195
Traditional theories of working memory and executive function,
when mapped in straightforward ways into the neural domain,
yield predictions that are only partly supported by the recent
neuroimaging studies. Neuroimaging studies suggest that
some constituent functions, such as maintaining information in
active form and manipulating it, are not discretely localized in
prefrontal regions. Some hypothesized executive processes,
such as goal management, have effects in several cortical
regions, including posterior regions. Such results suggest a
more dynamic and distributed view of the cortical organization
of working memory and executive functions.
Addresses
Center for Cognitive Brain Imaging, Department of Psychology,
Carnegie Mellon University, Pittsburgh, PA 15213, USA
Correspondence: Patricia A Carpenter; e-mail: carpenter+@cmu.edu
Current Opinion in Neurobiology 2000, 10:195–199
0959-4388/00/$ — see front matter
© 2000 Elsevier Science Ltd. All rights reserved.
Abbreviations
BA Brodmann’s area
DLPFC dorsolateral PFC
fMRI functional magnetic resonance imaging
PET positron emission tomography
PFC prefrontal cortex
SPECT single photon emission computed tomography
Introduction
Executive processes and working memory have proven to
be a fruitful test bed for neuroimaging studies of cognition.
Executive processes are implicated in complex cognition,
such as novel problem solving, which entails identifying
and coordinating the steps to a new goal, evaluating the
intermediate outcome, and modifying the plan as needed.
Executive processes are also associated with task-set con-
trol, modifying behavior as appropriate in light of changes
in the environment, such as inhibiting prepotent or previ-
ous responses. Executive processes are also associated with
the functions that are impaired as a consequence of frontal
lesions, particularly those related to the implementation of
schemas that organize behavior over time [1,2]. Working
memory has been operationalized primarily as the process-
es and structures that keep information available over a
relatively short time, such as postulated verbal and spatial
peripheral buffers [3,4
•
]. In this standard perspective,
executive processes manipulate the contents of the work-
ing memory buffers.
In the first part of this article, we describe how recent
advances in functional neuroimaging have been framed in
light of some classic questions about the organization of
working memory and executive processes. In the second
part, we examine how these same neuroimaging results
suggest that the classic questions might be reconstrued,
and then suggest some possible implications of these alter-
native hypotheses.
Classic issues
Perhaps the major theoretical issue concerning executive
processes is whether discrete regions (modules) of the pre-
frontal cortex (PFC) are dedicated to particular operations
and, if so, the characterization of those processes, including
their domain specificity [5
••
]. For example, Owen,
Petrides and their colleagues [6,7] proposed that the mid-
ventrolateral region (Brodmann’s area [BA] 45/47) supports
the organization of response sequences based on informa-
tion retrieved from posterior areas, whereas the
mid-dorsolateral region (BA 9/46) supports the active
manipulation or monitoring of information within working
memory. Another proposal by Goldman-Rakic [8] is that
PFC is organized by processing domain into object, spatial
position, and verbal PFC regions.
Another issue concerns the relation between the computa-
tions of the prefrontal regions and those of more posterior
regions in working-memory functions. For example,
Goldman-Rakic [8] proposed that, in working memory
tasks, the PFC regions operate conjointly with posterior
regions as multi-modal domain-specific networks. The
hypothesis that working-memory functions are accom-
plished by large-scale networks overlaps with other
theoretical proposals, including Mesulam’s framework [9].
Although the temporal relations among these various com-
ponents may be critically important [10], they are only
beginning to be addressed by neuroimaging studies [11].
A third issue is how to conceptualize the capacity con-
straints of working memory — a topic that was framed by
Miller’s classic chunking hypothesis [12]. When working
memory is equated with information maintenance, con-
straints can be conceptualized as temporal. When the task
is more complex, such as problem solving, these con-
straints may be conceptualized as limits in the complexity
of the computations or representations [13,14]. For exam-
ple, one proposal is that immediate thought varies in the
amount of concurrent processing demand relative to a sys-
tem’s resources, and that constraints emerge from system
throughput [15].
Neuroimaging results
The localization of working memory versus executive pro-
cessing [6,7] was addressed in a meta-analysis of several
positron emission tomography (PET) and functional mag-
netic resonance imaging (fMRI) studies of verbal and
spatial n-back tasks [16
•
,17
•
]. In an n-back task, sequen-
tially presented items (letters, spatial positions, or
Working memory and executive function: evidence from
neuroimaging
Patricia A Carpenter, Marcel Adam Just and Erik D Reichle