Human Bain Mapping 30:2008 2020 (2009) Neual Coelates of Pesonally Familia Faces: Paents, Patne and Own Faces Magot J. Taylo, 1 * Maie Asalidou, 2 Saah J. Bayless, 3 Dew Mois, 1 Jennife W. Evans, 1,4 and Emmanuel J. Babeau 5 1 Depatment of Diagnostic Imaging and Reseach Institute, Hospital fo Sick Childen, Toonto, Canada 2 Depatment of Psychology, Yok Univesity, Toonto, Canada 3 Depatment of Psychology, Univesity of Wincheste, Wincheste, England 4 Institute fo Medical Sciences, Univesity of Toonto, Toonto, Canada 5 Univesité de Toulouse, CeCo, UPS, CNRS, Toulouse, Fance Abstact: Investigations of the neual coelates of face ecognition have typically used old/new paadigms whee subjects lean to ecognize new faces o identify famous faces. Familia faces, howeve, include one s own face, patne s and paents faces. Using event-elated fmri, we examined the neual coelates of these pesonally familia faces. Ten paticipants wee pesented with photogaphs of own, patne, paents, famous and unfamilia faces and esponded to a distinct taget. Whole bain, two egions of inteest (fusifom gyus and cingulate gyus), and multiple linea egession analyses wee conducted. Compaed with baseline, all familia faces activated the fusifom gyus; own faces also activated occipital egions and the pecuneus; patne faces activated simila aeas, but in addition, the paahippocampal gyus, middle supeio tempoal gyi and middle fontal gyus. Compaed with unfamilia faces, only pesonally familia faces activated the cingulate gyus and the extent of activation vaied with face categoy. Patne faces also activated the insula, amygdala and thalamus. Regions of inteest analyses and lateality indices showed anatomical distinctions of pocessing the pesonally familia faces within the fusifom and cingulate gyi. Famous faces wee ight latealized wheeas pesonally familia faces, paticulaly patne and own faces, elicited bilateal activations. Regession analyses show expeiential pedictos modulated with neual activity elated to own and patne faces. Thus, pesonally familia faces activated the coe visual aeas and extended fontal egions, elated to semantic and peson knowledge and the extent and aeas of activation vaied with face type. Hum Bain Mapp 30:2008 2020, 2009. VC 2008 Wiley-Liss, Inc. Key wods: face pocessing; fmri; patne; paents; own; familia faces INTRODUCTION *Coespondence to: Magot J. Taylo, Diecto of Functional Neuoimaging, Depatment of Diagnostic Imaging, Hospital fo Sick Childen, 555 Univesity Ave., Toonto, Ont. M5G 1X8, Canada. E-mail: magot.taylo@sickkids.ca Received fo publication 11 Mach 2008; Revised 20 May 2008; Accepted 29 June 2008 DOI: 10.1002/hbm.20646 Published online 22 August 2008 in Wiley InteScience (www. intescience.wiley.com). Consideable pogess has been made in ou undestanding of face pocessing ove the last yeas though a vaiety of functional neuoimaging investigations. That faces ae paticula visual stimuli has been poposed fo many yeas [Bodame, 1947; Chacot, 1883; Wilband, 1892], but ecent technological advances have allowed the identification of the neual substates of face pocessing, leading to detailed anatomo-functional models [Allison et al., 2000; Gobbini and Haxby, 2007; Haxby et al., 2000]. VC 2008 Wiley-Liss, Inc.
Pesonally Familia Faces Although most neuoimaging face pocessing studies have used faces unfamilia to the paticipants to peclude effects of familiaity, othes have focused pecisely on the effect of familiaity. These latte use eithe old/new paadigms, in which paticipants have to ecognize faces they have just leaned ( old faces ) among distactos [ new faces ; e.g., Dubois et al., 1999; Ishai and Yago, 2006], o ask paticipants to ecognize famous among unknown faces [e.g., Elfgen et al., 2006; Gobbini et al., 2004; Ishai et al., 2005]. Fewe studies on familia face pocessing have used stimuli with the ecological validity of being pesonally familia. Pesonally familia faces can ange fom casual acquaintances to colleagues, close family, and loves to one s own face, a highly familia face. All pesonally familia faces have associations with an accumulation of expeiences and social inteactions. As they ae pocessed fequently and epeatedly, in a ange of conditions, this extensive expeience is likely eflected in neual pocesses [Balas et al., 2007]. We wished to detemine whethe pesonally familia faces ae pocessed similaly to unfamilia faces and whethe thee ae distinct neual activation pattens acoss the diffeent types of pesonally familia faces. Face Peception and Identification In thei behavioual cognitive model, Buce and Young [1986] descibed the putative stages equied fo face identification, postulating that a face is fist ecognized as known and is then matched to a paticula individual, befoe etieval of multimodal and semantic knowledge about the peson. This model has poven vey useful and has been updated with ecent data [Been et al., 2001; Schweinbege and Buton, 2003]. Face identification in these geneal models is thought to ely on a posteio to anteio vental-tempoal axis, pedominant in the ight hemisphee [De Renzi et al., 1994; Kanwishe et al., 1997; McCathy et al., 1999; Puce et al., 1996; Rossion et al., 2000], with posteio egions pocessing visuospatial aspects of faces and anteio egions pocessing integated modality independent and social aspects of faces. Ealy pocessing of facial featues is associated with activation of the infeio occipital gyi [Haxby et al., 2000]. Infomation is then poposed to follow two main pathways: one though the supeio tempoal sulcus, specialized in changeable aspects of faces such as eye gaze, expession, and lip movement [e.g., Allison et al., 2000]; the othe though the fusifom gyus up to the anteio tempoal pole, a pathway specialized fo face identification and multi-modal integation with voice, name and biogaphical elements [Gono-Tempini et al., 2001; Joubet et al., 2004, 2006; Leveoni et al., 2000; Palle et al., 2000]. The model of Haxby et al. [2000] was ecently evised to include ecognition of pesonally familia faces [Gobbini and Haxby, 2007]; the authos poposed that these faces ecuit a distibuted netwok including aeas of visual pocessing such as the fusifom, infeio occipital gyi and posteio supeio tempoal sulcus as well as aeas associated with both peson knowledge and emotion. They suggest that full ecognition of a peson includes not only ecognizing the visual appeaance and etieving biogaphical infomation, but also etieving pesonal taits, attitude, mental states and intentions, involving aeas such as the anteio paacingulate, the posteio supeio tempoal sulcus and the pecuneus. Lastly, they agued that social inteactions and familiaity ecognition modulate the emotional esponse to familia faces, implicating aeas such as the insula and amygdala. Thus, face ecognition depends on a netwok of bain aeas [Babeau et al., 2008; Ishai, 2008; Leveoni et al., 2000]; we hypothesize that aspects of this netwok ae modulated accoding to the type of face viewed. Identification of Pesonally Familia Faces Most of us ae able to ecognize hundeds of faces; howeve, pesonally familia faces shae extended exposue and inceased peson knowledge attained though social inteactions. Although diffeences in pesonal elevance of these faces ae impotant, thee ae also obvious distinctions among these familia faces. Fo example, paents faces ae the fist to be leaned and ecognized in infancy, theeby having pesonal elevance fo the longest peiod of time. They ae the visual stimuli stoed when visual and memoy systems wee in the ealiest stages of matuation, and paents faces undego steady updating with age. Fo most couples, a patne s face is one of the faces seen most often, evey moning, evening and week-ends. A patne s face also holds emotional valence and peson knowledge attibuted to pesonal closeness. Along the same line, one s own face is seen as a mioed eflection many times a day, often because we need to act on it (e.g., fo bushing teeth, shaving, combing hai) and it may be a cue fo self-knowledge o awaeness, a special kind of peson knowledge. Behavioual studies Behavioal investigations have examined possible hemispheic asymmeties in pocessing pesonally familia faces. Some authos have found a ight-hemisphee advantage fo one s own face [Keenan et al., 1999, 2000], wheeas othes found a left-hemisphee advantage [Bady et al., 2004]. When viewing one s own face was compaed with viewing othe familia faces such as colleagues o famous faces, the evese hemispheic effect was found [Bady et al., 2004; Keenan et al., 1999]. Howeve, some of these studies used mio images of paticipant s own face wheeas othes did not, which may impact latealization of own face pocessing. In split-bain patients, the initial studies by Spey et al. [1979] found that both hemisphees wee able to ecognize one s self-image, wheeas Tuk et al. [2002] found a left hemisphee advantage and Keenan et al. [2003] found a ight hemisphee advantage. As the esults fom these vaious studies ae contadictoy, eseach using contolled potocols is waanted. 2009
Taylo et al. Neuoimaging studies A few ecent PET and fmri studies have investigated possible diffeences in the neual substate undelying pocessing of pesonally familia faces, although none investigated the pocessing of paents faces (except Gobbini et al. [2004] but combined with othe pesonally familia faces). Investigations of mothes viewing photogaphs of thei child found diffeential bain activations in occipito-tempoal aeas and the fusifom gyus as well as obito-fontally, that coelated with mothes mood, eflecting positive emotional aousal elated to thei childen s faces [Nitschke et al., 2004]. In anothe epot, inceased activation was found in mothes viewing thei own child (compaed with thei child s fiends) in the amygdala, the insula and the supeio tempoal sulcus and anteio paacingulate gyus, aeas thought to eflect emotional esponse, attachment, and empathy [Leibenluft et al., 2004]. In studies of viewing one s own face, some epot inceased activity of the left fusifom gyus [Kiche et al., 2001; Sugiua et al., 2000, 2005], while the ight fusifom gyus usually shows inceased activity to othe face stimuli. In contast, Platek et al. [2004] and Devue et al. [2007] identified a ight-sided netwok fo own faces. A late study found a bilateal netwok of bain aeas associated with subject s own faces [Platek et al., 2006], which was thought to be simila to the netwok engaged in selfawaeness and the executive aspects of own face pocessing. Uddin et al. [2005] showed paticipants digital mophs between thei own face and a familia face and found that activation of the ight infeio paietal lobule, infeio fontal gyus and infeio occipital gyus inceased as the stimuli contained moe of one s own face and less of the familia face. Thus, accoding to the above studies, viewing one s own face could tigge a stongly latealized self netwok, but with no consensus as to whethe it is ight o left latealized. In contast to one s paents and own face, one s spouse o patne s face should activate aeas associated with pevious omantic, emotional and social inteactions. Two functional imaging studies examined neual esponses in intensely in love young adults with mean elationship lengths of 0.61 yeas [Fishe et al., 2005] and 2.4 yeas [Batels and Zeki, 2000]. Fishe et al. [2005] found inceased activation in the ight vental tegmental aea and ight posteio-dosal body of the caudate nucleus; some of thei paticipants also showed activity in the ight insula, and ight anteio and posteio cingulate cotex. The latte findings ae in geneal ageement with the study of Batels and Zeki [2000], who found activity bilateally in the medial insula, the anteio cingulate, the caudate nucleus, and the putamen. Kiche et al. [2001] found highe activity in the ight insula fo patne faces. Batel and Zeki [2000] also epoted deactivation in the cingulate gyus, the amygdala as well as pefontal, paietal, and middle tempoal cotex. Oveall, these studies suggest that patne s faces in ealy stages of a elationship activate aeas in the limbic system [Fishe et al., 2005], which include aeas associated with peson knowledge at late stages in a elationship [Batel and Zeki, 2000]. Relationships, howeve, last much longe; thus to fully chaacteize pocessing elated to patnes/spouses, it is essential to investigate the neual coelates of patne faces well past the stage of the initial in love peiod. Although we eadily ecognize famous faces such as politicians o actos, we do not typically have pesonal expeience with them. This is likely eflected in the pocessing of famous faces, which only implicates the infeio occipital and fusifom gyi, the supeio tempoal sulcus bilateally and the amygdala [Ishai et al., 2002]. Howeve, thee is disageement whethe ight o left hemisphee activation to famous faces is stonge [Denkova et al., 2006; Ege et al., 2005; Ishai et al., 2005; Poutois et al., 2005]. Thus, a numbe of functional imaging studies have investigated the neual substates of pesonally familia faces, but little convegence is found among the epoted esults, pehaps due to dissimila potocols. The maked discepancies suggest that thee ae vaiables such as the types of pesonally familia faces, which could affect the esults. Hee we wished to answe the following questions: (i) ae pesonally familia faces pocessed with the same functional mechanisms as unfamilia faces? and (ii) ae thee activation diffeences acoss types of pesonally familia faces? Bain esponses to the pesentation of paents, patne, own and famous faces wee compaed with unknown faces. We expected that pesonally familia faces would poduce wide activation pattens than unfamilia faces, paticulaly including fontal stuctues. Unlike one s own face and paents faces, patne faces wee expected to ecuit aeas associated with emotional contol, such as the insula and the amygdala. METHODS Paticipants We studied 10 paticipants, 4 males, mean age 35.4 yeas (67.7 yeas SD), who met the following citeia: they had lived with thei patne fo at least 2 yeas (aveage elationship length 9.1 yeas (65.01 yeas), they gew up in the same house as both paents and both wee still alive, and they could povide ecent high quality photogaphs of themselves, thei patne and thei paents. Stimuli Mateial Each subject povided his/he own digitized photogaphs of his/he patnes, paents and own face, following a standadized potocol contolling fo neutality (no 2010
Pesonally Familia Faces Figue 1. An example of the stimulus set used in the expeiment fo one subject. The task was to pess a button each time the ghost (the supeimposition of all faces) appeaed. emotion on the face), gaze diection (looking staight at the camea), and light conditions. Subject s own face was evesed ight/left, as when viewing one s face in a mio. All subjects also povided a shot list of famous people, men and women, who they would easily ecognize if they saw a pictue of them, fom which we chose two (a male and a female). We also included photogaphs of two unfamilia subjects (male and female). Thus, the set of eight photogaphs was unique fo each subject. The set was pocessed to emove all infomation unelated to the faces themselves (backgound, paaphenalia, etc.). The photogaphs wee conveted to gayscale. Figue 1 shows an example of a set of photogaphs fo a single subject. A ghost image was pepaed fo each set by supeimposing all photogaphs and adjusting levels of tanspaencies. Stimuli wee epeated such that at least 40 tials of each face type (paent, patne, own, famous, unfamilia, and scambled) wee pesented. The ode was pseudo-andom such that one stimulus type did not epeat without at least fou intevening faces. The faces wee pesented fo 500 ms and ISI was jitteed between 1.7 and 2.0 s. The ghost image occued with a pobability of 8% and was included to maintain subject s attention. Paticipants wee asked to focus on the photogaphs and pess a button to the ae ghost image. Pocedue Paticipants gave infomed witten consent and the Reseach Ethics Boad at the Hospital fo Sick Childen appoved all pocedues. Afte standad MR sceening, paticipants completed the task in the scanne, followed by a shot questionnaie including questions such as How many minutes do you look at youself (in a mio) a day on aveage? and Fo how many yeas have you been with you patne?. All MR imaging was conducted on a 1.5T Signa Twin EXCITE3 scanne (GE Medical Systems, WI; softwae ev.12m4) with a standad quadatue head coil. Foam padding comfotably esticted head motion. A set of high-esolution T1-weighted 3D SPGR images coveing the whole bain was acquied (TE/TR/a 59 ms/4.2 ms/158, 116 slices, voxel size 5 1 3 1 3 1.5 mm 3, 2 NEX, 7 min) as an anatomical efeence pio to the acquisition of functional images. Functional images wee acquied with a standad gadient-ecalled echo-plana imaging sequence (TE/TR/a 5 40 ms/2,000 ms/908, voxel size 5 3.75 3 3.75 3 5mm 3 ) ove 27 contiguous axial slices with inteleaved acquisition. Face stimuli wee displayed on MR compatible goggles (CinemaVision, Resonance Technology, CA). Subjects esponded to ghost tials using an MR compatible keypad (Lumitouch, Photonics Contol, CA). Stimuli wee contolled and esponses ecoded using the softwae Pesentation (Neuobehavioal Systems, CA) on a pesonal compute. A TTL tigge pulse fom the scanne ensued stimulus synchonization with image acquisition. Data pocessing and analyses Data analyses wee caied out in AFNI [Cox, 1996], using motion coection, 8-mm spatial blu (FWHM), signal intensity nomalization fo pecent signal change and deconvolution using a fixed haemodynamic esponse function, fo all subjects. Images wee spatially nomalized to the MNI N27 bain in Talaiach steeotaxic space and esampled to 3-mm cubic voxels. Goup images wee analyzed using a andom effects analysis of vaiance. The esults wee thesholded at P < 0.01 [coected fo cluste size; Xiong et al., 1995] fo the whole bain and fo the egions of inteest (ROIs) in fusifom and cingulate gyi. These ROIs wee chosen as they wee the only two egions activated by all pesonally familia faces in the whole-bain esults. The fusifom gyus was active in contasts with the baseline, and the cingulate gyus activated in contasts with unfamilia faces. The selection of ROI masks was anatomically based on the Talaiach stuctual template defined in AFNI [MNI N27 bain in TLRC space; Eickhoff et al., 2007]. The pobability map was thesholded at P 5 0.01 (t 5 3.25) fo familia faces vesus baseline and P 5 0.01 (t 5 2.88) fo pesonally familia faces vesus unknown faces. The ROI masks wee then applied to the thesholded data. To contol fo multiple compaisons in the whole bain, fusifom and cingulate ROI analyses, we pefomed 1,000 Monte Calo iteations at an uncoected P-value of 0.01 on the 43,349 voxels, 596 voxels, and 1,878 voxels in the espective masks. This yielded a minimum volume of 5.70 cc (210 voxels), 0.49 cc (18 voxels), and 1.6 cc (59 voxels) at a P-value of 0.05 fo the whole bain, the fusifom gyus, and the cingulate gyus, espectively. A lateality index (LI: [Left 2 Right]/[Left 1 Right]) of activated voxels was calculated fo the whole 2011
Taylo et al. TABLE I. Aeas of significant activation duing ecognition of all faces types minus baseline Anatomical egion Bodmann aea Hemisphee x y z (t)-value Own faces minus baseline Fusifom gyus BA 37 R 42 248 214 5.507 L 240 247 214 4.695 Infeio occipital gyus BA 19 R 43 277 24 5.184 L 234 276 24 4.678 Lingual gyus BA 18 R 17 278 23 3.641 L 25 279 23 3.837 Cuneus BA 17/18 R 24 286 13 4.640 L 222 282 18 4.339 Pecuneus BA 7 R 20 270 41 3.821 Patne s faces minus baseline Fusifom gyus BA 37 R 47 245 216 5.262 L 238 246 216 6.417 Lingual gyus BA 18 R 12 276 23 3.398 L 217 276 26 4.587 Cuneus BA 18 R 19 285 18 3.510 Paahippocampal gyus BA 36 R 14 42 26 4.202 Middle tempoal gyus BA 39 R 45 264 20 5.209 Supeio tempoal gyus BA 13 R 45 243 20 3.394 Pecuneus BA 7 L 26 258 37 3.589 Middle fontal gyus BA 9 L 237 26 29 4.664 Paent faces minus baseline Fusifom gyus BA 37 R 40 249 216 4.933 L 238 260 211 3.970 Famous faces minus baseline Fusifom gyus BA 37 R 40 255 214 4.329 Unfamilia faces minus baseline Anteio cingulate BA 32 R 7 30 25 24.217 L 23 33 24 23.837 Cingulate gyus BA 24 R 6 210 28 23.772 L 25 13 30 24.754 bain and each ROI. LI > 0.20 was deemed left dominant and LI < 20.20 as ight dominant, values in-between wee consideed bilateal. Thee multiple linea egession models wee pefomed on activity elated to own, patne, and paent faces based on the questionnaie. Pedictos of the activity elated to own faces included age, time spent looking at one s self in the mio in minutes pe day, and a ating out of 10 of how comfotable they wee with thei own face. Age, hous spent with patne pe day and elationship length in yeas wee the pedictos fo activity elated to patne faces. Activity elated to paent faces was egessed on age, time it would take to each the paents home, and days spent with paents pe yea. Simila to above, clustewise multiple compaison contols wee used at a P-value of 0.01 yielding a minimum volume of 5.1 cc (188 voxels) fo own faces, 2.2 cc (81 voxels) fo patne faces, and 2.3 cc (85 voxels) fo paent faces with a P-value of 0.05. RESULTS To examine the neual coelates of face type ove the whole bain, each face type was compaed to est/baseline (the fixation coss; Table I) and unfamilia faces (Table II). Based on the stingent cluste size theshold, the epoted egions appea as pat of clustes, which ae equal to o lage than 210 voxels (Fig. 2a,b). Compaed with baseline, patne faces activated bilateal (L/R) fusifom gyus, ight (R) lingual gyus, cuneus (R), paahippocampal gyus (R), middle tempoal gyus (R), supeio tempoal gyus (R) and left (L) pecuneus and middle fontal gyus (L). Paent faces, compaed to baseline, only activated the fusifom gyus (L/R). Pictues of the paticipant s face (i.e., own face) activated fusifom gyus (L/R), infeio occipital gyus (L/R), lingual gyus (L/R), cuneus (L/R) and pecuneus (R). Famous faces activated the fusifom gyus (R) and unfamilia faces showed deceased activation of the anteio cingulate (L/R) and cingulate gyus (L/R). The LI esults ove the whole bain showed that, compaed to baseline, unfamilia faces activation was bilateal (LI 5 0.002; Fig. 2e). All othe faces wee ight latealized with patne faces the least and famous faces the most ight latealized [LI 520.288 fo patne; LI 520.311 fo paent; LI 520.373 fo own; and LI 521.0 fo famous faces]. All pesonally familia faces wee contasted with unfamilia faces. Results showed that patne faces had the most extended activations, including the anteio cingulate (L/R), cingulate gyus (L/R), medial fontal gyus (L/R), middle fontal gyus (L), infeio fontal gyus (BA 47; L), 2012
Pesonally Familia Faces TABLE II. Aeas of significant activation duing ecognition of pesonally familia minus unfamilia faces Anatomical egion Bodmann aea Hemisphee x y z (t)-value Own faces minus unfamilia faces Anteio cingulate BA 32 R 3 35 20 3.930 L 21 36 20 3.857 Cingulate gyus BA 32/24 R 7 18 30 3.946 L 21 20 30 3.716 Medial fontal gyus BA 9 R 18 27 31 3.698 L 215 29 30 4.272 Middle fontal gyus BA 46 L 238 36 17 3.342 Patne s faces minus unfamilia faces Anteio cingulate BA 32 R 5 33 24 4.133 L 23 31 24 3.377 Cingulate gyus BA 32 R 5 31 29 3.639 L 22 31 28 4.032 Medial fontal gyus BA 10 R 7 59 8 3.787 L 26 49 8 3.450 Middle fontal gyus BA 8 L 223 36 38 4.279 Infeio fontal gyus BA 47 L 222 17 214 4.074 Middle tempoal gyus BA 20/21 L 249 211 214 3.322 Insula BA 13 L 228 15 24 3.300 Amygdala BA 34 L 225 22 210 3.218 Thalamus L 220 226 5 3.303 Paahippocampal gyus BA 34/28 L 214 29 214 3.314 Pecuneus BA 7 L 26 256 41 3.375 Paent faces minus unfamilia faces Cingulate gyus BA 32/24 R 5 26 30 3.659 L 21 21 30 3.901 Supeio fontal gyus BA 6/8 R 6 22 56 3.831 middle tempoal gyus (L), paahippocampal gyus (L), amygdala (L), insula (L), thalamus (L) and pecuneus (L). The paent faces only activated the cingulate gyus (L/R) and the supeio fontal gyus (BA 6; R). Own faces compaed with unfamilia faces activated the anteio cingulate (L/R), cingulate gyus (L/R), medial fontal gyus (L/R) and middle fontal gyus (L). The LI esults showed that compaed with unfamilia faces, patne faces wee left latealized (LI 5 0.205), paent faces wee ight latealized (LI 5 20.413), and own faces wee bilateal (LI 520.112; Fig. 2f). The ROI fo the fusifom gyus showed bilateal activation fo pesonally familia faces, while famous faces activated only the ight fusifom (Fig. 2c). Activation associated with own and patne faces extended most anteioly in both hemisphees, while own faces also extended into infeio pats of the fusifom gyus. Activation elated to paent faces occupied pimaily cental egions of the fusifom gyus, ovelapping with both own and patne faces. The lateality index showed that familia and all pesonally familia face types showed highe voxel counts in the ight fusifom (own, LI 5 20.220; patne, LI 5 20.251; paents, LI 520.316; famous, LI 520.566), with inceasing asymmety fom own to famous faces (Fig. 2e). The ROI analysis of the cingulate gyus showed geate activation fo pesonally familia faces than unfamilia faces, with the most extensive activation seen to own faces, and the least to paents (Fig. 2d). Paent faces activated mostly the ight cingulate gyus. Activation associated with own and patne faces extended most anteioly in the left hemisphee, wheeas own and paent faces extended most anteioly in the ight hemisphee. Activation of own and patne faces ovelapped in posteio pats of the cingulate gyus. The lateality index confimed the initial obsevation of slightly moe voxels activated in the ight cingulate gyus fo paents faces (LI 520.194) and own face (LI 520.222) and moe voxels activated in the left cingulate gyus fo patne faces (LI 5 0.288). Famous faces did not elicit significant activation in this aea. Significant pedictos of activation elated to patne faces wee time spent daily with patne and elationship length, and included activity in ight supeio tempoal gyus and left insula (see Table III). The oveall egession model was significant, suggesting that these pedictos adequately explained the data. All pedictos of own faces (age, time spent in font of a mio, ating of own face) coelated significantly with medial fontal gyus activation (Table III). Howeve, the oveall model did not fully explain the data, suggesting that additional pedicto vaiables ae needed to descibe activation elated to own faces. Activation elicited by paents faces did not coelate with age, distance fom paents, o days spent with paents pe yea. DISCUSSION We examined the neual substates associated with pocessing patne, paents and own faces, each in compaison 2013
Taylo et al. Repesentation of the egions that showed significant BOLD esponses fo (a) contast fo all face categoies > Baseline, (b) contast fo pesonally familia faces > unfamilia faces, (c) ROI fo the fusifom gyus fo all face types, and (d) ROI fo the cingulate gyus fo pesonally familia faces. Lateality indices (LI) Figue 2. ae pesented on gaphs (e) fo all faces vesus baseline in the fusifom gyus and whole bain analysis and (f) fo pesonally familia vesus unfamilia faces in the cingulate gyus and the whole bain analysis. with baseline and unknown faces. We found that pesonally familia faces all activated the fusifom and the cingulate gyi, with othe aeas activated depending on the face. Notably, patne faces showed the most extensive activation. Paent faces, which have been known fo the longest time, showed the least activation, an intiguing finding 2014
Pesonally Familia Faces TABLE III. Aeas of significant activation as function of pedictos fo patne and own faces Pedicto Anatomical egion Diection Bodmann aea Hemisphee x y z (t)-value Patne faces Time spent togethe with patne daily Pecental gyus ; BA 6 R 48 210 34 24.10 L 254 25 30 24.24 Infeio fontal gyus ; BA 9 R 60 7 26 24.59 Postcental guus ; BA 4 L 254 214 31 24.52 Relationship length Pecental gyus ; BA 42/43 R 56 210 12 24.17 L 255 210 13 24.83 Supeio tempoal gyus ; BA 22 R 59 1 4 24.86 Tansvese tempoal gyus ; BA 42 R 56 213 12 24.01 Insula ; BA 13 L 239 210 12 24.38 Own faces Age Paacental lobule ; BA 6 R 7 229 64 26.23 Postcental gyus ; BA 3 R 28 229 63 25.01 Pecuneus ; BA 7 R 4 247 59 24.31 Time spent in font of mio daily Medial fontal gyus : BA 9 R 10 48 23 4.22 L 23 50 18 4.53 Supeio fontal gyus : BA 9 L 28 41 33 11.81 Own face ating Supeio fontal gyus : BA 9 R 6 56 23 5.08 L 23 57 24 6.74 Supeio fontal gyus : BA 10 R 19 57 21 3.73 Medial fontal gyus : BA 9 R 2 48 21 4.81 L 23 48 21 4.03 Pecuneus : BA 7 L 24 253 61 4.45 Paacental gyus : BA 6 R 6 232 66 5.33 Postcental gyus : BA 3 R 31 229 63 5.58 Aows show the diection of coelation: : is positive, ; is negative. consideing that one could expect that these faces to tigge a netwok of bain aeas involved in autobiogaphical memoy. Lateality index analyses evealed that, egading whole bain o egional analyses, pesonally familia faces wee less ight latealized than famous faces. Compaed with unfamilia faces, patne faces wee left latealized while own faces wee bilateal. These esults, the fist to distinguish long-tem patne and paents face activations, indicate that the netwok model of familia face pocessing [Gobbini and Haxby, 2007; Haxby et al., 2000] is modulated accoding to the type of familia face. These esults contibute to ou undestanding of some featues pesented by posopagnosic patients. These patients usually ecognize the faces of close elatives bette than othe faces. This has usually been explained by highe fequency and deepe memoy taces fo these faces. Howeve, these esults suggest that it could be elated to distinct netwoks which ae latealized diffeently fo pesonally familia faces than fo othe faces. Also, these findings make impotant pedictions fo futue studies, paticulaly in neuophysiology, as components associated with face pocessing (e.g., N170-P2) which ae usually lage ove the ight hemisphee, should be modulated by whethe the face is famous, the patne s o one s own face. All Face Types Vesus Baseline As expected, activations elicited by all faces compaed to a fixation coss ecuited aeas associated with the visual ecognition of faces [e.g., McCathy et al., 1997; Puce et al., 1996] such as the fusifom gyus. The patne, paent, and own faces evoked bilateal activation in the fusifom gyus; famous faces activation was significant only in the ight hemisphee. Patne and own faces also activated bilateal lingual gyus and cuneus; infeio occipital gyus activation was only seen fo own faces. In addition to the coe visual system [e.g., Gobbini and Haxby, 2007], futhe aeas wee involved fo pocessing patne and own faces; both activated the pecuneus, but patne faces activated the left while own faces activated ight. In the face pocessing liteatue, the pecuneus is associated with peson knowledge pocessing [Gobbini and Haxby, 2006, 2007], but moe geneally the pecuneus is linked with visuospatial analysis of objects [Faillenot et al., 1999], and a ecognition netwok suppoting etieval [Nagahama et al., 1999; Rebe et al., 2002]. Thus, the pecuneus may well be involved in etieval of peson knowledge. The intiguing latealizations may be attibuted to the diffeences between peson knowledge and self-knowledge, and meit futhe study. 2015
Taylo et al. Patne faces also geneated activation in the middle and supeio tempoal gyi, the paahippocampal, and the middle fontal gyi (BA 9). The supeio tempoal gyus is pat of the netwok linked with the social cognition of faces [Allison et al., 2000]. The paahippocampal gyus is involved in encoding visual infomation [e.g., Kichhoff et al., 2000; Rombouts et al., 1999], which in face studies is elated to the ecognition of familia faces [Babeau et al., 2008; Leveoni et al., 2000] and memoy etieval of pesonal and social knowledge [Sugiua et al., 2005]. Memoy etieval is also likely to be suppoted by activity in the middle fontal gyus, which has been linked with cognitive functions such as woking memoy and attention [e.g., Chistoff and Gabielli, 2000; Petides, 1996]. Activation in this aea was associated with pesonally familia faces as well [Batel and Zeki, 2000; Gobbini and Haxby, 2006; Platek et al., 2006]; we suggest that in this study, the middle fontal gyus was involved in holding the peson infomation in mind. Thus compaed to baseline, all pesonally familia and famous faces activated the coe visual system fo pocessing faces, with an inteesting lateality diffeentiation in the pecuneus in the compaisons of patne and own faces. Patne faces activated an extensive netwok of egions, likely linked with the mnemonic and semantic attibutes of the patne. Pesonally Familia Vesus Unfamilia Faces Patne, paent and own faces elicited significantly moe cotical activation than unfamilia faces. All pesonally familia faces evoked activation in the cingulate gyus bilateally; while patne and own faces also activated the anteio cingulate bilateally. The cingulate gyus is a multimodal aea [Tuak et al., 2002]. In face pocessing studies it shows a stonge esponse to familia faces [Platek et al., 2006], self-esembling faces [i.e., kin ecognition; Platek et al., 2008], and it is likely involved in the integation of infomation elicited by the face [Devue et al., 2007]. Significant cingulate gyus activation was evident in the compaisons between unfamilia and pesonally familia faces, but not with famous faces. Anteio cingulate egions ae associated with emotional pocessing and posteio egions associated with cognitive pocessing [Bush et al., 2000]. The poposed emotional and cognitive subdivisions of the cingulate gyus have ecipocal inteconnections with aeas including the amygdala and lateal pefontal cotices, espectively [Bush et al., 2000], aeas also activated by pesonally familia faces in the cuent study. Theefoe, given the link between the cingulate gyus, and peson knowledge (including both emotional and semantic infomation), we suggest that the cingulate activation to pesonally familia faces is elated to shaed expeiences at a pesonal level. In addition, patne and own faces elicited activation in bilateal medial fontal and left middle fontal gyi; the infeio fontal gyus esponded only to patne faces. Pefontal egions undelie cognitive functions such as memoy and attention, although the exact natue of thei involvement emains a matte of debate. The middle fontal and medial fontal gyi have been linked to manipulation and monitoing of infomation effotfully held in mind [Chistoff and Gabielli, 2000], while the infeio fontal gyus (BA 47) is associated with pocessing syntax and speech [De Cali et al., 2007], as well as maintaining one o a few items in mind [Chistoff and Gabielli, 2000]. The infeio fontal gyus has also been elated to pocessing of emotional stimuli and maintains stong connections with infeio tempoal egions and limbic aeas such as the paahippocampal gyus [Petides and Pandya, 2002], linked to the active judgement of stimuli. Thus, as a patne s face is likely to evoke specific memoies o events with some emotional ovelay, we popose that pefontal activation elated to viewing a patne s face eflects these types of infomation being eviewed o monitoed. Besides the anteio cingulate and pefontal activity, patne faces activated aeas associated specifically with emotional pocessing o emotional valence [Bitton et al., 2006] such as the insula, amygdala and thalamus. The amygdala have been shown to elicit a stonge esponse to faces compaed to pictues [Segeie et al., 2008], and the ecipocal functional connections between the amygdala and the vental pefontal cotex impact cognitive judgement [Nomua et al., 2004] and the matuation of cognitive and affective contol [Hae and Casey, 2005]. Insula activity has been epoted to patne faces in the ealy in love stage of a elationship [Batels and Zeki, 2000; Fishe et al., 2005] and has been associated with pocessing emotions as pat of face ecognition [Gobbini and Haxby, 2007]. Thus, although ou paticipants had expeienced much longe elationships than paticipants in pevious studies, the limbic activation is consistent with these othe epots, and suggests some similaities between elationships that stat well and those that last fo many yeas. The supeio fontal gyus was activated uniquely to paent faces. Pevious face imaging studies have linked the supeio fontal gyus with a self-awaeness netwok [Platek et al., 2006], kin ecognition [Platek et al., 2005], pesonal choice [Tuk et al., 2004] and self-elated pocessing [Goldbeg et al., 2006]. So why would paent faces activate the supeio fontal gyus moe than any of the othe faces? The ight supeio fontal gyus has been implicated in a netwok of aeas associated with facial esemblance, linked with detection of facial familiaity and kin ecognition [Platek et al., 2005]. Paents would be associated with self-awaeness o self-esemblance, as we all see similaities and diffeences between ouselves and paents; these compaisons may contibute to ou self-image and selfelated pocesses and be evoked with seeing paents photogaphs. Oveall, these findings suggest that the anteio cingulate and cingulate gyus togethe with pefontal activations may facilitate the integation and monitoing of pesonal infomation pesent in both one s own face, paents 2016
Pesonally Familia Faces faces and patne s face, but not in unfamilia faces. Compaed to unfamilia faces, own faces ecuit aeas pimaily associated with peson knowledge (o self-knowledge), while patne and paents faces ecuited aeas associated both with peson knowledge, memoy, emotional pocessing and self-image. ROI Analyses and Lateality Indices Despite the fusifom gyus being elated to physiognomic pocessing of faces geneally, egions of inteest (ROI) analyses showed diffeences with pesonal familiaity. Aeas in the mid-fusifom, paticulaly in the ight hemisphee, wee activated fo all faces, but the extent vaied with the type of face (Fig. 2c). Patne and own faces activated the most anteio pats of the fusifom gyus bilateally, and own faces activation was also seen in supeio aeas in the left posteio fusifom. Paent faces activation was the most esticted, being simila to famous faces, except that it occupied cental aeas of the fusifom gyus bilateally, while activation to famous faces was mostly in the ight cental fusifom. Thus, the faces seen most often on a daily basis activated lage aeas of the fusifom. This is inteesting in light of developmental data which show that with inceased skill o familiaity with faces between 12-yea-olds and adults, the fusifom aea activated becomes smalle [e.g., Passaotti et al., 2003]. Clealy, the pesonal significance of one s own and one s patne s face impacts pocessing fom the ealy stages, poducing distinct and special activation pattens. The lateality index fo the fusifom (Fig. 2e), suggested that compaed to baseline, activation fo all faces is stongly latealized to the ight, with the most latealized being the famous faces and the least being own faces. A contast of these pattens may have poduced the appaent left-latealization epoted in the liteatue fo own faces. A lateality index of all faces compaed to baseline was also pefomed fo the whole bain, and esults wee vey simila to the index geneated fo the fusifom gyus. The similaity of the whole bain and fusifom gyus lateality indices suggests that fo faces, the lateality may be diven by activation in the fusifom gyus. These esults suggest that the accepted view that face pocessing elies pedominantly on the ight hemisphee may be modulated accoding to the type of face being pocessed. All pesonally familia faces activated lage pats of the cingulate gyus, compaed to unfamilia faces, and as with the fusifom, the ROI analyses showed that the extent diffeed with face type (see Fig. 2d). Own faces elicited the geatest spead of activation, shaing a lage anteio aea of ight cingulate gyus with paent faces and a lage anteio aea in the left with patne faces. Patne faces wee epesented moe dosally in both anteio and posteio aeas of the cingulate gyus. The cingulate cotex has been stongly linked with vaious cognitive functions elated to self-contol o self-eflection, and the lage activation to own faces is consistent with this. Also, the cingulate gyus is associated with monitoing emotions and empathy, aspects likely invoked by the faces of paents o patnes. The latealization of cingulate gyus activation may be elated to the duation of the familiaity. Paents and own faces have the longest extent of familiaity and tend to elicit moe activation in the ight cingulate. Patne s faces on the othe hand ae newe and ae epesented lagely in the left cingulate. Investigating the effect of duation of familiaity equies futhe eseach; howeve, the design would be complicated, and it would be difficult to patial out duation of the elationship while contolling fo pesonal familiaity. Unlike the fusifom, the lateality effects in the cingulate wee not closely linked to the whole-bain esults. Fo example, lateality indices showed that own faces wee ight latealized in the cingulate gyus, but the whole bain index suggested that own faces activate aeas bilateally (Fig. 2f). The latte is in ageement with a ecent epot that own face activations ae sustained by a bilateal netwok [Platek et al., 2006]. The type o aea of activation examined may lead to a moe ight- o left-sided patten and pehaps explain the discepant lateality epots in the liteatue [Bady et al., 2004; e.g., Keenan et al., 2000]. Pedictos of Activation Related to One s Own and Patne s Faces Pesonal familiaity is based on many factos, which would include not only the length of the elationship, but also the time spent togethe on a daily basis; both of these factos wee significantly elated to activation elicited by patne s faces. As time spent togethe daily inceased, activation in the infeio fontal, pecental, and postcental gyi deceased. Chistoff and Gabielli [2000] found that activation in the infeio fontal gyus was linked to monitoing items online; the pesent esult suggests that inceases in daily time togethe may educe the need fo effotful maintenance of infomation about the peson. Similaly, activation in the ight supeio and tansvese tempoal gyi and the left insula deceased as elationship length inceased. The insula is associated with pocessing of patnes faces [Batels and Zeki, 2000; Fishe et al., 2005] and pehaps the emotional component deceases as elationship length inceases. Pedictos of activation elated to one s own face, included age, time spent looking at one s self in the mio, and one s ating of one s own face. Activation in the ight pecuneus, paacental lobule and postcental gyus deceased with age. In contast, activation in the left pecuneus inceased with highe atings of one s own face. This is an inteesting distinction as it highlights a diffeence between a cuent efeence (i.e., how one ates one s face at a point in time), which is epesented in the left pecuneus, and a efeence acoss time fom chonological age in the ight pecuneus. The pecental gyus was peviously linked with activation elicited by own faces, consistent with Moita et al. [2008], who suggested that this 2017
Taylo et al. aea is specifically involved in own-face ecognition, although the activation they epoted was moe lateal and infeio than in the pesent study. The supeio fontal and medial fontal gyi (BA 9) activation inceased as both the time one spends in font of the mio and the ating of one s own face inceased. A numbe of studies have shown that the pefontal cotex, paticulaly on the ight, activated specifically unde cicumstances of self-evaluation o self-ecognition [Fossati et al., 2003; Moita et al., 2008; Uddin et al., 2005]. Oveall, evidence fom egession analyses indicate that expeiential pedictos such as time spent with one s patne o looking at one s self in the mio eveyday modulate neual activity. CONCLUSIONS This study detemined neual esponses to pesonally familia faces and extends pevious esults to diect compaisons of seveal types of pesonally familia faces, including patne, own and paent faces. Despite the small sample size, we pesent impotant data which suppot pio wok and give peliminay esults on the diffeentiation of the neual substates of pesonally familia faces. The cuent esults povide new evidence on the bain aeas implicated in pocessing paent faces, which wee the only ones to exhibit activation in the supeio fontal gyus. Patne faces engaged the most extensive cotical aeas including those associated with emotional pocessing and uniquely activated the infeio fontal gyus. The patten obseved in the ROI analyses of the fusifom and cingulate gyi showed anatomical distinctions in pocessing patne, paent and own faces. Lateality indices suggested that fusifom gyus activity, associated with the coe visual system, is ight latealized. Howeve, cingulate gyus activation associated with extended and emotional pocessing was moe complex; patne faces wee left-latealized, own faces wee ight-latealized, and paent faces showed bilateal activity with a ight bias. 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