Neuropsychologia 46 (2008) 409–418

Categorical and thematic knowledge representation in the brain: Neural

correlates of taxonomic and thematic conceptual relations
Olga Sachs a,∗ , Susanne Weis b , Timo Krings c , Walter Huber b , Tilo Kircher a
a Department of Psychiatry and Psychotherapy, RWTH Aachen University, Pauwelstr. 30, D-52074 Aachen, Germany
b Department of Neurology, RWTH Aachen University, Pauwelstr. 30, D-52074 Aachen, Germany
c Department of Neuroradiology, RWTH Aachen University, Pauwelstr. 30, D-52074 Aachen, Germany

Received 24 April 2007; received in revised form 8 August 2007; accepted 18 August 2007
Available online 25 August 2007

Abstract
Most current models of knowledge organization in the brain are based on hierarchical or taxonomic categories (animals, tools). Another important
organizational pattern is thematic categorization, i.e. categories held together by external relations, a unifying scene or event (car and garage).
We used fMRI to examine neural activation patterns as subjects performed a category construction task where these two category types were
contrasted. Subjects were visually presented with a target word followed by the presentation of two match words and had to choose by button press
one match that goes best with the target word. In the balanced or cross-categorization condition (Car/Garage Bus) both match words fit the target;
in the biased conditions only one match word fit the target either thematically (Car/Garage Brush) or taxonomically (Car/Bus Eraser).
We found that in the biased conditions, thematic and taxonomic categories recruited very similar cortical regions: left inferior frontal, middle
temporal and occipital regions. In the balanced condition subjects showed no behavioral preference for either thematic or taxonomic matches.
However, contrasting signal changes during a subjective taxonomic choice in the presence of a thematic alternative vs. a subjective thematic choice
in the presence of a taxonomic alternative required the additional recruitment of right middle frontal gyrus, left precuneus and left thalamus.
Our results suggest that thematic relations between objects are processed similarly to taxonomic relations, but require less cerebral processing
demand, providing validation for thematic categories as an alternative principle of conceptual organization.
© 2007 Elsevier Ltd. All rights reserved.

Keywords: Categorization; Thematic categories; Taxonomic categories; fMRI; Cross-categorization

How knowledge is represented and stored in the brain is still
an open question in cognitive neuroscience. The basic princi-
ple of organizing semantic information is categorization. Two
category types can be distinguished: (1) taxonomic categories
that include items of the same kind (e.g., superordinate category
vehicles with members such as car, bus, train, etc.) (2) the-
matic categories that include externally related items interacting
within scenes or events (e.g., car and garage, dog and leash).1
Most lesion and neuroimaging studies of knowledge rep-
resentation and categorization have focused on identifying
∗ Corresponding author. Tel.: +49 241 8089721; fax: +49 241 8082422.
E-mail address: osachs@ukaachen.de (O. Sachs).
1 Members of taxonomic categories are often referred to in the literature as
“categorically” or “semantically” related concepts. Members of thematic cate-
gories are often said to share an “associative” relationship and therefore referred
to as “semantic associates” or “associatively” related items. In this paper we
predominantly use the terms “taxonomic” and “thematic” categories.
brain areas involved in the processing of isolated taxonomic
categories, such as animals or tools. The intuitive idea that
distinct conceptual categories should also have their distinct
domain-specific localization in the brain (living vs. non-living
things, animals vs. tools) has gained support from the studies
of brain-damaged patients who have category-specific deficits
(e.g., Caramazza & Shelton, 1998; Warrington & McCarthy,
1983) and functional imaging studies of healthy subjects that
showed distinct areas of activation for natural kinds and artifacts
(e.g., Perani, Schnur, et al., 1999; Thompson-Schill, Aguirre,
D’Esposito, & Farah, 1999). For example, Perani, Schnur, et al.
(1999) have found that the left fusiform gyrus was involved in
the processing of pictures and words of living entities and the
left middle temporal gyrus was involved in the processing of
words and pictures of tools. Leube, Erb, Grodd, Bartels, and
Kircher (2001) have even found a hemispheric differentiation
between living and non-living entities: right frontal, temporal
and fusiform areas were activated when subjects judged words

0028-3932/$ – see front matter © 2007 Elsevier Ltd. All rights reserved.
doi:10.1016/j.neuropsychologia.2007.08.015
410 O. Sachs et al. / Neuropsychologia 46 (2008) 409–418
as belonging to the “living” category, whereas the processing
of the “non-living” category members was largely limited to
the left frontal and occipital areas. Several hypotheses were
proposed to account for the discovered anatomical segregation
of natural kinds and artifacts, ranging from the evolutionary
pressure to distinguish between living and non-living things to
the role of perceptual and functional properties in distinguish-
ing individual members of these categories. For example, the
sensory-motor account of conceptual representation suggests
that perceptual features play a greater role in the processing
of natural kinds, while functional features are more important
for artifacts. This difference is reflected in distinct brain areas
involved in the representation of natural kinds and artifacts found
in neuroimaging studies (Cappa, Perani, Schnur, Tettamanti, &
Fazio, 1998; Damasio, Grabowski, Tranel, Hichwa, & Damasio,
1996; Martin, Wiggs, Ungerleider, & Haxby, 1996).
The role of functional information in the conceptual represen-
tation of artifacts or “the manner in which an object interacts with
the environment and with the observer” is also emphasized in the
distributed conceptual structure theory of categorical represen-
tation (Tyler, Moss, Durrant-Peatfield, & Levy, 2000). Here it is
suggested that we know what something is only when we know
what it does and that the strong relationship between appear-
ance and function is particularly important for the conceptual
representation of artifacts. Tyler et al.’s theoretical claim was
supported in a later fMRI study where they found that objects and
their associated actions activated the same neural regions—left
fusiform gyrus, superior and middle temporal cortex (Tyler et
al., 2003). An earlier study conducted by Grabowski, Damasio,
and Damasio (1998) has also found a common area of activa-
tion for the retrieval of words denoting objects and words for
actions. These findings extend the traditional understanding of
what a category is by showing that categories are held together
not just by the internal correlations between their features, but
also by the external functional relationship between an object
and its environment.
We would like to take this idea a step further and investigate
the neural representation of categories that are held together
by strong external or complementary relations between objects
that co-occur or interact together in space and time (Lin &
Murphy, 2001). These categories are called thematic. They can
be formed when thematically related items, such as car and
garage, are grouped together in a conceptual sorting situation.
Unlike members of traditional similarity-based taxonomic cat-
egories that are usually organized in hierarchies or taxonomies
(e.g., artifacts → vehicles → cars → BMW), members of the-
matic categories do not share any perceptual features. However,
despite the absence of any perceptual similarity, members of
thematic categories share a very strong and salient functional
relationship. Members of taxonomic categories may also be
functionally related: for example, train and car are both used to
drive. However, their category membership is based on their per-
ceptual or functional similarity, whereas members of thematic
categories do not share any functional similarity, but comple-
ment each other through their function. In the absence of any
perceptual or functional similarity such functional relationship
is, in fact, the only connection that they have. Lin and Murphy
(2001) suggest that besides sharing a functional relationship,
thematic categories can often share other types of rela-
tions, e.g., spatial (roof–house), temporal (meal–bill) or causal
(electricity–light). However, in most of such cases a functional
relation remains strong also in the presence of other types of
relations.
The neural correlates of thematic categories are little under-
stood, although their importance and salience has been shown in
many different areas of cognitive psychology. Cognitive devel-
opment research suggests that children often show preference
for thematic categories (Nguen & Murphy, 2003; Smiley &
Brown, 1979). Anthropological research and studies of cate-
gorization and inductive reasoning among expert groups, such
as tree experts or commercial fishermen, indicate that people
often make knowledge-based thematic groupings that are not
based exclusively on similarity, but also on causal relations
between entities (Lopez, Atran, Coley, Medin, & Smith, 1997;
Medin, Lynch, Coley, & Atran, 1997; Shafto & Coley, 2003).
Finally, even the most typical research subjects, U.S. college
students, make thematic categories and use them to make induc-
tive inferences (Lin & Murphy, 2001; Ross & Murphy, 1999).
For example, Lin and Murphy (2001) have demonstrated that
when given both a taxonomic and a thematic match (e.g. lion
and litter box) and asked which one of them forms a better cate-
gory with cat, young educated adults performed thematic (litter
box–cat) rather than taxonomic categorization (lion–cat) most
of the time. These results held even when subjects were given
a taxonomic, similarity-based definition of a category as part of
the instructions for constructing categories. This shows that they
formed thematic categories fully aware of the fact that they were
supposed to form categories and not to sort objects according
to some other principle. This set of results also highlights the
increasing recognition in the current behavioral categorization
research that items can be cross-classified into more than one
category type (Lin & Murphy, 2001; Nguen & Murphy, 2003;
Ross & Murphy, 1999). Besides taxonomic categories (e.g. ani-
mals, plants, vehicles), people use a variety of other category
types that may include the same items, such as thematic cate-
gories (e.g. dog and leash, car and garage, picture and frame),
as well as ad hoc or goal-derived categories (e.g. things to carry
out of the burning house; Barsalou, 1983), or script categories
(e.g. breakfast foods; Ross & Murphy, 1999).
However, it is important to note that classical taxonomic
categories that combine “things of the same kind” differ from
thematic categories in many important ways. One crucial differ-
ence lies in how they are labeled: we use the same word to refer to
members of taxonomic categories and not to things that co-occur
together. One way to interpret the differences between thematic
and taxonomic conceptual links is to only call things of the
same kind “categories”, and to refer to items that are related the-
matically as “semantic associates” (accepted terminology in the
neuroimaging literature). Another possibility is to treat thematic
and taxonomic relations as two different, perhaps independent,
but equally important ways of organizing semantic knowledge
into groups that both can be called “categories” (more com-
mon in the cognitive literature). We use the terminology of the
cognitive literature because thematic relations have been inves-
 O. Sachs et al. / Neuropsychologia 46 (2008) 409–418
tigated there much more extensively than in the neuroimaging
categorization research.
Relatively good predictions for our study can be derived
from a large body of semantic priming studies that investigated
the structure of the lexical–semantic network by contrasting
the processing of categorical relations between prime target
pairs (what we refer to as taxonomic relations between cate-
gory members) with associative relations between prime target
pairs (what we refer to as thematic relations). For example,
Kotz, Cappa, von Cramon, and Friederici (2002) have compared
the neuroanatomical correlates of categorical and associative
relations between prime target pairs using an auditory primed
lexical decision task. They found that compared to the asso-
ciative relations (e.g., key–chain), the processing of categorical
relations (e.g. cow–dog) recruited additional brain areas in the
right precuneus, the isthmus gyrus cinguli and the cuneus,
thus suggesting a more effortful processing of categorical rela-
tions in the right hemisphere. Evidence from a number of
other semantic priming studies that used either visual fields
or ERPs to establish the hemispheric difference in the prim-
ing of different semantic relationships (Beeman, Friedman,
Grafman, & Perez, 1994; Chiarello & Richards, 1992) also
points toward the right-hemisphere advantage for the processing
of categorical or taxonomic relations between concepts. Finally,
a lesion study by Hagoort, Brown, & Swaab, 1996 showed
that patients with right hemisphere lesions produced a nor-
mal N400 priming effect while listening to associatively related
prime-target pairs (bread–butter), but a trend toward a reduced
N400 priming effect for categorically related prime-target pairs
(church–villa).
In the current study our goal is to (1) compare the neural corre-
lates of thematic and taxonomic categories and (2) to investigate
the issue of cross-categorization: what happens when the same
object is an equally good member of both a thematic and a tax-
onomic category. When we combine both lines of research that
were discussed above—neuroimaging categorization research
and semantic priming studies, we can develop fairly specific pre-
dictions regarding our research questions. Based on the results
of neuroimaging categorization studies that involved non-living
objects or artifacts (Perani, Schnur, et al., 1999) we hypothe-
size that both thematic and taxonomic categories made up of
such concepts should activate similar brain areas in the left mid-
dle temporal gyrus, as well as in the left frontal and occipital
areas. However, differences in the processing of taxonomic and
thematic relations between concepts found in semantic prim-
ing studies (Beeman et al., 1994; Chiarello & Richards, 1992;
Kotz et al., 2002) and in a lesion study by Hagoort et al. (1996)
suggest that we may also predict that taxonomic categories will
recruit additional right-hemispheric areas since they may require
a more effortful processing than thematic categories.
Besides investigating taxonomic and thematic categories in
isolation, our design allows us most importantly to distinguish
between cerebral activation during the subjective classification
into one or the other category (cross-categorization). We hypoth-
esize that when thematic and taxonomic categories compete,
there will be a greater additional activation during subjective tax-
onomic vs. thematic categorization reflecting increased demands
411
set on the evaluation of taxonomic categories when they compete
with the thematic categories.
1. Materials and methods
1.1. Participants
Fourteen healthy male subjects participated in the study. All of them were
right-handed according to the Edinburgh Inventory of Handedness (Annett,
1970). Their mean age was 28 (S.D. 7.5) years and they all spoke German as
their native language. All participants had normal or corrected-to-normal vision,
gave informed consent and were paid 10 Euros for participation in the study.
The study was approved by the Local Ethics Committee.
1.2. Stimuli and design
Subjects performed a forced-choice category construction task that is com-
monly used in categorization research (e.g., Lin & Murphy, 2001) and in
developmental classification studies (e.g., Smiley & Brown, 1979). In this task
subjects are first presented with a target word in the center of the computer
screen (e.g., Car), followed by the simultaneous presentation of two matches
(e.g., Garage, Bus). Subjects then have to decide which one of the two matches
goes best with the first target word to form a category. We used a categorical
design where all subjects were tested in three main experimental conditions
with 30 trials in each condition: (1) balanced condition where both matches
fit the target—Car/Garage Bus (one thematically, e.g., Car–Garage, and one
taxonomically, e.g., Car–Bus); (2) thematically biased condition where only
one match fits the target thematically—Car/Garage Brush; (3) taxonomically
biased condition where only one match fits the target taxonomically—Car/Bus
Eraser. The balanced condition was used for addressing the cross-categorization
question and the two biased conditions were used to investigate thematic and
taxonomic categories in isolation.
For each of the 30 German target words a thematic and a taxonomic match
were found that formed either a thematic or a taxonomic relation with the tar-
get. A pretest was conducted on a separate group of subjects to ensure that
thematic and taxonomic matches had an equally strong and salient relationship
with a target. Fifteen pretest participants were asked to rate the relationship
between potential thematic matches and their targets on a scale from 1 to 7
based on the definition of thematic categories in Lin and Murphy (2001), i.e.
how well they form an external or complementary relationship and co-occur
and interact together in space and time. A definition of taxonomic categories
was used to rate the relationship between potential taxonomic matches and their
targets, i.e. how similar they are in their perceptual, biological or functional
properties. All taxonomic matches were related to the target on the superor-
dinate level (e.g., bus and car are both vehicles). Based on the results of this
pretest, we selected only those thematic and taxonomic matches that scored
5 or higher. The selected thematic and taxonomic matches did not differ in
their average rating (Mthem = 6.69, S.D. = 0.32; Mtax = 6.60, S.D. = 0.49, t-test,
p = 0.13). The unrelated matches for the biased condition scored 2 or less on the
same test (Munr = 1.35, S.D. = 0.52). All selected stimuli belonged to the same
overall conceptual domain (all words depicted only artifacts), were concrete
and imageable (all words were concrete objects that are easy to visualize). The
CELEX lexical database (Baayen, Piepenbrock, & van Rijn, 1993) was used
to control for the average word frequency of German words used in taxonomic
and thematic trials (Mthem = 34.53, S.D. = 62.06; Mtax = 23.07, S.D. = 46.91, t-
test, p = 0.13). All stimuli were also controlled for word length (Mthem = 5.80,
S.D. = 1.24; Mtax = 5.63, S.D. = 1.30, t-test, p = 0.31). See Table 1 for examples
of the stimuli used in the experiment.
1.3. Procedure
An event-related design was used to present balanced, thematically biased,
taxonomically biased and baseline trials in a randomized order to participants.
Four randomized versions of the experiment were counterbalanced across sub-
jects to avoid a systematic effect of conditions on each other. Presentation of
stimuli was controlled by a computer using the Presentation 10.1 software pack-
age (Neurobehavioral Systems, http://www.neurobs.com/). Subjects lying in the
412 O. Sachs et al. / Neuropsychologia 46 (2008) 409–418
Table 1
Sample stimuli used in the experiment
Target word Thematic match Taxonomic match
German (English German (English German (English
translation) translation) translation)
Auto (Car) Garage (Garage) Bus (Bus)
Boot (Boat) Segel (Sail) Schiff (Ship)
Faden (Thread) Nadel (Needle) Seil (Rope)
Topf (Pot) Herd (Stove) Kessel (Kettle)
scanner viewed the computer screen through MR-compatible video goggles
(VisuaStim XGA, Resonance Technology, Inc., http://www.mrivideo.com/).
Each individual trial started with a fixation cross in the middle of the computer
screen presented for 500 ms followed by the presentation of the target word that
lasted 1000 ms. As soon as the target word disappeared two match words were
presented in the middle of the computer screen for 2000 ms. Match words were
presented one above the other to avoid saccadic eye movements. The presenta-
tion of match words was immediately followed by the # sign in the center of
the screen that lasted 2500–7500 ms to create jitter between individual trials.
Therefore, the average ISI was 5000 ms. Subjects could make their response at
any time point after the presentation of the two matches and before the next trial
began. They were instructed to press the top button of the parallel port device
when the top word was a better match to the target and to press the bottom
button when the bottom word was a better match. The pressing was done with
the middle and index finger of the left hand to avoid the motor-related activation
in the left hemisphere. The upper or lower position of thematic and taxonomic
matches in the balanced condition and the position of the related match in both
biased conditions were counterbalanced across trials.
A baseline task in this experiment was a letter categorization task used in
other categorization studies because it shares the same stimulus and response
characteristics as the main categorization task, but has no semantic component
(Devlin et al., 2002; Tyler et al., 2003). Participants were first presented with a
target letter string (e.g., Ccc), followed by the simultaneous presentation of two
matches (e.g., Gggggg, Cccc). Subjects then had to decide which one of the two
matches goes best with the target letter string. All letter strings were matched to
word stimuli in letter number.
1.4. MRI acquisition
All scanning was performed on a 1.5 T scanner (Gyroscan Intera, Philips
Medical Systems, Best, The Netherlands) using standard gradients and a cir-
cular polarized phase array head coil. For each subject, we acquired two series
of functional volumes of T2*-weighted axial EPI-scans including five initial
dummy scans parallel to the AC/PC line with the following parameters: number
of slices (NS) = 31; slice thickness (ST) = 3.5 mm; interslice gap (IG) = 0.35 mm;
matrix size (MS) = 64 × 64; field of view (FOV) = 240 mm × 240 mm; voxel
size = 3.75 mm × 3.75 mm; echo time (TE) = 50 ms; flip angle = 90; repetition
time (TR) = 2.8 s. After the functional run, for each participant an anatomical
scan was acquired using a high-resolution T1-weighted 3D-sequence consist-
ing of 140 transveral slices (TR = 7776 ms; TE = 3.56 ms; FoV = 256 mm; slice
thickness = 1 mm; inter slice gap = 1 mm, flip angle = 8). Three hundred twenty
two whole head scans were performed per subject divided into two runs. Each run
contained 158–166 whole head scans depending on the randomization version
the subject received.
1.5. fMRI data analysis
MR images were analyzed using Statistical Parametric Mapping software
(SPM2; www.fil.ion.ucl.ac.uk) implemented in MATLAB 6.5 (Mathworks Inc.,
Sherborn, MA).
After discarding the first five volumes, all images were realigned to the
first image to correct for head movement. Unwarping was used to correct for
the interaction of susceptibility artifacts and head movement. After realignment
and unwarping, the signal measured in each slice was shifted relative to the
acquisition time of the middle slice using a sinc interpolation in time to correct
for their different acquisition times. Volumes were then normalized into standard
stereotaxic anatomical MNI-space by using the transformation matrix calculated
from the first EPI-scan of each subject and the EPI-template. The default settings
for normalization in SPM2 with 16 non-linear iterations and the standard EPI-
template supplied with SPM2 were used. Afterwards, the normalized data with
a resliced voxel size of 4 mm × 4 mm × 4 mm were smoothed with a 10 mm
FWHM isotropic Gaussian kernel to accommodate intersubject variation in brain
anatomy. The time series data were high-pass filtered with a high-pass cut-off
of 1/128 Hz. The first-order autocorrelations of the data were estimated and
corrected for.
For each subject individually, responses during the balanced conditions were
sorted into taxonomic matches and thematic matches. These individual data were
used for modeling the expected hemodynamic response at stimulus onset for each
event-type (taxonomic match in the balanced condition, thematic match in the
balanced condition, taxonomic match in the bias condition, thematic match in
the biased condition, as well as baseline trials). The hemodynamic response was
modeled by two response functions, a canonical hemodynamic response function
(HRF; Friston et al., 1998) and its temporal derivative. The temporal derivative
was included in the model to account for the residual variance resulting from
small temporal differences in the onset of the hemodynamic response, which is
not explained by the canonical HRF alone. The functions were convolved with
the event-train of stimulus onsets to create covariates in a general linear model.
Subsequently, parameter estimates of the HRF regressor for each of the different
conditions were calculated from the least mean squares fit of the model to the
time series. Parameter estimates for the temporal derivative were not further
considered in any contrast.
An SPM2 random-effects group analysis was performed by entering param-
eter estimates for all conditions into a within-subject one-way ANOVA. We
expected the differences between processing of taxonomic as opposed to the-
matic categories to be quite small. Still, we were interested in studying effects
across the whole brain instead of restricting the search volume to pre-defined
regions of interest. Therefore, we chose to employ Monte-Carlo simulation of
the brain volume to establish an appropriate voxel contiguity threshold (Slotnick,
Moo, Segal, & Hart, 2003). This correction has the advantage of higher sensitiv-
ity to smaller effect sizes, while still correcting for multiple comparisons across
the whole brain volume. Assuming an individual voxel type I error of p < 0.001,
a cluster extent of 10 contiguous resampled voxels was indicated as neces-
sary to correct for multiple voxel comparisons at p < 0.05. The reported voxel
coordinates of activation peaks were transformed from MNI space to Talairach
and Tournoux (1988) atlas space by non-linear transformations (www.mrc-
cbu.cam.ac.uk/Imaging/mnispace.html). This was done to be able to use the
Talairach and Tournoux (1988) atlas to identify the anatomical brain region for
the activation peaks.
2. Results
2.1. Behavioral data
2.1.1. Choice data
In biased conditions where only one of the two matches could
form a category with the target, subjects made this choice cor-
rectly 97% (S.D. = 3.8) of the time for the thematically biased
condition and 95% (S.D. = 4.3) of the time for the taxonomi-
cally biased condition with no significant difference between
the two (p = 0.09), as revealed by a t-test (t-tests were also
used for other comparisons in the behavioral data section). Only
correct choices were included in the analysis of the imaging
data. In the balanced condition where each one of the matches
could form either a thematic or a taxonomic category with
the target, subjects made a thematic choice just as often as
they made a taxonomic choice (Mthem = 50.02%, S.D. = 10.19;
Mtax = 49.98%, S.D. = 10.19, p = 0.49). All but three subjects
have shown this choice pattern (within one standard deviation
above and below 50%), two out of the three remaining sub-
 O. Sachs et al. / Neuropsychologia 46 (2008) 409–418 413

Table 2
Activation peaks for the biased conditions with their localization
Anatomical region BA Coordinates z-Value No. voxels

x y z

Thematically biased condition > baseline

Inferior frontal gyrus (L)* 47, 9 −51 [−52] 21 [20] 25 [28] 6.26 275
Middle temporal gyrus (L)* 21 −59 [−60] −35 [−36] 2 [0] 5.04 88
Cuneus (L) 18 −8 [−8] −84 [−88] 23 [20] 3.85 59
Angular gyrus (L)* 39 −32 [−32] −56 [−60] 36 [36] 4.42 40
Precentral gyrus (L) 6 −48 [−48] −2 [−4] 44 [48] 4.69 29
Parahippocampal gyrus (L)* 18 −12 [−12] −47 [−48] 2 [0] 3.94 22
Cuneus (R)* 18 16 [16] −97 [−100] 9 [4] 4.00 21
Medial frontal gyrus (L)* 6 0 [0] −8 [−12] 63 [68] 3.94 15
Lingual gyrus (R)* 18 12 [12] −50 [−52] 3 [100] 3.61 11
Taxonomically biased condition > baseline
Inferior frontal gyrus (L)* 47, 45 −51 [−52] 20 [20] 21 [24] 6.94 400
Angular gyrus (L)* 39, 7 −32 [−32] −56 [−60] 36 [36] 4.41 59
Medial frontal gyrus (L)* 6 0 [0] 6 [4] 51 [56] 3.91 55
Cuneus (R)* 18 20 [20] −97 [−100] 9 [4] 4.91 40
Parahippocampal gyrus (L)* 19 −12 [−12] −47 [−48] 2 [0] 4.43 38
Lingual gyrus (L) 17 −16 [−16] −94 [−96] −5 [−12] 4.66 35
Middle temporal gyrus (L)* 21 −63 [−64] −43 [−44] 2 [0] 3.99 27
Middle frontal gyrus (R) 9 51 [52] 21 [20] 25 [28] 3.82 26
Lingual gyrus (R)* 18 16 [16] −50 [−52] 3 [0] 3.83 14
Thalamus (L) −24 [−24] −27 [−28] −2 [−4] 4.05 13
Insula (R) 13 40 [40] 16 [16] 3 [4] 3.86 12

Significance level and the size of the respective activation cluster (number of voxels) at p < 0.05, corrected. Coordinates are listed in Talairach and Tournoux (1988)
atlas space with MNI coordinates in []. BA is the Brodmann area nearest to the coordinate and should be considered approximate. Areas that are present in both
contrasts are marked with asterisk (*).

jects had a taxonomic preference (72% vs. 28% and 63% vs.
37%) and one subject showed a thematic preference (66% vs.
34%). These subjects were not excluded from further analy-
ses since it was expected that some participants may be biased
in one or other direction, reflecting a normal variability in the
population.
2.1.2. Reaction time data
Reaction time was measured from the moment the two
matches were presented till the subject made a correct response.
In the biased conditions it took subjects on average 1154 ms
(S.D. = 179 ms) to make a choice in the thematically biased
condition and 1191 ms (S.D. = 184 ms) to make a choice in
the taxonomically biased condition. As in the choice data,
there was no significant difference between the two (p = 0.29).
A similar pattern was observed in the balanced condition. It
took subjects on average 1189 ms (S.D. = 165 ms) to make a
thematic choice and 1186 ms (S.D. = 172 ms) to make a tax-
onomic choice. Again, there was no significant difference
between the time it took to make the two types of choices
(p = 0.49).
2.2. Imaging data
2.2.1. Thematically and taxonomically biased condition
Comparing the activation corresponding to the correct the-
matic choices in the thematically biased condition against the
baseline, we found the largest cluster of activation (275 vox-
els) in the left inferior frontal gyrus. Other areas of activation
included left middle temporal gyrus, left and right cuneus, left
angular and precentral gyrus, left parahippocampal gyrus and
left medial frontal gyrus. Finally, there was also a small area of
activation in the right lingual gyrus.
Very similar areas were recruited for correct taxonomic
choices in the taxonomically biased condition. The largest clus-
ter of activation (400 voxels) was again in the left inferior frontal
gurus spreading into left medial frontal gyrus. Left angular
gyrus, right cuneus, left parahippocampal gyrus and left mid-
dle temporal gyrus were activated similarly to the thematically
biased condition. However, lingual gyrus was activated in this
condition bilaterally, and there were two more significant clus-
ters of activation in the right hemisphere—one in the right middle
frontal gyrus and one in right insula. Finally, left thalamus was
activated as well. For exact coordinates, z-statistics and cluster
sizes see Table 2.
None of the differences in the areas recruited in the processing
of isolated thematic and taxonomic categories remained signifi-
cant in the subtraction of thematically biased condition from the
taxonomically biased condition or the other way round.
2.2.2. Balanced condition, thematic choice
For the analysis of the imaging data from the balanced tri-
als all responses were grouped according to the type of the
response—thematic or taxonomic and analyzed as separate con-
ditions.
Comparing the activation corresponding to the thematic
choice in the balanced condition to the baseline, we found five
clusters of activation again located in the left inferior frontal
414 O. Sachs et al. / Neuropsychologia 46 (2008) 409–418

Fig. 1. The areas activated in the balanced cross-categorization condition. Panel A: Regions activated for the contrast of thematic choices in the balanced condition
minus baseline. Panel B: Regions activated for the contrast of taxonomic choices in the balanced condition minus baseline. Panel C: Regions activated for the contrast
of taxonomic choices in the balanced condition minus thematic choices in the balanced condition. The panels have different scaling and scaling steps. The activation
map is shown overlaid onto selected sections of a canonical single-subject T1-weighted volume. The position of selected sections corresponds to the biggest clusters
of activation corrected at p < 0.05 across the whole brain. Scale bar shows t-values for contrasts.

gyrus, spreading to the left superior frontal gyrus. Left middle
temporal gyrus and right cuneus were activated as in the themat-
ically biased condition. However, there was also a small area of
activation in the right hemisphere–right inferior frontal gyrus.
See Fig. 1A and Table 3 for a summary of activations in this
condition.
2.2.3. Balanced condition, taxonomic choice
Comparing the activation corresponding to the taxonomic
choice in the balanced condition to the baseline, the main and
biggest clusters were again in left inferior frontal gyrus (459
voxels) and in the left middle temporal gyrus (170). However,
the area of activation in the right inferior and middle frontal
gyrus was larger than in the thematic choice (118 voxels com-
pared to 11 voxels). Except for the activation of right thalamus,
all other areas of activation corresponding to the taxonomic
choice in the balanced trials were limited to the left hemisphere:
left medial frontal gyrus, left precuneus, left cuneus, left lin-
gual gyrus and left parahippocampal gyrus (see Fig. 1B and
Table 3).
 O. Sachs et al. / Neuropsychologia 46 (2008) 409–418 415

Table 3
Activation peaks for the balanced condition with their localization
Anatomical region BA Coordinates z-Value No. voxels

x y z

Thematic choice > baseline

Inferior frontal gyrus (L)* 45, 47 −52 [−52] 24 [24] 21 [24] 6.71 478
Middle temporal gyrus (L)* 21 −59 [−60] −39 [−40] −1 [−4] 5.52 144
Superior frontal gyrus (L) 6 −4 [−4] 18 [16] 47 [52] 5.45 121
Cuneus (R) 18 20 [20] −97 [−100] 1 [−4] 4.41 13
Inferior frontal gyrus (R)* 47 48 [48] 19 [20] −4 [−4] 3.82 11
Taxonomic choice > baseline
Inferior frontal gyrus (L)* 47, 45 −51 [−52] 24 [24] 21 [24] 7.41 459
Middle temporal gyrus (L)* 21 −59 [−60] −35 [−36] 2 [0] 6.12 170
Inferior frontal gyrus (R)* 9, 45 48 [48] 24 [24] 21 [24] 4.81 118
Medial frontal gyrus (L) 6 0 [0] 14 [12] 47 [52] 5.01 114
Precuneus (L) 7, 19 −36 [−36] −68 [−72] 40 [40] 4.68 101
Cuneus (L) 17, 18 −16 [−16] −77 [−80] 15 [12] 3.86 38
Thalamus (R) 4 [4] −8 [−8] 4 [4] 3.76 37
Lingual gyrus (L) 17 −16 [−16] −94 [−96] −5 [−12] 5.52 26
Parahippocampal gyrus (L) 19 −16 [−16] −47 [−48] 2 [0] 4.55 20

Significance level and the size of the respective activation cluster (number of voxels) at p < 0.05, corrected. Coordinates are listed in Talairach and Tournoux (1988)
atlas space with MNI coordinates in []. BA is the Brodmann area nearest to the coordinate and should be considered approximate. Areas that are present in both
contrasts are marked with asterisk (*).

Table 4
Activation peaks for taxonomic choices in balanced condition > thematic choices in balanced condition with their localization
Anatomical Region BA Coordinates z-Value No. voxels

x y z

Precuneus (L) 7 −8 [−8] −68 [−72] 40 [40] 4.01 24

Middle frontal gyrus (R) 46 48 [48] 28 [28] 24 [28] 3.96 22
Thalamus (L) −20 [−20] −16 [−16] 4 [4] 4.14 14

Significance level and the size of the respective activation cluster (number of voxels) at p < 0.05, corrected. The opposite contrast (thematic choices > taxonomic
choices) revealed no difference in activation. Coordinates are listed in Talairach and Tournoux (1988) atlas space with MNI coordinates in []. BA is the Brodmann
area nearest to the coordinate and should be considered approximate.

2.2.4. Comparing taxonomic and thematic choice in the
balanced condition
When balanced trials where the thematic choice was made
were subtracted from balanced trials where the taxonomic choice
was made, three significant clusters of activation were found: in
left thalamus, right middle frontal gyrus and left precuneus (see
Fig. 1C and Table 4). All of these areas were also present in
the activations for the taxonomic choice of the balanced condi-
tion compared to the baseline. The opposite contrast (thematic
choice > taxonomic choice) revealed no differences.
Since the activations that were found in the taxo-
nomic > thematic contrast were relatively small, we investigated
them more thoroughly to rule out that the differences that we
found are not due to deactivations in the thematic condition
compared to the baseline, but represent true increases of the
BOLD response in the taxonomic condition. By plotting param-
eter estimates for each one of the conditions we established that
in all three clusters of activation there was an increase of sig-
nal in the taxonomic condition compared to the baseline and to
the thematic condition (see Fig. 2). It is worth noting that the
parameter estimate for the taxonomic condition in the thalamus
is weaker than in other regions. However, the overall pattern
across conditions in the thalamus is qualitatively (if not quan-
titatively) very similar to the patterns in the right MFG and the
precuneus. All of these regions have increased activation for the
taxonomic choice and a deactivation for the thematic choice.
3. Discussion
The main objective of this study was to (1) compare the neural
correlates of thematic and taxonomic categories and (2) to inves-
tigate the issue of cross-categorization. In response to the first
objective we found that, when investigated in isolation, thematic
categories recruited the same cortical network as the processing
of “classical” similarity-based taxonomic categories: left infe-
rior frontal, middle temporal and occipital regions that many
other studies have found to be implicated in the processing of cat-
egories (Devlin et al., 2002; Perani, Schnur, et al., 1999; Spitzer
et al., 1998; Tyler et al., 2003; Vitali et al., 2005). It is worth men-
tioning that unlike thematic categories, constructing taxonomic
categories involved activation in the right middle frontal gyrus
and in the right insula, although this difference was not signifi-
cant after the subtraction of thematic trials from the taxonomic
ones.
416 O. Sachs et al. / Neuropsychologia 46 (2008) 409–418
Fig. 2. Parameter estimates for the contrast of taxonomic choices in the bal-
anced condition minus thematic choices in the balanced condition. Plots of
mean parameter estimates with ± standard error showing response amplitude in
precuneus (top), middle frontal gyrus (center) and thalamus (bottom) for each
condition: thematically biased, taxonomically biased, thematic choice in the
balanced condition, taxonomic choice in the balanced condition and baseline.
Labeling of the y-axis is in arbitrary units.
Investigating the issue of cross-categorization, the second
objective of our study, has shown that there is no difference
in the reaction time or choice patterns when subjects had to
pick either a taxonomic or a thematic match to make a category.
When subjects had the free choice they were just as likely to
form a thematic category in the presence of a taxonomic alterna-
tive as they were to form a taxonomic category in the presence
of a thematic alternative. This pattern of results fits with the
imaging data from the biased conditions showing no differences
between thematic and taxonomic categories when they have no
competing alternatives. However, the brain activation data sug-
gest that making a subjective taxonomic choice in the presence
of a thematic alternative requires the recruitment of an additional
cortical network, i.e. thalamus, right middle frontal gyrus and
left precuneus. In contrast, making a thematic choice in the pres-
ence of a taxonomic alternative did not lead to the recruitment
of this network or any additional brain regions. This finding
supports our hypothesis of increased effort in the processing of
taxonomic categories that can be reflected in the involvement
of the right-hemispheric regions. Moreover, in line with our
predictions, this difference between brain areas involved in the
construction of taxonomic and thematic categories became sig-
nificant only when the two categories were pitted against each
other in the cross-categorization condition. Our results are all
the more interesting considering that the stimuli that subjects
saw while making both types of categories were identical. The
only difference is in the choice they made—either matching
objects thematically or taxonomically according to their subjec-
tive preference. The fact that matching objects taxonomically
required additional recruitment of the thalamus which is asso-
ciated with rule-based categorization (Grossman, Smith, et al.,
2002) or selective attention (Frith & Friston, 1996) suggests that
constructing taxonomic categories might require more effort.
Indeed, thematic concepts that are highly associated might be
activated with less effort than taxonomic concepts that are related
at the superordinate level and share a more abstract relationship.
Several categorization studies have also proposed that detec-
tion of similarity between taxonomically related entities takes a
deeper level of analysis (Lin & Murphy, 2001; Mandler, 1983).
The activation of right frontal cortex that was found to be impli-
cated in processing abstract nouns and verbs (Grossman, Kenig,
et al., 2002; Perani, Cappa, et al., 1999) supports the idea that
forming taxonomic categories requires assessing a more abstract
relationship between the target and the match word. This process
can also be more effortful and require the additional recruitment
of the right hemisphere. The greater activation of precuneus in
the taxonomic choice condition may indicate that additional
mental imagery processing was necessary to decide if taxo-
nomic matches were similar enough to the target to make up
a category. The activation of precuneus could also be associated
with increased demands set on the retrieval of episodic mem-
ory and contextual associations that were necessary to make the
taxonomic choice (Fletcher et al., 1995; Lundstrom, Ingvar, &
Petersson, 2005).
Outside of the competition or cross-categorization context,
processing of taxonomic and thematic categories in isolation
involved markedly similar brain regions—left inferior frontal
gyrus, left middle temporal gyrus, cuneus, precuneus, lingual
and angular gyri. Many previous studies have found these areas
to be implicated in explicit semantic tasks, semantic processing
and categorization. More specifically, the left inferior frontal
cortex is typically associated with lexical retrieval of seman-
tic knowledge, executive aspects of semantic processing, such
 O. Sachs et al. / Neuropsychologia 46 (2008) 409–418
as semantic working memory, semantic search, or comparing
semantic concepts in working memory (Bookheimer, 2002).
Since most of our stimuli were artifacts, the involvement of mid-
dle temporal gyrus, usually activated for tools or tool-associated
actions is not surprising either (Beauchamp, Lee, Haxby, &
Martin, 2002; Martin, Ungerleider, & Haxby, 2000; Tyler et al.,
2003). Finally, the activation of such visual areas as left and right
cuneus, precuneus and lingual gyrus has been previously asso-
ciated with the involvement of a visualization strategy during
semantic tasks and inspecting visual images (Fletcher, Shallice,
Frith, Frakowiak, & Dolan, 1996; Perani, Schnur, et al., 1999;
Vitali et al., 2005). This function can be equally important for
forming and evaluating both thematic and taxonomic categories
made up of artifacts.
Several caveats must be kept in mind regarding our results.
First, our results are necessarily driven by the semantic pro-
cesses involved in completing the cognitive task that we used.
These task demands can interact with the semantic knowl-
edge and influence the neural activation pattern that we found
(Grossman et al., 2006). Second, thematic categories often share
more than one type of thematic relations and are often con-
founded by a spatial relationship or a part-whole relationship
(that is also functional—otherwise there would be no need for
the part). Although in our stimuli development we controlled
for this factor and made sure that our thematic categories share
only either a functional or a part-whole relationship, there is
still a possibility that not all functional relations are 100%
pure.
In conclusion, our results show that on the neural level exter-
nal correlations that build thematic relations between objects
are processed in a way that is very similar to the process-
ing of similarity-based feature correlations. External correlation
between objects like closet and dress has obviously compen-
sated for their perceptual dissimilarity to lead to the same
neural activation pattern as the one elicited by closet and
dresser—members of the same taxonomic category furniture.
Thus, our results have implications for the general question
of semantic knowledge organization in the brain. Taxonomic
and thematic groupings seem to be two equally important ways
of recognizing commonalities among objects. It can be taken
as evidence that classical taxonomic categorization is not the
only way of organizing semantic knowledge and that thematic
associations create equally salient conceptual groups. Whether
such groups can be called categories in neuroimaging research
is an open question—our results suggest that there is no rea-
son why they should not. While the current study only begins
to tackle this issue, further research of thematic and other
ways of semantic organization can improve our understanding
of the fundamental issue motivating this research–knowledge
representation in the brain and the principles of conceptual
organization.
Acknowledgements
The study was supported by a grant from the Interdisciplinary
Center for Clinical Research “BIOMAT” within the Faculty of
Medicine at the RWTH Aachen University (IZKF VV N68)
417
and by the START-Program of the Faculty of Medicine, RWTH
Aachen.
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