Brain (2001), 124, 2087–2097
Identification of famous faces and buildings
A functional neuroimaging study of semantically unique
items
M. L. Gorno-Tempini and C. J. Price
Wellcome Department of Cognitive Neurology, Institute of
Neurology, London, UK
Correspondence to: M. L. Gorno-Tempini, UCSF
Department of Neurology, 350 Parnassus Avenue,
Suite 800, Box 1207, San Francisco, CA 94143, USA
E-mail: marilu@itsa.ucsf.edu
Summary
Several functional imaging experiments have clearly
established that the fusiform gyri are preferentially
responsive to faces, whereas the parahippocampal/lingual
gyri are more responsive to buildings. Other studies have
demonstrated that famous faces additionally activate the
anterior temporal cortex relative to unfamiliar faces,
animals, tools, body parts and maps. One explanation for
this apparent specialization for known people could be
that famous faces are ‘semantically unique items’. In
other words, they carry unique semantic associations that
are not shared by other perceptually similar category
members. If this hypothesis is correct, the anterior
temporal cortex should also respond to other semantically
unique items, such as famous buildings. In this PET study,
we investigated the effect of fame (famous relative to nonfamous) on activation elicited by famous and non-famous
faces and buildings during a same–different matching
task. We found that, when the task was held constant,
category-specific activations in the fusiform and
parahippocampal/lingual areas were not modulated by
fame. In contrast, in the left anterior middle temporal
gyrus there was an effect of fame that was common
to faces and buildings. These results suggest that the
identification of famous faces and buildings involves
category-specific perceptual processing in the fusiform
and parahippocampal/lingual regions, respectively, and
shared analysis of unique semantic attributes in the left
anterior temporal cortex.
Keywords: faces; buildings; semantic and lexical processes; PET; fusiform
Abbreviations: BA ⫽ Brodmann area; FB ⫽ famous buildings; FF ⫽ famous faces; FFA ⫽ fusiform face area; MTG ⫽
middle temporal gyrus; NFB ⫽ non-famous buildings; NFF ⫽ non-famous faces; RT ⫽ reaction time; SB ⫽ scrambled
buildings; SF ⫽ scrambled faces
Introduction
Psychological studies have suggested that the task of fully
identifying and naming a famous person is achieved by a
cascade of sequential processing stages (Bruce and Young,
1986). Neuropsychological evidence supports this view, and
patients have been described with impairments at various
stages of the identification process, including: (i) the presemantic stage, when recognition of famous faces is impaired
only in the visual domain, i.e. prosopagnosia; (ii) the semantic
stage, when loss of biographical information about known
people (person-specific semantics) occurs regardless of the
stimulus modality; and (iii) the post-semantic lexical retrieval
stage, when name retrieval is impaired but semantic
information is retrieved correctly, i.e. proper name anomia.
The issue of whether these deficits reflect the existence of
face- or person-specific cognitive modules has been debated
© Oxford University Press 2001
since the earliest reports of prosopagnosia. In some cases,
the impairment appeared to be so strictly selective for either
faces (De Renzi, 1986; De Renzi et al., 1991; Sergent and
Signoret, 1992; McNeil and Warrington, 1993; Farah et al.,
1995), person-specific semantics (Hanley et al., 1989; Evans
et al., 1995) or people’s proper names (McKenna and
Warrington, 1980; Lucchelli and De Renzi, 1992) that the
existence of dedicated cognitive modules seemed possible.
In other cases, prosopagnosia extended to impaired
recognition of specific objects that, like faces, have many
visually similar neighbours, e.g. breeds of dogs, types of
flowers or cars (Lhermitte and Pillon, 1975; Damasio et al.,
1982, 1990), individual animals (Bornstein et al., 1969; Assal
et al., 1984), buildings and landmarks (Pallis, 1955; Landis
et al., 1986). Similarly, patients with deficits in retrieving
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M. L. Gorno-Tempini and C. J. Price
person-specific semantics or proper names can also have
difficulties with other objects that, in addition to being part
of a perceptually homogeneous category, also have unique
semantic and lexical associations. These stimuli are referred
to as ‘semantically unique items’. For example, a patient
might be able to recognize a stimulus as a dog but be unable
to identify it as Lassie, the collie that starred in the popular
TV series (Ellis et al., 1989). Other examples of semantically
unique items include famous buildings and landmarks. These
stimuli have often been reported as impaired not only in
prosopagnosia (see above) but also in patients with personspecific deficits at the semantic (Ellis et al., 1989; Kartsounis
and Shallice, 1996; McCarthy et al., 1996) and lexical
(Semenza and Zettin, 1988, 1989) levels. Such co-occurring
deficits are not consistent with the existence of personspecific modules, but do support the hypothesis that specific
deficits can arise from the different demands that faces and
proper names place on the object- and word-processing
systems. First, they need to be distinguished from other,
perceptually similar category members. Then they need to
be linked to unique semantic and lexical features.
The debate over the extent to which face-processing
engages specific modules has recently extended into the
functional neuroimaging literature. Most studies of facespecificity have focused on the perceptual level of processing
and have used unfamiliar faces that cannot be linked to
specific semantic or lexical representations. Viewing and
matching unfamiliar faces relative to other categories of
objects consistently activate a region of the lateral fusiform
gyrus bilaterally, but more consistently on the right, that has
recently been labelled the ‘fusiform face area’ [FFA; mean
Talairach coordinates across subjects: x ⫽ 40, y ⫽ –55,
z ⫽ –10 (Kanwisher et al., 1997)]. However, the response
in the FFA is not exclusive to faces (Ishai et al., 1999) and
the same area also responds to animals (Chao et al., 1999)
and animal faces (Maguire et al., 2001) and is activated
when visually similar objects are categorized at the
subordinate level, e.g. when distinguishing different types of
birds and cars, especially when the subject is an expert
(Gauthier et al., 1999, 2000). Interestingly, pictures of
buildings, which also need a subordinate level of perceptual
categorization, activate a more medial region in the ventral
temporo-occipital cortex when compared with other
categories of objects. This region has been reported to be
located in the right anterior lingual sulcus (coordinates in
Talairach space: x ⫽ 20.6 ⫾ 5.0, y ⫽ 53.8 ⫾ 6.7, z ⫽
–9.2 ⫾ 3.9) (Aguirre et al., 1998), bilateral medial fusiform
(x ⫽ –26, y ⫽ –57, x ⫽ –14 and x ⫽ 28, y ⫽ –57, z ⫽
–13) (Ishai et al., 1999) and parahippocampal gyri (x ⫽ 20,
y ⫽ –39, z ⫽ –5 and x ⫽ –28, y ⫽ –39, z ⫽ –6) (Epstein
et al., 1999). Even if the peak activations in these studies
were relatively close, it has recently been suggested that,
while the more anterior parahippocampal region might be
more involved in processing spatial layouts, such as rooms
and scenes (Epstein and Kanwisher, 1998; Epstein et al.,
1999), the more posterior fusiform/lingual region might be
more involved in processing single buildings (Haxby et al.,
2000; Maguire et al., 2001). We will refer to this buildingresponsive area as the ‘parahippocampal/lingual’ region.
To summarize, the currently available imaging and
neuropsychological data indicate that faces and buildings
place different demands on perceptual, presemantic
processing, although the specificity of these responses is not
absolute. The neuroimaging study reported in this paper
focuses on the identification of famous faces and buildings.
This is an area that has received little attention in the
neuroimaging literature and few studies have investigated
brain responses to either familiar (friends or associates),
newly learned (seen once or twice before scanning) or famous
(well-known celebrities) faces or buildings. Therefore, many
issues are still debated. For instance, the effect of fame on
the FFA and parahippocampal/lingual activations has not yet
been fully established. Differential responses to famous or
familiar faces in the FFA have been confounded by the task
and/or attentional demands (Sergent et al., 1992; Kim et al.,
1999; Leveroni et al., 2000). When the cognitive task was
held constant, we found no effect of fame on the fusiform
face activation (Gorno-Tempini et al., 1998). Similarly, the
effect of fame on the parahippocampal/lingual activation
remains unclear. Responses to famous buildings have never
been investigated and only one study has compared familiar
(campus landmarks for students) and unfamiliar landmarks
(Epstein et al., 1999). However, in this study, the analysis
was confined to the ‘parahippocampal place area’ (a region
that responds to scenes more than objects and buildings) and
no other brain areas were investigated.
Other studies investigating activation for famous faces have
highlighted the anterior temporal cortices, either bilaterally
(Sergent et al., 1992; Damasio et al., 1996; Leveroni et al.,
2000) or on the left only (Gorno-Tempini et al., 1998, 2000;
Henson et al., 2000). For instance, the left anterior middle
temporal gyrus (MTG) was more active for: (i) naming
pictures of famous faces (well-known celebrities) relative to
five other categories of objects (animals, artefacts, body parts,
maps and colours) (Gorno-Tempini et al., 2000); and (ii)
matching famous proper names (e.g. Marilyn Monroe) relative
to object names constructed from two words (e.g. wheelbarrow) (Gorno-Tempini et al., 1998). However, this region
has not been observed for familiar (friends and associates)
relative to unknown faces, which activated the right temporal
pole (Nakamura et al., 2000).
Several questions therefore remain. First, are the fusiform
and parahippocampal/lingual regions modulated by fame?
Secondly, does the left anterior MTG respond to famous
faces (e.g. Prince Charles) more than famous buildings (e.g.
Buckingham Palace)? Thirdly, is the right anterior temporal
cortex responsive to faces and buildings and is it also
influenced by fame? To address these questions, we engaged
subjects in a same–different judgement on visually presented
pairs of pictures depicting famous, non-famous and scrambled
faces and buildings. We chose universally famous items
rather than personally familiar stimuli because semantic and
PET study of famous faces and buildings
lexical associations should be more stable and less influenced
by personal experience and individual differences. We chose
the same–different matching task because it allowed us to
keep the task constant while famous, non-famous and
scrambled stimuli were compared with each other. The same
task and face stimuli were used in our previous study
(Gorno-Tempini et al., 1998). Therefore, when investigating
the effect of fame, which was manipulated in both
experiments, we combined the old and new data in a single
statistical analysis (see Methods).
We predicted that (i) the FFA and the parahippocampal/
lingual regions would be more active for faces than buildings
(Kanwisher et al., 1997; Aguirre et al., 1998; Chao et al.,
1999; Ishai et al., 1999; Maguire et al., 2001); (ii) the
anterior temporal cortex would be more responsive to
famous faces than to non-famous faces and scrambled stimuli
(Gorno-Tempini et al., 1998); and (iii) the FFA response
would not be affected by fame (Gorno-Tempini et al., 1998).
Our open questions were (i) whether the parahippocampal/
lingual area would respond differently to famous and nonfamous buildings; and (ii) whether the left and/or right
anterior temporal cortex would show a common or differential
response to famous faces and famous buildings, when
compared with their non-famous counterparts.
Methods
Subjects
Fifteen male subjects (age range 18–38 years) took part in
the experiment. They were all right-handed, native English
speakers, healthy, on no medication and free from any history
of neurological or psychiatric illness. Subjects gave informed,
written consent to the study, which was approved by the
local hospital ethics committee and the Administration of
Radioactive Substances Advisory Committee (UK).
Experimental design and procedure
Each subject underwent 12 PET relative perfusion scans
while deciding whether two visual stimuli, displayed
simultaneously as a pair in the centre of a computer screen,
were the same or different. Whereas the task was constant
across conditions, the stimuli were pairs of famous or nonfamous faces or buildings. The experiment was therefore of
a 2 ⫻ 2 factorial design, with category (faces and buildings)
and fame (famous and non-famous) as the two factors. A
low-level visual control condition was also introduced, in
which subjects had to perform a same–different task on pairs
of scrambled stimuli. Therefore, six experimental conditions
were created: famous faces (FF), non-famous faces (NFF),
scrambled faces (SF), famous buildings (FB), non-famous
buildings (NFB) and scrambled buildings (SB). Subjects saw
10 stimulus pairs (six same and four different) per scan and
responded with a key-press: right button for same and left
2089
button for different pairs. Reaction times (RTs) and accuracy
were recorded. Each pair of stimuli was presented for 5 s at
the rate of one stimulus every 6 s. Each of the six conditions
was repeated twice with different stimuli. There were no
stimulus repetitions within subjects and condition order
was counterbalanced between and within subjects. Since all
stimuli were novel to the subjects at the time of scanning,
after the PET session, the famous faces and buildings were
shown again to ensure that correct identification had
occurred.
Stimuli
The famous faces came from a pool of 200 black-and-white
photographs of celebrities in different professional categories.
Their familiarity was determined by a behavioural study
conducted on 20 normal male subjects (age range 18–
33 years) who were shown each face on a computer screen
for 5 s and had to name the person. Only those faces that
were named within the 5 s by at least 19 subjects were
included in the PET study. The same procedure was used to
choose the famous buildings. Twenty normal male subjects
(age range 20–35 years) viewed 120 black-and-white pictures
of famous buildings and landmarks and only stimuli that
were named within 5 s by at least 19 subjects were used
during the scanning sessions. The names of the famous
stimuli are listed in Appendix I.
The non-famous faces were matched to the famous faces
for mean age, sex and facial expression. The non-famous
buildings were chosen to match their famous counterparts
for general category (e.g. churches and arches). For instance,
the non-famous stimulus corresponding to the Leaning Tower
of Pisa was an unknown tower (Fig. 1).
The scrambled control stimuli were obtained by scrambling
each face and building stimulus in the same way. To maintain
a constant spatial frequency power density spectrum in the
scrambled stimuli, the manipulation was on the phases of
each spatial frequency in the image. The phase of each lower
frequency component, starting from the lowest frequency,
was swapped with the phase of a corresponding higherfrequency component, starting with the highest. A pattern
was obtained that was no longer recognizable as a face.
Faces were framed with a black oval mask to avoid
differences in the picture background. The same masking
procedure eliminated as much as possible of the surrounding
scene from the building stimuli. The framed faces and
buildings were then paired to obtain a single stimulus that
measured 9.4 ⫻ 13.6 cm. The pairs comprised two stimuli
displayed either one next to the other on a black background
(all faces and half of the building pairs) or one above the
other (half of the building pairs). The corresponding
scrambled stimuli were framed and paired similarly. Since
the building and face conditions were never compared directly
without also being compared with their respective scrambled
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M. L. Gorno-Tempini and C. J. Price
stimuli, the different rotation was matched across type of
stimulus. The buildings, faces and scrambled conditions were
also matched for mean luminance.
structural MRI was obtained with a 2 T Magnetom Vision
scanner (Siemens, Erlangen, Germany).
Data analysis
PET and MRI scanning
Each subject underwent 12 PET relative perfusion scans over
a period of 2 h. Scans were obtained using a Siemens/CPS
ECAT Exact HR⫹ (model 962) PET scanner (Siemens/CTI,
Knoxville, Tenn., USA) with collimating septa retracted.
Participants received a 20 s intravenous bolus of H2150 at a
concentration of 55 MBq/ml and a flow rate of 10 ml/min
through a forearm cannula. For each subject, a T1-weighted
Data were analysed by statistical parametric mapping, using
SPM99 software from the Wellcome Department of Cognitive
Neurology, London, UK (http//www.fil.ion.ucl.ac.uk/spm)
implemented in Matlab (Mathworks. Sherborn, Mass., USA)
using standardized procedures (Friston et al., 1995a, b),
including realignment for head movements, spatial
normalization to the Montreal Neurological Institute template
brain (Cocosco et al., 1997) in the space of Talairach and
Fig. 1 Examples of the stimuli used in Experiments 1 and 2 for the face conditions and in Experiment 2 for the building conditions.
PET study of famous faces and buildings
Tournoux (Talairach and Tournoux, 1988) and smoothing.
The smoothing kernel was a 3D Gaussian filter of 16 mm.
Condition and subject effects were estimated according to
the general linear model at each voxel. Our design matrix
specified two groups of subjects. One comprised nine subjects
who performed the same–different matching paradigm
described above on face (FF, NFF and SF) and building (FB,
NFB and SB) stimuli. The other group comprised six subjects
who performed the same–different matching task on the face
but not the building stimuli. The data from the group of six
subjects have been reported previously (Gorno-Tempini et al.,
1998) and are referred to here as Experiment 1. The new
data (group of nine subjects) are referred to as Experiment
2. Both were included in the present study to (i) increase
sensitivity to the effect of fame; and (ii) allow comparisons
to be made between studies. However, differential effects of
category and category ⫻ fame interactions were estimated
using data from Experiment 2 only.
Three main comparisons were performed in order to
identify regions that were commonly or differentially
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activated by category (faces or buildings) or fame (famous
or non-famous).
Main effect of category irrespective of fame
(Experiment 2 only)
The effect of faces relative to buildings was identified with
a conjunction of FF–FB and NFF–NFB. Using the inclusive
masking option in SPM99, we also eliminated any areas that
were not active for FF–SF or NFF–SF at P ⬍ 0.001. The
effect of buildings relative to faces was based on a conjunction
of the reversed contrasts (FB–FF and NFB–NFF), inclusively
masked with FB–SB and NFB–SF at P ⬍ 0.001.
Once the face- and building-specific areas had been
identified, we also investigated the effect of fame within
these areas, i.e. FF–NFF and NFF–FF in the face area and
FB–NFB and NFB–FB in the building area.
Main effect of fame irrespective of category
(Experiment 1 and 2)
This was identified with a conjunction of three contrasts: (i)
FF–NFF (Experiment 1); (ii) FF–NFF (Experiment 2); and
(iii) FB–NFB (Experiment 2). Inclusive masking, with
FF–SF (Experiment 1 and 2) and FB–SB (Experiment 2) at
P ⬍ 0.001, ensured that activation was also present above
controls in both experiments.
The same procedure was performed in order to identify
regions specific to non-famous stimuli.
Activations specific to famous faces or famous
buildings (Experiment 2 only)
Fig. 2 Corrected means and standard errors of the RTs for each of
the six experimental conditions (Experiment 2)
These were identified using the inclusive masking procedure
to find areas that were activated for (i) FF–NFF; (ii)
FF–SF; and (iii) the interaction of faces and fame (FF–
NFF) – (FB–NFB).
Likewise, regions more involved in processing FB were
Table 1 Coordinates and Z values of the brain regions identified by our main analysis
(see Results)
Contrast
Area (BA)
Faces ⬎ buildings
Buildings ⬎ faces
Right fusiform (37/20)
Bilateral parahippocampus/
lingual/fusiform (36/19/37)
FF–NFF ⫹ FB–NFB
FF–NFF ⬎ FB–NFB
Left MTG (21)
Right MTG (21)
*P ⬍ 0.001 uncorrected.
Talairach coordinates
Z
x
y
z
48
⫺26
⫺26
⫺34
28
30
⫺48
⫺44
⫺38
⫺48
⫺48
⫺40
⫺22
⫺14
⫺18
⫺12
⫺12
⫺16
4.9
6.0
5.0
4.5*
5.5
4.5*
64
62
0
⫺2
⫺16
⫺14
4.7
3.2*
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M. L. Gorno-Tempini and C. J. Price
PET study of famous faces and buildings
those more active for (i) FB–NFB; FB–SB; and (ii) the
interaction of fame with buildings (FB–NFB) – (FF–NFF).
Thresholds for both experiments
We accepted a level of significance of P ⬍ 0.05 corrected
for multiple comparisons for all our main contrasts, except
for the interaction, which we report at P ⬍ 0.001 uncorrected.
When claiming the absence of an effect, we lowered the
threshold to P ⬎ 0.1 uncorrected.
Results
Behavioural data
All subjects performed the experimental task at ceiling level,
making fewer than three errors. Reaction time data were
collated for each experimental condition and any score that
deviated ⬎2.5 SD from the mean was replaced with the
mean of that condition. A factorial ANOVA (analysis of
variance) was used to identify main effects of category (faces
and buildings), fame (famous and non-famous) and the
interaction between these variables (Fig. 2). No significant
effect was found, indicating no difference in difficulty and
attentional demands between the four experimental conditions. However, when a one-way ANOVA including the
scrambled condition as a variable was performed, a significant
difference in RTs was found (P ⬍ 0.05). A post hoc Scheffé
test indicated that same–different matching for the scrambled
conditions took significantly longer than for the faces or
buildings. Reaction times for FF, NFF and SF in Experiment
1 have been reported elsewhere (Gorno-Tempini et al., 1998).
There was no significant difference between the FF and NFF,
but matching took significantly longer for scrambled stimuli
than for both the face conditions. Thus, the behavioural data
were consistent across experiments.
After scanning, subjects were presented with each face or
building from the famous set and asked to name it within
5 s. No subject named fewer then 26 out of 28 faces and 25
out of 28 buildings. This confirmed that subjects were
able to establish semantic and lexical associations for the
famous stimuli.
Scanning data
Main effect of category irrespective of fame
Faces versus buildings. When famous and non-famous
faces were compared with the corresponding building
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conditions, a region in the right fusiform gyrus [Brodmann
area (BA) 37/20], corresponding to the FFA, was activated
at a corrected level of significance (P ⬍ 0.05) (Table 1).
Both FF and NFF evoked activation in the FFA and there
was no significant effect of FF–NFF or NFF–FF (P ⬎ 0.1).
Figure 3A illustrates that the FFA response to buildings
(relative to controls) was very small (P ⬍ 0.05 uncorrected).
Buildings versus faces. When FB and NFB were
compared with the corresponding face conditions, an area
in the medioventral occipitotemporal cortex was activated
bilaterally (BA 36/19/37) at a corrected level of significance
(P ⬍ 0.05). The peak of the activation was in the superior
lingual gyrus but the cluster spread to the parahippocampus
and, on the left side, also to the medial fusiform gyrus.
Because of the resolution and smoothing of the PET data,
we will refer to this activation as the parahippocampus/
lingual region. This area was activated by both FB and NFB
and there was no effect of FB–NFB or NFB–FB (P ⬎ 0.1
uncorrected). Figure 3B illustrates that the response in these
regions was almost exclusive to buildings, with no effect of
faces versus controls (P ⬎ 0.9 uncorrected).
Main effect of fame irrespective of category
Famous versus non-famous stimuli. The conjunction
analysis demonstrated that only one region showed a
significant effect of fame (P ⬍ 0.05 corrected). This was the
left anterior MTG (BA 21; P ⬍ 0.05 corrected; 240 voxels).
The high level of significance of the conjunction analysis
reflects the consistency of this activation across contrasts
(FF–NFF and FB–NFB) and across experiments (Experiments
1 and 2). Figure 3C illustrates an equivalent effect of famous
versus non-famous faces and buildings in Experiments 1 and
2 (P ⬍ 0.005 to P ⬍ 0.001). Furthermore, in this region
there was no interaction between fame and category (P ⬎ 0.1
uncorrected).
Non-famous versus famous stimuli. No significant
effect was found for non-famous versus famous stimuli.
Areas specific to famous faces or famous
buildings
A small region in the right anterior MTG (BA 21) showed
a significant interaction (P ⬍ 0.001 uncorrected). In this area
there was a small effect of FF–NFF (Z ⫽ 2.7; P ⬍ 0.01
uncorrected) but no effect of FB–NFB (P ⬎ 0.1 uncorrected)
Fig. 3 From top to bottom, this figure illustrates areas of activation and parameter estimates for regions that were more activated for (A)
faces than for buildings, (B) buildings than for faces, (C) famous than for non-famous faces and buildings, and (D) famous than for nonfamous faces only. In the left column, all activations are superimposed on axial slices of the mean of the nine subjects’ normalized
structural MRIs and thresholded at P ⬍ 0.001 (uncorrected). In the right column, the plots indicate the value of the normalized regional
cerebral blood flow at the indicated voxel (y-axis) for each of the experimental conditions in Experiments 1 and 2 (x-axis).
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M. L. Gorno-Tempini and C. J. Price
(Fig. 3D). Figure 3 illustrates that in this region there was also
a small effect of FF versus NFF in Experiment 1 (Z ⫽ 2.4;
P ⬍ 0.005 uncorrected) and the conjunction of FF–NFF in
Experiments 1 and 2 raised the joint significance of this
effect to P ⬍ 0.001 uncorrected (Z ⫽ 3.2).
Summary of results
As predicted, we demonstrated that (i) the FFA and the
parahippocampal/lingual region were more active for faces
and buildings, respectively; (ii) the anterior MTG was more
active for famous faces relative to non-famous and scrambled
faces; and (iii) there was no effect of fame on the FFA. In
addition, we found that (i) there was no effect of fame on
the parahippocampal/lingual region; and (ii) the left anterior
MTG was equally active for famous faces and famous
buildings. In the right anterior MTG, there was a small effect
of fame for faces but not for buildings.
Discussion
In this study, we investigated brain responses to famous and
non-famous faces and buildings in order to determine whether
(i) the fusiform and parahippocampal/lingual regions, which
have been associated previously with processing of unfamiliar
faces and buildings, would respond differently to famous
and non-famous stimuli when the cognitive task was kept
constant; and (ii) the left and right anterior temporal regions,
previously associated with famous face naming, would be
commonly or differentially responsive to another category of
unique items: famous buildings. The results show categoryspecific effects in the right fusiform and bilateral parahippocampal/lingual gyri for faces and buildings, respectively, but
no effect of fame. In contrast, the left anterior MTG showed
an effect of fame for both faces and buildings, but no effect
of category. The responses of the right MTG were less clearcut, with only a small effect of fame for faces and no effect
for buildings. We first discuss the anterior temporal and then
the fusiform and parahippocampal/lingual activations.
Anterior temporal activations
Many previous functional neuroimaging studies have
demonstrated that activation in the left anterolateral portions
of the middle and inferior temporal gyri (BA 21/20) increases
with the demands on semantic processing, particularly when
specific information must be retrieved. For instance, this
occurs when subjects categorize objects (Devlin et al., 2000)
and match words according to specific semantic features
(Mummery et al., 1998) or associations (Vandenberghe et al.,
1996). Interestingly, the left MTG is also activated when
famous faces and proper names are matched (Gorno-Tempini
et al., 1998), with greater activation for naming famous faces
than objects from other categories (Damasio et al., 1996;
Gorno-Tempini et al., 2000). The naming and matching of
objects are usually associated with activation in more
posterior regions of the left inferior or middle temporal cortex
(Bookheimer et al., 1995; Martin et al., 1995; Perani et al.,
1995; Damasio et al., 1996; Moore and Price, 1999). Our
question concerns why the left anterior temporal lobe is more
active for the naming of famous faces than for object-naming.
One explanation might be that there is a discrete region in
the left anterior temporal lobe that is specific to personspecific semantic or lexical attributes. This would be
consistent with patients having anterior temporal damage
and loss of person-specific semantics, but not with the
neuroimaging studies that have shown activation of the
anterior MTG when retrieving specific semantic features
related to objects. A more likely explanation is that naming
or matching famous faces elicits more activation than objectnaming in areas associated with the retrieval of semantic
features. This might occur because the semantic associations
evoked by famous faces are unique and are not shared by
other items of the same category. We tested this hypothesis
by measuring the response to famous buildings, which, to be
correctly identified, also need to evoke specific semantic and
lexical associations that are not shared with other category
members. We found equivalent responses for famous faces
and famous buildings in the left anterior MTG, thereby
confirming that matching semantically unique items increased
the demand on a semantic processing area, even when subjects
were not explicitly required to identify them. However, when
normal subjects were exposed to a very famous stimulus,
identification and lexical retrieval occurred together automatically. Therefore, the anterior left MTG may also play a
role in linking semantic and lexical information and we
cannot exclude the possibility that this region is involved in
pure lexical retrieval processes (Damasio et al., 1996). Future
studies, combining neuropsychological and neuroimaging,
are necessary to solve this issue (Gorno-Tempini et al., 2001)
and to investigate whether responses could be affected by
familiarity or the age of acquisition with the object/face
concepts.
We were also interested in how the right anterior temporal
cortex responded to famous faces and buildings. Indeed,
neuropsychological patients with person-specific semantic
deficits often suffer from anterior temporal pathologies, such
as epilepsy, neurodegenerative diseases and herpes simplex
encephalitis, which are most commonly bilateral. Furthermore, patients have been described with mainly left, bilateral
or mainly right damage (De Renzi et al., 1987; Ellis et al.,
1989; Hanley et al., 1989; Evans et al., 1995; Kroll et al.,
1997; Hodges and Graham, 1998; Kitchener and Hodges,
1999), again suggesting that person-specific semantic knowledge is distributed bilaterally. However, although three
functional neuroimaging studies have shown bilateral
(Sergent et al., 1992; Leveroni et al., 2000) responses to
famous faces, activations have been left-lateralized in others
(Gorno-Tempini et al., 1998, 2000; Henson et al., 2000).
The only right-lateralized anterior temporal activation was
found for familiar faces and scenes (Nakamura et al., 2000).
In our study, responses in the anterior temporal cortex were
PET study of famous faces and buildings
much weaker in the right than the left hemisphere and were
only observed for famous faces. It is possible that the
differential anterior temporal lateralization in our study and
that of Nakamura and colleagues (Nakamura et al., 2000) is
determined by the degree of familiarity with the stimuli, which
influences the amount of semantic and lexical information that
can be retrieved. The faces and buildings in our experiment
were famous and well known to each subject, whereas in the
study of Nakamura and colleagues the degree of familiarity
was not controlled across subjects or stimuli. Another possibility is that semantic information can be retrieved by
activation of either the left or the right anterior temporal
cortex. This would explain why semantic impairments do
not usually occur when anterior temporal damage is limited
to one hemisphere. Further evidence, combining functional
imaging and lesion studies, is therefore required to determine
the different roles of the left and right anterior temporal lobes.
Fusiform and parahippocampal/lingual
activations
Converging evidence has indicated that unfamiliar faces and
scenes or buildings evoke differential responses in the fusiform
and parahippocampal/lingual gyri when compared with each
other and with other categories of objects (Aguirre and
D’Esposito, 1997; Kanwisher et al., 1997; Aguirre et al., 1998;
Chao et al., 1999; Epstein et al., 1999; Epstein and Kanwisher,
1998, 2000; Ishai et al., 1999; Kanwisher et al., 1999;
Nakamura et al., 2000; Maguire et al., 2001). We confirmed
these results by showing a preferential response to faces in a
region in the right fusiform gyrus and an almost exclusive
response to buildings in bilateral parahippocampal/lingual
areas. The results of the present study also demonstrate that,
when cognitive task and attentional load were controlled (no
difference in RT between conditions), both these categoryspecific regions were unaffected by fame. Previous studies that
found a differential effect of fame in the FFA did not control
for task (Sergent et al., 1992) or used passive viewing (Henson
et al., 2000) or familiarity decision (Leveroni et al., 2000)
paradigms, in which it is likely that subjects would have been
more engaged in the task when the stimuli were familiar.
Indeed, it has been established clearly that the FFA is modulated
by attentional demands (Wojciulik et al., 1998). In contrast,
we demonstrate that it is unaffected by fame when subjects
attend to the perceptual features of unfamiliar as well as
famous faces.
Our results are consistent with a pre-semantic perceptual role
for the FFA in processing faces and for the parahippocampal/
lingual region in processing buildings. Although the
identification of the specific mechanism responsible for these
activations is beyond the aim of our study (for discussion, see
Gauthier, 2000; Haxby et al., 2000; Kanwisher, 2000), the role
of the FFA in the perceptual stage of the face-recognition
process is consistent with lesion studies. In fact, temporooccipital lesions, which include the FFA, usually cause the
2095
‘apperceptive’ type of prosopagnosia, where the defective
recognition of familiar faces is accompanied by defective
processing of unfamiliar faces (De Renzi et al., 1991). On the
other hand, more anterior lesions in the temporal lobes are
more likely to cause the ‘associative’ type of prosopagnosia,
in which the deficit seems to be selective for familiar faces
(Gentileschi et al., 1999) and can also progress to the loss of
person-specific semantics (Evans et al., 1995; Kitchener and
Hodges, 1999). A similar argument could apply to the
parahippocampal region. This area and the surrounding
fusiform/lingual regions have been shown previously to
respond preferentially to stimuli that depict spatial and
topographical information about the environment, e.g. rooms,
buildings, houses, scenes and landscapes. Our results support
the hypothesis that, within this region, the more posterior
lingual focus is the most responsive to single buildings (Haxby
et al., 2000). The fact that the parahippocampal/lingual area did
not respond differently to famous and non-famous buildings
suggests a presemantic perceptual role in the analysis. This
would then be followed by further analysis in more anterior
regions, depending on the task required, for instance, in the
right hippocampus for active space navigation (Maguire et al.,
1998) and the left anterior temporal cortex for retrieval of
semantic information.
In conclusion, we demonstrate that distinct regions within
the ventral occipitotemporal cortex are involved in the
category-specific perceptual processing of face and building
stimuli. In contrast, we found that a semantic area in the left
anterior MTG was engaged by both famous faces and
famous buildings. Future studies are necessary to establish
unequivocally the role of this region in semantic and/or
lexical retrieval processes.
Acknowledgement
This work was funded by The Wellcome Trust.
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Received January 22, 2001. Revised May 15, 2001.
Accepted June 11, 2001
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Appendix I
Famous faces
Famous buildings
Bob Geldof
Michael Caine
Prince Charles
Tom Cruise
Princess Margaret
Queen Elizabeth
Eddie Murphy
Elvis Presley
John Major
John F. Kennedy
Joan Collins
Marilyn Monroe
Mel Gibson
Princess Diana
Bill Clinton
Arnold Schwarzenegger
Albert Einstein
Sarah Ferguson
Julia Roberts
Prince Andrew
Meryl Streep
John Travolta
Margaret Thatcher
Tony Blair
Ronald Reagan
Neal Kinnock
Gary Lineker
Twiggy
Millennium Dome
Clock Tower, Houses of Parliament
Taj Mahal
Leaning Tower of Pisa
The Louvre
Sydney Opera House
Buckingham Palace
Tower of London
The White House
Eiffel Tower
Arc de Triomphe
Wellington Arch
Brooklyn Bridge
Tower Bridge
Royal Albert Hall
Kew Gardens
Colosseum
Kensington Palace
BT Tower
World Trade Centre (twin towers)
Canary Wharf Tower
Empire State Building
Westminster Abbey
St Paul’s Cathedral
Capitol, Washington, DC
Globe Theatre
Battersea Power Station
Notre Dame, Paris