Seeing John Malkovich: the Neural Substrates of Person Categorization

www.elsevier.com/locate/ynimg NeuroImage 24 (2005) 1147–1153

Seeing John Malkovich: the neural substrates of person categorization

David J. Turk,a,* Amy C. Rosenblum,a Michael S. Gazzaniga,a and C. Neil Macraeb aDepartment of Psychological and Brain Sciences, Center for Cognitive Neuroscience, Dartmouth College, 6162 Moore Hall, Hanover, NH 03755, USA bSchool of Psychology, University of Aberdeen, Aberdeen AB24 2UB, UK

Received 25 May 2004; revised 25 October 2004; accepted 28 October 2004 Available online 8 December 2004

Neuroimaging data have implicated regions of the ventral temporal the basis of findings such as these, it has been suggested that it is cortex (e.g., fusiform gyrus) as functionally important in face not stimulus type that modulates activity in visual association recognition. Recent evidence, however, suggests that these regions are cortex, but rather the level of categorical specificity at which not face-specific, but rather reflect subordinate-level categorical objects are identified (see Haxby et al., 2000; Tarr and Gauthier, processing underpinned by perceptual expertise. Moreover, when 2000). people possess expertise for a particular class of stimuli (e.g., faces), Any object, including a person, can be identified at multiple subordinate-level identification is thought to be an automatic process. To investigate the neural substrates of person construal, we used levels of abstraction (e.g., bird vs. canary; male vs. Al Pacino—see functional magnetic resonance imaging (fMRI) to contrast brain Rosch et al., 1976). It is generally assumed, however, that objects activity while participants judged faces at different levels of semantic are first identified at what is termed the entry level of categorical specificity (i.e., identity vs. occupation). The results revealed that representation (Jolicoeur et al., 1984). This is the level at which a participants were quicker to access identity than occupational knowl- name can be generated or matched most rapidly to an object. edge. In addition, greater activity was observed in bilateral regions of Although such descriptions dominate the recognition process, in no the fusiform gyrus on identity than occupation trials. Taken together, sense is this the only level at which items can be identified; these findings support the viewpoint that person construal is charac- sometimes, people prefer subordinate or exemplar-based descrip- terized by the ability to access subordinate-level semantic information tions of stimuli. Given this observation, Gauthier and colleagues about people, a capacity that is underpinned by neural activity in have speculated that the apparent specificity of the fusiform gyrus discrete regions of the ventral temporal cortex. D 2004 Elsevier Inc. All rights reserved. for faces is a byproduct of differences in the nature of face processing and object recognition (Gauthier et al., 1997, 1999, 2000). Specifically, whereas object recognition typically unfolds at Keywords: Face processing; fMRI; Fusiform gyrus; Semantic knowledge; the basic level of categorical abstraction (e.g., dog rather than Person perception spaniel), face recognition is automatized at the individual or exemplar-based level (e.g., Sean Connery rather than human). Supporting this viewpoint, Tanaka (2001) has demonstrated that An extensive literature has demonstrated that regions of the adults identify familiar faces more often and as rapidly at the ventral temporal cortex (e.g., fusiform gurus) play a critical role in individual-level than the basic level of abstraction. Critically, person perception, notably the registration and recognition of however, this downward shift in the specificity of identification has others (Haxby et al., 2000; Hoffman and Haxby, 2000; Kanwisher also been observed when canine experts view dogs (Tanaka and et al., 1997, 1999; McCarthy et al., 1997). While it was initially Taylor, 1991) and when people respond to artificial objects (i.e., suggested that this effect may reflect the domain specificity of face Greebles) for which they have received extensive prior training processing in the brain (Kanwisher et al., 1997; McCarthy et al., (Gauthier et al., 1999). As Tanaka (2001) contends, bface expertise, 1997), subsequent work has revealed that activity in ventral like object expertise, promotes a downward shift in recognition to temporal cortex is modulated by a wide range of stimuli (Chao et more subordinate levels of abstractionQ (p. 534). al., 1999; Haxby et al., 2001; Ishai et al., 1999). This includes, but But what is the functional significance of activity in regions of is not restricted to faces, such that activation in the fusiform gyrus ventral temporal cortex during subordinate-level identification? has been reported when bird and car experts view exemplars from While activity in this region is acknowledged to index fine- their respective areas of interest (Gauthier et al., 1999, 2000). On grained perceptual discrimination (Gauthier et al., 1997; Haxby et al., 2001; Mason and Macrae, in press), it is worth noting that * Corresponding author. Fax: +1 603 646 1181. portions of the fusiform gyrus, particularly in the left hemisphere, E-mail address: [email protected] (D.J. Turk). have also been associated with the generation of semantic Available online on ScienceDirect (www.sciencedirect.com). knowledge about objects (Gorno-Tempini and Price, 2001;

Gorno-Tempini et al., 1998; Kan et al., 2003; Thompson-Schill, ments of a person’s occupation (see Sergent et al., 1992). To 2003; Thompson-Schill et al., 1999; Zelkowicz et al., 1998). It is explore this possibility, we utilized a variant of the visual- possible, therefore, that this cortical region may also be sensitive identification task adopted by Gauthier et al. (1997).Ina to the semantic specificity of person categorization (Martin, matching paradigm, participants were required to report if a face 2001). An emerging literature has demonstrated that conceptual and verbal label matched at the occupational or identity level. knowledge is grounded in the neural architecture that supports Brain activity was measured during the performance of these perceptual processing (Kan et al., 2003; Martin, 2001; Martin et tasks. al., 1995, 2000; Thompson-Schill, 2003). As such, one might expect activity in visual association cortex to be modulated by the specificity of semantic construal, in much the same way that these Method regions are responsive to the specificity of perceptual identification (Gauthier et al., 1997; Haxby et al., 2001). Participants and design Generally speaking, categorization level and perceptual processing are related such that, the finer the level of Thirteen participants (6 males, mean age = 24 years) identification, the more perceptual information that is required completed the study for course credit or $10. All participants to support the judgment under consideration (e.g., pelican vs. were right-handed, reported no significant abnormal neurological bird, Gauthier et al., 1997; Sergent et al., 1992; Tarr and history, and had normal or corrected-to-normal visual acuity. Gauthier, 2000). Unlike object processing, however, faces can be Informed written consent for all participants was obtained prior classified not only with respect to their physical properties (e.g., to the experiment in accordance with the guidelines established sex, race, emotion), but also with regard to their applicable by the Committee for the Protection of Human Subjects at semantic categorizations (e.g., politician, humanitarian). More- Dartmouth College. The experiment had a single factor (level of over, because there is nothing in the physical structure of a face abstraction: occupation or identity) repeated measures design. that specifies membership in any of these nonvisually derived categories (or indeed the possession of a particular name), Stimulus materials and procedure semantic judgments at different levels of specificity (e.g., occupation vs. identity) are matched for perceptual difficulty. Color photographs of 84 celebrities were used. Each facial That is, the same perceptual information is required to compute photograph was resized and placed on a black background that was the occupation or name of a person. It should also be noted that 200 Â 200 pixels (2 Â 2 in.) in size. The stimulus set comprised 42 occupational judgments do not necessitate the prior establishment singers and 42 actors, with an equal number of male and female of a person’s identity. On many occasions, people are able to exemplars in each set. Each participant performed two tasks: an provide biographical details about a person, yet be unable to identity task and an occupation task. In the identity task, name the individual in question (Young et al., 1985, 1986). Given participants were required to report, via a button press, whether a these observations, famous faces provide a useful means to photograph matched a simultaneously presented name (e.g., a investigate the neural activity that accompanies semantic judg- picture of John Malkovich together with the name dJohn ments at different levels of specificity. MalkovichT or a gender-matched name from the same occupational If, therefore, regions of ventral temporal cortex are sensitive group, e.g., Sean Connery). In the occupation task, the name was to the specificity of semantic categorization, identity judgments replaced by an occupation (e.g., a picture of John Malkovich should be accompanied by greater neural activity than judg- together with the word dactorT or dsingerT; see Fig. 1). The stimulus

Fig. 1. Schematic representation of a single functional run. D.J. Turk et al. / NeuroImage 24 (2005) 1147–1153 1149 pairs (face–name or face–occupation) were presented for 2 s, with Results a 500-ms interval between trials. In total, participants completed 12 blocks (21 trials per block) of each trial type distributed across Behavioral data collected during scanning showed no difference three functional runs with equal numbers of congruent and in the accuracy [t(12) b 1, ns] of participants’ responses [respective incongruent stimulus pairs in each run. Participants completed a M’s: occupation = 82.7%; identity = 82.3%]. There was, however, total of 252 trials in each of the two conditions. a difference in the time taken to make a correct response [t(12) = 3.16, P b 0.01], such that responses were faster on identity than Image acquisition occupation trials (respective M’s: 765 ms vs. 853 ms). To examine the neural correlates of person construal, we Imaging was performed on a 1.5-T whole body scanner compared the BOLD response associated with identity judgments (General Electric Medical Systems Signa, Milwaukee, WI) with to that associated with occupation judgments (i.e., identity N a standard head coil. Visual stimuli were generated using an Apple occupation). This contrast revealed an increased BOLD signal in G3 laptop computer running PsyScope software (Cohen et al., several areas in the frontal, parietal, and occipital cortices (see 1993). Stimuli were projected to participants with an Epson (model Table 1). Of greater theoretical interest, however, were observed ELP-700) LCD projector onto a screen positioned at the head end differences in activity in regions of the ventral temporal cortex and of the bore. Participants viewed the screen through a mirror. A limbic system. As expected, bilateral regions of the fusiform cortex fiber-optic light sensitive key-press, interfaced with the PsyScope were more active during identity than occupation trials (see Fig. 2). Button Box (New Micros, Dallas, TX), was used to record In addition, a greater BOLD signal was observed for identity than participants’ behavioral performance. Cushions were used to occupation judgments in the left parahippocampal gyrus (BA 28/ minimize head movement. 34). There were no brain areas that demonstrated significantly Anatomical images were acquired using a high-resolution 3-D greater activity during occupation than identity judgments (i.e., spoiled gradient recovery sequence (SPGR; 124 saggital slices, occupation N identity). TE = 3.2 ms, TR = 8 ms, flip angle = 158, voxel size = 1 Â 1 Â 1.2 mm). Functional images were collected in runs using a gradient spin-echo, echo-planar sequence sensitive to BOLD General discussion contrast (T2*) (TR = 2500 ms, T2* evolution time = 35 ms, flip angle = 908, 3.75 Â 3.75 in-plane resolution). During each The current behavioral results corroborate the contention that, at functional run, 240 sets of axial images (25 slices; 4.5-mm slice least for famous faces, the entry point for person recognition is at thickness, 1-mm skip between slices) were acquired allowing the level of a unique identifier (Tanaka, 2001; Tarr and Gauthier, complete brain coverage. 2000). Participants were quicker to access the identity than the occupation of a collection of famous individuals, thereby suggest- Image analysis ing that person construal is automatized at the exemplar level of categorical abstraction. This result is at odds with serial models of All data were analyzed using SPM99 software (Wellcome face processing. The influential Bruce and Young (1986) model, Department of Cognitive Neurology, London, UK; Friston et for example, postulates that access to a person’s name follows (and al., 1995). Functional data were corrected for differences in is contingent upon) the prior retrieval of semantic knowledge (e.g., acquisition time between slices for each whole-brain volume, occupation). More recent models of face processing, however, can realigned within and across runs to correct for head movement, accommodate the current findings. In the Interactive Activation and coregistered with each participant’s anatomical data. and Competition (IAC) model (Burton, 1994; Burton et al., 1990), Functional data were then transformed into a standard information from face recognition units (FRUs), early visual anatomical space (2-mm isotropic voxels) based on the ICBM processing, and name input units (NIUs) link directly to a unique 152 brain template (Montreal Neurologic Institute), which person identity node (PIN). Thus, concurrent name and face inputs approximates Talairach and Tournoux’s (1988) atlas space. (as in our identity-level condition) should activate a single PIN and Normalized data were then spatially smoothed (6-mm full- prompt the elicitation of a rapid response. Responding should be width-at-half-maximum [FWHM]) using a Gaussian kernel. The impeded, however, when concurrent semantic and face inputs are normalized and smoothed images were then used for statistical present (as in our occupation-level condition) as several PINs (e.g., analysis. Clint Eastwood, Brad Pitt, Mel Gibson) can share the same For each participant, a general linear model, incorporating task semantic unit (e.g., actor). This therefore adds noise to the system effects (modeled as a box-car function convolved with the and impairs (i.e., slows down) responding. canonical hemodynamic response function), a mean, and a linear Along with rapid response times, identity matching prompted trend were used to compute parameter estimates (b) and t- increased activity in bilateral regions of the fusiform gyrus contrast images (containing weighted-parameter estimates) for (Gauthier et al., 1997; Tarr and Gauthier, 2000). This then supports each comparison at each voxel. A random-effects analysis (one- the notion that activity in the fusiform gyrus is modulated by the sample t test, hypothesized mean = 0) was then applied to the specificity of semantic construal. In a related positron emission individual subject t contrast images to create mean t images tomography (PET) investigation, Sergent et al. (1992) presented (thresholded at P = 0.0001, uncorrected). An automated peak participants with a series of famous faces and contrasted the neural search algorithm identified the location of peak activations and activity associated with gender and occupational judgments. As in deactivations based on z value and cluster size (10 mm3). This the current inquiry, the results revealed significant increases in analysis enables individual trial types to be directly compared activity in bilateral areas of the fusiform gyrus (BA 36 and 37). (e.g., identity N occupation) to identify activations that differ While suggestive that this activity may index the specificity of between conditions. semantic construal, judgments of identity demand greater percep- 1150 D.J. Turk et al. / NeuroImage 24 (2005) 1147–1153

Table 1 Summary of the significant results for the identity N occupation contrast Region Left hemisphere Right hemisphere Talairach coordinates Talairach coordinates K value xyzZmax K value xy z Zmax Occipital cortex Medial/superior occipital cortex (BA18) 143 À26 À89 32 5.40 24 26 À82 21 4.61 Inferior occipital cortex/cuneus 108 À28 À89 4 4.60 Lingual gyrus (BA 18) 31 À12 À88 À9 4.14 Lingual gyrus (BA 18) 14 À15 À77 6 4.11 Cuneus (BA 19) 11 À6 À82 24 4.18 24 8 À91 12 4.61 Superior occipital cortex (BA 19) 34 34 À80 26 4.23 Inferior longitudinal fasciculus (BA 18) 15 12 À77 11 4.15 Inferior medial occipital cortex (BA 19) 194 28 À76 33 5.18 Medial occipital cortex (BA 19) 26 34 À71 22 4.37

Parietal cortex Parietal lobe (BA 7) 12 24 À75 55 4.04 51 32 À65 59 4.26 24 28 À60 38 4.54 Precuneus (BA 7/31) 21 À12 À52 45 4.35

Temporal cortex Fusiform gyrus (BA 19) 188 À34 À78 À15 4.96 14 40 À72 À6 4.00 Fusiform gyrus (BA 37) 50 À38 À57 À12 4.79 50 46 À59 À9 4.25 Fusiform gyrus (BA 36) 12 34 À7 À32 4.40 Parahippocampal gyrus (BA 28/34) 20 À14 À14 À18 4.25

Frontal cortex Inferior frontal gyrus (BA 45/46) 35 40 17 29 4.37 Threshold, P b 0.0001. tual processing than assessments of a person’s gender or the more Baker et al., 2001; McDermott et al., 2003). In a recent study, basic-level categorization of objects (Gauthier et al., 1997; Mason Mitchell et al. (2002) reported activity in the fusiform gyrus when and Macrae, in press); thus, the precise functional significance of participants made abstract semantic judgments about unfamiliar the effects reported by Sergent et al. (1992) is unclear. Rectifying persons. What this suggests is that semantic judgments about other this ambiguity, the current results demonstrated that, when tasks people (both familiar and unfamiliar) recruit areas of visual are matched for perceptual difficulty, activity in the fusiform gyrus associative cortex that also support the perceptual aspects of face tracks with the specificity of semantic construal (i.e., identity N processing. That semantic processing is grounded in the reactiva- occupation). Thus, mirroring the effects of perceptual discrim- tion of other neural systems may explain why increased activity ination (Gauthier et al., 1997; Haxby et al., 2001), areas of ventral was also observed in the left parahippocampal gyrus during temporal cortex appear to be sensitive to the specificity of semantic identity judgments. This region receives significant inputs from the judgments (Thompson-Schill, 2003). fusiform and lingual cortices (Van Essen and Anderson, 1990) and It is interesting to note that subordinate-level semantic is believed to play a significant role in the person perception processing elicited greater activity in the left than right fusiform process. As Sergent et al. (1992, p. 30) noted, bthe para- gyrus (see Fig. 2). This finding is consistent with the demonstration hippocampal gyrus...plays a pivotal role by reactivating the that while the generation of object knowledge is associated with connections between afferent information and semantic, episodic increased activity in the fusiform gyrus bilaterally, activity is and emotional information...What makes this area particularly typically stronger in the left hemisphere than in the right crucial in face recognition comes from the fact that faces...can be hemisphere (Gorno-Tempini and Price, 2001; Gorno-Tempini et identified as representing unique individuals only if they reactivate al., 1998; Kan et al., 2003; Thompson-Schill, 2003; Thompson- a minimum of biographical information related to the individualQ. Schill et al., 1999; Zelkowicz et al., 1998). Thompson-Schill et al. In other words, accessing semantic knowledge about others (1999), for example, observed significant activity in the left involves the reactivation of perceptual, affective, and memorial fusiform gyrus during a property-verification task, but only when representations in an interacting network of neural regions (Haxby conceptual knowledge was assessed. This then supports the idea et al., 2001). that areas of visual association cortex are activated during semantic The functional imaging data obtained in this study are processing (Kan et al., 2003; Martin et al., 1995). To access consistent with the neuropsychological literature on semantic conceptual knowledge about other people, perceivers must activate processing (Damasio et al., 2004; Galton et al., 2001; Rogers the relevant person-specific perceptual representations (see Bar- and McClelland, 2003, Mummery et al., 2000). Galton et al. alou, 1999; Thompson-Schill, 2003). (2001), for example, examined patterns of temporal lobe atrophy in As well as supporting person-specific effects, the fusiform Semantic Dementia (SD) and related these findings to the cognitive gyrus may also be implicated in abstract semantic processing (see profile of SD patients. They found that in contrast with normal D.J. Turk et al. / NeuroImage 24 (2005) 1147–1153 1151

Fig. 2. Regions of the ventral temporal cortex (fusiform gyrus) showing significantly greater BOLD response in the identity N occupation contrast ( P b 0.0001). These BOLD differences are illustrated graphically for each of the four main clusters in the fusiform gyrus (A–D). controls, SD patients showed significant volumetric loss bilaterally and Hodges, 2000; Hodges et al., 1992). Recent neuropsycho- in the temporal poles, and left lateralized atrophy in the amygdala, logical and computational accounts of temporal lobe function parahippocampal gyrus, fusiform gyrus, and inferior and middle suggest that the temporal poles may be responsible for the amodal temporal gyri. Furthermore, performance on semantic association integration of information from modality-specific processing tasks correlated only with the size of the left fusiform gyrus. The regions in posterior temporal areas from which categorization current fMRI study also found significantly greater BOLD signal emerges (Barsalou et al., 2003; Damasio, 1989; see also Rogers in left fusiform and parahippocampal regions for identity judg- and McClelland, 2003). So, why did activity in this functionally ments, lending further support to the functional significance of important region not emerge in the current investigation? Our these regions in person construal. intuition is that characteristics of the current paradigm together Interestingly, increased activity was not observed in the with technical limitations of fMRI may account for the apparent temporal poles in the current investigation. This is perhaps lack of activity in the temporal poles. In the current task, surprising given the acknowledged importance of this region in participants were not required to generate verbal responses (either the processing and representation of semantic information (Garrard identity or occupational judgments) to the presented faces, but 1152 D.J. Turk et al. / NeuroImage 24 (2005) 1147–1153 rather to match the stimuli (i.e., face plus verbal item) at either the Devlin, J.T., Russell, R.P., Davis, M.H., Price, C.J., Wilson, J., Moss, H.E., occupational or identity-based level of semantic abstraction Matthews, P.M., Tyler, L.K., 2000. Susceptibility-induced loss of (Gauthier et al., 1997; Kan et al., 2003). As lesions in the temporal signal: comparing PET and fMRI on a semantic task. NeuroImage 11, poles typically impede naming or category generation (Damasio et 589–600. Friston, K.J., Holmes, A.P., Worsley, K.J., Poline, J.-P., Frith, C.D., al., 2004), it is perhaps not surprising that this region was not Frakowiak, R.S.J., 1995. Statistical parametric maps in functional differentially activated in a category-verification task (Kan et al., imaging: a general linear approach. Hum. Brain Mapp. 2, 189–210. 2003). Similarly, the amodal integration of different types of Galton, C.J., Patterson, K., Graham, K., Lambon-Ralph, M.A., Williams, information to support a semantic judgment (e.g., occupation or G., Antoun, N., Sahakian, B.J., Hodges, J.R., 2001. Differing patterns identity) may be maximized when participants are required to of temporal atrophy in Alzheimer’s disease and semantic dementia. generate a semantic response. As verbal information was provided Neurology 57, 216–225. to participants on each trial, this likely minimized the contribution Garrard, P., Hodges, J.R., 2000. Semantic dementia: clinical, radiological, played by the temporal poles in the current task. Instead, semantic and pathological perspectives. J. 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