WINDOWS TO THE BRAIN

Working Memory Models: Insights From Neuroimaging

Robert D. Shura, Psy.D., Robin A. Hurley, M.D, and Katherine H. Taber, Ph.D.

FIGURE 1. The Atkinson and Shiffrin Model of memory is diagramed in the gray panel. Sensory information is perceived and suppressed via selective
attention processes, then briefly held in the limited-capacity modality-specific sensory register. Information enhanced by selective attention enters
into short term memory (STM). The process of rehearsal (maintenance) can sustain information in STM. Information in STM is automatically encoded
(stored, consolidated) into long term memory (LTM).1 Although flawed, this model continues to provide a simple and easy approach to understanding
memory. Assumptions that proved to be inconsistent with research included that any information in STM will transfer to LTM and that STM is required

for access to LTM. Additionally, the low level of inter-

ference when concurrent tasks utilized different senses
indicated that STM was not a unitary store. The multi-
component model of working memory (WM) essentially
reconceptualized STM by expanding it into more than one
component (yellow) and adding a set of active processes.2
Three buffers support short term processing of visual-
spatial (VSSP), auditory-verbal (PL), and multimodal (EB)
information, under the control of the central executive
(CE, green). The CE directs all other aspects of memory
processes and attention. CE processes overlap with tasks
commonly referred to as executive functions.

FIGURE 2. A recent series of studies utilized voxel-based lesion-symptom mapping (each voxel designated as intact or lesioned) to delineate the
neural architecture underlying the major types of intelligence (general, fluid) and major components of WM (domain-specific information mainte-
nance, information monitoring, information manipulation, cognitive flexibility).3–6 Visually summarized are the cortical areas most associated with
fluid intelligence and several aspects of WM in the final model.6
Fluid intelligence (red) was associated with a primarily right
lateralized network of frontal and parietal areas. Domain
specific WM lateralized by domain, with verbal/numeric WM
(green) primarily associated with areas in the left hemisphere
and spatial WM (blue) primarily associated with areas in the
right hemisphere. Areas associated with information manipu-
lation aspects of WM (yellow) partially overlapped with both
general intelligence (not illustrated) and verbal/numeric WM in
the left hemisphere and fluid intelligence in the right hemi-
sphere. The information monitoring aspect of WM (purple) was
associated with a primarily right hemisphere network of cor-
tical areas, but overlapped the fluid intelligence network only
in inferior parietal cortex.

COVER AND FIGURE 3. Several recent meta-analyses of

functional imaging studies reported bilateral activation of a
network of frontal, parietal, temporal, cerebellar, and thalamic
areas across WM tasks and stimuli types.7–10 Illustrated here are
data from one study.7 Main effect of WM (gold) was associated
with the expected wide spread network of areas (see COVER for
insula and thalamus). Increasing WM load (pink) was associated
with increased activations in almost all the same areas, sug-
gesting a network supporting domain-general executive control
processes. Areas most associated with object identity (green)
were in a similar region of prefrontal cortex but ventral to areas
most associated with object location (blue), supporting the
proposed dorsal/ventral gradient in prefrontal functions.7

neuro.psychiatryonline.org J Neuropsychiatry Clin Neurosci 28:1, Winter 2016


emory, the ability to maintain access to in-
M formation in the absence of sensory input, is
fundamental to all goal-directed behavior. Early
models of memory were grounded in the logic
models of the rising world of computer science and in-
formation theory.11 One popular model (Atkinson and Shiffrin
Model) developed at that time proposed three stages of de-
clarative memory: sensory memory (sense-specific registers),
short term memory (STM), and long term memory (LTM).
Information was lost if not successfully passed to the next
stage (Figure 1).1 Although flawed, this model continues to
provide a simple and easy approach to understanding mem-
ory. One assumption that proved to be inconsistent with re-
search was that there is a single STM system. The low level of
interference when concurrent tasks utilized different senses
indicated that STM was not a unitary store. The multi-
component model of working memory (WM) essentially re-
conceptualized STM by expanding it into more than one
component and adding a set of active processes (Figure 1).2
This model of WM continues to be the most widely used
conceptual map.12–17 The original publication has been cited
nearly 10,000 times in more than 40 years since its initial
presentation.16 Due to the importance of intact WM for daily
functioning, foundational understanding of WM theory, tests,
and recent neuroimaging is of value for both researchers and
clinicians working in neuroscience-related fields.
Cognitive Psychology
The concepts of STM and WM are related, as both represent
processes that provide temporary storage of information
(i.e. maintenance of information for a short period of time in
the absence of perceptual input). A key difference is that
the concept of WM adds processes that support manipula-
tion of the information, providing a basis for goal-directed
behaviors.2,12,13,16,17 The multicomponent model of WM con-
tinues to evolve, but currently includes: the phonological loop
(PL), the visuospatial sketchpad (VSSP), the episodic buffer
(EB), and the central executive (CE) (Figure 1). The PL and
VSSP provide temporary maintenance and manipulation of
domain-specific information under the attentional control of
the CE. The PL provides access to auditory-verbal information,
and is perhaps the most researched aspect of this model. The
VSSP provides access to visual information. The need for two
components was based on research indicating far less in-
terference between different types of domain-specific in-
formation during WM tasks than would be expected if there
were a single component. Other domains of information
(e.g. tactile, smell, taste) are much less studied. The EB
component of WM was added to the model much later to
provide the capacity to integrate information across domains
and temporarily maintain and manipulate multidimensional
representations.2 The EB is also thought to be responsible for
the process underlying the cognitive strategy of collapsing
related data together (“chunking” information) to increase at-
tentional capacity. The CE component provides higher-order
J Neuropsychiatry Clin Neurosci 28:1, Winter 2016
SHURA ET AL.
control processing. It has been dissociated into multiple in-
dependent but related processes, such as attentional control,
multitasking, retrieval strategy switching, selective inhibition,
and mental manipulation.2 There are two major views of the
CE. The first conceptualizes the CE from a model of general
intellectual ability; the second views the CE from the per-
spective of information manipulation. It is important to em-
phasize that the multicomponent model of WM is a cognitive
model that does not propose biological substrates. Delineation
of the neural substrates of the WM components is now an area
of active investigation.
Neuropsychology
The concept of intelligence provides a context to understand
other cognitive abilities. Intelligence is generally defined as
general mental capacity. There are numerous theories of in-
telligence, with many including WM as a component. The
Cattell-Horn-Carroll (CHC) model of intelligence, a compre-
hensive and widely accepted theory, is hierarchical.18,19 Gen-
eral intellectual ability (g) encompasses the comprehensive
representation of global intelligence. Within g are various
broad abilities (e.g. fluid intelligence [mental efficiency] and
crystallized intelligence [acquired knowledge]), each of which
can be further segmented into numerous narrow abilities.20
Although the model encompasses a STM component, WM is
not directly incorporated. However, abilities such as general
sequential reasoning (under the fluid intelligence factor) and
memory span (under the general memory and learning factor)
require use of WM. Thus, WM is represented in (but not
actually part of ) the model. Numerous studies have found
moderate to high correlations between WM and g.21 Two
mainstream intelligence tests (Stanford-Binet and the Wechsler
Intelligence Scales [WIS]) directly incorporate WM into their
batteries, comprising a full scale intelligence quotient (IQ score).
The relationship between WM and intelligence is partially
mediated by how intelligence is measured. Of particular note,
the WIS Working Memory Index includes Digit Span, Arith-
metic, and Letter-Number Sequencing, three commonly used
tests in both clinical and research settings. As the WIS are some
of the most popular tests of intellectual ability in current use,
many IQ estimates will inevitably be greatly influenced by WM
ability.
Although conceptualized as an active STM process, WM
tasks are commonly classified as attentional in neuropsy-
chology. One major text introduces WM within the more
global memory paradigm but describes tests of WM as the
combination of storage and manipulation of information.22
WM tests are included among tests of attention, not with
tests of memory. Commonly used WM tests do not directly
correlate to the multicomponent WM model. WM tests are
classified generally, or at most, divided by modality (visual
versus verbal). In general, WM tests can be grossly divided
into two categories: complex span and complex tracking.
Additionally, there is a division between those tests com-
monly used in research and those commonly used clinically.
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WORKING MEMORY MODELS
One study assessed whether research WM tests (operation
span, listening span, and N-back) were consistent with those
used clinically (WIS and Wechsler Memory Scale tests).23
The three research tests and WAIS-III Letter-Number Se-
quencing were the most strongly correlated. The model
containing those four measures provided the best fit to the
latent WM variable. Thus, WIS Letter-Number Sequencing
(a complex span test) may be a common metric for trans-
lation across the settings.
Two commonly used tests of complex tracking are Paced
Auditory Serial Addition Test (PASAT) and N-back. During
the PASAT, single-digit numbers are presented at various
time intervals and the respondent must add each number to
the immediately preceding number, though there are a
number of versions of the test available.24 The PASAT is
frequently included in fixed multiple sclerosis batteries due to
its sensitivity to impairments common to that disorder.25,26
Psychometrically, the task seems to load heavily on processing
speed, attention, and executive functions.27 Although the test
is difficult and many find it aversive, it was reported to be the
third most popular test of attention among neuropsycholo-
gists according to one survey.28 The N-back paradigm is a
category of complex tracking tests that is frequently used in
neuroimaging studies. A series of stimuli (verbal or nonverbal)
are presented, requiring the respondent to signal when the
new stimulus matches the one presented “n” items before.
Thus, this test requires both WM and vigilance. Increasing
the span (or “n”) increases the complexity of the test. The
stimuli can be numbers, letters, shapes, or location of a
stimulus in space.
In contrast, complex span tests focus more on attentional
capacity. A commonly used test is Digits Backward. Strings of
digits are presented with the respondent reciting the digits in
reverse order. On subsequent trials the length of the string
(digit span) is increased. The test is heavily researched, easy to
administer, and a regular subtest in the WIS. Current versions
combine Digits Backward with the simpler Digits Forward
and the Digit Sequencing tests. However, factor analyses in-
dicate that simple span and complex span tasks are related but
not equivalent.29 Digits Backward is an ideal test for use as
part of a bedside exam due to its ease and speed of adminis-
tration. A Digits Forward span should be completed first. The
typical forward span for adults is seven digits; the typical
backward span is four or five digits correct.22,30,31 Numerous
other complex span tasks are in use (e.g. reversed spelling,
mental arithmetic, WIS Letter-Number Sequencing). The
Corsi block tapping test, has been computerized, making it
useful for neuroimaging studies.32 The stimulus is an array of
3D blocks arranged in space. The examiner taps the blocks in
a particular order, with the respondent then tapping the
blocks in the same (or reverse) order. On subsequent trials,
the number of taps increases (similar to digit span). A series of
dual procedure and suppression experiments using the Corsi
forward and backward paradigms found that the Corsi test
employs VSSP and CE but not PL, suggesting that the Corsi
test fits well with the multicomponent model of WM.33 The
2 neuro.psychiatryonline.org
importance of this finding is that most WM tests employ PL
and CE, with fewer tests available to elicit VSSP.
Neurobiology
The classic approach for identifying the neural architecture
supporting particular aspects of function is the study of pa-
tients with focal brain lesions.2,13 Studies that establish cor-
relations between morphological and performance variables
(i.e. lesion-deficit analyses) provide a basis for identifying
areas of the brain that support specific abilities. As intelli-
gence and WM are strongly interrelated, the most informative
studies include a sufficiently broad range of performance
measures to support assessment of both. A recent series of
studies utilized voxel-based lesion-symptom mapping (each
voxel designated as intact or lesioned) to delineate the neural
architecture underlying the major types of intelligence (gen-
eral, fluid) and major components of WM (domain-specific
information maintenance, information monitoring, informa-
tion manipulation, cognitive flexibility).3,4,6 Both general and
fluid intelligence were associated with widely distributed
networks of cortical areas (ventrolateral and dorsolateral
prefrontal [PFC], primary motor and somatosensory, inferior
and superior parietal, and superior temporal cortices) and
associated major white matter tracts. General intelligence was
primarily left lateralized, whereas fluid intelligence was pri-
marily right lateralized (Figure 2).3,6 The areas associated
with general intelligence were similar in locations but greater
in extent than reported in a previous study.34 Domain specific
WM lateralized by domain (Figure 2), with verbal/ numeric
primarily associated with areas in the left hemisphere (dor-
solateral PFC, superior parietal and superior temporal corti-
ces, angular gyrus) and spatial primarily associated with areas
in the right hemisphere (dorsolateral PFC and superior pa-
rietal cortices).6 Both overlapped with areas associated with
intelligence in dorsolateral PFC and superior parietal corti-
ces.3,6 Cognitive flexibility and the information manipulation
aspect of WM were both associated with primarily left
hemisphere networks of cortical areas (Figure 2). Cognitive
flexibility overlapped areas associated with general intelli-
gence throughout that network.3,4 Some of the left hemi-
sphere areas associated with information manipulation
overlapped with areas associated with general intelligence
(superior parietal and superior temporal cortices), whereas
others (angular gyrus) did not.3,6 In addition, some areas in
the right hemisphere associated with fluid intelligence were
also associated with information manipulation (dorsolateral
PFC). The information monitoring aspect of WM was as-
sociated with a primarily right hemisphere network of cor-
tical areas (ventrolateral PFC, inferior parietal and middle
temporal cortices) that overlapped the fluid intelligence
network only in inferior parietal cortex.6
Studies in healthy individuals are also informative. A
meta-analysis of the association between PFC volume and
thickness and tests of executive functioning found a mod-
erate effect size for volume and a weak effect size for
J Neuropsychiatry Clin Neurosci 28:1, Winter 2016

thickness.35 In the regional analysis, both lateral and medial
PFC volumes were significantly correlated with executive
functioning.35 Similarly, larger gray matter volume was as-
sociated with better executive performance both at the
global (whole brain) and regional (PFC, insular and cere-
bellar clusters) levels in another study.36 A study quantified
several measures of cortical morphology (gray matter vol-
ume, cortical surface area, cortical thickness) to identify
areas supporting aspects of intelligence (fluid, crystalized,
spatial) and basic cognitive processing (WM capacity, ex-
ecutive updating, attention, processing speed).37 Statistical
parametric mapping of gray matter volume indicated that all
three intelligence factors were associated with partially
overlapping clusters in right dorsolateral PFC. Spatial in-
telligence was also associated with clusters in right inferior
temporal cortex (fusiform gyrus) and caudate. WM capacity
was associated with a cluster in right dorsolateral PFC that
overlapped fluid and crystalized intelligence. WM was also
associated with clusters in right primary somatosensory
cortex and left frontal and occipital cortices. Attention was
associated with clusters within frontal, temporal and oc-
cipital cortices.37 A study that estimated size of visual cortex
using both structural (landmark based parcellation) and
functional (retinotopic mapping) measures found that the
size of V1 (first stage of visual processing) correlated strongly
with visual WM capacity.38 As noted by the authors, these
findings suggest that individual differences in the size of
domain-specific cortices may contribute to individual dif-
ferences in domain-specific WM performance.
Functional imaging provides a range of methods for
probing the neural architecture supporting particular as-
pects of brain function. Task-activated functional imaging
studies are most commonly used to identify areas of the
brain that are more active during performance of a particular
task. Most functional imaging studies utilize functional
magnetic resonance imaging (fMRI), which provides an in-
direct measure of neuronal activity based on local changes in
blood flow. The majority of studies compare signal intensity
of relatively large groups of voxels (cluster or region) across
conditions (univariate analyses). Multivariate pattern ana-
lytic (MVPA) techniques have recently been introduced that
utilize machine learning techniques to identify patterns of ac-
tivated voxels that encode task-related information. Compar-
isons across types and phases of tasks have the potential to
provide insight into the brain areas supporting different
components of WM. One example is comparing tasks that
require similar manipulations of information but utilize
different information domains. Processes related to main-
taining domain-specific information (e.g. visual, verbal)
might be expected to localize differently, whereas domain-
general processes (e.g. remembering rules or goals, ignoring
distractors) might be expected to localize similarly.
Several recent meta-analyses of functional imaging
studies reported bilateral activation of a network of frontal,
parietal, temporal, cerebellar, and thalamic areas across WM
tasks (e.g. N-back, Sternberg, delayed matching to sample,
J Neuropsychiatry Clin Neurosci 28:1, Winter 2016
SHURA ET AL.
prospective memory) and stimuli types (Figure 3).7–10 The
meta-analysis that compared by gender also found a few
differences, such as greater activation of limbic areas in
females and parietal areas in males.8 Increasing WM load
(increasing task difficulty or adding the presence of dis-
tractors) was associated with increased activations in almost
all areas (exceptions were rostral lateral PFC and thalamus),
suggesting a network supporting domain-general executive
control processing (Figure 3).7 A study utilizing MVPA re-
ported successful cross-modal (visual WM task, verbal WM
task) prediction of WM-load-sensitive areas, further sup-
porting a common domain-general neural substrate.39 A
study utilizing graph-based modularity analysis found that
WM tasks activated multiple areas within the executive
control (lateral PFC, superior parietal lobule) and salience
(anterior insular and dorsal cingulate cortices) networks.40
Connectivity within the executive control network and be-
tween the executive control network and salience network
increased with increasing WM load.40
Some areas, most commonly within lateral PFC and pa-
rietal cortex, display sustained activation during the delay
period in WM tasks.12,13,17,41,42 One interpretation has been
that this represents domain specific information being
maintained in an active state. However, fMRI studies uti-
lizing MVPA have reported that patterns of activity in lateral
PFC contain higher order information required for task
performance (e.g. task rules, stimulus-response mapping,
categories based on multiple domains of information) rather
than domain specific information.12,13,41,42 These findings
support the alternative view that sustained activity in lateral
PFC and parietal cortex indicates allocation of cognitive
resources (e.g. attention) to maintain domain specific in-
formation elsewhere (top-down modulation). Congruent
with this view, MVPA studies identified patterns of activity
in sensorimotor cortices during delay periods consistent
with population encoding of domain specific information
even in the absence of elevated activity.12,13,41,42 Patterns
were predictive across memory types (WM, LTM), sug-
gesting a common neural substrate for WM and LTM. An
fMRI study using MVPA reported dynamic shifts in patterns
of activity when a retrocue (cue presented when the stim-
ulus is no longer available) signaled a change in the stimulus
dimension required for task performance.43 This suggests
that what is encoded is not a static representation of all
information contained in the stimulus, but rather specifically
favors task-relevant aspects.
The multicomponent model of WM is a conceptual
model, informed by and developed based on cognitive psy-
chology research. Advances in identifying neural correlates
of processes that contribute to WM have provided insights
into the underlying neural architecture. For example, in the
multicomponent model WM and LTM are functionally dis-
tinct systems. There is growing evidence from neuroimaging
research, particularly MVPA studies, that WM and LTM are
neuroanatomically colocated. This apparent disparity has
been reconciled in state-based models of WM, which
neuro.psychiatryonline.org 3
WORKING MEMORY MODELS
postulate that these types of memory differ by activation
state rather than location in the brain.12,13,15,17,41 In these
models, attention is the key factor determining the accessi-
bility of memory content. Terminology and number of acti-
vation states varies, but the basic postulates are similar. WM
contains information that is the focus of attention (region of
direct access). Other information ranges from less accessible
(latent WM, activated LTM) to not directly accessible
(LTM). The hierarchical process model also incorporates
timescale and information complexity to distinguish the
nature of memory processing in domain-specific areas from
that occurring in higher order areas.15
Conclusions
WM is a critical cognitive ability that has emerged as an im-
portant domain in comprehensive neuropsychological evalu-
ation. WM can be conceptualized as a set of active processes
in declarative memory systems that occur between the sen-
sory register and stable, theoretically permanent memory
stores (LTM). The multicomponent model of WM was de-
veloped to address aspects of the passive model of STM that
did not match research findings. Conceived of as a component
of general intellectual ability, it is also classified as an atten-
tional skill. Thus, WM has become a complicated and multi-
faceted construct of mental ability. Tests of WM typically
involve either complex span or complex tracking. Many tests
do not translate well to the bedside exam and tests used in
research often differ from those commonly used clinically.
WM tests utilizing mental arithmetic and reverse span are the
most practical to employ in bedside or less-formal evaluation.
Imaging studies frequently place WM abilities in dorsolateral
PFC and in frontoparietal networks. There is a relatively
consistent lateralized effect between the verbal (PL) and
visual-spatial (VSSP) aspects of WM. As imaging studies con-
tinue to inform WM conceptualization, cognitive models will
likely continue to evolve.
AUTHOR AND ARTICLE INFORMATION
Dr. Shura is affiliated with the Mental Health Service Line and the
Veterans Affairs Mid Atlantic Mental Illness Research, Education, and
Clinical Center at the W.G. Hefner Veterans Affairs Medical Center in
Salisbury, North Carolina and the Department of Psychiatry at Wake
Forest School of Medicine in Winston-Salem, North Carolina. Drs. Hurley
and Taber are affiliated with the Research and Academic Affairs Service
Line and the Veterans Affairs Mid Atlantic Mental Illness Research, Edu-
cation, and Clinical Center at the W.G. Hefner Veterans Affairs Medical
Center in Salisbury, North Carolina. Dr. Hurley is affiliated with the De-
partments of Psychiatry and Radiology at Wake Forest School of Medicine
in Winston-Salem, North Carolina, and the Menninger Department of
Psychiatry and Behavioral Sciences at Baylor College of Medicine in
Houston, Texas. Dr. Taber is affiliated with the Division of Biomedical
Sciences at the Via College of Osteopathic Medicine in Blacksburg, Virginia,
and the Department of Physical Medicine and Rehabilitation at Baylor
College of Medicine in Houston, Texas
Corresponding author: Send correspondence to Dr. Hurley; e-mail:
Robin.Hurley@va.gov.
J Neuropsychiatry Clin Neurosci 2016; 28:A4–5; doi: 10.1176/appi.
neuropsych.15120402
4 neuro.psychiatryonline.org
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