Evidence for Integrated Visual Face and Body - IA de Investigación especializada en el MIT

Evidencia de integración visual del rostro y el cuerpo
Representations in the Anterior Temporal Lobes

Bronson B. Harry1,2, Katja Umla-Runge3, Andrew D. Lawrence3,
Kim S. Graham3, and Paul E. Downing2

Abstracto

■ Research on visual face perception has revealed a region in
the ventral anterior temporal lobes, often referred to as the
anterior temporal face patch (ATFP), which responds strongly
to images of faces. Hasta la fecha, the selectivity of the ATFP has been
examined by contrasting responses to faces against a small
selection of categories. Aquí, we assess the selectivity of the
ATFP in humans with a broad range of visual control stimuli
to provide a stronger test of face selectivity in this region. En
Experimento 1, participants viewed images from 20 stimulus
categories in an event-related fMRI design. Faces evoked more
activity than all other 19 categories in the left ATFP. In the right
ATFP, equally strong responses were observed for both faces
and headless bodies. To pursue this unexpected finding, en

Experimento 2, we used multivoxel pattern analysis to examine
whether the strong response to face and body stimuli reflects a
common coding of both classes or instead overlapping but dis-
tinct representations. On a voxel-by-voxel basis, face and whole-
body responses were significantly positively correlated in the
right ATFP, but face and body-part responses were not. Este
finding suggests that there is shared neural coding of faces and
whole bodies in the right ATFP that does not extend to individ-
ual body parts. A diferencia de, the same approach revealed distinct
face and body representations in the right fusiform gyrus. Estos
results are indicative of an increasing convergence of distinct
sources of person-related perceptual information proceeding
from the posterior to the anterior temporal cortex. ■

INTRODUCCIÓN

fMRI studies of humans, Old World monkeys (macaques),
and New World monkeys (marmosets) have uncovered
several face-selective regions in the occipital and tem-
poral lobes (Hung et al., 2015; Tsao & Livingstone, 2008;
Kanwisher & Yovel, 2006; Haxby, Hoffman, & Gobbini,
2000). Although cross-species homology has not yet been
clearly established, this network of face-selective regions
shows a strikingly similar organization across human and
nonhuman primates (Hung et al., 2015; McMahon, Russ,
Elnaiem, Kurnikova, & Leopold, 2015; Rajimehr, Joven,
& Tootell, 2009; Tsao, Moeller, & Freiwald, 2008) y
consists of several ventral regions spanning the occipital
corteza, inferior temporal lobes, and STS.

An influential theoretical perspective (Haxby et al.,
2000), based on human functional imaging studies, di-
vides face-selective regions into a “core” system compris-
ing extrastriate nodes for the visual analysis of faces and
an “extended” system incorporating additional neural
regions that work in concert with the core system to ex-
tract various types of social information from faces. El
core regions include the occipital face area (OFA; Pitcher,
Dilks, sajonia, Triantafyllou, & Kanwisher, 2011; Gauthier
et al., 2000), the fusiform face area (FFA; Weiner &

1Western Sydney University, 2Bangor University, 3Universidad de Cardiff

Grill-Spector, 2010; Kanwisher & Yovel, 2006), y el
posterior STS (Pitcher et al., 2011; Chip, alison, Bentín,
Sangre, & McCarthy, 1998). OFA and FFA are proposed to
process static facial form, with the OFA more engaged in
part-based processing and the FFA more engaged in pro-
cessing the configuration of individual parts (harris &
Aguirre, 2010; Schiltz, Dricot, Goebel, & rossión, 2010;
Liu, harris, & Kanwisher, 2009; Yovel & Kanwisher, 2005),
whereas posterior STS processes changeable aspects of
faces (p.ej., eye gaze; Haxby et al., 2000). A diferencia de, el
extended system is proposed to include regions such as
the amygdala and the anterior temporal cortex, areas
that are argued to be important in appraising emotional
facial expressions (Calder, lorenzo, & Joven, 2001; pero
see Mende-Siedlecki, Verosky, Turk-Browne, & todorov,
2013) and encoding person-specific semantic knowledge
(Quiroga, Kreiman, Koch, & Frito, 2008; Thompson et al.,
2004), respectivamente.

Recent evidence prompts consideration of whether
there are anterior temporal regions that should also be
included as a part of the core system (duchaína & Yovel,
2015; collins & Olson, 2014; Haxby & Gobbini, 2011). A
number of reports (Ku, Tolias, Logothetis, & Goense, 2011;
Nestor, Plaut, & behrman, 2011; Pinsk et al., 2009;
Rajimehr et al., 2009; Tsao et al., 2008) provide evidence
of at least one face-selective region in the anterior tempo-
ral lobes in humans and macaques. In macaques, electrical

© 2016 Instituto de Tecnología de Massachusetts. Published under a
Creative Commons Attribution 3.0 no portado (CC POR 3.0) licencia

Revista de neurociencia cognitiva 28:8, páginas. 1178–1193
doi:10.1162/jocn_a_00966

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stimulation of the anterior temporal face patches (ATFPs)
selectively induces activity in the posterior network
(Moeller, Freiwald, & Tsao, 2008), suggesting that these
areas are functionally connected. Además, the most
ATFP is unique in that neurons in this region respond
invariantly to different face views (Meyers, Borzello,
Freiwald, & Tsao, 2015; Freiwald & Tsao, 2010), sugerir-
ing that it forms higher-level representations that are
needed for identification (for similar evidence in human
Participantes, see Yang, Susilo, & duchaína, 2016; Anzellotti,
Fairhall, & Caramazza, 2014). In support of the view that
human ATFP captures a similarly abstract representation,
Nasr and Tootell (2012) found in human participants that
fMRI activity in the ATFP closely mirrored changes in rec-
ognition performance brought about by image manipula-
tions such as face inversion and contrast reversal.

A limitation of previous studies examining visual selec-
tivity in the human ATFP has been the use of relatively
few visual control categories (p.ej., Nasr & Tootell, 2012;
Rajimehr et al., 2009; Tsao et al., 2008). A wide assess-
ment of responses to items from a range of categories is
vital for determining the selectivity of a region’s response
profile (Desimone, Albright, Bruto, & bruce, 1984) y
thus for making inferences about its functional role(s).
There have been multiple-category surveys of the inferior
temporal cortex (Mur et al., 2012; Vul, Ejército, Hsieh,
Golland, & Kanwisher, 2012; Downing, chan, Peelen,
Dodds, & Kanwisher, 2006), but these studies only exam-
ined responses in posterior temporal regions.

This study attempts to resolve the aforementioned
limitations by measuring the profile of responses in the
functionally defined human ATFP to a wide range of visu-
ally presented stimulus categories. A further aim is to
compare this profile with that of more posterior face-
selective regions (OFA, FFA) in an effort to reveal how
categorical information (particularly about people)
emerges over the span of the temporal lobes.

To begin addressing these aims, in Experiment 1, nosotros
used a blocked-design functional localizer to first identify
the ATFP, as well as OFA and FFA, in individual partici-
pants. We used a simultaneous odd-one-out visual dis-
crimination (“oddity”) task as a localizer. This task was
selected on the grounds that it has been found to be
effective in previous fMRI research at selectively acti-
vating anterior temporal regions (Barense, Henson, Sotavento,
& graham, 2010; O’Neil, Cate, & Köhler, 2009; Sotavento,
Scahill, & graham, 2008) and that performance on this
paradigm is sensitive to selective lesions of anterior
temporal regions (es decir., perirhinal cortex [PrC]) in mon-
keys (Buckley, Booth, Rolls, & Gaffán, 2001) and humans
(Barense, Gaffán, & graham, 2007; Lee et al., 2005). En
the main experiment, the same participants were pre-
sented with images of items of 20 different kinds in an
event-related design, while they performed a 1-back task
to maintain attention to the stimuli. In this way, we were
able to assess in detail the selectivity profile of ATFP and
compare it with more posterior face-selective regions.

EXPERIMENT 1

Métodos

Participantes

Twenty healthy postgraduate volunteers (edad media =
25 años, range = 22–30 years; 13 women) were recruited
from Bangor University. All participants were screened
for MRI exclusion criteria and gave written informed
consent for participation in the experiment, which was
approved by the research ethics committee of the School
of Psychology at Bangor University, Reino Unido.

Materials

Stimuli for the localizer runs (oddity task) consistió en
96 grayscale images of faces, natural scenes, and com-
mon handheld objects (Cifra 1, arriba). Images were orga-
nized into 32 triplets for each category (Barense et al.,
2010). Each triplet was presented in a triangular forma-
ción, consisting of a pair of foil images and a target image,
en un 1200 × 840 pixel white background. The foil images
were two pictures of the same face, scene, or object taken
from different viewpoints. The target was another image
from the same category as the foil pair and was selected
to be highly similar in appearance to the other pictures.
Thirty-two triplets consisting of three black squares were
also constructed to appear as an active baseline condition.
One of the squares (the target) was slightly larger or smaller
than the other two shapes.

Stimuli for the main experimental runs (1-back task)
consistió en 48 color images from each of 20 diferente
categories (Cifra 1, abajo). Categories consisted of
birds, (headless) human bodies, carros, chairs, clothes,
crystals, faces, fish, flowers, fruit and vegetables, insects,
instruments, (nonhuman) mammals, prepared food, rep-
tiles, spiders, herramientas, armas, indoor scenes, and outdoor
escenas (Downing et al., 2006). These stimuli were selected
to capture a range of object category distinctions (es decir.,
animate vs. inanimate, large vs. pequeño, natural vs. manmade)
that modulate responses in the ventral temporal lobes
(Konkle & Oliva, 2012; Mahon & Caramazza, 2011). Estímulos
were centred on a white 400 × 400 pixel background,
except for scenes, which were cropped to completely fill
the image dimensions.

Procedimiento

To localize functional ROIs, participants completed four
runs of an oddity task, each comprising 21 blocks of 15 segundo.
Blocks 1, 6, 11, 16, y 21 were fixation-only rest con-
ditions. Each of the four stimulus blocks (faces, escenas,
objects, and shapes) was presented once between each
pair of rest blocks. Stimulation blocks consisted of three
oddity trials, cada 5 sec in duration. Participants indicated
the location of the target stimulus (the odd item out) por
pressing one of three buttons. Block order for each set of
stimulation conditions was randomly determined between

Harry et al.

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and Events 22–24 were assigned to fixation-only rest
condiciones. The full sequence was divided into eight
separate runs. For Runs 2–8, the final item from the pre-
ceding run was presented at the beginning of the next
run to reestablish sequence context (haciendo 145 events
per run). Participants were assigned to one of five counter-
balanced sequences. Stimulus and target events were pre-
sented for 300 msec followed by an ISI of 1200 msec that
consisted of a central fixation cross. For rest events, a fixa-
tion cross appeared for 1500 mseg. Fixation-only rest
bloques (duration = 16 segundo) were presented at the begin-
ning and end of each run.

Localizer runs were interspersed throughout the scan-
ning session so that participants completed one run of
the oddity task after every two runs of the main experi-
mental task.

Brain images were acquired with a Philips Achieva 3.0-T
scanner with a 32-channel head coil. BOLD contrast func-
tional images were collected with a T2*-weighted, gradient
EPI sequence (repetition time = 2000 mseg, echo time =
35 mseg, flip angle = 90°, campo de visión = 240 mm × 240 mm,
acquisition matrix = 96 × 96, in-plane resolution =
2.5 mm × 2.5 mm, slice thickness = 2.5 mm, no slice gap).
Volumes consisted of 28 slices angled −30° from the AC–
PC plane to maximize signal over the medial-temporal
lobes. Volumes were positioned to completely cover the
temporal and occipital lobes at the expense of the dorsal
parietal cortex. A high-resolution T1-weighted anatomical
image was also acquired for each participant (3-D magne-
tization prepared rapid gradient-echo sequence; 175 slices,
voxel size = 1 mm isotropic, campo de visión = 256 mm ×
256 mm, repetition time = 8.4 mseg, echo time = 3.8 mseg,
flip angle = 8°). Stimuli were displayed on a Cambridge
Research Systems BOLDScreen located behind the
scanner bore and were viewed via a mirror fixed to the
head coil. Presentation of the stimuli was controlled by
Psychtoolbox (Brainard, 1997) running on MATLAB (El
MathWorks, Natick, MAMÁ).

Cifra 1. Example of stimuli presented in Experiment 1: localizer task
(arriba) and main experimental task (abajo).

Image Preprocessing and Analysis

runs and counterbalanced within runs according to a Latin
square design.

For the main experimental runs, all images from each
del 20 stimulus categories were presented once in a
rapid, event-related design. Participants completed a
1-back task by pressing a button whenever a stimulus
was immediately repeated. Stimulus order was deter-
mined with a first-order counterbalanced, optimized,
norte = 24, Tipo 1, Index 1 secuencia (Aguirre, 2007). Este
procedure generated a sequence of 1153 events, incluir-
ing an initial event to establish sequence context. El
20 stimulus categories were assigned to Event types
1–20 for each participant. Event 21 was assigned to tar-
get events, whereby the previous item was repeated,

Functional MRI data were preprocessed with SPM8
(Wellcome Department of Imaging Neuroscience, Londres,
Reino Unido; www.fil.ion.ucl.ac.uk/spm/software/spm8/ ) and in-
cluded rigid body realignment, coregistration, tissue seg-
mentation, normalization to the Montreal Neurological
Instituto (MNI) 152 template with DARTEL (Ashburner,
2007) and spatial smoothing (6-mm FWHM Gaussian
kernel).

We localized face-selective regions for each individual
with data collected from the oddity task. Estimates of the
BOLD response in each voxel and category were derived
by entering the boxcar function of stimulation that was
convolved with the canonical hemodynamic response
into a fixed effects general linear model. Face selectivity
in each voxel was calculated by contrasting activity evoked
by faces against the average of scenes and objects.

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Revista de neurociencia cognitiva

Volumen 28, Número 8

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Mesa 1. Mean MNI Peak Coordinates for Each ROI

Mean MNI Coordinates

42.4

−41.1

43.3

−41.9

37.3

−38.5

−78.0

−80.9

−49.6

−51.6

−14.0

−13.2

−6.1

−7.63

−20.5

−21.0

−38.5

−33.0

6.7

6.4

3.8

3.1

5.0

4.3

Dakota del Sur

6.1

4.8

5.7

4.9

2.5

5.8

4.2

5.9

4.2

2.7

7.6

7.4

rOFA

lOFA

rFFA

lFFA

rAFP

lAFP

Face-selective ROIs were localized by finding the most
face-selective voxel within expected regions of cortex
(OFA, inferior or mid-occipital gyrus; FFA, mid-fusiform
gyrus; ATFP, anterior occipito-temporal sulcus or anterior
collateral sulcus) near to typical MNI coordinates identi-
fied in previous studies ( Julian, Fedorenko, Webster, &
Kanwisher, 2012; right OFA [rOFA]: 44, −76, −12; izquierda
OFA [lOFA]: −40, −76, −18; right FFA [rFFA]: 38,
−42, −22; left FFA: −40, −52, −18; Axelrod & Yovel,
2013; right ATFP [rATFP]: 34, −10, −39; left ATFP:
−34, −11, −35). ROIs were defined by selecting all sig-
nificant ( pag < .001, uncorrected), contiguous voxels cen- tered around the peak voxel closest to the coordinates provided above. For analyses of response profiles in the main experiment, ROI size was limited to 50 voxels be- cause previous studies have shown that regions larger than this do not fully capture category selectivity (Mur et al., 2012). Estimates of the response to each of the 20 categories presented in the 1-back task were modeled separately as instantaneous neural events (i.e., duration = 0 msec) convolved with the canonical hemodynamic response. An additional nuisance regressor of no interest was included to model responses to the initial trial and to all target trials. The values of the beta estimates for each category were averaged over all voxels included in each ROI. Results Behavioral Performance Average performance on the oddity task was 81% (SEM = 3%) correct for faces, 83% (SEM = 3%) correct for scenes, 82% (SEM = 3%) correct for objects, and 76% (SEM = 3%) correct for shapes. One-way ANOVA showed no sig- nificant differences in performance between categories of stimuli ( p > .2). Average performance on the 1-back task
era 83% (5%).

ized in 15 de 20 Participantes. Localizing the ATFP is
problematic because of signal loss in the anterior tem-
poral lobes (because of proximity to air-filled spaces such
as the ear canal and the sinus cavity), and finding this
region in 60–75% of participants is consistent with pre-
vious studies that used a single-session protocol (Axelrod
& Yovel, 2013; Rajimehr et al., 2009). FFA was localized
bilaterally in all 20 Participantes, whereas lOFA or rOFA
was localized in 19 de 20 Participantes (bilaterally in 18 par-
ticipants). Significar (±SD) peak coordinates for each ROI
are presented in Table 1. Individual MNI coordinates
for the peak voxels within the ATFP are provided in
Mesa 2. Cifra 2 illustrates the location of the ATFP in
four representative participants.

Event-related Response Profiles

Del 20 stimulus categories tested in the main experi-
mento, faces evoked the maximal response in all of the
independently defined functional ROIs (OFA, Cifra 3;
FFA, Cifra 4; ATFP, Cifra 5). In line with previous fMRI
studies of category selectivity (Mur et al., 2012; Downing

Mesa 2. Individual MNI Peak Coordinates for the Left and
Right AFP

Left AFP

Right AFP

Participant

−40

−12.5 −35

−37.5 −20

−22.5

−

−17.5 −22.5

−15

−37.5

−17.5 −22.5

−12.5 −30

32.5 −12.5 −42.5

−22.5 −32.5

−7.5 −32.5

−37.5 −17.5 −25

−12.5 −45

−

−15

−

−37.5

−42.5

−10

−32.5

37.5 −12.5 −50

−17.5 −32.5

37.5 −10

−40

−5

−47.5

37.5 −12.5 −42.5

−37.5 −12.5 −32.5

−37.5

−7.5 −42.5

−

−12.5 −40

−

27.5 −12.5 −40

−42.5 −17.5 −22.5

−5

−35

−

−12.5 −40

−17.5 −27.5

37.5 −17.5 −42.5

−

−40

−30

−35

−45

−35

−40

−35

−45

−40

Definition of ROIs

The right-hemisphere ATFP was localized in 13 de 20 par-
ticipants, and the left anterior face patch (AFP) was local-

Significar

Dakota del Sur

−38.5 −13.2 −33.0

37.3 −14.0 −38.5

4.3

5.8

7.4

2.5

7.6

Harry et al.

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Cifra 2. Location of the AFP
(rojo; faces > scenes + objects,
pag < .001). D o w n l o a d e d f r o m l l / / / / j f / t t i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e d o u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 8 / 8 2 8 1 / 1 8 7 / 8 1 1 1 9 7 5 8 1 / 6 1 3 7 3 8 o 5 c 2 n 0 _ 8 a / _ j 0 o 0 c 9 n 6 6 _ a p _ d 0 0 b 9 y 6 g 6 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j / t . f u s e r o n 1 7 M a y 2 0 2 1 et al., 2006), we expected significantly stronger responses to faces compared with all other categories. To test the selectivity of each ROI, we compared the response to faces against the response to the next most effective stimulus category in a 3 × 2 × 2 repeated-measures ANOVA with ROI, Hemisphere, and Stimulus category as factors. This analysis revealed a significant three-way interaction be- tween ROI, Hemisphere, and Stimulus category (F(2, 18) = 4.5, p < .05), a significant two-way interaction between ROI and Stimulus category (F(2, 18) = 6.03, p < .01), and significant main effects for ROI (F(2, 18) = 29.83, p < .001), Hemisphere (F(1, 9) = 11.70, p < .01), and Stimulus category (F(1, 9) = 8.19, p < .05). To interpret the three-way interaction, we carried out two separate 3 × 2 repeated-measures ANOVAs for each hemisphere with ROI (OFA, FFA, ATFP) and Stimulus category (face vs. next best category) as factors. For the left hemisphere, this analysis found only significant main effects of ROI (F(2, 26) = 22.75, p < .001, Bonferroni- corrected) and Stimulus category (F(1, 13) = 9.34, p < .05, Bonferroni-corrected). Analysis of the right hemi- sphere revealed a significant interaction between ROI and Stimulus category (F(2, 24) = 8.1, p < .01, Bonferroni- corrected) and significant main effects of ROI (F(2, 24) = 17.97, p < .001, Bonferroni-corrected) and Stimulus cate- gory (F(1, 12) = 10.18, p < .01, Bonferroni-corrected). Simple effects analysis revealed a main effect of stimulus category in the rOFA (F(1, 18) = 7.2, p < .05, Bonferroni- corrected) and FFA (F(1, 19) = 7.2, p < .05, Bonferroni- corrected), but not in the rATFP (F(1, 13) = 0.15). Post hoc tests revealed that the response averaged across faces and bodies in the rATFP was significantly higher than the next most effective category (spiders; F(1, 13) = 9.3, p < .01). A final analysis examined whether the structure of nonpreferred responses in the ATFP was similar to that found in the posterior face-selective regions. It is a known property of the latter regions that they generally respond more strongly to animate compared with inani- mate categories of stimuli (Wiggett, Pritchard, & Downing, 2009; Downing et al., 2006). Thus, for each ROI, we com- pared the response averaged over all nonhuman ani- mate categories (birds, fish, insects, mammals, reptiles) against the response averaged over all inanimate categories (cars, chairs, clothes, crystals, flowers, fruit and vegetables, instruments, prepared foods, tools, weapons). To test the preference of each ROI, we compared the response to animates and inanimates in a 3 × 2 × 2 repeated-measures ANOVA with ROI, Hemisphere, and Animacy (animate, inanimate) as factors. This analysis revealed significant two- way interactions between ROI and Animacy (F(2, 18) = 18.7, p < .001) and between Hemisphere and Category (F(2, 9) = 10.4, p = .01) and significant main effects for ROI (F(2, 18) = 29.01, p < .001), Hemisphere (F(1, 9) = 6.8, p < .05), and Animacy (animates > inanimates, F(1, 9) =
32.6, pag < .001). To interpret the interaction effects, we carried out three separate 2 × 2 repeated-measures ANOVAs for each ROI (OFA, FFA, ATFP) with Hemisphere and Animacy as fac- tors. This analysis found only significant main effects for Animacy in the FFA (F(1, 19) = 59.13, p < .001, Bonferroni- corrected) and ATFP (F(1, 10) = 8.7, p < .05, Bonferroni- corrected). In contrast, analysis of the OFA revealed a significant main effect of Hemisphere (F(1, 18) = 11.84, 1182 Journal of Cognitive Neuroscience Volume 28, Number 8 p < .01, Bonferroni-corrected) and Animacy (F(1, 18) = 62.3, p < .001, Bonferroni-corrected) and a two-way inter- action between Hemisphere and Animacy (F(1, 18) = 11.13, p < .01, Bonferroni-corrected). Simple effects anal- ysis revealed a significant effect of Animacy in both the rOFA (F(1, 18) = 54.6, p < .001, Bonferroni-corrected) and lOFA (F(1, 18) = 40.0, p < .001, Bonferroni-corrected). Discussion Face-selective regions across the temporal lobes showed a similar profile of activity, across a wide range of stimu- lus kinds, consistent with a model in which these regions cooperate functionally (Moeller et al., 2008). There was variation, however, in the pattern of responses across the regions tested; in particular, the right-hemisphere ATFP showed a strong response to both faces and bodies (statis- tically different from the response to spiders, the next most effective category) despite the fact that the ROI was local- ized independently on the basis of a contrast of faces versus scenes and objects. This pattern of colocalized sig- nificant response to bodies and faces, along with a weaker response to other kinds of objects, is highly similar to that found in the right fusiform gyrus. In that region, strong fMRI responses to faces and bodies have been found to overlap closely ( Weiner & Grill-Spector, 2010; Peelen & Downing, 2005). One account of this general finding is that it reflects the common co-occurrence of faces and bodies in the visual input and the need to jointly process the socially relevant information they provide (Peelen & Downing, 2007). Several studies have attempted to determine whether the fusiform face- and body-selective responses reflect a sin- gle neural system or rather two distinct ones. Schwarzlose, Baker, and Kanwisher (2005) used high-resolution imag- ing to show that, in many participants, alongside “shared” voxels that respond to both categories, it is possible to identify distinct, but adjacent, highly selective patches for faces and bodies, referring to these accordingly as FFA and “fusiform body area” (FBA). Another approach Figure 3. Selectivity profile for rFFA (top) and lFFA (bottom). Error bars show the SEM. D o w n l o a d e d f r o m l l / / / / j f / t t i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e d o u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 8 / 8 2 8 1 / 1 8 7 / 8 1 1 1 9 7 5 8 1 / 6 1 3 7 3 8 o 5 c 2 n 0 _ 8 a / _ j 0 o 0 c 9 n 6 6 _ a p _ d 0 0 b 9 y 6 g 6 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j f t . / u s e r o n 1 7 M a y 2 0 2 1 Harry et al. 1183 Figure 4. Selectivity profile for rOFA (top) and lOFA (bottom). Error bars show the SEM. D o w n l o a d e d f r o m l l / / / / j t t f / i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e d o u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 8 / 8 2 8 1 / 1 8 7 / 8 1 1 1 9 7 5 8 1 / 6 1 3 7 3 8 o 5 c 2 n 0 _ 8 a / _ j 0 o 0 c 9 n 6 6 _ a p _ d 0 0 b 9 y 6 g 6 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j f . t / u s e r o n 1 7 M a y 2 0 2 1 tests for distinct neural systems at the pattern level, with- out requiring that they be identified in a binary fashion with separate sets of voxels. The logic of this method is that overlapping voxels (at whatever resolution) need not reflect shared neural processes—an assumption com- monly made in fMRI research (Peelen & Downing, 2007). For example, in a region where there are overlapping but functionally distinct face and body representations, local patterns of selectivity to these two categories should be uncorrelated (or negatively correlated). That is, con- sidered across a set of voxels, variability in the selectivity for bodies would not be expected to relate systematically to variability in the selectivity for faces. In contrast, where there are two overlapping and integrated representa- tions, the variability in selectivity to these two categories would be expected to be related across voxels: Strong selectivity to one category should tend to predict strong selectivity to the other. This would result in a positive correlation between the local patterns of selectivity evoked by each category. Studies taking this approach have found evidence for independent fusiform face- and body-selective representations (Kim, Lee, Erlendsdottir, & McCarthy, 2014; Weiner & Grill-Spector, 2010; Peelen, Wiggett, & Downing, 2006; see also Downing, Wiggett, & Peelen, 2007). Thus, motivated by these previous findings in the extra- striate cortex and by our present results in the right- hemisphere ATFP, in Experiment 2, we used the multivoxel approach described above to examine whether faces and bodies recruit overlapping or segregated representations in the anterior temporal lobes and in the fusiform gyrus. EXPERIMENT 2 We localized face- and body-selective regions (FFA, ATFP, FBA) in the right-hemisphere mid-fusiform and anterior temporal regions with a blocked 1-back design. Only responses in the right hemisphere were examined, as the rATFP demonstrated strong responses to both faces and bodies in Experiment 1. We opted for a block 1184 Journal of Cognitive Neuroscience Volume 28, Number 8 design 1-back localizer task in Experiment 2 because a pilot oddity task including headless bodies was too dif- ficult for participants to complete. Then, we assessed the functional responses in each ROI to six different conditions (faces, whole bodies, body parts, mammals, foods, and tools) with an event-related design. These categories were selected to assess responses across animate, natural, and manmade objects. Moreover, body parts were included to examine the breadth of body responses in ATFP. We improved the experimental pro- tocol with a coronal slice orientation, which, compared with axial orientation (as used in Experiment 1), has been shown to maximize signal over the anterior temporal lobes (Axelrod & Yovel, 2013), and we obtained higher- resolution images (voxel size = 2 mm3) to mitigate partial voluming effects. Bangor University. All participants were screened for MRI exclusion criteria, after which they gave written informed consent for participation in the experiment, which was approved by the research ethics committee of the School of Psychology at Bangor University, United Kingdom. Materials Stimuli for the localizer consisted of 40 images each of faces, bodies, and chairs (Downing et al., 2006). Stimuli for the main experiment consisted of 24 images each of faces, (headless) bodies, body parts, (nonhuman) mam- mals, food, and tools (Figure 6). All images were pre- pared in a similar manner as Experiment 1. None of the images presented in the localizer task appeared in the event-related runs. Methods Procedure Ten healthy postgraduate volunteers (mean age = 25 years, range = 24–29 years; six women) were recruited from Participants completed five runs of a 1-back localizer task, each consisting of 25 blocks. Blocks 1, 5, 9, 13, 17, 21, Figure 5. Selectivity profile for right (top) and left (bottom) AFP. Error bars show the SEM. D o w n l o a d e d f r o m l l / / / / j f / t t i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e d o u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 8 / 8 2 8 1 / 1 8 7 / 8 1 1 1 9 7 5 8 1 / 6 1 3 7 3 8 o 5 c 2 n 0 _ 8 a / _ j 0 o 0 c 9 n 6 6 _ a p _ d 0 0 b 9 y 6 g 6 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j / t f . u s e r o n 1 7 M a y 2 0 2 1 Harry et al. 1185 Figure 6. Example stimuli presented in Experiment 2. D o w n l o a d e d f r o m l l / / / / j t t f / i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e d o u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 8 / 8 2 8 1 / 1 8 7 / 8 1 1 1 9 7 5 8 1 / 6 1 3 7 3 8 o 5 c 2 n 0 _ 8 a / _ j 0 o 0 c 9 n 6 6 _ a p _ d 0 0 b 9 y 6 g 6 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j / t . f u s e r o n 1 7 M a y 2 0 2 1 and 25 were fixation-only rest conditions lasting 10 sec in duration. Each of the three stimulus blocks (faces, bod- ies, and chairs) was presented once between each pair of rest blocks. Stimulation blocks were composed of 15 stimulus exemplars drawn from a pool of 40 images. Stimuli were presented sequentially and appeared for 300 msec followed by a 700-msec ISI. Repetitions oc- curred twice per block. For the main experimental runs, all images from each of the six stimulus categories were presented in an event-related design; each stimulus was presented once per run. Participants performed a 1-back task. Stimulus order for each run was determined with a first-order counterbalanced, optimized, n = 8, Type 1, Index 1 se- quence. This procedure generated a sequence of 193 events of eight types, including an initial event to estab- lish sequence context. Event types 1–6 were assigned to the six stimulus classes, Event type 7 was assigned to target events (1-back, stimulus repetition), and Event type 8 was assigned to fixation-only rest condition. Par- ticipants completed three runs of the main experiment, resulting in 72 trials per condition. Stimulus and target events were presented for 300 msec followed by an ISI of 1700 msec consisting of a fixation cross. For rest events, a fixation cross was presented for 2000 msec. Fixation blocks (duration = 16 sec) were presented at the beginning and end of each run. Runs of the localizer and the main experiment were completed in an alter- nating sequence. Brain images were acquired with a Philips Achieva 3.0-T scanner with a 32-channel head coil. BOLD contrast functional images were collected with a T2*-weighted, gradient EPI sequence (repetition time = 2500 msec, echo time = 35 msec, flip angle = 90°, field of view = 240 mm × 240 mm, acquisition matrix = 120 × 120, in-plane resolution = 2 mm × 2 mm, slice thickness = 2 mm, no slice gap). Volumes were composed of 28 slices in coronal orientation that were split into two sep- arate stacks of 14 slices to cover the anterior temporal lobes and the mid-fusiform gyrus (Axelrod & Yovel, 2013). A high-resolution T1-weighted anatomical image was also acquired for each participant. All other as- pects of the experimental setup were the same as in Experiment 1. Image Preprocessing and Analysis Preprocessing was similar to Experiment 1 except that, to better preserve the local spatial patterns of brain activity, images were not normalized and spatial smoothing was performed with a 3-mm Gaussian kernel. Responses to each category in the blocked localizer and event-related runs were derived in a similar manner as Experiment 1. To evaluate mean univariate response profiles in selective regions similar to Experiment 1, the FFA, ATFP, and FBA were defined from the localizer by contrasting each cate- gory against chairs (faces > chairs, bodies > chairs; pag < .001, uncorrected). For the pattern analysis, localizer data were further used to identify two functional ROIs in both the mid- fusiform and anterior temporal lobes for the purpose of pattern analyses. First, a broad “human form” selective ROI was defined as the union of all face- and body- selective voxels within the mid-fusiform and collateral sulcus. This combined ROI was examined to ensure that the results of the pattern analysis would not be biased toward one stimulus category owing to unbalanced voxel selection. Second, we examined only the face-selective voxels corresponding to the ATFP. For this analysis, we defined the ATFP, as in Experiment 1, as the 30 most face-selective voxels (i.e., faces > bodies + chairs, pag < .001) that were contiguous with the peak voxel residing within the collateral sulcus. For both ROIs, an indepen- dent measure of face, whole body, and body part selec- tivity was calculated from the main experiment data by contrasting the response of each of these conditions against the average of mammals, food, and tools. The resultant t values were extracted for all voxels residing within each ROI. For each participant and ROI, the ex- tracted pattern of t values quantifying face selectivity was correlated with the corresponding pattern of t values for whole bodies and body parts. 1186 Journal of Cognitive Neuroscience Volume 28, Number 8 Results Behavioral Performance Average performance in the localizer scans was 75.1% (SEM = 4.2%), and average performance in the main ex- periment was 78.8% (SEM = 7.5%). Univariate Analysis of Main Experiment The three independently defined ROIs (rFFA, right FBA, and rATFP) all showed maximal responses to the ex- pected categories: faces in FFA and ATFP and whole bodies in FBA (Figure 7). To examine whether the pre- ferred category evoked significantly more activity than all other stimuli, the responses to the preferred category and the next most effective category were entered into a 3 × 2 repeated-measures ANOVA with ROI (FFA, FBA, ATFP) and Stimulus category (preferred category vs. next most effective) as factors. For the FFA and the ATFP, re- sponses to faces were compared against mammals and bodies, respectively, whereas in the FBA, bodies were Figure 7. Profile of responses to six stimulus categories, for the rFFA (top), right FBA (center), and right AFP (bottom). Error bars show the SEM. D o w n l o a d e d f r o m l l / / / / j f / t t i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e d o u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 8 / 8 2 8 1 / 1 8 7 / 8 1 1 1 9 7 5 8 1 / 6 1 3 7 3 8 o 5 c 2 n 0 _ 8 a / _ j 0 o 0 c 9 n 6 6 _ a p _ d 0 0 b 9 y 6 g 6 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j / t f . u s e r o n 1 7 M a y 2 0 2 1 Harry et al. 1187 D o w n l o a d e d f r o m l l / / / / j f / t t i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e d o u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 8 / 8 2 8 1 / 1 8 7 / 8 1 1 1 9 7 5 8 1 / 6 1 3 7 3 8 o 5 c 2 n 0 _ 8 a / _ j 0 o 0 c 9 n 6 6 _ a p _ d 0 0 b 9 y 6 g 6 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j t / . f u s e r o n 1 7 M a y 2 0 2 1 Figure 8. Correlation between face and body selectivity (left) and face and body-part selectivity (right) for the union of all independently defined face- and body-selective voxels in the mid-fusiform and anterior temporal lobes. Correlations shown were obtained for a range of thresholds for defining face- and body-selective voxels (x axis). These findings suggest that, in the anterior temporal cortex, face and body representations are integrated, in contrast to the fusiform gyrus, where they appear to remain distinct. Error bars show the SEM. compared with faces. This analysis revealed only a signif- icant main effect of Stimulus category (F(1, 7) = 53.69, p < .001), indicating that faces and whole bodies evoked more activity than the next most effective stimulus cate- gory in face- and body-selective regions, respectively. Pattern Analysis We performed a pattern analysis to examine the relation- ship of face- and body-selective populations in both the right-hemisphere mid-fusiform and anterior temporal lobes. In the first analysis, we examined all voxels that showed a preference for either faces or bodies by taking the union of the face- and body-selective regions as de- fined by the localizer runs. These regions were defined for a range of thresholds (tROI > 3.5, 3, 2.5, 2) to ensure
that the results of the pattern analysis were not depen-
dent on how stringently voxels were selected. Separately
for the mid-fusiform and anterior temporal ROIs, voxel-
wise patterns of selectivity in the main experiment for
faces and for whole human bodies (faces > mammals +
alimento + herramientas, bodies > mammals + alimento + herramientas) eran
correlated for each participant. Single-sample t tests
comparing the Fisher-transformed correlations in each
región (Cifra 8, izquierda) showed that face and body selec-
tivity were negatively correlated in the mid-fusiform ROI
(significantly at thresholds tROI > 3, pag = .0321; TROI > 2,
pag < .01, Bonferroni-corrected). This suggests, consistent with previous findings (Kim et al., 2014; Peelen et al., 2006), that face and body representations remain distinct in the fusiform gyrus—faces and bodies elicit distinct patterns of local activity. In contrast, significantly positive correlations between face and body selectivity were observed in the anterior temporal lobes (rs range = 0.24–0.36, all ps at different selection thresholds < .05, Bonferroni-corrected), suggestive of a shared face and body representation. In principle, however, the positive pattern correlation between face and body selectivity in the anterior tem- poral lobe could be because of general factors affecting responses in this region (i.e., signal limitations, use of common baseline), rather than a specific property of face and body representations. To exclude this possibility, we performed the same analysis comparing patterns of face and body-part selectivity (Figure 8, right). This analysis revealed that face and body-part selectivity were nega- tively correlated in the mid-fusiform region (all ps < .01, Bonferroni-corrected). Critically, face and body-part selectivity were not significantly correlated in the anterior temporal lobes (all ts < 1). Furthermore, in the anterior temporal region, the correlation between face and whole- body selectivity was significantly greater than the correla- tion between face and body-part selectivity ( ps < .05, Bonferroni-corrected). Therefore, it is not simply the case that response patterns evoked by any and all visual stimuli are positively correlated within the anterior tem- poral region. Although examining the union of all face- and body- selective voxels is an unbiased method of voxel selection, this approach is less directly comparable with previous studies of the human ATFP. Therefore, in a second pat- tern analysis, we limited the analyzed region to only the 30 most face-selective voxels (selected from the localizer data) located within the collateral sulcus. Overall, the cor- relations observed for the ATFP were similar to those found in the first pattern analysis. The mean correlation between face and whole-body selectivity was significantly above zero (rmean = .38, t = 4.73, p < .01, Bonferroni- corrected), whereas the mean correlation between faces and body parts was not (rmean = .08, t = 0.86). Moreover, 1188 Journal of Cognitive Neuroscience Volume 28, Number 8 the correlation between face and whole-body selectivity was greater than between faces and body parts (t = 8.05, p < .0001, Bonferroni-corrected), indicating that the spatial organization of face- and body-selective re- sponses observed for the combined ROI is also present in the ATFP. Discussion In Experiment 2, we did not replicate the finding of Experiment 1 that faces and headless bodies drive activity in the rATFP equally well, perhaps because, in this study, we scanned at a higher spatial resolution and optimized slice selection, resulting in more precise localization of highly face-selective voxels in the anterior temporal cor- tex. The multivoxel pattern analysis showed, however, that, in this region, there is a significant positive correla- tion between the activity patterns evoked by face and whole-body stimuli. Importantly, this was significantly greater than the relationship between faces and indi- vidual body parts, which was not greater than chance, indicating that the positive correlation was not because of general properties of the responses observed in this region. Furthermore, these results were distinct from the findings from the mid-fusiform region, where patterns of face and body selectivity were negatively correlated, indicating independent (or at least less integrated) encod- ing of these stimuli. Taken together, these results suggest that part of the right anterior temporal lobe, in contrast to the posterior fusiform gyrus, encodes an integrated rep- resentation of the visual appearance of the face and of the whole body—a representation that does not extend to isolated body parts. GENERAL DISCUSSION Our study aimed to survey the response profile of the ATFP (Rajimehr et al., 2009) to different categories of stimuli. The profile of ATFP across multiple categories appeared, to a large extent, to mirror that of the FFA and OFA. A notable finding from Experiment 1 was the high response to human bodies (without faces) in the independently defined right-hemisphere ATFP ROI. In previous work on human anterior temporal face repre- sentations, Tsao and colleagues (2008) found a reliable response to bodies that was nonetheless significantly lower than the response to faces. These findings of colo- cated strong responses to faces and bodies are another potentially important similarity between the visual rep- resentations of the anterior and posterior inferior tem- poral lobes, where face- and body-evoked activations are intertwined in the fusiform gyrus (Kim et al., 2014; Weiner & Grill-Spector, 2010; Peelen & Downing, 2005; Schwarzlose et al., 2005). Therefore, in Experiment 2, we used a pattern analysis to examine the spatial organization of face and whole- body representations in the anterior and posterior tem- poral regions. In the fusiform gyrus, the pattern of face and body selectivity was negatively correlated, consistent with functionally distinct representations. In contrast, in the anterior temporal region, the correlation between patterns evoked by bodies and faces was positive, sug- gestive of integrated, whole-person representations. This interpretation is supported by the finding that face- evoked activity in the rATFP was more similar to whole bodies compared with isolated body parts, suggesting that anterior person-selective responses are primarily driven by whole-person form information. These findings, considered alongside a recent study showing super-additive responses to combined face and whole-body stimuli in the ATFP of the macaque (Fisher & Freiwald, 2015), provide support for the existence of integrated whole-agent processing regions in the primate visual system. In principle, integrated processing is not required to form whole-agent representations (Afraz, 2015), as whole-agent information is also present in the distributed response across face- and body-selective re- gions in the mid-fusiform. Evidence of super-additive re- sponses and joint selectivity, however, confirms that integrated representations are indeed formed in the ante- rior temporal cortex and likely play a key role in visual recognition (Lehky & Tanaka, 2016). More specifically, these and other findings provide some evidence for a hierarchically organized chain of face- and body-form representations along the length of the ventral occipito-temporal cortex (Grill-Spector & Weiner, 2014; Taylor & Downing, 2011; Minnebusch & Daum, 2009) that become progressively more integrated with each other (Figure 9). In the posterior occipito- temporal cortex, face and body representations (OFA, EBA) are anatomically distinct (Pitcher et al., 2009) and appear to emphasize the representation of component parts (e.g., Schiltz et al., 2010; Liu et al., 2009; Taylor, Wiggett, & Downing, 2007). More anteriorly, the FFA and FBA encode more holistic properties of their pre- ferred stimuli (Brandman & Yovel, 2016; Harris & Aguirre, 2010; Schiltz et al., 2010; Liu et al., 2009; Taylor et al., 2007; Schiltz & Rossion, 2006). A question under active inves- tigation is whether, in the fusiform regions, there is fur- ther integrated processing across faces and bodies (Bernstein, Oron, Sadeh, & Yovel, 2014; Kaiser, Strnad, Seidl, Kastner, & Peelen, 2014; Song, Luo, Li, Xu, & Liu, 2013; Schmalzl, Zopf, & Williams, 2012), with some evi- dence both for (e.g., Bernstein et al., 2014) and against (Fisher & Freiwald, 2015; Kaiser et al., 2014) this proposal. The current results argue for even closer integration of face and whole-body representation in the anterior-most reaches of the temporal cortex. On this view, the anterior temporal cortex can be seen as a core region in a broadly defined person-form processing pathway that combines domain-specific representations that are constructed in the extrastriate cortex. This perspective integrates the current findings with previous behavioral work and suggests how the very Harry et al. 1189 D o w n l o a d e d f r o m l l / / / / j t t f / i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e d o u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 8 / 8 2 8 1 / 1 8 7 / 8 1 1 1 9 7 5 8 1 / 6 1 3 7 3 8 o 5 c 2 n 0 _ 8 a / _ j 0 o 0 c 9 n 6 6 _ a p _ d 0 0 b 9 y 6 g 6 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j / . t f u s e r o n 1 7 M a y 2 0 2 1 Figure 9. Schematic illustration of a proposed hierarchical organization of human form information in the ventral temporal lobes. EBA and OFA, which are anatomically distinct, engage in domain-specific part-based processing of their preferred categories. FFA and FBA are closely overlapping and form domain-specific holistic representations of faces and bodies, respectively. Finally, face and body information is functionally integrated in the anterior temporal lobes (including the region typically identified as the AFP) to contribute to whole-person representation. Red clusters, faces > chairs,
pag < .001; green clusters, bodies > chairs, pag < .001. different kinds of visual cues provided by bodies and faces are brought together to represent what must be the real object of interest for social perception: whole people (Macrae, Quinn, Mason, & Quadflieg, 2005). Evi- dence from perceptual studies shows that judgments of identity, emotion, and gender from faces can be strongly influenced by the state of the body (Rice, Phillips, Natu, An, & O’Toole, 2013; Aviezer, Trope, & Todorov, 2012). Moreover, adaptation to pictures of bodies presented in isolation can alter the perception of subsequently viewed faces (Ghuman, McDaniel, & Martin, 2010), suggesting that faces and bodies share processing mechanisms. A very important open question is to what extent the pathways described here contribute to the integrated processing of different facial and bodily cues. Notably, extensive work by de Gelder and colleagues (2006) points to other, largely subcortical routes that are involved in rapidly extracting and integrating emotional information from faces and bodies (Meeren, van Heijnsbergen, & de Gelder, 2005). Here, we propose that the ATFP forms part of a ventral temporal pathway involved in person perception and identification that integrates static form cues from across the face and the body. Evidence of joint face and whole-body selectivity in the ATFP also aligns with paired object associative responses found in the PrC (Fujimichi et al., 2010). PrC is an ante- rior temporal region located in the anterior collateral sulcus (rhinal sulcus; Suzuki & Naya, 2014), which has been argued to occupy the highest point in the ventral visual processing pathway (Murray, Bussey, & Saksida, 2007). PrC receives dense input from visual area TE (Suzuki & Naya, 2014) and also receives input from regions of STS containing body-form selective neurons (Suzuki & Naya, 2014; Oram & Perrett, 1994). In the macaque, paired-associate learning involving two objects alters the selectivity of neurons in the PrC such that neu- rons that are selective for a particular object also become selective for a paired associate when repeatedly pre- sented together (Fujimichi et al., 2010). Similar to the present findings, these “unitized” representations also show a hierarchical organization, with a gradient of in- creasingly overlapping responses found spanning area TE and regions within the PrC (areas A36 and A35; Hirabayashi et al., 2014; Fujimichi et al., 2010). Given that the ATFP was primarily observed in the anterior collateral sulcus, it is possible that the current evidence of integrated face and body processing reflects a specific instance of a general PrC process that forms unitized representations of highly relevant objects via paired-associate coding mechanisms. Such a hierarchical scheme accords with proposals arising from the literature on memory and conceptual representations. These hold that the PrC is involved in forming complex, conjunctive object representations by combining feature-based representations derived in the extrastriate cortex (Clarke & Tyler, 2014; Graham, Barense, & Lee, 2010; Barense et al., 2005, 2007; Bartko, Winters, Cowell, Saksida, & Bussey, 2007; Buckley & Gaffan, 2006; Lee et al., 2005). According to this account, functionally integrating domain-specific representations serve to help buffer the perceptual system against inter- ference, which has a larger impact on the simpler, part- based representations found in the extrastriate cortex ( Watson & Lee, 2013; Barense et al., 2012; Bartko, 1190 Journal of Cognitive Neuroscience Volume 28, Number 8 D o w n l o a d e d f r o m l l / / / / j t t f / i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e d o u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 8 / 8 2 8 1 / 1 8 7 / 8 1 1 1 9 7 5 8 1 / 6 1 3 7 3 8 o 5 c 2 n 0 _ 8 a / _ j 0 o 0 c 9 n 6 6 _ a p _ d 0 0 b 9 y 6 g 6 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j f / t . u s e r o n 1 7 M a y 2 0 2 1 Cowell, Winters, Bussey, & Saksida, 2010; Fujimichi et al., 2010). To summarize, this study demonstrated that the ATFP shares several similarities with other face-selective re- gions found in the extrastriate cortex. Differences were evident, however, in the way that face- and body-selective responses are organized in these distributed temporal lobe brain areas, with evidence for integrated selectivity of faces and whole bodies found only in the ATFP. In that sense, the present findings are consistent with models that posit a posterior-to-anterior gradient in perceptual representations of objects that is built on increasingly complex combinations of features (Lehky & Tanaka, 2016). Acknowledgments We thank Morgan Barense for providing the stimuli used in the oddity task and Richard Ramsey for comments on an earlier draft. This work was supported by the Biotechnology and Biological Sciences Research Council grant BB/1007091/1. Reprint requests should be sent to Bronson B. Harry or Paul E. 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