Neuropsychologia 44 (2006) 2149–2157

What have we learned about cognitive development from neuroimaging?

Sarah Durston a,b,∗ , B.J. Casey a
a Sackler Institute for Developmental Psychobiology, Weill Medical College of Cornell University,
1300 York Avenue, Box 140, New York, NY 10021, USA
b Rudolf Magnus Institute of Neuroscience, Department of Child and Adolescent Psychiatry,

University Medical Center Utrecht, Heidelberglaan 100, 3584 CX, Utrecht, The Netherlands
Received 6 June 2005; received in revised form 22 September 2005; accepted 15 October 2005
Available online 21 November 2005

Abstract
Changes in many domains of cognition occur with development. In this paper, we discuss neuroimaging approaches to understanding these
changes at a neural level. We highlight how modern imaging methods such as functional magnetic resonance imaging (fMRI) and diffusion
tensor imaging (DTI) are being used to examine how cognitive development is supported by the maturation of the brain. Some reports suggest
developmental changes in patterns of brain activity appear to involve a shift from diffuse to more focal activation, likely representing a fine-tuning
of relevant neural systems with experience. One of the challenges in investigating the interplay between cognitive development and maturation
of the brain is to separate the contributions of neural changes specific to development and learning. Examples are given from the developmental
neuroimaging literature. The focus is on the development of cognitive control, as the protracted developmental course of this ability into adolescence
raises key issues. Finally, the relevance of normative studies for understanding neural and cognitive changes in developmental disorders is discussed.
© 2005 Elsevier Ltd. All rights reserved.

Keywords: Cognition; Development; Functional MRI; DTI; ADHD

Over the last decade, investigators have become increasingly
interested in how cognitive development is supported by the
maturation of the brain (Casey, Tottenham, Liston, & Durston,
2005). Investigators interested in the interplay between the two
have used a host of paradigms and methods to investigate this
association. In the following paper, we review some of the meth-
ods that have been used to examine the behavioral and neural
basis of the development of cognition. This paper is not intended
as an exhaustive review of the literature, but rather as a discussion
of how the study of development may benefit from neuroimaging
approaches. We will focus largely on cognitive control, as the
protracted developmental course of this ability into adolescence
is relevant to a number of developmental disorders, such as atten-
tion deficit hyperactivity disorder (ADHD), childhood-onset
schizophrenia, and childhood-onset obsessive–compulsive dis-
order. Illustrative examples from the developmental literature
of other cognitive functions will also be discussed. In the final
section of this manuscript, we will discuss some points from the
∗ Corresponding author. Tel.: +31 30 250 8161; fax: +31 30 250 5444.
E-mail address: S.Durston@azu.nl (S. Durston).
ADHD-literature, as an example of how atypical development
may be relevant to our understanding of typical developmental
processes.
Cognitive control may be defined as the flexible regula-
tion of thoughts and actions in the presence of competing
ones, and is involved in many cognitive functions, including
motor inhibition, interference inhibition, cognitive flexibility
and attentional control. In recent years, the ability to flexibly
adapt behavior has played a critical role in theories of cogni-
tive development. These theories have characterized immature
cognition as being particularly susceptible to interference (e.g.,
Brainerd & Reyna, 1993; Dempster, 1992; Munakata, 1998),
with younger children having difficulties in overriding compet-
ing attentional or behavioral responses. This susceptibility to
interference during childhood is illustrated by studies showing
that performance on Stroop, flanker, and go no-go tasks contin-
ues to develop over childhood and does not reach full maturity
until roughly 12 years or later (e.g., Carver, Livesey, & Charles,
2001; Diamond, Cruttenden, & Nederman, 1994; Diamond &
Taylor, 1996; Enns & Akhtar, 1989; Enns & Cameron, 1987;
Enns, Brodeur, & Trick, 1998; Gerstadt, Hong, & Diamond,
1994; Luria, 1961; Passler, Isaac, & Hynd, 1985; Ridderinkhof

0028-3932/$ – see front matter © 2005 Elsevier Ltd. All rights reserved.
doi:10.1016/j.neuropsychologia.2005.10.010
2150 S. Durston, B.J. Casey / Neuropsychologia 44 (2006) 2149–2157
& Van der Molen, 1997; Ridderinkhof, Van der Molen, & Band,
1997; Tipper, Bourque, Anderson, & Brehaut, 1989; Van der
Meere & Stemerdink, 1999). This work shows a developmen-
tal trend in the ability to suppress information and actions over
the ages of 4–12 years, as indexed by mean reaction times and
accuracy rates. Importantly, age-related differences in accuracy
are not observed on these tasks in the absence of interfering
information (e.g., Enns et al., 1998) illustrating the differential
developmental trajectories for cognitive control relative to more
simple, target detection tasks.
1. Structural brain development
Although pediatric neuroimaging is a relatively young field,
several studies have begun to track the anatomical course of
normal brain development (see, Durston et al., 2001, for a
review). These structural MRI studies have shown an increase
in brain volume over the first few years and then relative sta-
bility in brain volume by later childhood (Caviness, Kennedy,
Richelme, Rademacher, & Filipek, 1996; Courchesne et al.,
2000; Giedd et al., 1996, 1999; Reiss, Abrams, Singer, Ross,
& Denckla, 1996; Thompson et al., 2000). The relative stabil-
ity of total brain volume in later development belies a dynamic
interplay of simultaneously occurring progressive and regressive
events, with different regions following different time courses.
Significant changes occur in regional cortical and subcortical
gray matter volumes (Caviness et al., 1996; Giedd et al., 1996,
1999; Jernigan, Trauner, Hesselink, & Tallal, 1991; Sowell,
Thompson, Holmes, Jernigan, & Togan, 1999; Sowell et al.,
2003). Perhaps the most informative studies to date have been
those tracking changes over time within individuals. These
studies have shown differential development between different
regions of the brain, where increases in cortical gray matter fol-
lowed by volume loss apppear to occur first in sensori-motor
and last in higher-order association cortices such as the dorso-
lateral prefrontal cortex (see Giedd, 2004 for a review; Gogtay
et al., 2004). In another example of this approach, Sowell et al.
(2004) tracked changes in cortical gray matter thickness longitu-
dinally in preadolescent children, and showed regionally specific
increases in classical language regions, accompanied by more
widespread thinning in other cortical areas. Interestingly, they
were also able to show a relationship between cortical thinning
and improvements in language ability.
Simultaneous with ongoing changes in gray matter, white
matter shows linear increases in volume during childhood
and into adulthood, with changes appearing to follow a pat-
tern from caudal to more rostral (Giedd et al., 1999; Jernigan
et al., 1991; Paus et al., 1999; Pfefferbaum et al., 1994).
Diffusion tensor imaging (DTI), a relatively new technique
offers greater detail than conventional structural MRI tech-
niques, as it provides information on the directionality and
regularity of myelinated fiber tracts. Initial studies using this
method have confirmed changes in cortical white matter path-
ways during development (e.g., Klingberg, Vaidya, Gabrieli,
Moseley, & Hedehus, 1999). These changes presumably reflect
the ongoing myelination of axons by oligodendrocytes, enhanc-
ing neuronal conduction and communication. The direct appli-
cation of this methodology to examining development of cog-
nitive control will be discussed in a subsequent section of
this paper.
2. Structural brain changes and cognitive development
By definition, developmental changes in cognition coincide
with changes in the brain. In order to inform us about the func-
tional organization of the brain, investigators have begun to
link neuroanatomical development to cognitive changes. For
example, the basal ganglia have a number of projections to and
from the prefrontal cortex and both brain regions mature rel-
atively late (Sowell et al., 1999). As this circuitry has been
implicated in goal-directed actions, development of this cir-
cuitry has been suggested to relate to the development of cog-
nitively driven actions throughout childhood and adolescence.
Reiss et al. (1996) have shown that the largest increase in
prefrontal volume, in fact, takes place during this period of
development consistent with this claim. However, these par-
allel changes could be coincident. A number of studies have
begun to link developmental changes in brain anatomy to behav-
ior (e.g., Olesen, Nagy, Westerberg, & Klingberg, 2003; Sowell
et al., 2004). In one such example, Casey, Castellanos et al.
(1997) showed correlations between performance on measures
of impulse control and volumetric measures of the prefrontal
cortex and basal ganglia in healthy children relative to children
with ADHD. Such correlational analyses are intriguing as they
suggest a possible causal relationship between brain and behav-
ior. However, they provide only an indirect link between the
two, whereas advances in current imaging methods allow for
more direct assessment of this relationship using non-invasive
measures.
3. Functional brain changes and cognitive development
More direct investigations of structure–function associations
and their development have been performed using functional
magnetic resonance imaging (fMRI). These studies suggest dif-
ferent developmental trajectories of brain regions involved in
cognitive control. A number of investigators have examined
differential maturation of this function in frontal and parietal
cortices (e.g., Booth et al., 2003; Luna et al., 2001; Rubia et al.,
2000). For example, Adleman et al. (2002) used a Stroop task
to show that increases in activation associated with improved
cognitive performance occurred by adolescence for the parietal
lobe, whereas increases in prefrontal activation continued into
adulthood. Similar findings have been reported for the develop-
ment of working memory, where increased capacity is supported
by higher levels of activation in prefrontal and parietal cortices in
adolescents compared to children (Casey et al., 1995; Klingberg,
Forssberg, & Westerberg, 2002). A number of investigators have
suggested that maturation is not only associated with enhanced
activation in areas that are critical for task performance, but also
with attenuation of activation in non-critical areas. For exam-
ple, Tamm, Menon, & Reiss (2002) showed that maturation was
associated with more focal activation in areas critical for task
performance on a go no-go task.
 S. Durston, B.J. Casey / Neuropsychologia 44 (2006) 2149–2157
Other studies have shown differential recruitment of sub-
cortical as opposed to cortical regions with development (e.g.,
Booth et al., 2003; Bunge, Dudukovic, Thomason, Vaidya, &
Gabrieli, 2002; Rubia et al., 2000). For example, Casey, Thomas,
Davidson, Kunz, & Franzen (2002) examined development of
neural systems involved in developing new stimulus-response
mappings, while overriding pre-existing ones. Here, the extent of
activation in subcortical structures (hippocampus for new learn-
ing, striatum for overriding old responses) was far greater for
children than adults, while their performance was significantly
worse, suggesting that as these functions mature the pattern
of activation associated with them becomes more focal. This
pattern occurred in parallel with greater recruitment of corti-
cal regions with maturation. Others (e.g., Booth et al., 2004)
have shown that better performance is not necessarily associ-
ated with greater activation in development, as although greater
fronto-striatal activation was associated with better performance
on a go no-go task, they found that better selective attention per-
formance was associated with less activation in parietal areas
in a visual search task. Studies investigating cognitive develop-
ment in other domains have illustrated that changes in patterns of
activation are not unique to studies involving cognitive control
(e.g., Hertz-Pannier et al., 1997; Nelson et al., 2003; Schlaggar
et al., 2002). For example, Thomas et al. (2004) showed dif-
ferential patterns of subcortical–cortical activation in a study of
motor-sequence learning.
Taken together, developmental neuroimaging studies of cog-
nitive control, as well as other functions, suggest that cognitive
development is supported by changes in patterns of brain acti-
vation, including enhancement of activation in critical areas,
attenuation in others, and changes in the extent of activation as
well as shifts in lateralization (e.g., Booth et al., 2003, 2004;
Bunge et al., 2002; Gaillard, Balsamo, Ibrahim, Sachs, & Xu,
2003; Hertz-Pannier et al., 1997; Luna & Sweeney, 2004; Nelson
et al., 2003; Rubia et al., 2000; Thomas et al., 2004). In some
studies, developmental changes in patterns of brain activation
could be conceptualized as a shift in patterns of activation from
diffuse to more focal, where diffuse refers to larger or more
areas of activation, and focal indicates smaller areas of activa-
tion, with greater magnitude of signal change (Casey, Trainor
et al., 1997; Casey et al., 2002; Gaillard et al., 2000; Hertz-
Pannier et al., 1997; Holland et al., 2001; Luna et al., 2001;
Monk et al., 2003; Moses et al., 2002; Nelson et al., 2003;
Schlaggar et al., 2002; Tamm et al., 2002; Turkeltaub, Gareau,
Flowers, Zeffiro, & Eden, 2003) similar to changes seen in adult
learning (Karni et al., 1995, 1998). These changes may repre-
sent a fine-tuning of relevant neural systems (Johnson, Halit,
Grice, & Karmiloff-Smith, 2002), or related developmental
changes, such as new brain regions coming online and reduced
involvement of others, and may be related to shifts in cognitive
strategy, in some studies. Differences between developmental
studies with respect to which areas are seen to come online
with development illustrate that neural changes observed using
functional imaging are critically dependent on the task used.
Certain brain regions may become more involved in one aspect
of cognition with development, while becoming less involved
in others.
2151
4. Distinguishing between differences in performance
with age and developmental changes
As previously discussed, development and learning coincide
during childhood. As such, it is a challenge to separate the
contributions of each, and examine neural changes specific to
development, learning and developmental learning. This para-
dox may be addressed in part by functional imaging studies with
paradigms designed to assess which areas are critically related
to task performance. In particular, studies equating performance
between groups have the potential to address such issues. In
developmental studies, younger children will often have poorer
performance on a paradigm than an older comparison-group.
This potentially confounds developmental changes, as differ-
ences in MR results may then be related to differences in task
performance or difficulty.
Investigators have addressed differences in performance
across developmental age groups in a number of ways. One
approach is to consider subgroups at different ages that are
matched on performance. For instance, Schlaggar et al. (2002)
compared adults and children, aged 7–10 years on a single
word processing task and found differences in the recruitment
of prefrontal and extrastriate regions. As these differences could
have been attributable exclusively to performance discrepancies,
they selected a subgroup of their subjects, matched for perfor-
mance and showed that some of the brain regions examined
showed differences attributable to age, independent of perfor-
mance (Schlaggar et al., 2002).
An alternative approach is to incorporate different levels of
task difficulty in the paradigm design by using a parametric
manipulation. This allows the post hoc comparison of groups
at different levels of task difficulty, while matching for per-
formance. For instance, we previously developed a task incor-
porating a parametric manipulation of a go no-go paradigm,
where no-go trials were preceded by 1, 3 or 5 go trials (Durston,
Thomas, Worden, Yang, & Casey, 2002; Durston, Thomas, Yang
et al., 2002). Subjects showed an increase in errors with increas-
ing interference from the preceding go-trials. We used this task
to investigate the development of the neural basis of response
inhibition (the stopping on no-go trials) and found that success-
ful response inhibition was associated with greater activation
of prefrontal and parietal regions for children than for adults.
In adults, activation in ventral prefrontal regions increased with
increasing interference from go-trials. However, unlike adults,
the circuitry appeared to be maximally activated in children
when suppressing a behavioral response regardless of the num-
ber of preceding responses. These findings confirm that imma-
ture cognition is more susceptible to interference, as children
make more errors than adults. Furthermore, they show that this
diffference is paralleled by maturational differences in under-
lying fronto-striatal circuitry (Durston, Thomas, Yang et al.,
2002).
A third possible approach is to examine correlations between
neural activity and performance and age. Performance and age
can both be examined independently by covarying the other. Sev-
eral groups have used this approach to tease apart performance-
based versus age-based differences in developmental studies
2152 S. Durston, B.J. Casey / Neuropsychologia 44 (2006) 2149–2157
(e.g., Casey, Trainor et al., 1997; Casey et al., 2002; Thomas
et al., 2004).
5. Developmental progressions versus single snap-shots
in time
The previously reviewed studies illustrate that if we are to
explain changes in a dynamic system, it is essential to exam-
ine developmental progressions in behavior and the brain, rather
than examining behavior at single snap-shots in time. The impor-
tance of examining developmental trajectories is perhaps most
obvious in understanding and determining whether atypical
development is due to a developmental delay or deficit. A num-
ber of behaviors may be completely appropriate at one age but
inappropriate at another. Clinical disorders, such as attention
deficit hyperactivity disorder, may reflect exaggerated and/or
residual processes that do not necessarily diminish or change
with maturity. Understanding the normal progression in a spe-
cific behavioral or neural system will have a significant impact
in determining the biological substrates of clinical disorders and
targeting effective treatments and interventions.
Although the majority of imaging studies to date have
examined developmental progressions using a cross-sectional
approach, there are a number of advantages to tracking changes
within individuals using a longitudinal approach. This is espe-
cially important in biological systems, as there is an extreme
degree of individual variability in brain structure, relative to
developmental variability, during childhood and adolescence
(Caviness et al., 1996). There are several examples of longi-
tudinal approaches in Section 1 (see, Giedd, 2004, for a review).
In a recent functional imaging study, we used a version of the
previously described go no-go paradigm to track the develop-
ment of cognitive control longitudinally in a sample of children
as they reached adolescence (Durston et al., in press). We showed
a developmental shift in patterns of cortical activation from
diffuse to focal activity. Here, we showed that sensori-motor
regions uncorrelated with task performance were recruited less,
while a region in the ventral prefrontal cortex showed enhanced
recruitment. Activity in this region has been shown to be cor-
related with performance on this task in prior studies (Casey,
Trainor et al., 1997; Durston, Thomas, Worden et al., 2002;
Durston, Thomas, Yang et al., 2002; Konishi, Nakajima, Uchida,
Sekihara, & Miyashita, 1998; Konishi et al., 1999) and in the
current study (Durston et al., in press). In contrast, activa-
tion in structures such as the primary motor cortex remained
unchanged for the simple comparison of responding (go) versus
not responding (no-go) during performance of the task. Inter-
estingly, the significant negative correlation between accuracy
and activation in ventral prefrontal cortex was accompanied by
a trend-level positive correlation between age and activation
in this region. This suggests that at this crucial developmen-
tal stage, subjects may in fact be responding more impulsively,
related to faster responses and therefore need more top-down
control in overriding a response. This finding ties in with anec-
dotal evidence of increased impulsive and risk-taking behavior
in adolescence, and illustrates the complexity of the relation-
ships between development, performance and patterns of brain
activation. A parallel cross-sectional analysis based on a sec-
ond group of children, the same ages as the longitudinal group,
showed less specific results, confirming added sensitivity in lon-
gitudinal approaches.
6. Brain connectivity and cognitive development
The developmental shift from diffuse to focal cortical
activation may reflect the functional consequences of synaptic
pruning and other regressive processes in combination with
strengthening of relevant connections. However, few studies to
date have related maturational changes in brain connectivity
to their functional correlates. New imaging techniques such as
DTI are beginning to allow us to do exactly that (see Fig. 1;
e.g., Klingberg et al., 1999; Olesen et al., 2003). DTI measures
water-diffusion in the brain, and can be used to estimate the
regularity of white matter, as an indirect measure of myelination
and connectivity (Mori, Crain, Chacko, & van Zijl, 1999; Paus et
al., 2001). For example, Liston et al. recently used this technique
to relate functional changes in frontostriatal networks with the
development of performance on the previously described cog-
nitive control task (see Section 4; Durston, Thomas, Worden et
al., 2002; Durston, Thomas, Yang et al., 2002) to changes in the
connectivity between these structures (Liston et al., in press). An
automated fiber-tracking algorithm was used to delineate white
matter fibers projecting from ventral prefrontal cortex to the
caudate nucleus. These two structures are known to contribute to
cognitive control based on previous functional imaging studies
(Casey, Trainor et al., 1997; Durston et al., in press). A posterior
tract, not expected to contribute to this ability, was also delin-
eated with this method as a control. Diffusion in prefrontal and
posterior tracts became more restricted between the ages of 7 and
30 years. This shift was paralleled by an age-associated increase
in efficiency in cognitive performance. Further, fronto-striatal
radial diffusivities that are sensitive to changes in myelination,
predicted individual differences in reaction time, when con-
trolling for age and accuracy. This pattern was not observed in
posterior tracts (Liston et al., in press). Collectively, these results
indicate that maturation of prefrontal white matter and enhanced
connectivity between fronto-striatal structures contributes to
a developing capacity and individual variability in cognitive
control.
7. Atypical development of cognitive control: attention
deficit hyperactivity disorder
The emergence of clinical disorders of cognitive control
throughout childhood and adolescence underscores the impor-
tance of understanding the neurobiological basis of such control
(see Durston et al., 2001 for a review). Attention Deficit Hyper-
activity Disorder (ADHD) is currently the best-characterized
childhood disorder associated with deficits in this ability, and
a discussion of this disorder may aid in our understanding
of typical development of this ability. ADHD first manifests
in early childhood, before the age of 7 and is characterized
by age-inappropriate levels of impulsive, hyperactive of dis-
tractible behavior (i.e., cognitive control deficits). Children with
 S. Durston, B.J. Casey / Neuropsychologia 44 (2006) 2149–2157 2153

Fig. 1. How diffusion tensor imaging can be combined with anatomical and functional MRI to investigate functional networks. Panel A: the caudate nucleus is
defined on an anatomical MRI scan, as a seedpoint for fiber tracking. Panel B: functional activation maps from a single subject during a cognitive control task,
overlayed on the anatomical MRI scan. Panels C and D: vectors calculated from the DTI scan, where the colors represent the direction of the vector, overlayed on
a section of the anatomical scan and, finally. Panel E: fibers tracked on the basis of the vectors in panel D, connecting both caudate nuclei with regions in frontal
cortex, including the frontal region activated in panel B.
2154 S. Durston, B.J. Casey / Neuropsychologia 44 (2006) 2149–2157
ADHD have poor performance on tasks that require inhibitory
control, such as go-no go and stop tasks. This finding has
lead some investigators to theorize that a deficit in inhibitory
components of cognitive control may be central to this disorder
(Barkley, 1997). This theory in turn implicates fronto-striatal
circuitry in ADHD given its involvement in cognitive control
(see Durston, 2003, for a review). As discussed previously, the
development of fronto-striatal circuitry is protracted as synaptic
pruning and myelination of prefrontal fibers proceeds slowly
throughout late childhood and adolescence (Casey, Castellanos
et al., 1997; Casey, Trainor et al., 1997; Conel, 1939–1967;
Durston, Thomas, Worden et al., 2002; Durston, Thomas,
Yang et al., 2002; Huttenlocher, 1979; Klingberg et al., 1999;
Paus et al., 2001; Sowell et al., 1999). Children’s capacity
for cognitive control develops across the first decade with
younger children more susceptible to interference on a variety
of tasks in this domain (Casey, Trainor et al., 1997; Casey et
al., 2002; Diamond, 1990; Munakata & Yerys, 2001). Children
recruit distinct and often larger, more diffuse fronto-striatal
regions when performing cognitive control tasks than adults do,
suggesting development within and refinement of projections
to and from these regions with maturation (Bunge et al., 2002;
Casey, Trainor et al., 1997; Casey et al., 2002; Luna et al., 2001).
The imaging literature on ADHD implicates fronto-striatal
circuitry. The first functional imaging studies used single pho-
ton emission computed tomography (SPECT) to look at cerebral
blood flow in children with ADHD. The authors published a
series of papers in which they used this method and found
evidence for reduced blood flow in striatal regions in this dis-
order (Lou, Henriksen, & Bruhn, 1984; Lou et al., 1989; Lou,
Henriksen, & Bruhn, 1990). Although these findings constituted
the first evidence of functional cerebral deficits in ADHD, the
spatial resolution of SPECT is low and ethical constraints meant
that adequate control subjects could often not be included in
these studies. A series of papers looking at cerebral metabolism
in adults and adolescents with ADHD using positron emis-
sion tomography (PET) at first suggested a reduction in global
metabolism in ADHD (Zametkin et al., 1990), although later
reports suggested that these findings might be more specific to
females with ADHD than to males (Ernst et al., 1994, 1998;
Zametkin et al., 1993). In line with suggestions of reduced base-
line levels of cerebral metabolism and perfusion in ADHD, sev-
eral studies have shown that administration of methylphenidate
increases cerebral metabolism and perfusion in frontal regions
(Kim et al., 2001; Matochik et al., 1993, 1994).
More recently, studies have been published using functional
MRI to investigate brain activity in ADHD during the perfor-
mance of cognitive control tasks (Vaidya et al., 1998; Rubia et
al., 1999; Bush et al., 1999; Durston et al., 2003). Vaidya et
al. showed that children with ADHD had higher frontal activa-
tion and lower striatal activation than control children during
response inhibition, whereas administration of methylphenidate
lead to improved performance associated with increased frontal
activation for both groups and an increase in striatal activation
for the children with ADHD, perhaps representing a normaliza-
tion of striatal function with medication (Vaidya et al., 1998).
Durston et al. (2003), in part, replicated these findings in younger
children with ADHD showing that the striatum was the most
robust region of difference between children with and with-
out the disorder. Other studies have shown reduced activation
of frontal circuitry in adolescents with ADHD (Rubia et al.,
1999) or abnormal recruitment of brain networks in ADHD, dur-
ing performance of cognitive control tasks (Bush et al., 1999;
Schweitzer et al., 2000).
In summary, this literature suggests local abnormalities in
cerebral activation in ADHD, with a hypo-perfusion of pre-
frontal and possibly striatal areas. Functional studies using cog-
nitive control paradigms show deficits in frontal and striatal
function, further implicating this circuitry. These results con-
firm the evidence from the developmental fMRI literature that
the development of cognitive control is supported by the mat-
uration of fronto-striatal circuitry, as poor cognitive control in
ADHD appears to be related to changes in this circuitry.
8. Conclusions
It is well established that, with development, significant
changes occur in cognitive ability, especially in the context of
overriding competing thoughts and actions. In this paper, we
discuss the added value of neuroimaging approaches to under-
standing these changes in cognitive development. We focus
on cognitive control, as its protracted developmental course
into adolescence raises key points. Changes in brain structure
occur in parallel with changes in cognition. However, to show
that these changes are not merely coincident, it is necessary
to directly link developmental changes in brain anatomy to
behavior.
Structural brain development involves a dynamic interplay
of simultaneously occurring progressive and regressive events,
with different regions following different time courses. Inves-
tigators using correlational analyses have suggested that these
maturational changes in brain anatomy may be related to cog-
nitive development. Current imaging methods including func-
tional MRI and DTI allow a more direct linking of the two. FMRI
investigations have confirmed early evidence from anatomical
studies implicating fronto-striatal circuitry in the development of
cognitive control. Generally, developmental changes in patterns
of brain activity appear to involve a shift from diffuse to more
focal activation, likely representing a fine-tuning of relevant neu-
ral systems. Here, the shift to focal activation may represent
increases in the magnitude or extent of activation in regions
critical to task performance, whereas the shift from diffuse acti-
vation represents attenuation of activation in other, less critical,
cortical areas, either in the number of regions activated, or in
the extent of activated tissue. Other developmental changes may
include new brain regions coming online, while the involvement
of others is reduced, and may be related to shifts in cognitive
strategies.
One of the challenges in investigating the interplay between
cognitive development and maturation of the brain is to sepa-
rate the contributions of neural changes specific to development
and learning. Training studies in children will be necessary to
tease these apart ultimately (e.g., Klingberg et al., 2005), but
imaging studies equating performance are critical in separating
 S. Durston, B.J. Casey / Neuropsychologia 44 (2006) 2149–2157
which changes are related to maturation versus differences in
performance between groups. Other, new and promising imag-
ing approaches include controlling for large variation between
individuals by tracking both cognitive and brain development
in the same individuals over time in longitudinal studies. Fur-
thermore, DTI is now being used to relate the maturation of
white matter tracts to cognitive development, and may refine
our understanding of the role of local changes in white matter
to cognitive development (see Fig. 1). Finally, investigations of
developmental disorders such as ADHD have the potential to
inform us on the role of circuitry affected in these disorders, by
relating neural to cognitive changes in affected individuals.
Acknowledgements
The authors gratefully acknowledge Drs. R.M. Mandl and
M.J. Mulder for their assistance in constructing Fig. 1, and
two anonymous reviewers for their helpful suggestions. The
work described in this paper was supported in part by an NWO
VENI-grant to SD, and P01 MH62196, R01 MH63255, and R21
DA15882 to BJC.
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