al. 1990. Wedareparticularly to be elicited under dell 1994). Thus. it has been claimed that sub-cortical regions are necessary for spontaneous emotional displays but not controlled ones. approach unlikely in humans.the disappointmentofthe youngerchildrenwas far easier to detect (Saarni 1984).235. B (2003) This content downloaded from 14. Thus.theyturn towardsthe speaker. event and also a potenincludebothan emotional display If there is no observer. Bloom 2002). Weddell 1994. therepresentation be appropriate of the emoter'sintenthas been shown to be crucial. 1987. 1992).The infant will look infant's discovery willbe theprimary and their behaviour towards caregiver emotionaldisplay. ing when the an emotional caregiver response object displays thechildwilllookat thecaregiver to determine to another. the learning to the learning of names fornovel of thought similarly a newword. Trans. elicitsan unconexpression. Individuals are capable of intentionally manipulating their emotional displays. Frontal environment Phil. Smith et al. theemotional tialobserver. 1996. The US is themother macaque's expression offear. However. R. whichshe showsto the CS. 1996).

a sub-cortical has yet to find pathway(retinocollicular-pulvinar-amygdalar) 2000. task conditions that pathway has been provided by work with G.bilateral regions in the lateral fusiform glia receives inputs from both the amygdala and other gyrus and the posterior superior temporal sulcus structures processing emotional information. at ca. Figure1. 2002). many imaging studies the neural response to emotional expressions changes in visual input and hence produce quite distinct investigating to the facial and neutral mask. The suggestionis that the fusiform gyrusis more involved in the processing of facial identitywhereas the superior temporal sulcus is more involved in the processing of 3. Morris et al.Later neuro-imagingwork with G. between the emotions of emotional expressions to the amygdala that can then fear and happiness afteronly 120 ms in the orbital frontal cortex of a patient (Kawasaki et al.Y. and neuralsubstratesR. R. 1999). Thus. 2001). 2000. as single unit responses. 160 ms. duringauditoryfearconditioningcan be mediated by proHalgren et al. This would sugmodulate the processing of information through the corticalpathway (Pizzagalli et al. Halgren et al. Indeed. A schematicof regionsknownto be involvedin of emotionalexpressions. 2001). information on conditioned stimuli expressions (Pizzagalli et al.However. This 'blindsight' patient showed some abilityto discriminate (by guessing) between different facial expressions in his blind hemifield. a patient with a long-standing increase attention to emotional expressions result in right-sided hemianopia afteroccipital frontal cortex Phil. This literature or by a corticalpathway (retinogeniculostriate-extrastriate-evidence of early amygdala activitythat the sub-cortical route should predict. 2000). 2001). 1999. Neurons in the superior by feedback from structures such as the amygdala colliculus are capable of responding to much more rapid (Adolphs 2002). The suggestion is that the sub-cortical pathway is fast However. Trans. Campeau & Davis 1995). 2002). Although (Kanwisher et al. 1999). From there have been suggestions that information on the ca. 1999). amygdala basal ganglia for processing systems expressions responses to fear conditioned faces exhibit conditionstimuli emotional specific covariations with neural activityin the posterior thalamus and superior colliculus (Morris et al. 2002). Haxby et al. R. 1999. 1999. Pizzagalli et al. Adolphs 2002). The basal ganeral occipital gyri. 50 ms after information sufficientto dis1999). 2000. Soc. which occurs withinthe relatively al. in the vinar. EXPRESSIONS OF OTHERS Recent event-related potential and magnetoencephalogTwo dissociable routes have been shown to be involved studies have allowed considerable specification of raphy in processing fear conditioning (Armony et al..b. the earliest activityseen is fusiform) (de Gelder et al. 2000).Facial expressions. tomographyimaging) have been demonstratedin the pul1999). such responses fail to elicit conscious experience. Adolphs 2002). perhaps as a consequence of the amygdala activity. In addition. demonstrated differential amygdala responses to fearful versus happy expressions when these were presented to both the blind and seeing hemifields. Lond. 1998. 1984.thereare strongsuggestionsof a dissociation in funcamygdala display spontaneous to et al. RESPONDING TO THE EMOTIONAL social communication (Haxby et al. the time-course for the processing of emotional LeDoux 2000). This again suggeststhe possibilitythat response slow response time of primary visual cortex neurons to emotional stimuli in the temporal cortex is modulated (Macknik & Livingstone 1998). activityis seen in the fusiformgyrus and emotional expressions of otherscan be conveyed eitherby superior temporal sulcus (Streit et al. striateand fusiformactivity only occurred in response to stimuli andother motor amygdala programmes presented to the seeing hemifield. 1999. 2000. Analogously. The earliest activitythat discriminates between emotional facial expressions is seen in midjections to the amygdala fromeitherthe auditorythalamus line occipital cortex from between 80 to 110 ms postor auditory cortex (LeDoux et al. 1997. 2002. 1999. Moreof lesions do the reduce over.235. emotional displays (Anderson & Phelps 2000). they displays do not affectthe production of controlled fearful or other sulcus (Hasselmo et al. neuro-imaging studies have indicated that cortex is likelyto be crucial forrepresenting threespecificareas are involved in face processing: the latgoals to either show or suppress an emotional expression. Critchley et al.positive covariationsof cerebral blood flow (as measured by positron emission poral cortex after the amygdala activation (Streit et al. Additional support for the suggestion of a sub-cortical neutral expressions (Phillips et al. Streitet al. 1997. Hoffman & Haxby 2000). Moreover. the production The cortical route involves regions of occipital and posterior temporal visual cortex (Haxby et al.3 on Sat. Iidaka et al. 1999. there has been a report of neuronal discrimiand allows immediate automatic access of information on nation. occurred ca. 220 ms in the rightamygdala (Streit et al.In addition. stimulus (Pizzagalli etal. J. 1989. Visual masking is assumed to be a result of interference between the induction of neural activityby the tinguishfaces fromother objects was available (Sugase et stimulus and the mask. 1999).139. Howhave responses expression reported greatersuperior temporal sulcus and fusiform gyrus activityto emotional expressions relative to ever. Blair 563 communication lobe damage at the age of 8 years (de Gelder et al. In particular. Romanski & LeDoux 1992a. In gest a sub-cortical route to orbital frontalcortex.Y. Halgren et al. 1999. 13 Jul 2013 02:05:56 AM All use subject to JSTOR Terms and Conditions . superior colliculus and amygdala in response to Indeed. fearful novel tion between the fusiformgyrus and superior temporal objects (Prather 2001). There appear to be further activations of superior temsupport of a sub-corticalpathway. a recentstudyexaminingsingle unit activity masked facial expressions of anger that had been pretemporal visual cortex in monkeysfound that information sufficientto distinguish different emotional expressions viously associated with an aversivestimulus (Morris et al. B (2003) This content downloaded from 14.

2001. Breiter et al.given the suggested role of fearful. Activity parietal cortex.564 R. 1996. a recent review of patient performance across studies. B (2003) 4. they allow the rapid transmissionof taste aversions. 2000). NEUROTRANSMITTERINVOLVEMENTIN RESPONDING TO THE EXPRESSIONS OF OTHERS There is a growingbody of data indicatinga degree of differential neurotransmitter involvement in systems responsible for the processing of emotional expressions. Blair et al. Schmolck & Squire 2001). 1999. 2001b. 1996. Thus. Streit et al. with a few exceptions (Kesler-West et al. In particular. it should be noted. serotonergicmanipulations have been found to This content downloaded from 14. 1999. Soc. 1997.to my knowledge. 2001). 1996. 1999. Everitt et al. As stated in the beginningof this paper. 2000. Crucially. Sprengelmeyer et al. are activatedby angryexpressionsand response reversalas a function of contingencychange (Sprengelmeyeret al. pharmacological interventions can alter the communicatory salience of emotional expressions. two studies investigatingwhich cortical regions. 1999. 1998) and patientswithdamage to the insula present with selective impairmentforthe recognitionof disgusted expressions (Sprengelmeyer et al. Blair et al. similar areas of lateral orbital frontalcortex Interestingly.or at least the proximal region of superior temporal sulcus. associaInstead. 1999. modulate amygdala activity(Schneider et al. Cools et al. 1999. it is far less clear that the angry expression is a basic reinforcer. Trans. They appear to serve to informthe observerto stop the currentbehavioural action ratherthan to convey any information as to whetherthat action should be initiatedin the future. R. Kesler-West et al. 2002). 1997. 1996. amygdala lesions have been consistently ted with impairment in the recognition of fearful expressions (Adolphs et al. Amygdala lesions rarelyresult in impairmentin the recognition of happy expressions (Adolphs et al. that did observe amygdala activationby angryexpressionsfound veryweak activationthat was significantly less than that seen to fearfulexpressions et al. 1994. Nakamura et al.most neuro-imaging dala activationto angryexpressions (Sprengelmeyeret al. 2001) although. Morris et al. Small et al. In studies do not observeamygaddition. 1998. 1998. 1993. Two additional cortical areas that have been linked to the processing of emotional expressions are bilateral frontalcortex and inferior parietal corregions of inferior tex. 1998. this may reflect the ease with which happy expressions are recognized (Ekman & Friesen 1976).235. the suggestionis that the disgusted expressions of others activate in particular the insula allowing taste aversion (disgust expression USnovel food CS associations) to occur. a fundamental assumption of this paper is that emotional expressionsare communicatorysignals that serve specific purposes. Functional imaging studies have consistentlyshown that disgusted expressions engage the insula and putamen (Phillips et al. These areas are of potentialinterest mal areas are activatedwhen eitheran individual is initiating a movement or when they are observing another initiate the same movement (Iacoboni et al. Importantly. but are used Disgusted expressions are also reinforcers most frequently to provide information about foods (Rozin et al. J. Moreover. Schmolck & Squire 2001). KeslerWest et al.In other words. reportedthat ca. 1999). Impairment in the processing of sad expressions is not uncommonly found in patients with amygdala lesions (Adolphs et al. have sad and happy expressionsall generallyfound that fearful. Blair & Cipolotti 2000). thatneuro-imagingstudhappy expressionsas reinforcers. Blair et al. Gorno-Tempini et al. 2000). 1997. 1999. Kesler-West et al. (Whalen 2001). ies. Vuilleumier et al. It is thus sad and unsurprising. 2001a. 2002). as proxi1996. when damaged. 13 Jul 2013 02:05:56 AM All use subject to JSTOR Terms and Conditions . sad and happy expressions can all be fearful. Baird et al. 50% of patientswith Phil. Morris et al. 2000) and expression processing (Phillips et al. 1999). increased superior temporal sulcus and fusiformgyrus activity(Narumoto et al. most effected expression recognition stressed the importance of the inferiorparietal cortex (Adolphs et al. Pizzagalli et al. is frequently implicated in the processing of face stimuli (Haxby et al. Halgren et al. Drevets et al. The only study. insula lesions have been found to block the acquisition and expression of taste aversion learning(Cubero et al. 1997. 1999. R. 1993). 2001). 1994. many other studies in the inferior have not. Phillips et al. Thus. forresponsereverangryexpressionscan be seen as triggers sal (Blair et al.3 on Sat. 2000. In contrastto the expressionsconsidered above. 1999. Three neuro-imaging studies have observed inferior to emotional expressions (George et frontalcortex activity al. 1996. However.The amygdala has been implicated in aversive and appetitive conditioningincluding instrumentallearning (Killcross et al. There have been occasional suggestions that amygdala damage leads to general expression recognition impairment but these reports are typicallyfrom patients whose lesions extend considerablybeyond the amygdala (Rapcsak et al. Fine & Blair 2000). LeDoux 2000). 2001. The claim is that this perspectiveallows an understandinginto specific patterns of activation seen for specific emotions. though it should be noted that happy expressions have been reported to both increase and decrease amygdala activity (Breiter et al. communication and neuralsubstrates amygdala damage presentwith impairmentfor the recognition of sad expressions (Fine & Blair 2000). Calder et al. 1996). 1999. Orbitalfrontal cortexis cruciallyimplicatedin response reversal(Dias et al. Angryexpressions are known to curtail the behaviour of othersin situationswhere social rules or expectationshave been violated (Averill 1982). Pessoa et al. Calder et al. 2001. For example. Indeed. that modulate the probabilitythat a viewed as reinforcers particularbehaviour will be performedin the future.139. Lond. 2000). 2000). 2002). 1998. the observeris warned not to approach the food that the emoter is displayingthe disgust reaction to. O'Doherty et al. Blair Facial expressions. 2001). The neuropsychologicalliteraturesupports the neuro-imaging literatureabout the importance of the amygdala in the processing of fearful expressions. 1996. Experimental investigations in macaques have shown that there is a primarytaste cortical region in the anteriorinsula (Rolls 1997) and neuro-imagingstudies in humans have also shown the insula to be involved in the representationof taste (O'Doherty et al. 2001). This has prompted suggestionsthatrespondingto anotherindividual's expression relies on the activation of motor programmes that the individual uses for the production of expressions (Preston & de Waal 2003).

Bremner et al.to my knowledge. Leary et al.3 on Sat. a smile control. In addition. 2003). 1987. although the central nucleus of the amygdala which projects to autonomic centres in the brain stem is densely innervatedby GABA neurons. Reference was made to a crucial determinantof whetheran expression will be elicited: the presence of others (Jones & Raag 1989. For example. Trans. contains only scattered GABA neurons (Swanson & Petrovich 1998). Lond. Lewis et al.communication and neuralsubstratesR. 1992) and the impact of noradrenergic manipulations of the amygdala's role in the augmentation of episodic memory is well known (Cahill & McGaugh 1998. J. which only involves the zygomatic major muscle action that pulls the corners of the lips upwards. a non-Duchenne smile. as a functionof its interconnections with cortical regions.sad and happy expressions as unconditioned stimuli for aversive and appetitive conditioning and instrumental learning. 1991). 1991). B (2003) 6. R. speech disturbances. ACKNOWLEDGINGOTHER INDIVIDUALS' EXPRESSIONS: CLOSING THE COMMUNICATORYLOOP In this paper the communicatoryfunctionof emotional expressions has been stressed. Phil. More recent work has demonstrated that embarrassment display unfolds in the followingreliable sequence. while diazepam abolished the increase in lateral orbital frontal cortex activityas a function of increased angry the increase in amygdala activityas expression intensity. Keltner & Anderson 2000). head movements down. (1996) presented evidence that people are actually motivatedto convey embarrassmentto others as a way of repairingtheir social image.235. Keltner & Buswell 1997) have suggested that embarrassment serves an important social function by signalling appeasement to others. The basic idea is that embarrassment serves to aid the restoration of relationshipsfollowingsocial transgressions(Keltner & Buswell 1997). which is a lower facial action that potentiallyinhibitsthe smile. in the social referencing example provided above.This mightsuggestthatindividuals should stop displaying emotional expressions when the audience has demonstratedthat they have registeredthe display of the emoter. 5. which occurred ca. This studythus adds support to the suggestionthat GABAergic manipulations impact the role of orbital frontal cortex in modulating the response to interpersonalsignals of conflictsuch as anger. Fridlund 1991. is more involved in unafexpressionsand thus relatively respondingto fearful fected by GABAergic manipulations. Certainly. DEVELOPMENTAL PSYCHOPATHY AND ACQUIRED SOCIOPATHY If emotional expressions serve a communicatory function. Individuals typically display expressionswhen thereis an audience to witness these expressions. At presentonly one study. as I have been arguing. Leary & Meadows (1991). Given these differential effectsone might predict that the serotonergic and affecting noradrenergic manipulations are differentially the amygdala's role in responding to fearful. the basolateral nucleus of the amygdala. for example. Semin & Manstead (1982) found that people reacted more positivelyto others aftera social transgressionif the were visiblyembarrassed. Keltner & Buswell 1997. This involves gaze aversion. a second smile control. affectthe processing of fearfuland happy differentially expressions (Harmer et al. projecting to cortical regions. was not a functionof increased fearful expressionintensity affected by diazepam. Gilbert 1997. Three ways in which develwill be discussed below with referopment can be affected ence to the neuro-psychiatric conditions of autism. Embarrassmentis associated with gaze aversion. face touches. PATHOLOGICAL EXPRESSION PROCESSING: THE CASES OF AUTISM. Zangara et al. Interestingly. Miller 1996. noradrenergicmanipuaffect the processing of sad lations to differentially expressions (Harmer et al. R. I although this would intuitively know of no empirical literaturedemonstratingit to be so.139. There are high concentrationsof benzodiazepine receptor sites in both amygdala and the frontalcortex (Dennis et al. eye positions. 1988. 25% of the time (Keltner 1995). There is a good deal of empirical evidence to support this 'appeasement' or remedial functionof embarrassment from studies of both humans and non-human primates (Leary & Meadows 1991. 13 Jul 2013 02:05:56 AM All use subject to JSTOR Terms and Conditions . Thus.has examof these pharined the neural underpinningsof the effects macological agents (Blair et al. It is plausible that the basolateral nucleus. Gilbert 1997. embarrassment displays may be initiated by an individual following an emoter's display of anger: ifthe individual's behaviour was unintentional or the angryobserver is of high status. Jones et al. developmental psychopathyand acquired sociopathy. 2002). Blair & processing angryexpressions(Borrill Curran 1999. This investigated the impact of diazepam on the neural response to morphed angry and fearful expressions. slouched posture (Goffman 1967. Cahill 2000). appear to be the case. Asendorpf 1990. Chovil 1991. However.When a person's untoward behaviour threatens his/her standing in an important social group. 2001 a).we mightexpect that atypical responding to the emotional expressions of others would adverselyaffectdevelopment. 2001b) whereas dopaminergic affect the and GABAergic manipulations differentially et of al.it is known that there is considerable serotonergicand noradrenergicinnervationof the amygdala (Amaral et al. whereas GABAergic manipulations impact the role of orbital frontalcortex in modulating the response to interpersonal signals of conflictsuch as anger. Leary argues that embarrassmentdisplays diffusenegative social evaluations and the likelihood of retaliation. Blair 565 Facial expressions. Soc. the caregivershould stop to displayfearwhen the infantdemonstratesthatthey will now not approach the aversivenovel object. shifting a nervous smile and a rigid. (1996) and others (Keltner 1995. 2000). One particular case where there are clear indications that the audience demonstratesthat they have registered the display of the emoter is seen during embarrassment displays. Autism is a severe developmental disorder described by the American Psychiatric Association's diagnostic and statisticalmanual (DSM-IV) as 'the presence of markedly abnormal or impaired development in social interaction and communication and a markedlyrestrictedrepertoire of activities and interests' (American PsychiatricAssoci- This content downloaded from 14. Leary transgressors et al. visible signs of embarrassmentfunction as a nonverbal acknowledgementof shared social standards. and then face touching. However. In other words.

139. wrote 'these children have come into the world with an innate inabilityto formthe usual. children with autism have usually been found to be unimpaired in facial affectrecognition (Ozonoff et al. Frith forthe 1989). Bormann-Kischkelet al.566 and neuralsubstrates R. All four of these studies have reportedreduced orientationto the caregiverby the childrenwithautism althoughthiswas only in the lower abilitysample in the Bacon et al. A series of studies has examined the behavioural reactions of individualswith autism when the child has been playing with the experPhil. However.3 on Sat. 1997b.usually distress(Sigman et al. 1998. J. Howard et al. 1990. Dissanayake et al. Psychopathy. Thus. It impulsivity is identifiedin childrenwith the antisocial process screening device (Frick & Hare 2001) and in adults with the revised psychopathychecklist (Hare 1991). who originallydescribed the disKanner. Adolphs et al. 1990. the idea has for people with autism is existed that the main difficulty an inabilityto enter into emotional relationships. Social referencing. Soc. the learning of emotional valence fornovel objects.235. 2000) witha recentclaim suggestingthatthisis specific forfearful expressions (Howard et al.Many have reportedthat childrenwithautismhave difficulty recognizingthe emotional expressions of others (Hobson 1986. More recently.These criteriamust be evident before 3 years of age. the individual does not learn to avoid committingbehaviours that cause harm to others and will commit them if. to one object when the caregiver displays an emotional response to another. in turn. The above argument generates further predictions about emotion in autism. However. 1997a). B (2003) This content downloaded from 14.is shown by less than 1% of individuals in mainstreameducation (Blair & Coles 2000) and less than 30% of individuals incarcerated in forensicinstitutions(Hart & Hare 1996). will attempt to represent the intended cue that elicited the emoter's expression. without the emoter's referent representing theymay associate valence to novel objects inappropriately or not at all. theirthoughts. When they are. 2001). autism is interesting well-documented impairmentin theoryof mind shown by patients with this disorder (Frith 2001). 1997b. One account of psychopathyhas linked the disorderto early amygdala dysfunctionand consequent impairment in processing fearfuland sad expressions (Blair 1995. Impairmentin theoryof mind is interesting communicatory role of emotional expressions. A child with autism presented with another individual in distress will show aversive autonomic arousal to the other's distress (Blair 1999) and. should be impaired in childrenwith autism. a healthyindividual. Baron-Cohen et al. 1995). 1996. this does not reflect ness to other individuals' emotion. That is.If it is. 2001. Blair Facial expressions. Trans. 1995. when witnessingthe emotional display of another individual. Thus. biologicontact with other people. and poor behavioural control (Hare 1991). just as cally provided affective other children come into the world with innate physical or intellectual handicaps' (Kanner 1943. we mightpredict anomalous behavioural reactionsto the emotional displays of other individuals in children with autism given their theory-of-mind impairment. 2000). Bacon et al.and that this deficit in social interaction leads to their profound difficulties (Hobson 1993). p. In addition. this disorder is not equivalent to the psychiatricdiagnoses of conduct disorder or antisocial personality disorder (American PsychiatricAssociation 1994). 250). he receives reward (Blair 1995). The child with autism should fail to use the emoter's gaze directionto calculate the correct object to associate the valence elicitedby the emoter's display in the same way that theyfailto use a speaker's gaze directionduring novel word use to calculate the speaker's referent(Baron-Cohen et al. duringsocial referencing. intentionsand knowledge (Premack & Woodruff 1978. Corona et al. Leslie 1987. Because of this lack of specification. I would therefore argue that autism does not represent a disorderwhere thereis atypicalrecognitionof emotional because of the expressions. These psychiatric diagnoses are relatively poorly specified and concenon the antisocial behaviour shown by tratealmost entirely the individual ratherthan any formof functionalimpairment. Baron-Cohen et al. the above only applies to studies where the groups have not been matched on mental age. R. Many studies have investigatedthe abilityof individuals with autism to recognize the emotional expressions of others. As long as autism has been recognized.the child will look at the caregiverto determine the direction of their attention (Moses et al. Because of these high rates of diagnosis. 13 Jul 2013 02:05:56 AM All use subject to JSTOR Terms and Conditions .e. However. popuwiththese diagnostictools are highly lations identified heterogeneous and also include many individuals with other disorders. Adolphs et al. Theory of mind refers to the abilityto representthe mental states of others. So. as has been argued above. communication imenter and the experimenterhas feigned an emotional reaction. 1990. The main criteriafor the diagnosis in DSM-IV can be summarized as qualitative impairmentin social communication and restrictedand repetitivepatterns of behaviour and interests. a diminished capacity forremorse. R. The basic suggestion is that psychopathicindividualsrepresentthe developmentalcase where sad and fearful expressions are not aversiveunconditioned stimuli. 2001). childrenwith autism present with no impairment in expression recognition (Ozonoff et al. (1998) a lack of responsivestudy. Theory of mind should be involved in the representation of the emoter's intention.desires. i. several studies have found the emotion processing impairmentto be pronounced only when the emotion is a complex 'cognitive' emotion such as surpriseor embarrassment (Capps et al. beliefs. psychopathicindividuals have been found to presentwith reduced amygdaloid volume relative to comparison indi- ation 1994. for ifthe child is attending example. Bormann-Kischkel et al. Indeed. This. p. this is exactly what is seen in children with autism.In particular. 1999). As a consequence of this. 1992.rates of diagnosis of conduct disorder reach up to 16% of boys in mainstream education (American PsychiatricAssociation 1994) and rates of diagnosis of antisocial personalitydisorder are over 80% in forensicinstitutions(Hart & Hare 1996). 2001). Prior et al. Blair et al. 66).we should see a reductionin the usual orientationresponse to the emoter to calculate the elicitingstimulus. predicts that children with autism may present with very unusual emotional reactions to objects. the psychiatrist order in 1943. Baron-Cohen et al. 1992. by doing them. Prior et al. 1993. 1990. Lond. is a disorder characterized Psychopathy developmental in part by callousness. In line with this theory. Importantly.in contrast. 1998).it has been suggested that autism is due to an innate impairmentin the abilityto perceive and respond to the affective expressions of others.

patients with acquired sociopathy followinglesions of the orbital frontalcortex presentwithgenerallyimpairedexpressionrecognitionbut this impairment is particularly marked for angry expressions (Hornak et al.sadness preferentially and happiness). Importantly.139.This leads to a failure in socialization. patients with acquired sociopathy present with frustrationor threat-inducedreactive aggressionwhether theiracquired lesion of the orbitalfrontalcortex occurs in childhood (Pennington & Bennetto 1993. 1997). the processing of other individuals' sadness and fear is particularlyaffected. Phil. The consequences of impairment in being able to adequately process the emotional displays of others can be severe. 1996. Although there have been suggestions that developmental psychopathy and acquired sociopathy might be formsof the same disorder (Damasio 1994). B (2003) The strong suggestion is thereforethat this impairment underlies their socially inappropriatebehaviour. Althoughsystemsgenerallyinvolved in processing facial stimuli. Finally.it does not decrease the probabilityof antisocial behaviour in childrenwho presentwith the emotional dysfunctionof psychopathy (Wootton et al. and when it is not. in particular. the emoter's emotional displays are a function of the presence of observers and the observer will attemptto determine the referentof the emoter's display. it is highlylikelythat they fail to adequately process the emoter's referent and that they therefore process the disbecause of their impairmentin theoryof play incorrectly mind. although it has been repeatedly shown that the use of empathyinducing positive parentingstrategiesby caregivers decreases the probability of antisocial behaviour in healthydeveloping children.fulfil teria for 'sociopathic disorder' (American Psychiatric Association 1980). Indeed. 1996.235. Blair 567 viduals (Tiihonen et al. 1997) and. Blair & Cipolotti 2000). 1999) or adulthood (Grafman et al. 13 Jul 2013 02:05:56 AM All use subject to JSTOR Terms and Conditions . 1996). their display can be intentionally manipulated. Presumably. reflecting with theirabilityto be socialized (Cornell et al. counterAcquired sociopathy representsan interesting point to developmental psychopathy. 2001) and aversive conditioning tasks (Veit et al. particularly to these ability recognize expressions (Blair et al. If we assume that emotional expressions serve a communicatory function. a region involved in taste aversion. we must predict that they will be more likelyto be displayed when a potential observer is present. The psychopathic individual does not learn to avoid actions that cause harm to others. appropriate to display emotional expressions. this different now appears unlikely(Blair 2001). The orbital frontal cortex is implicated in the response to angryexpressions (Sprengelmeyeret al. In addition.Blumer & Benson (1975) had used the term 'pseudo-psychopathy' to refer to patients with frontallobe lesions presentingin this manner. In psychopathicindividuals. In contrast. social rules that stipulate when it is. Lond. communication and neuralsubstratesR. Crucially. Children learn display rules. the communicatoryfunctionof emotional expressions is reflected in the partlydissociable neural systemsthat are additionally involved in processing emotional expressions.This is indeed the case. the emoter's intent modulates the frontal lobe-basal ganglia circuitry thathas been implicated in the production of emotional expressions. followingacquired lesions of the orbitofrontal the DSM-III diagnostic cricortex. Blair & Cipolotti 2000). This remains to be investigated. As such they engage the insula. I have argued forthe communicatoryrole of angryand embarrassment expressions in regulating social hierarchical interactions. Moreover. Assuming the observer will have been accomplishes this. the argument is not that the display of expressions implies that the emoter intended to convey a specific message to the observer. during an emotional memory task (Kiehl et al. (1990) to characterizeindividualswho. 2002). In contrast. CONCLUSIONS In this paper.3 on Sat. Soc. relative to comparison individuals. 2001). the role of angry behavioural action and expressionsin stoppingthe current the role of embarrassmentdisplays in communicating a lack of intent to commit the action that has resulted in that an indisocial disapproval. Psychopathic individuals present with pronounced levels of goal-directed instrumentalaggression and antian impairmentthat interferes social behaviour. Anderson et al. R. Although emotional expressions are not intentional communications. 2000) and reduced amygdala activation.the augmentationof startle reflexby visual threatprimes or arousal to the anticipation of punishmentare all impairedin psychopathicindividuals (Blair 2001).Facial expressions. Blair et al. Also in line with the theory. I have stressed the communicatoryfunction of emotional expressions. R.'Acquired sociopathy' was a term introduced by Damasio et al.psychopathic individuals show pronounced impairment in processing sad and fearfulexpressions. Trans. theyare used most frequently mation about foods. such as the occipital cortex. Although disgusted expressions are also to provide inforreinforcers. we must predict that the display of the expression will be terminated when the observer has shown clear indication that they have received the communication. Kesler-West et al. I have argued that although individuals with autism may be able to recognize the expressions of others. Previously. fusiformand the superior temporal sulcus process expressions.individuals withthe developmental disorder of psychopathyand individuals with acquired sociopathy followinglesions of the orbital frontalcortex fail to respond appropriately to specific expressions.Thus we can learn to intentionallymask or alterour expressionsas a functionof these display rules. in childhood. 1998. developmental psychopathyand acquired sociopathy present very differently. expressions that serve as positive or negative reinforcers activate the amygdala (fearfulness. in functions that recruitthe amygdala such as aversiveconditioning and instrumental learning. impaired Blair et al. Thus. 1999. They show reduced autonomic responses to these expressions (Aniskiewicz 1979. J. Interestingly. This content downloaded from 14. then. 2001). We mightexpect therefore vidual whose response to angry/embarrassment should presentwith impaired expressions is dysfunctional modulation of their social behaviour. it is simply that emotional expressions serve a communicatory function. appropriate information transferred fromthe emoter to the observer. Angry expressions initiate response reversal and activate regions of orbital frontal cortex that are involved in the modulation of behavioural responding. 7. Thus. their socialization is markedly impaired.

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