Introduction
Patients with dorsolateral prefrontal cortex (DLPFC)
lesions commonly experience deficits on CR tasks
that involve maintaining or shifting hypotheses in
working memory in response to feedback.
1–4
Similar
executive function deficits have been detected in
patients with subcortical lesions involving the
basal ganglia,
5,6
thalamus
7,8
and cerebellum.
9–11
These
lesion studies suggest that common, yet apparently
disparate, neural systems may mediate CR task
performance.
Non-human primate studies have demonstrated
the existence of multiple parallel frontal-subcortical
circuits.
12
One such circuit involving the DLPFC,
basal ganglia (caudate nucleus) and thalamus (ante-
rior and mediodorsal nuclei) has been implicated in
such complex behaviors as working memory and
rule-based learning.
13
More recently, anatomical
interconnections have been demonstrated between
the DLPFC and the lateral cerebellum via the ventral
segment of the dentate nucleus and the mediodorsal
nucleus of the thalamus.
14
The present study was designed to examine the
possible participation of these frontal-subcortical
circuits in intact humans while performing a CR task.
Whole-brain fMRI was used to detect regional
changes in blood oxygenation associated with
increased neural activity.
15,16
In clinical settings, CR
is typically assessed with the Wisconsin Card Sorting
Task (WCST). The WCST can not be easily adapted
to fMRI scanning, since problem solving does not
occur in response to a finite number of time-locked
stimulus presentations. Additionally, the ratio of
positive to negative feedback is not fixed across
subjects and the solutions are defined a priori in a
fixed order, thereby creating problems for repeat
testing. To circumvent these limitations, we used an
alternative task
17
with demonstrated sensitivity to CR
deficits in patients with prefrontal
2
and basal
ganglia
5,18
lesions.
Subjects and Methods
Eleven normal volunteers (four males and seven
females; ages 19–45 years, mean 29 years; mean
education 16.5 years) participated in the study. All
subjects were strongly right-handed (mean laterality
quotient 87.8 on the Edinburgh Handedness
Inventory
19
). Potential subjects were excluded if they
Brain Imaging
1
1
1
1
1
p
© Rapid Science Publishers Vol 8 No 8 27 May 1997 1987
LESIONS involving the dorsolateral prefrontal lobes may
produce deficits on conceptual reasoning (CR) tasks in
humans. Such deficits can also occur with subcortical
lesions involving the basal ganglia, thalamus, or cere-
bellum, suggesting a common, yet widespread, neural
network supporting this executive function. Here we
report the results of a whole brain functional magnetic
resonance imaging (fMRI) experiment in healthy volun-
teers while performing a CR task. Compared to a senso-
rimotor control condition, the CR task resulted in
discrete subcortical activation sites primarily involving
the right basal ganglia, right thalamus and left lateral
cerebellum. Cortical activation was present in multiple
systems, including the dorsolateral prefrontal and infe-
rior frontal/insular areas; posterior parietal, superior
extrastriate, and premotor areas; inferior extrastriate
and middle temporal regions; and midline pre-supple-
mentary motor and anterior cingulate regions. Our find-
ings provide strong evidence that CR is mediated by
interacting neural systems involving the cerebral cortex,
basal ganglia, thalamus, and cerebellum.
Key words: Basal ganglia; Cerebellum; Conceptual
reasoning; Functional magnetic resonance imaging (fMRI);
Parietal cortex; Prefrontal cortex; Thalamus
Functional MRI
evidence for subcortical
participation in
conceptual reasoning
skills
Stephen M. Rao,
1,2,3,4,CA
Julie A. Bobholz,
1
Thomas A. Hammeke,
1,3
Allyson C. Rosen,
1
Scott J. Woodley,
1
Joseph M. Cunningham,
1
Robert W. Cox,
4
Elliot A. Stein,
3,4
and Jeffrey R. Binder
1,2,4
Departments of
1
Neurology,
2
Cellular Biology
and Anatomy, and
3
Psychiatry and Behavioral
Medicine, and
4
Biophysics Research Institute,
Medical College of Wisconsin, 9200 West
Wisconsin Avenue, Milwaukee, WI 53226, USA
CA,1
Corresponding Author and Address
NeuroReport 8, 1987–1993 (1997)