Introduction
Working memory refers to a brain system that
provides temporary storage and manipulation of
the information necessary for such complex cogni-
tive tasks as language comprehension and learning.
This brain system requires the simultaneous storage
and processing of information. The function of
the phonological loop, which stores and rehearses
speech-based information
1
is one of the working
memory systems. Broca’s area (Brodmann 44/45 in
the left inferior frontal lobe) was found to be involved
in the phonological loop in studies using positron
emission tomography (PET)
2
and in the covert
speech tasks using functional magnetic resonance
imaging (fMRI).
3–5
Recently, in the spatial working
memory task, the sustained activity after the visual
presentation was detected in Broca’s area using
fMRI.
6
The activation in Broca’s area in the spatial
task may have been due to verbalization that the
subjects may have used in the spatial memory task.
Unless this component can be controlled or excluded
one cannot conclude that Broca’s area is involved
in spatial memory. In psychology it is difficult to
define the termination of a ‘sustained’ phonological
process. In particular, in working memory studies
using multiple-task trials during an experiment,
6
it is
possible to alter naturally responses of the memory
processing.
Animal studies involving hippocampal lesions
suggested that the hippocampal formation is involved
in spatial working memory.
7,8
The hippocampal
formation is located in the medial part of the temporal
lobe of the human brain. The hippocampus is
connected directly and indirectly, in both afferent
and efferent directions, with various other parts of
the brain, and thus can be considered the center
of a more extended system.
9
Clinical lesion studies
and neuropathological examinations have shown that
the human hippocampal formation is involved in
learning and memory function.
10,11
Human functional
imaging studies showed activation in the hippocampal
formation during memory task period.
12,13
However,
a sustained response in human hippocampus has not
been observed in any studies. We investigated the
hypothesis that human hippocampus is involved
in working memory. The sensitivity of fMRI, espe-
cially at high fields,
14–17
has advanced sufficiently to
permit the examination of this hypothesis.
In the present study, blood oxygen level depen-
dent (BOLD)
17
based fMRI at 4 Tesla was performed
in conjunction with a word memory task on seven
healthy right-handed volunteers. The echo planar
imaging (EPI) time courses of cerebral responses
during and after a single task period were monitored.
Cognitive Neuroscience and Neuropsychology
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© Rapid Science Ltd Vol 9 No 6 20 April 1998 1041
TEMPORAL behavior of activation associated with the
neural substrate of human memory function was inves-
tigated during and after an auditorily instructed word
memory task using multislice functional magnetic reso-
nance imaging. The hippocampal formation, which is
involved in human memory function, displayed a long-
term sustained response that persisted significantly
(~90 s) beyond the duration of the memory task. This
sustained period was ~two-fold longer than the dura-
tion of the post-task activation observed in auditory
areas and Broca’s area, which are involved in the phono-
logical loop of the verbal working memory. These obser-
vations suggest that the hippocampal memory
processing involves sustained activation in the transi-
tional function for the long-term memory over the
working memory period. NeuroReport 9: 1041–1047
© 1998 Rapid Science Ltd.
Key words: Brain; Cerebral multiphasic sustained response;
Echo-planar imaging (EPI); Functional magnetic resonance
imaging (fMRI); Hippocampus; Humans; Long-term
sustained response; Memory; Network
Human hippocampal
long-term sustained
response during word
memory processing
Toshinori Kato,
CA
Peter Erhard,
Yoshihiro Takayama, John Strupp,
Tuong Huu Le, Seiji Ogawa
1
and Kamil Ugurbil
Center for Magnetic Resonance Research,
Department of Radiology, University of
Minnesota, 385 East River Road, Minneapolis,
MN 55455, USA;
1
Bell Laboratories, Lucent
Technologies, Murray Hill, NJ 07974, USA
CA
Corresponding Author
Website publication 27 March 1998 NeuroReport 9, 1041–1047 (1998)