Restorative Neurology and Neuroscience 27 (2009) 405–419 405

DOI 10.3233/RNN-2009-0492
IOS Press

Training of the executive component of

working memory: Subcortical areas mediate
transfer effects
Erika Dahlina,b,e,∗ , Lars Bäckmanc, Anna Stigsdotter Neelyd and Lars Nyberg a,b,e
a
Department of Integrative Medical Biology, Umeå University, Umeå, Sweden
b
Department of Radiation Sciences, Umeå University, Umeå, Sweden
c
Aging Research Center, Karolinska Institutet, Stockholm, Sweden
d
Department of Psychology, Umeå University, Umeå, Sweden
e
Umeå Center for Functional Brain Imaging (UFBI), Umeå University, Umeå, Sweden

Abstract. Purpose: Several recent studies show that training can improve working memory (WM) performance. In this review,
many issues related to WM training, such as neural basis, transfer effects, and age-related changes are addressed.
Method: We focus on our own studies investigating training on tasks taxing the executive updating function and discuss our
findings in relation to results from other studies investigating training of the executive component of WM.
Results: The review confirms positive behavioral effects of training on working memory. The most common neural pattern
following training is fronto-parietal activity decreases. Increases in sub-cortical areas are also frequently reported after training,
and we suggest that such increases indicate changes in the underlying skill following training. Transfer effects are in general
difficult to demonstrate. Some studies show that older adults increase their performance after WM training. However, transfer
effects are small or nonexistent in old age.
Conclusions: The main finding in this review is that sub-cortical areas seem to have a critical role in mediating transfer effects
to untrained tasks after at least some forms of working memory training (such as updating).

Keywords: Fronto-parietal changes, subcortical changes, executive training, fMRI, transfer effects, aging

1. Background functions through guided interventions. We discuss

Previous studies have found training-related changes
after teaching strategies (Nyberg et al., 2003), coaching
programs (Van Puyenbroeck and Maes, 2006), and edu-
cation in compensatory strategies (Boman et al., 2004).
People with limited cognitive capacity are often affect-
ed in their everyday life due to problems with executive
functions (Mateer, 1999) and therefore, it is important
to increase our knowledge about the modifiability of
executive functions and related working memory (WM)
∗ Corresponding
author: Erika Dahlin, IMB, Physiology Section,
Umeå University, SE-901 87 Umeå, Sweden. Tel.: +46 90 786 51
86 connection 13; Fax: +46 90 786 66 83; E-mail: erika.dahlin@
physiol.umu.se.
what is known about specific functions and neural cor-
relates underlying increased performance after training
of executive functions and WM. The discussion will
lead to some conclusions about how underlying neural
mechanisms mediate effects to untrained tasks.
WM refers to the ability to hold and monitor infor-
mation during a short delay and involves simultaneous
storage and manipulation (Baddeley, 1996). According
to Baddeley’s classical model, WM consists of three
main components. One is specialized for maintenance
of speech-based information (the phonological loop),
and one is specialized for maintenance of visual and
spatial information (the visuospatial sketchpad). In ad-
dition to these two “slave” systems, the model also in-
cludes a control structure termed the central executive,

0922-6028/09/$17.00  2009 – IOS Press and the authors. All rights reserved
406 E. Dahlin et al. / Training of the executive component of working memory

which is considered responsible for the regulation of

cognitive processes (i.e., executive functions). Miyake
et al., (2000) provided evidence for a division of ex-
ecutive functioning into three separate, but moderately
correlated, functions: updating, shifting and inhibition.
Miyake and colleagues defined updating as a function
that update and monitor working memory representa-
tions, shifting as a function responsible for attention or
task switching, and inhibition as the ability to deliber-
ately inhibit dominant, automatic, or prepotent respons-
es. The central executive component of WM is almost
certainly the most important component in terms of its
general impact on cognition (Baddeley, 1996; Eysenck,
2000; Repovs and Baddeley, 2006), but nevertheless
remains the least studied component.
In the spirit of the lab-review format, we focus on our
own studies investigating training on tasks taxing the
executive updating function (see Fig. 1 for description
of training tasks) and discuss our findings in relation to
results from other studies investigating training of the
executive component of WM. Four of the training tasks
included in our studies had a similar structure. In these
tasks, single items (letters, numbers, spatial locations,
or colors) were presented and the participants were in-
structed to monitor and update the four last presented
items during list presentation. In the remaining train-
ing task (which was performed twice at each session),
words from various semantic categories were present-
ed. The participants were instructed to mentally place
the words into categories (animals, clothes, countries,
relatives, sports, professions) and continuously update
the content and remember the last presented word in
each category (see Dahlin et al., 2008 for a detailed
description of the training program). Here we describe
training-related performance changes, neural changes,
and effects on untrained tasks (transfer). With the ex-
ception of the last section, where data from elderly par-
ticipants are discussed, we only include studies involv-
ing young healthy individuals.
Fig. 1. Training tasks included in each training session in the updat-
ing training described by Dahlin, Nyberg et al., (2008). In four prac-
2. Training effects ticed tasks participants were instructed to recall the last four items
in correct order (letters, numbers, spatial locations, or colors). The
remaining training task was performed twice each session. Partici-
In this section, we discuss previous WM training pants were instructed to mentally place words into categories (indi-
studies and compare different interventions in terms of cated by boxes at the bottom of the screen), continuously update the
content, and remember the last presented word in each category. In
training-related changes. A training-related change can this example; first box/category ANIMALS, second SPORTS, and
be indicated either by a change in accuracy, change in third CLOTHES.
reaction time, or a change in error rate and is defined
inconsistently in previous studies. Participants’ health tors for training-related changes. Nonetheless, factors
status, gender, and life style as well as format differ- like these have not attracted much interest in previ-
ences between training programs may be important fac- ous WM training studies. We suspect that especially
 E. Dahlin et al. / Training of the executive component of working memory
the amount of training is a critical factor for training-
related changes, and will highlight this factor. Sever-
al studies have revealed that it is possible to increase
performance (Brehmer et al., 2008; Jaeggi et al., 2008;
Olesen et al., 2004; Westerberg and Klingberg, 2007),
reduce reaction times (Karbach and Kray, in press), and
decrease error rate (Hempel et al., 2004) on WM tasks
after longer (> 1 week) training periods. It has also
been found that it is possible to reduce reaction times
(Bherer et al., 2005; Garavan et al., 2000; Jansma et al.,
2001) and error rate (Landau et al., 2004) after shorter
(< 1 week) periods of WM training. In all studies list-
ed above, performance continued to change throughout
the training period, which raises questions about the
optimal length of the training period. At present, lit-
tle is known about optimal lengths of training periods,
number of training sessions, and number of training
tasks. The fact that different dependent measures are
used in various studies does not facilitate comparisons.
That said, we estimated the relation between length
of training period and the size of training-related
changes (see Table 1). To do so, we divided 22 studies
into three groups; 1 (Effect Size, ES
1.5), 2 (ES >
1.5 and ES < 3.0), and 3 (ES  3.0), based on cal-
culated effect sizes on the dependent measures ((post
training – baseline)/standard deviation at baseline). All
were current studies that investigated training of the
executive component of WM to some extent. In studies
where several dependent measures for a given task were
used, we calculated effect sizes on the measure which
revealed the most pronounced effect. Further, if sever-
al criterion tasks were included, the most executively
demanding task was analyzed.
A comparison of the length of training periods in
studies listed in groups 1 and 3 revealed that higher lev-
els of training-related changes were reported in studies
where participants received longer periods of training
compared to studies where participants received short-
er periods of training. Hence, from the estimation of
ES in Table 1 it seems as if the length of the training
period is a critical factor for the magnitude of the train-
ing gains. However, it is a simplification to only dis-
cuss length of training period, as type of training tasks
and number of training sessions are presumably also
important factors. Not surprisingly, the general pattern
in Table 1 is that more sessions of training resulted in
larger effect sizes. The fact that almost all of the stud-
ies in Table 1 that used long training periods also used
many training sessions makes it difficult to draw strong
conclusions about the relative importance of these two
factors. However, it is interesting to note that both
407
Kramer (1999) and Karbach and Kray (in press) used
4 sessions of training on task switching (the trained
tasks were not identical, albeit similar), but the ES in
Kramer (1999) were < 1.5 whereas the ES in Karbach
and Kray (in press) were  3. The main difference
between the studies was the length of the training pe-
riod. In Kramer (1999) participants trained for one
week, whereas in Karbach and Krays (in press) they
trained for four weeks. This is one example where the
same number of training sessions can produce different
training-related effects depending on the total length
of the training period. Studies that investigate this is-
sue in a systematic way are warranted, but a tentative
conclusion we can draw is that longer training periods
and more training sessions in yields larger effects on
trained tasks.
In line with the impression that higher levels of train-
ing lead to larger training-related gains, a recent study
(Jaeggi et al., 2008), where participants trained on an n-
back task for 8, 12, 17, or 19 days, found that groups re-
ceiving more training showed larger gains compared to
groups receiving less training. Interestingly, the pattern
was dose-dependent; performance increased gradually
with the number of days practiced. We found a similar
pattern of dose-dependent effects when we calculated
effect sizes across training weeks (Dahlin et al., 2008).
The average performance during training session 1–3
(week 1), session 4–6 (week 2), session 7–9 (week 3),
session 10–12 (week 4), and session 13–15 (week 5)
were subtracted from pre-test performance (baseline)
and divided by the standard deviation at pre-test. We
found that the ES for week 1 = 0.97; ES week 2 =
1.81; ES week 3 = 2.25; ES week 4 = 2.51; and ES
week 5 = 2.76 (ES for post-training = 3.00). If these
patterns reflect number of training sessions or a longer
training period remains unknown.
As previously mentioned, even brief periods of train-
ing can lead to behavioral changes in the criterion task.
In fact, the most common pattern when testing healthy
participants on a task more than once is a change, i.e.
a test-retest effect, on the repeated task (Levine et al.,
2004). To control for such effects, it is imperative to
include control groups in all training studies. There are
however a lot of training studies not including control
groups (Table 1) even if the importance of such groups
have been frequently discussed over the years. We in-
cluded both trained and untrained participants in a re-
cent study where we examined training-related changes
after five weeks (a total of 15 sessions) of updating
training (Dahlin et al., 2008; Dahlin, Stigsdotter Neely
et al., 2008). Even if the change in performance was
408 E. Dahlin et al. / Training of the executive component of working memory
more pronounced for the trained participants, we found
a clear change from pre- to post-test for both groups.
By including a control group in this study, we avoided
to misinterpret at test-retest effect as a training-related
effect.
A further aspect of WM training is that the proficien-
cy of the underlying cognitive skill differs, between in-
dividuals and finding an optimal level of task difficulty
for each individual can be critical for successful train-
ing. One study divided the participants based on im-
provement scores and revealed increased performance
after five days of training on an auditory WM task for a
strong learning group but not for a weak learning group
(Gaab et al., 2006). One way to regulate task demands
is to use adaptive training adjusted to individuals’ per-
formance. Only six of the studies listed in Table 1 used
training tasks with adjusted difficulty levels depend-
ing on the participants’ performance (Brehmer et al.,
2008; Dahlin et al., 2008; Dahlin, Stigsdotter Neely
et al., 2008; Jaeggi et al., 2008; Olesen et al., 2004;
Westerberg and Klingberg, 2007). In our estimation
of training-related gains, all studies that used adaptive
training procedures revealed medium to high levels of
improvement (Table 1). Thus, our conclusions is that
it is important to ensure that the training has an optimal
degree of difficulty for each individual during the entire
training period.
After finding training-related effects, it is of rele-
vance to ensure the efficacy of training. One means
of doing so is to investigate long-term maintenance of
gains. Long-term follow-up assessments are often in-
cluded in other types of cognitive interventions, such
as training of encoding strategies in episodic memory
(Bottiroli et al., 2008; Stigsdotter Neely and B äckman,
1993) and training of fluid intelligence in relation to
everyday functional outcomes (Willis et al., 2006), but
are rarely addressed in WM training studies (Table 1).
To investigate long-term effects after updating train-
ing, we invited trained and control participants for a
follow-up test session 18 months post training (Dahlin
et al., 2008). Performance on the trained task was
decreased at follow-up compared to the post-training
session, but still significantly higher than at baseline
for trained participants. By contrast, performance at
follow-up did not differ from that at baseline for con-
trols. The maintenance effect after 18 months suggests
a rather robust influence of training. Another study
listed in Table 1 (Kramer et al., 1999) found long-term
effects 2 months post training. One goal in this review
was to delineate factors important for long-term main-
tenance of gains. The fact that only two WM training
studies (Dahlin et al., 2008; Kramer, 1999), differing
in several respects (number of training sessions, num-
ber of months between post-test and follow-up test, the
dependent measure used), have investigated long-term
maintenance makes it difficult to draw firm conclusions.
In the next section, we will further elaborate on changes
in underlying functions and networks when addressing
neural correlates of training-related cognitive changes.
3. Training-related neural changes
Anatomically, Baddeley (1996) defined the central
executive as a system located in the frontal lobes. Imag-
ing studies have also revealed that the executive com-
ponent of WM draws on frontal as well as parietal cor-
tex integrity (e.g., Cabeza and Nyberg, 2000a; Col-
lette and Van der Linden, 2002; Prabhakaran et al.,
2000). In addition, recent work indicates that WM
functioning is also dependent on different subcortical
areas (e.g., Cools et al., 2008; Marklund et al., 2007;
McNab and Klingberg, 2008). These patterns of ac-
tivation in tasks tapping the executive component of
WM was also found in a recent study from our lab
(Nyberg et al., 2009), where frontal, parietal, and sub-
cortical brain areas exhibited a load-dependent activa-
tion pattern during WM performance. In this study,
we investigated brain activation patterns in young low-
performers, young high-performers, and elderly per-
sons during n-back performance. A task with three lev-
els of loads (1/2/3) was used, and capacity-constraints
in working memory between load 1–2 for the elder-
ly and between 2–3 for young low-performers relative
to young high-performers were found. Constraints in
neural activity also followed this pattern by showing
a monotonically increasing response in parietal cortex
and thalamus for young high-performers, whereas ac-
tivity leveled off at 1-back for the elderly and at 2-back
for young low-performers in these brain regions. The
response in frontal cortex followed a similar pattern of
capacity-constraints, although the old showed greater
activation at the lowest level compared to the younger
groups (Fig. 2; Nyberg et al., 2009). Hence, training-
related activation changes may not be located in iso-
lated parts of the brain but rather seem to be widely
distributed. In the following sections, we discuss key
regions in which altered patterns of brain activity post
training have been documented.
 E. Dahlin et al. / Training of the executive component of working memory
Fig. 2. Load-sensitive activation patterns in fronto-paroetal and
sub-cortical areas. (a) All groups showed a decrease in number
of correct responses as a function of load. The pattern suggests
that capacity was heavily taxed at 1-back for the elderly adults and
between 2-back and 3-back for young low-performers. Error bars
represent standard errors around the means. (b) The old adults
showed greater activation at the lowest load level compared to both
younger groups, and activity decreases with increasing load in a left
frontal area. A capacity-constrained response was seen for young
low-performers between 2- and 3-back in the same region. (c) A ca-
pacity-constrained response was seen for old between 1- and 2-back
and for young low-performers between 2- and 3-back in a right infe-
rior parietal region. (d) A capacity-constrained response was found
between 1- to 2-back for the old and between 2- and 3-back for young
low-performers in thalamus.
3.1. Fronto-parietal training-related changes
Both fronto-parietal increases and decreases after
WM training have been reported (Table 1), and it is not
409
clear how to interpret such divergent training-related
brain changes with respect to neural efficiency. Increas-
es in activation may reflect recruitment of additional
cortical units with practice. Alternatively, increases
may stem from a strengthening of the BOLD response
within a particular region (Poldrack, 2000). Increas-
es in fronto-parietal areas have been found after both
longer (> 1 week) visuo-spatial WM training (Olesen
et al., 2004; Westerberg and Klingberg, 2007) as well
as after shorter training on an n-back task (Hempel et
al., 2004).
However, the most common fronto-parietal pattern
after both repeated exposure to WM tasks (Chein and
Schneider, 2005) and more intensive training (Kelly et
al., 2006; Kelly and Garavan, 2005) is decreased acti-
vation. Such a pattern of functional activation changes
are reported after longer (> 1 week) periods of train-
ing on a complex auditory task (Gaab et al., 2006),
a visuospatial WM task (Brehmer et al., 2008), as
well as after shorter (< 1 day) periods of training on
a visuospatial WM task (Garavan et al., 2000) and
an n-back task (Hempel et al., 2004). We found de-
creased fronto-parietal activation for the trained group
in our recent study (Dahlin et al., 2008), where training-
related changes after five weeks of updating training
were investigated. The decreased training-related ac-
tivations were located in right anterior prefrontal cor-
tex, right somatosensory association cortex and right
supramarginal gyrus. All these areas are important for
higher-order cognitive processing and attentional mod-
ulation (Cabeza and Nyberg, 2000a). Especially the
anterior prefrontal cortex is thought to be critical to
strategic processes involved in executive functions and
memory retrieval. The decreased fronto-parietal acti-
vation following training may reflect that the task was
initially difficult and required intense executive control.
That is, participants may initially have generated task
strategies that involved brain regions associated with
attentional functions. Conceivably, with practice, the
task became less effortful and performance required
less control. The reduction in executive control follow-
ing training led to reductions of activation in areas im-
portant for attention, particularly regions in the fronto-
parietal cortex (Chein and Schneider, 2005; Poldrack,
2000). In line with our results, a recent study (Brehmer
et al., 2009) reported marked fronto-parietal decreases
after five weeks of spatial and verbal WM training. By
contrast, as noted above, Olesen (2004) reported in-
creased fronto-parietal activation after an almost iden-
tical intervention procedure. A possible reason for dis-
crepant findings could be whether the training program
410 E. Dahlin et al. / Training of the executive component of working memory
is adaptive or not. Adaptive training makes it neces-
sary to implement new strategies during the training
period, which could increase the executive demands,
and hence result in greater fronto-parietal activation.
However, both Olesen et al., (2004) and Brehmer et al.,
(2008) used an adaptive training regimen, and the rea-
son for inconsistent results must therefore be searched
for elsewhere.
Another possible reason for inconsistency in brain
activation changes between studies is variations in the
amount of training. In Hempel et al., (2004), a limit-
ed amount of training on an n-back task led to fronto-
parietal increases, whereas more extensive training led
to fronto-parietal decreases. Even though the amount
of training in terms of number of days did not differ in
the studies discussed above, a more extensive program
was used in Brehmer et al. (2009; 7 training tasks)
compared to Olesen et al., (2004; 3 training tasks).
Consistent with this assertion, in Dahlin, Stigsdotter
Neely et al., (2008) participants practiced on 6 tasks
during each training session. It is reasonable to assume
that the probability of observing training-related de-
creases is greater following more extensive training, al-
though training-related fronto-parietal decreases have
also been reported after less extensive training (see Ta-
ble 1). To further understand training-related brain ac-
tivation changes, it is of interest to relate the fronto-
parietal changes to subcortical changes in activation
patterns.
3.2. Sub-cortical training-related changes
Subcortical areas such as the basal ganglia are impor-
tant in human cognition (for review, see Seger, 2006).
Indeed, there are studies reporting subcortical activa-
tion changes following WM training (Table 1). The
most common pattern in these studies is subcortical in-
creases following training, and only two studies (Lan-
dau et al., 2004; Tomasi et al., 2004) demonstrated de-
creased subcortical activation following training. Lan-
dau et al., (2004) showed activation decreases in pari-
etal, occipital, and precentral gyrus as well as in anteri-
or thalamus and left insula during encoding after prac-
ticing a face WM task. Tomasi et al., (2004) showed
activation decreases in several regions of prefrontal cor-
tex, cerebellum as well as in the pulvinar. Importantly,
the amount of practice in both Landau et al., (2004)
and Tomasi et al., (2004) was limited to the scanning
session. In studies where participants receive more
training, increased subcortical activation changes are
more common. For example, training-related increas-
es in right striatum and left thalamus were found af-
ter five weeks of WM training (Olesen et al., 2004),
and increased bilateral activity in striatum following
five weeks of updating training was found in our study
(Dahlin et al., 2008). Hence, more training seems to re-
sult in increased subcortical activity. One possibility is
that more training produces more substantial changes in
the underlying skill, which could be reflected in subcor-
tical activity increases. Hence, the increased training-
related activity we observed in striatum following up-
dating training likely reflects strengthening of the up-
dating skill (which was upheld 18 months after com-
pletion of training). However, to more fully understand
the neural reorganization that takes place after train-
ing, it is critical to draw attention to how cortical and
subcortical areas interact.
3.3. Neural networks involved in WM training
As noted, we found decreased activation in fronto-
parietal areas after updating training, whereas activa-
tion in subcortical areas increased post training. These
results support the views that training does not result
in a monotonic increase or decrease in neural activity
(Kelly and Garavan, 2005; Landau et al., 2004), and
that training-related activation changes are not restrict-
ed to an isolated part of the brain. To better understand
the neural reorganization that takes place after training,
it is critical to identify neural networks underlying these
activity changes. In a model describing a network crit-
ical to cognitive functioning, Alexander et al., (1986)
delineated the parallel organization of functionally seg-
regated circuits linking basal ganglia and cortex. A
network that is strongly implicated in cognitive training
is the dorsolateral prefrontal circuit, which consists of
at least two distinct basal ganglia-thalamocortical cir-
cuits that selectively influence separate prefrontal ar-
eas. According to this model, key regions include the
dorsal prefrontal cortex (BA 10) and posterior parietal
cortex (BA7). These areas are highly overlapping with
the fronto-parietal areas where we found decreased ac-
tivation following training (Fig. 3; Dahlin, Stigsdotter
Neely et al., 2008). This circuit continues to the globus
pallidus and the substantia nigra. In turn, these regions
project to the thalamic nuclei, which project back to
prefrontal cortex.
The connection between cortical areas and the basal
ganglia have also been described in several compu-
tational models (Frank et al., 2001; O’Reilly, 2006).
Frank et al., (2001) postulated that the frontal cortex
is responsible for active maintenance of information,
 E. Dahlin et al. / Training of the executive component of working memory
Fig. 3. The network underlying cognitive performance described by
Alexander et al., (1986) in relation to training-related changes ob-
served by Dahlin Stigsdotter Neely et al., (2008). The brain map at
the top shows dorsal prefrontal cortex (BA 10) and posterior pari-
etal cortex (BA7) exhibiting decreased activation following updat-
ing training. According to Alexander et al., (1986) projections arise
from these areas (blue dotted line from BA 7, green dotted line from
BA 10) to striatum. The brain map at the bottom (z = −4) shows
associative striatum, exhibiting increased activation following updat-
ing training. The circuit then continues to the globus pallidus and
substantia nigra, which projects to thalamus, which in turn projects
back to prefrontal cortex. The brain maps are thresholded at p <
0.005 (cluster size > 10) n = 22.
whereas the basal ganglia serve a selective, dynamic
gating function that enables frontal memory represen-
tations to be rapidly updated. Analogously, in a re-
cent review of biologically-based computational mod-
els of high-level cognition, O’Reilly (2006) presented
evidence that rapid updating requires a mechanism for
gating or switching between on and off states. The ex-
tensive connectivity between PFC and the basal ganglia
is thought to be critical for this gating mechanism. In
relation to the training-related fronto-parietal decreas-
es and corresponding striatal increases that we found
following updating training, these models suggest that
the fronto-striatal network was involved. Even at the
411
end of our experiment, the frontal cortex maintained
some degree of involvement, although much decreased
compared to pre training. With practice, subcortical
brain regions took an increasingly more prominent role.
The basal ganglia gating mechanism may have be-
come more efficient after training, and thus, the fronto-
parietal cortex did not have to be involved to the same
extent. This notion is in line with a theory about rule
learning that argues that prefrontal cortex is responsible
for initial rule or association induction (Packard and
Knowlton, 2002). Subsequently, the striatum mediates
rule application during task performance. This theo-
ry predicts that frontal activity should be high early in
training and decrease when rules are learned, where-
as striatal activity should increase concomitantly with
increases in learning.
In addition, the pattern of fronto-parietal decreases
and subcortical increases after training have also been
found after motor learning, which is more extensively
studied compared to cognitive training. Less fronto-
parietal activation and more subcortical activation is in-
terpreted to reflect lower demands on cognitive control,
that is, a more effective system in motor learning (Will-
ingham, 1998; see also Poldrack et al., 2005). We in-
terpret the decreased cortical activation as an indication
of more automatized task performance following train-
ing, and the increased striatal activation as a change in
the underlying skill. A consequence of change in the
underlying skill may be that untrained tasks drawing on
the same skill also can be affected. Along these lines,
we have argued that subcortical areas may have an im-
portant role in mediating effects on untrained tasks fol-
lowing training of updating WM (Dahlin, Stigsdotter
Neely et al., 2008). This topic will be elaborated on in
the next section.
4. Transfer effects
One goal after cognitive training is to demonstrate
effects on trained as well as untrained tasks. However,
the typical pattern in studies investigating effects on
untrained tasks, i.e., transfer effects, is one of limit-
ed generalizability, despite large improvement on the
criterion task (Table 1).
A frequently discussed factor in the context of trans-
fer effects is the relation between trained and untrained
tasks. As early as in 1901 Thorndike and Woodworth
(Thorndike and Woodworth, 1901) found that training
in one task caused improvement in performing another
untrained task. The function trained was that of esti-
 412
Table 1
Overview of 21 WM training studies
ES Study Tr task Tr Period Groups Transfer Maintenance Fronto- Sub-
tr/ctrl parietal cortical

1.5 Tomasi et al., (2004) visual attention during scanning 25/0 – – ⇓ ⇓
NeuroImage

1.5 Weissman et al., (2002) visual attention during scanning 15/0 – – ⇑⇓ ⇑
Cognitive Brain Res

1.5 Minear & Shah, (2008) Predictable or random 2 S, 2 D 62
/31 Predictable and random task switching – – –
Mem Cognition task switching

1.5 Kramer et al., (1999) swithing 4 S, 1 W 52
# /0 – 2M – –
Acta Psychol

1.5 Sohn et al., (2006) comparison of stimuli 3 S, 3 D 49
/0 comparison of stimuli – – –
Learn individ differ

1.5 Hempel et al., (2004) n-back daily training, 4 W 9/0 – – ⇑⇓ –
Am J Psychiatry
1.5-3 Garavan et al., (2000) visuospatial WM during scanning 17
/0 – – ⇓ –
Microsc Res Tech
1.5-3 Landau et al., (2004) face WM during scanning 10/0 – – ⇓ ⇓
NeuroImage
1.5-3 Milham et al., (2003) stroop (inhibition) during scanning 11/0 – – ⇓ –
NeuroImage
1.5-3 Sayla et al., (2006) object or spatial WM 2 scanning sessions 10/0 – – ⇓ *
Cereb Cortex
1.5-3 Jansma et al., (2001) verbal WM tot 45 min, 4 S 15/0 – – ⇓ *
J Cogn Neurosci
1.5-3 Gaab et al., (2006) pitch memory 5 S, 5 D 14
/10 – – ⇓ ⇑
NeuroImage
1.5-3 Erickson et al., (2007) dual-task processing 5S, 2–3 W 16/15 – – ⇓ ⇒
Cereb Cortex
E. Dahlin et al. / Training of the executive component of working memory

1.5-3 Jaeggi et al., (2008) n-back 8,12,17 or 19 D 35/35
fluid intelligence – – –

PNAS
1.5-3 Brehmer et al., (2008) spatial and verbal WM daily, 5 W 12# /0 – – ⇓ *
Poster CAC
1.5-3 Olesen et al., (2004) spatial WM ∼ 20 D, 5 W 11/11 Stroop, span board, Raven’s matrices – ⇑ ⇑
Nat Neurosci
 Table 1, continued
ES Study Tr task Tr Period Groups Transfer Maintenance Fronto- Sub-
tr/ctrl parietal cortical
3 Karbach & Kray, switching 4 S, 4W 42
/14 switching, verbal and spatial WM, interfer- – – –
(in press) ence control, fluid intelligence
Dev Sci
3 Bherer et al., (2005) auditory discrimination 5 S, 3 W 24
# /12# auditory discrimination visual discrimination – – –
Psychol Aging visual discrimination
3 Bherer et al.,(2008) visual discrimination 5 S, 3 W 32# /12# auditory discrimination visual discrimination – – –
Exp Aging Res
3 Dahlin et al., (2008) updating 15 S, 5 W 16# /16# updating 18 M – –
Psychol Aging
3 Dahlin et al., (2008) updating 15 S, 5 W 16# /7# updating – ⇓ ⇑
Science
3 Westerberg et al., visual WM, backward ∼ 25 D, 5 W 3/11 visuospatial WM, reasoning – ⇑ *
(2007) digit span, letter-span
Physiol Behav
M = Months; D = Days; S = Sessions; ⇑ = increased training-related activity; ⇓ = decreased training-related activity; ⇒ = no significant training-related changes; Tr = training;
Ctrl = Controls; # Also other groups than young included (only young in table);
More than one group; –not investigated; ∗ no finding.
E. Dahlin et al. / Training of the executive component of working memory
413
414 E. Dahlin et al. / Training of the executive component of working memory
mating areas of a specific shape from 10 to 100 cm 2
and a clear improvement was seen in the trained task.
Effects on untrained tasks were also found but the effect
was more pronounced when areas were similar in shape
to those in the training series compared to when new
shapes were used. Hence, the gains could be attribut-
ed to common elements in the trained and untrained
tasks, and the improvement differed depending on level
of similarity between the trained and untrained tasks.
Following this seminal paper, the terms near- and far-
transfer were introduced in the literature. Near-transfer
tasks are untrained tasks that are similar to the trained
task, whereas far-transfer tasks are untrained tasks that
have less in common with the trained task.
To investigate whether shared processes in the
trained and untrained tasks were central for finding
transfer effects we built on the model of executive func-
tions devised by Miyake et al., (2000). Our prediction
was that five weeks of training on a specific execu-
tive function, updating, may result in transfer effects
to other untrained executive tasks taxing updating. We
tested one untrained task (n-back), which also required
updating, and one untrained task (Stroop) assumed to
depend on the untrained executive function inhibition.
We found transfer to n-back but not to Stroop (Dahlin,
Stigsdotter Neely et al., 2008). Hence, our results sug-
gest that the ability trained was improved, not only the
trained task. In another recent paper (Dahlin, Nyberg
et al., 2008), we tested trained and untrained partici-
pants on a wider battery of untrained tasks before and
after five weeks of updating training. Here, we did
not find effects to other transfer tasks despite an ex-
tensive battery of non-trained tasks assessing perceptu-
al speed, reasoning, episodic memory, semantic mem-
ory, and working memory. Based on these findings,
our conclusion is that transfer effects are difficult to
demonstrate and, when demonstrated, they reflect im-
provement in a specific ability commonly underlying
task performance.
In a study by Jaeggi et al., (2008) where participants
trained on n-back for 8, 12, 17, or 19 days, the degree
of gain in a fluid intelligence transfer task critically de-
pended on the amount of training. Although all groups
showed transfer of learning, the magnitude of improve-
ment increased with more training in a dose-dependent
fashion. Most likely, more training leads to more robust
changes in the underlying skill, which in turn should
increase the likelihood of obtaining transfer effects.
We have argued that a key factor for transfer effects
is shared underlying processes required for successful
performance in trained and untrained tasks. In line with
this notion, the transfer effects from n-back training to
fluid intelligence discussed above (Jaeggi et al., 2008)
may reflect the fact that WM and fluid intelligence share
a common capacity constraint and attentional control
processes (Halford et al., 2007; Gray et al., 2003).
Further evidence for our view on transfer is that the WM
training studies documenting transfer effects (Table 1)
mainly report effects to tasks similar to the trained task.
Knowledge about neural correlates of transfer may
shed further light on underlying mechanisms. How-
ever, little is known about neural correlates of transfer
effects. Following Thorndike and Woodworth (1901),
it has been hypothesized that overlapping networks
or brain areas for the trained and the untrained tasks
are necessary for producing transfer effects (Jonides,
2004). To our knowledge, only our recent study has
investigated neural correlates of transfer (Dahlin, Stigs-
dotter Neely et al., 2008). We found overlapping
fronto-parietal activation before training for the trained
updating task (letter memory), the untrained updat-
ing task (n-back), and the untrained inhibition task
(Stroop). Further, we found overlapping activation in
striatum before training in letter memory and n-back,
but not in Stroop. This pattern is interesting to view in
relation to the behavioral results of transfer effects to
n-back, and indicates that pre-training activity in stria-
tum is important for finding transfer effects following
updating training. When investigating training-related
changes we found overlapping training-related increas-
es for both letter memory and n-back in the associa-
tive striatum that was also activated in both tasks pre
training. Hence, the striatal complex was a key com-
ponent mediating transfer effects after training on WM
updating. Previous studies have proposed that striatum
is a gating structure selecting specific regions in frontal
cortex to be updated or to maintain already existing in-
formation (O’Reilly, 2006). We interpret the increased
striatal activations as an indication that the updating
skill was more effective following training.
Another key aspect when discussing training and
transfer effects is differences between individuals, or
groups of individuals. Individual differences in cog-
nitive ability have been found to influence the degree
of transfer after training on a visual comparison task
(Sohn et al., 2006). If differences between individu-
als exist within a group, there are most certainly al-
so differences between groups with different cognitive
ability such as younger versus older adults.
 E. Dahlin et al. / Training of the executive component of working memory
Fig. 4. Letter memory performance for training and control groups.
Error bars are standard errors around the means. The dotted line
indicates baseline performance for young participants.
5. Working memory training in elderly adults
Cognitive interventions have revealed that it is pos-
sible to increase performance in children with ADHD
(Klingberg et al., 2002), patients with traumatic brain
injury (Kim et al., 2008), and schizophrenic patients
(López-Luengo and V ázquez, 2003). However, the
most studied group in cognitive interventions, apart
from young adults, is healthy older adults. Episodic
memory, working memory, and executive functioning
decline in normal aging (e.g., Craik and Jennings, 1992;
Kliegl et al., 2001, Bäckman, 1999; Light, 1991), and
several studies have tried to improve performance on
tests of these functions in older adults by means of var-
ious training procedures. Cognitive interventions in el-
derly persons have often focused on the effects of teach-
ing strategies (the method of loci) to enhance episodic
memory performance and found performance gains af-
ter training (e.g., Nyberg et al., 2003; Verhaeghen and
Marcoen, 1996; Verhaeghen et al., 1992). Training of
WM is not investigated to the same extent in elderly
adults, but healthy older persons can increase perfor-
mance after more than one week of training in spatial
and verbal WM tasks (Brehmer et al., 2008), switching
tasks (Karbach and Kray, in press; Kray and Epplinger,
2006), as well as exercises aimed at improving bottom-
up processing of auditory information (Mahncke et al.,
2006). We investigated updating training in healthy el-
derly adults between 65–70 years of age, using a train-
ing program identical to the one described above for
young adults. Results showed training-related gains in
the letter-memory updating task of about the same mag-
nitude as observed for young adults (Dahlin, Nyberg et
al., 2008; Dahlin, Stigsdotter Neely et al., 2008).
415
A key finding was that older persons performed
above the baseline level of young persons after five
weeks of updating training (Fig. 4). Long-term mainte-
nance of gains 18-months after training was also found;
in fact, older adults performed above the baseline lev-
el of young even one and a half year post training.
Hence, the training effect was quite robust and five
weeks of computer-based training made elderly adults
perform as they would be 40 years younger. At the
same time, the elderly persons performed below their
younger counterparts both initially and after training
and, unlike the young, they exhibited no transfer ef-
fects. This pattern of (a) cognitive plasticity in old age;
and (b) age-related constraints in plasticity character-
izes much intervention research comparing individu-
als across the adult life span (e.g., Verhaeghen et al.,
1992; Baltes and Kliegl, 1992; Jones et al., 2006). The
age-related differences observed by Dahlin, Nyberg et
al., (2008) likely reflect alterations in the fronto-striatal
dopaminergic system (for reviews, see B äckman et al.,
2006; Prull et al., 2000).
To investigate training-related brain activity changes,
all elderly participants were fMRI scanned before and
after the training period. As for young adults, we found
fronto-parietal activation before the intervention during
the trained letter-memory updating task. However, in
contrast to young, we found increased activation fol-
lowing training in fronto-parietal areas for older adults.
This may reflect that task performance was not automa-
tized to the same extent as for the young and that the task
was executively demanding even after five weeks of
training for older persons. Further, we found increased
striatal activation post training in letter-memory for old-
er participants, although the striatum was not activat-
ed before training. Conceivably, these findings reflect
age-related changes in the striatal dopamine system,
which serves as a powerful mediator of the cognitive
losses that occur during the aging process (B äckman
et al., 2000; Bäckman et al., 2006). In particular, the
striatal dopamine system plays a role in age-related dif-
ferences in executive functioning (Erixon-Lindroth et
al., 2005). Hence, age-related changes in the striatal
dopamine system may have constrained the training, so
that the underlying updating skill was not strengthened
to the same extent as for young persons. These changes
may also account for the lack of transfer effects in older
adults.
6. Directions for future research
The research discussed above clearly indicates that
it is possible to train WM functions. The overall data
416 E. Dahlin et al. / Training of the executive component of working memory
pattern suggests that more training brings about larg-
er training-related gains. However, it is complicated
to compare the amount of training between studies us-
ing different training regimens and different dependent
measures. A first direction for future research is there-
fore to conduct more studies like that of Jaeggi et al.,
(2008), varying the amount of training. Hence, studies
are warranted that compare training-related changes af-
ter varying the number of sessions, spacing of sessions,
duration of training, and number of tasks trained. Of
interest is also whether the patterns of dose-response
effects vary across the adult life span. To make it pos-
sible to design optimal training regimens in the future,
it is also important to investigate effects of different
lengths of training periods and number of training ses-
sions in a systematic way. Today it is not known if five
weeks of 15 training sessions is more or less effective
compared to one week of 15 training sessions.
A second direction for future research is to conduct
more studies investigating transfer effects systemati-
cally. As noted, transfer effects are generally limited
and restricted to tasks similar to the criterion task. The
question about transfer effects is complex and sever-
al issues remain unresolved. One important aspect in
this context is shared processes in the trained and un-
trained tasks. As discussed above, we found transfer
to n-back after five weeks of training on other tasks
requiring updating. We interpret this as an effect on a
general updating skill. An interesting avenue for future
research would be to test the generalizability of this as-
sumption by examining whether n-back training results
in improved letter-memory performance. Of interest
is also to investigate the boundaries of transfer across
the three major components of executive functioning
outlined by Miyake et al., (2000): updating, switching,
and inhibition. In particular, we need knowledge about
whether the limited degree of transfer from updating
training (e.g., null effects in the inhibitory Stroop task)
holds for training in switching and other updating tasks
as well.
A third direction for future research is that more stud-
ies should investigate neural correlates of WM training
and transfer. A particularly intriguing line of inquiry
is to further investigate the striatal blood flow increas-
es observed following training in both letter memo-
ry and n-back in our recent study (Dahlin, Stigsdot-
ter Neely et al., 2008). It is well known that the
striatum is a region in which the dopaminergic inner-
vation is particularly dense (Alexander et al., 1986;
Björklund and Dunnett, 2007), and pharmacological
imaging in rodents suggests that striatal blood flow
is largely driven by dopamine agonism (Knutson and
Gibbs, 2007). Some recent studies have found evidence
for dopamine release in frontal cortex and hippocam-
pus during working-memory performance (Aalto et al.,
2005) and in striatum during both card-sorting (Monchi
et al., 2006) and sequential learning (Badgaiyan et al.,
2007). Also, Schott et al., (2008) recently showed a
sizable relationship between a PET-derived measure of
dopamine release and an fMRI-based measure of blood
flow in ventral striatum during reward-related learn-
ing. To determine whether the training-related increas-
es in striatal blood flow observed by Dahlin, Stigsdot-
ter Neely et al., (2008) are driven by increased release
of dopamine as a function of training, it would be of
great interest to employ a similar multimodal imaging
approach.
Acknowledgments
This work was funded by grants from the Swedish
Research Council and the Joint Committee for Nordic
Research Councils in the Humanities and the Social
Sciences for a Nordic Center of Excellence (NCoE) in
Cognitive Control to L.N., by grants from the Swedish
Research Council and Swedish Brain Power to L.B,
and by grants from the Swedish Council for Working
Life and Social Research to A.S-N.
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