Child Development, July/August 2009, Volume 80, Number 4, Pages 1147–1162

Neurodevelopmental Correlates of Theory of Mind in Preschool Children

Mark A. Sabbagh, Lindsay C. Bowman, Lyndsay E. Evraire, and Jennie M. B. Ito
Queen’s University at Kingston

Baseline electroencephalogram (EEG) data were collected from twenty-nine 4-year-old children who also com-
pleted batteries of representational theory-of-mind (RTM) tasks and executive functioning (EF) tasks. Neural
sources of children’s EEG alpha (6–9 Hz) were estimated and analyzed to determine whether individual dif-
ferences in regional EEG alpha activity predicted children’s RTM performance, while statistically controlling
for children’s age and EF skills. Results showed that individual differences in EEG alpha activity localized to
the dorsal medial prefrontal cortex (dMPFC) and the right temporal–parietal juncture (rTPJ) were positively
associated with children’s RTM performance. These findings suggest that the maturation of dMPFC and rTPJ
is a critical constituent of preschoolers’ explicit theory-of-mind development.

The facility with which we both create and negoti-
ate our social worlds is owed in part to having
a theory of mind—the understanding that people’s
observable actions are motivated by internal men-
tal states (e.g., beliefs, desires) that are related to,
but ultimately distinct from, reality (Perner, 1991;
Wellman, 1990). A key entailment of a mature
theory-of-mind understanding is that mental
states—particularly epistemic mental states such as
beliefs and knowledge—are person-specific repre-
sentations of the world. This understanding is
sometimes called a ‘‘representational’’ theory of
mind (RTM) and is typically indexed by a canonical
battery of tasks, including the ‘‘false belief’’ task in
which children are asked to predict how they or
another would act on the basis of a belief that does
not comport with reality (e.g., Perner, 1991). Recent
research using looking-time paradigms has sug-
gested that an implicit appreciation of RTM may
be present in infants as young as 12 months old
(e.g., Onishi & Baillargeon, 2005; Surian, Caldi, &
Sperber, 2008). However, an explicit understanding
of RTM seems to undergo a more protracted time
We thank the families from the Kingston, Ontario community
who graciously volunteered their participation. We also thank
Phil Zelazo, Don Tucker, Nathan Fox, and Rebecca Saxe for help-
ful advice and discussion of the methodology, statistical analy-
ses, and interpretation of the data. This research was supported
by a CFI New Opportunities award and NSERC Discovery Grant
awarded to Sabbagh, and by NSERC undergraduate summer fel-
lowships awarded to Bowman and Evraire. Bowman is now at
University of Michigan, and Evraire is now at University of Wes-
tern Ontario.
Correspondence concerning this article should be addressed to
Mark A. Sabbagh, Psychology Department, Queen’s University,
Kingston, Ontario, Canada, K7L 3N6. Electronic mail may be
sent to sabbagh@queensu.ca.
course, emerging between the ages of 3 and 5 years
old (Wellman, Cross, & Watson, 2001).
The timetable of explicit RTM development is
roughly the same across cultures; although there
are some subtle and important variations, 3-year-
olds typically fail RTM tasks whereas 5-year-olds
show near-ceiling performance (Callaghan et al.,
2005; Liu, Wellman, Tardif, & Sabbagh, 2008; Sab-
bagh, Moses, & Shiverick, 2006). One known excep-
tion to this stereotyped developmental timetable
comes from autism, a neurodevelopmental disorder
in which performance on marker tasks of explicit
RTM development is particularly impaired (Baron-
Cohen, 2001; Peterson, Wellman, & Liu, 2005).
These findings of cross-cultural synchrony, com-
bined with the relatively specific impairments seen
in autism suggest that specific neuromaturational
factors may be associated with explicit RTM devel-
opment. The goal of this study was to gain an ini-
tial characterization of the neurodevelopmental
correlates of RTM in preschoolers.
Electroencephalogram Measures of Neurocognitive
Development
Electrophysiological measures provide one of the
most widely used and reliable windows on young
children’s neurocognitive development. Of particu-
lar interest has been developmental changes in the
‘‘alpha band’’ (6–9 Hz) of children’s electroenceph-
alogram (EEG). From infancy through the preschool
 2009, Copyright the Author(s)
Journal Compilation  2009, Society for Research in Child Development, Inc.
All rights reserved. 0009-3920/2009/8004-0015
1148 Sabbagh, Bowman, Evraire, and Ito
years, alpha gradually becomes the highest ampli-
tude resting rhythm over all regions of the scalp
(Marshall, Bar-Haim, & Fox, 2002). Alongside the
changes in alpha amplitude (power), there are also
important regional changes in the EEG alpha coher-
ence, a measure of the extent to which spectral EEG
is correlated at any two electrode sites (Thatcher,
1994; Thatcher, Walker, & Guidice, 1987), In gen-
eral, increases in EEG coherence reflect increases in
synchronized neural firing either within or across
neural populations (Nunez, 1995). When applied in
the developmental context, increases in resting
alpha coherence are generally thought to reflect
functional, maturational changes in the organiza-
tion of neurocognitive systems (Thatcher, 1992).
Recent advances in EEG analysis have made it
possible to use the cross-spectral matrix (essentially
a matrix of all possible pairwise coherence measures
within an EEG recording montage), to estimate the
intracerebral sources of spectral EEG power. One
such technique is standardized low-resolution elec-
tromagnetic tomography (sLORETA) that computes
current density throughout the solution space (6,237
5 mm3 voxels within gray matter and hippocampus;
Pascual-Marqui, 2002). The sLORETA method dis-
tinguishes itself from some other source localization
solutions such as brain electrical source analysis, in
that it does not seek a fixed number of sources of
scalp activity; instead, it estimates the current den-
sity at all points in the solution space. The general
sLORETA technique has been used extensively and
validated in event-related potential (ERP) studies
(see Pascual-Marqui, Esslen, Kochi, & Lehmann,
2002, for a review). More relevant to the present arti-
cle, recent research with adults has used a version
of the sLORETA method to show how group differ-
ences in regional current density estimated from
spectral EEG (including alpha) predicts aspects of
cognitive and affective information processing (Piz-
zagalli, Peccoralo, Davidson, & Cohen, 2006; Pizza-
galli, Sherwood, Henriques, & Davidson, 2005).
Because of the previously described relations
between resting EEG alpha and functional neural
development, we reasoned that regional increases
in current density estimates can be taken to reflect
increased synchronous activity within associated
neural assemblies, which in turn can be attributed
to ongoing neurodevelopmental processes
(Thatcher, 1992). Thus, in the present study we
sought to investigate the neurodevelopmental cor-
relates of RTM development by examining the
extent to which individual differences in preschool-
ers’ regional sLORETA current density estimations
(based upon resting EEG alpha activity) were posi-
tively associated with their performance on a bat-
tery of RTM tasks.
Neurodevelopmental Bases of RTM Development:
Predictions
There is now a considerable body of literature
investigating the neural bases of RTM reasoning in
adults. An in-depth review of this literature is
beyond the scope of this report. In brief, this litera-
ture has identified several neural regions that are
most consistently associated with the kind of RTM
reasoning that children acquire during the pre-
school year (e.g., false-belief reasoning), including
the dorsal medial prefrontal cortex (dMPFC), the
right and left temporal–parietal juncture (TPJ), and
the precuneus (PC; Gallagher & Frith, 2003; Saxe,
2006). Intriguingly, some recent research suggests
that these same regions (along with some other
regions) provide the neural substrates for RTM rea-
soning in school-aged children (Kobayashi, Glover,
& Temple, 2007). Accordingly, a straightforward
developmental hypothesis is that preschoolers’
RTM development is associated with the functional
maturation of these regions.
There are a couple of reasons, however, to ques-
tion whether this straightforward hypothesis would
bear out. First, and perhaps most fundamentally, it
is not clear whether it is appropriate to assume that
the functional neural specializations seen in expert
adults (or older children) would be the same as
those seen in preschoolers who are only acquiring
an explicit RTM reasoning. Karmiloff-Smith (1997)
noted that it is not possible to determine solely
from research with adults whether observed con-
tent-specific functional specializations are present
from the outset of development or whether they are
the outcome of a more protracted developmental
process. A second related concern is that even if
RTM reasoning is associated with generally homol-
ogous regions in adults and children, it is possible
that their contributions would not be as domain-
specific in children as they appear to be in adults.
These regions may perform more domain-general
computations and their specialization for RTM rea-
soning might emerge with development.
While these concerns might apply broadly, they
can currently be most well articulated with respect
to the role that dMPFC might play in RTM devel-
opment. A recent review suggests that medial
frontal regions, extending into the dMPFC, are
especially involved in some aspects of executive
functioning (EF; Ridderinkhof, Ullsperger, Crone, &
Nieuwenhuis, 2004). In particular, medial frontal

regions are recruited when people are required to
negotiate situations in which there is either
response conflict or decision uncertainty. What
makes this particularly interesting from a develop-
mental perspective is that there is now extensive
evidence showing that response-conflict EF tasks
are strongly associated with preschoolers’ RTM
development (Carlson & Moses, 2001; Hala, Hug, &
Henderson, 2003; Hughes, 1998; Perner, Lang, &
Kloo, 2002). There are several reasons that
response-conflict EF skills associated with medial
frontal functioning might be intrinsically connected
with children’s abilities to reason about RTM (see
Moses, 2001, for a fuller treatment). In short, RTM
tasks typically have the methodological task
demand of requiring children to inhibit their habit-
ual tendency to refer to the true location of an
object to reference the falsely believed location. Fur-
thermore, once children are able to conceptualize
that epistemic states can mismatch reality, they
have to inhibit their usually correct strategy of
assuming that epistemic states match reality. In any
case, the fact that response-conflict EF and RTM are
associated in development, and both are associated
with medial prefrontal cortex, raises the possibility
that the contribution of dMPFC to RTM develop-
ment is mediated though its contribution to
response-conflict EF.
Thus, in this first investigation of the neurode-
velopmental bases of preschoolers’ RTM reasoning,
we included in our research design a battery of
response-conflict EF tasks that are typically associ-
ated with RTM development in preschoolers. This
allowed us to statistically control for children’s EF
skills and determine whether any potential associa-
tion between current density in dMPFC and RTM
development can be accounted for by a common
association with EF in development.
Method
Participants
Eighty-eight typically developing 4-year-olds
(range = 48–62 months, M = 53.57, SD = 3.71) were
recruited to the study. Although no systematic
demographic data were collected, participants were
from a predominantly European Canadian, middle-
class, and military background, reflecting the
demographics of the region in southeast Ontario,
Canada from which they were drawn. All children
were tested in two laboratory visits. The two visits
occurred within 2 weeks of each other. Participants’
age in months was calculated from their birth date
Neurodevelopment and Theory of Mind 1149
at the time of the first session. Parents reported that
all participants were born within 2 weeks of their
original due date, were developing typically, and
had no history of neuropsychological disease or
trauma.
Our criteria for retaining participants were that
they had to complete all of the focal tasks in the
battery, and contribute at least 50 s of artifact-free
EEG data for use in analyses. The EEG criterion is
comparable with that used in previous child EEG
studies (Fox, Rubin, Calkins, & Marshall, 1995). Of
the initial 88 participants, 10 children were excluded
because they failed to complete the behavioral
battery. An additional 49 were excluded because
they did not contribute sufficient EEG data. To help
explain why such a large number of children failed
to contribute sufficient EEG, we checked our records
and found that of our first 30 participants, 27 did
not meet the EEG criterion. This high exclusion rate
in the first third of the study was likely attributable
to our inexperience collecting EEG data with 4-year-
old children. After the first 30 participants, we were
more successful on this front and were able to retain
about half of all participants tested.
Thus, the final sample consisted of 29 children (9
boys and 20 girls). Ages ranged from 48 to
62 months (M = 54.00, SD = 3.70). The large num-
ber of exclusions raised questions about whether
our final sample might show a selection bias,
thereby limiting the generality of the findings. To
address this possibility, we report following analy-
ses comparing the included and excluded samples
on the behavioral measures.
Measures and Materials
All of the behavioral tasks that we used in this
study have been used several times in other stud-
ies. Thus, here we provide only a brief description
of each task and how it was scored for purposes of
the present analyses.
RTM Battery
Knowledge access (Wellman & Liu, 2004). Children
were shown a small wooden box and asked what
they thought was inside it. After children took a
guess, or responded that they did not know, they
were shown that the box contained a toy elephant.
Children were then asked if the puppet, ‘‘Tiger,’’
knew what was inside the box given that he had
never seen inside. Children passed the task if they
answered that Tiger did not know what was inside
the box (score = 0–1).
1150 Sabbagh, Bowman, Evraire, and Ito
False belief: Contents (Gopnik & Astington, 1988). Chil-
dren were shown a Smarties box and asked what
they thought was inside. After children responded
that they thought the box contained Smarties or
chocolates, they were shown that the box contained
crayons. Children were then asked what ‘‘Mickey
Mouse’’ (a puppet) would think was inside the
Smarties box given that Mickey had never seen
inside the box. Children passed the task if they
answered that Mickey would think the box con-
tained Smarties (score = 0–1).
False belief: Location (Wimmer & Perner, 1983). Children
were shown a scenario in which one puppet,
‘‘Heidi,’’ hid a plane under her bed, and then
another puppet, ‘‘Andy,’’ moved the plane from
the bed to the toy box whereas Heidi was not
looking. Children were then asked where Heidi
would look for her plane. Children passed the task
if they answered that Heidi would look for her
plane under the bed where she left it (score = 0–1).
Appearance–reality (Flavell, Green, & Flavell, 1986). Chil-
dren were shown a sponge that was painted to look
like a rock and asked what they thought it was.
After children responded that they thought it was a
rock, they were shown that it was a sponge. Child-
ren were then asked again what the object looked
like. Children passed the task if they correctly
answered that it looked like a rock (score = 0–1).
EF Battery
Grass–snow Stroop (Carlson & Moses, 2001). Although
grass is typically associated with the color green,
and snow is typically associated with the color
white, children were instructed to reverse these
associations and point to a green card when the
experimenter said ‘‘snow’’ and to a white card
when the experimenter said ‘‘grass.’’ The final score
was the proportion (percentage) of correct
responses over 16 trials. Sometimes children made
multiple responses on a single trial, but only their
first responses were scored.
Bear–dragon (Reed, Pien, & Rothbart, 1984). Children
were instructed to follow the commands given by a
bear puppet (e.g., ‘‘touch your nose’’) but to ignore
any commands from a dragon puppet. All com-
mands required hand actions only. On the ‘‘dragon
trials,’’ children received a score of 0 if they fully
complied to the dragon’s command, a score of 1 if
they performed a partial movement in response to
the dragon’s command, a score of 2 if they per-
formed a movement that did not correspond to the
dragon’s command, a score of 3 if they did not
respond to the dragon’s command but used a strat-
egy to help them avoid compliance (such as saying
‘‘no’’ to the dragon or sitting on their hands), and a
score of 4 if they did not move at all in response to
the dragon’s command and did not use any strat-
egy (range = 0–20).
Dimensional-change card sort (Zelazo, 2006). Chil-
dren were instructed to sort cards that varied on
two dimensions: (a) color (red and blue) and (b)
shape (boats and rabbits). First, children were
instructed to sort the cards according to shape (i.e.,
boats in one basket, rabbits in the other). Then, they
were asked to switch and sort the cards according
to color (i.e., red in one basket, blue in the other).
Children were given a score based on the number
of postswitch sorts that clearly demonstrated they
were sorting based on the second dimension (i.e.,
color; range = 0–3).
Less is more (Carlson, Davis, & Leach, 2005). Children
were shown two trays: one containing a large
amount of candy (i.e., five jelly beans) and one con-
taining a small amount of candy (i.e., two jelly
beans). Children were told that when they pointed
to a tray, the candy in that tray would go into
‘‘Naughty Monkey’s’’ cup, and they would get the
candy in the other tray to put in their cup. Children
received a score of 0 if they pointed to the tray with
the larger amount, a score of 1 if they pointed
directly to the smaller amount. Scores were
summed across 16 consecutive trials (range = 0–16).
EEG Recording
Rocket ship–spiral line video. During EEG record-
ing, participants watched a 6-min video that had
two components: (a) a still picture of a rocket ship
and (b) an animation of a green line that mapped
out a spiral (alternating expanding and contracting
spirals with each presentation). The rocket ship and
spiraling line components were each 30 s in length
and were each presented six times, in alternation.
The video began with an expanding spiraling line
clip. Because it is difficult for preschoolers to sit
still, the alternating video allowed us to instruct
children that it was okay to move a little during the
spiraling line segments, but to do their best to stay
still while the picture of the rocket ship was on the
screen. This method was adopted from previous
research (Fox et al., 1995) and is the most well-
established protocol for collecting baseline or rest-
ing EEG alpha with preschoolers.
EEG acquisition and analysis. Electroencephalogram
was recorded from the scalp with a 128-channel
Geodesic Sensor Net (EGI, Eugene, OR). The net
consisted of 128 carbon-fiber electrodes knitted into

an elastic geodesic tension structure that when
applied, distributed electrodes evenly over the
scalp. Each electrode was positioned relative to the
vertex electrode in a geodesic montage that
stretches around the sphere of the head. Before
application, the net soaked for 5 min in a mild elec-
trolyte solution to aid conductance.
Electroencephalogram activity at all channels
was recorded referenced to the vertex electrode
(Cz), sampled at 500 Hz, and digitally filtered
between 0.01 and 200 Hz (time constant = 1 s). EEG
was recorded continuously throughout the video,
and any necessary electrode readjustments were
made during sections of the video for which no
data analyses were planned. Impedances were
maintained below 30 kX throughout recording.
The raw EEG recordings were filtered (60-Hz
notch) and edited to include only recordings
that were made while participants quietly viewed
the still photograph of the rocket ship. These
data were then further divided into smaller 2-s
segments, which were in turn submitted to a soft-
ware algorithmic artifact rejection program (Vision
Analyzer; Brain Vision GMBH, Gilching, Germany)
that combed the data for evidence of artifact (gradi-
ent threshold = ±50 lV in 100 ms, amplitude
threshold = ±200 lV, global maximum difference
threshold = ±300 lV). Hand-coding of 25% of the
EEG records confirmed that these criteria reliably
identified artifact due to blink, eye movement, and
participant movement. Segments that contained
artifact were removed from further analyses. Only
participants who contributed at least 25 good seg-
ments of EEG (i.e., 50 s of data) were considered
for analysis. The artifact-free EEG segments were
then transformed to average reference to ensure
accurate source-localization.
Cross-spectral matrices were created for the
single-subject average-referenced data in 0.5-Hz
frequency intervals from 5.5 to 9.5 Hz, thereby
ensuring we captured activity in the preschoolers’
alpha band (i.e., 6–9 Hz). The nine resulting cross-
spectral matrices were then used to compute
three-dimensional distributions of the standardized
current density using standardized estimates of the
minimum norm inverse solution as applied by sLO-
RETA (see Pascual-Marqui, 2002, for details).
The sLORETA transformation results in activation
values for 6,237 ‘‘voxels’’ (5 mm3) located within
cortical gray matter and hippocampus, as defined
by the Probability Atlas from the Montreal Neuro-
logical Institute. Preliminary analyses showed that
participants’ sLORETA current density estimations
across the nine frequency bins were highly inter-
Neurodevelopment and Theory of Mind 1151
correlated at all voxels (Cronbach’s a = .88–.99,
M = 0.95), and so the sLORETA activation values
were collapsed across the nine frequency bins to
create a final reliable measure of the regional
current-source activation values in the 6–9 Hz
(alpha) band at each voxel for each participant.
Procedure
This study was part of a larger investigation of
the neurodevelopmental correlates of children’s
social-cognitive development, and thus, in addition
to the RTM and EF batteries described above, the
larger study included a number of additional tasks
that were not analyzed in this study. Most promi-
nently, there were a number of behavioral measures
of children’s pretend play including two structured
role playing activities, two versions of the ‘‘Moe’’
task (Lillard, 1993), a self-pretense task (Mitchell &
Neal, 2005), and a measure of children’s narrative
absorption (Rall & Harris, 2000). We also included
two tasks that are considered precursors to RTM
understanding, including children’s understanding
of diverse desires and diverse beliefs (Wellman &
Liu, 2004). Finally, we included a standard measure
of children’s vocabulary development (PPVT–III;
Dunn & Dunn, 1997). Given the number of
behavioral measures, children were asked to visit
an on-campus laboratory for two sessions. We fol-
lowed standard practice for individual differences
studies and administered the tasks in a fixed order
to all participants (see Carlson & Moses, 2001).
Session 1
After obtaining informed consent, we recorded
electrophysiological data (EEG) from the scalp using
the presoaked sensor net. Participants sat in a stable
chair while an experimenter applied the net and
adjusted it to ensure correct placement and good
contact between the electrodes and the scalp. The
final net placement was photographed and later
examined to verify that net placement was correct at
the outset of the session. Participants were then
given the instructions for watching the video
described above, the lights were dimmed for opti-
mal video viewing, and EEG recording began. Par-
ents were allowed to remain in the room with their
child during recording, and both parent and child
were asked to be as quiet as possible throughout the
procedure. Net placement, adjustment, instructions,
and recording took approximately 20–25 min.
After EEG recording, the experimenter removed
the net, and participants moved to a different room
1152 Sabbagh, Bowman, Evraire, and Ito

for behavioral testing. For the first session, behavioral
tasks were administered in the following order: (a)
false belief: contents, (b) diverse desires, (c) diverse
beliefs, (d) knowledge access, (e) grass–snow, (f)
role-play activity, (g) narrative task, and (h) Moe
task.
Session 2
Only behavioral data were collected in Session 2.
After obtaining informed consent, participants were
tested with a single experimenter in the same room
as Session 1. The tasks were administered in the
following order: (a) bear–dragon, (b) false belief:
location, (c) card sort, (d) appearance-reality, (e)
role play activity, (f) self-pretense, (g) Moe task, (h)
less is more, and (i) PPVT. Session 2 took approxi-
mately 45 min.
Results
Behavioral Data
For the RTM battery, preliminary analyses
showed that performance was in line with previous
research that has used these tasks (see Table 1).
Scale reliability analyses showed that the RTM bat-
tery had moderate internal consistency (a = .623),
which is also comparable with theory-of-mind bat-
teries used in previous research. Children’s perfor-
mance on the final battery was associated with
children’s age in months, r(27) = .444, p = .016.
For the EF battery, preliminary analyses revealed
ceiling level performance and low battery correla-
tion for the bear–dragon task. Upon closer inspec-
tion of the preschool EF literature, we discovered
that very strong performance on bear–dragon is
common for children in this age group (see Carlson,
2005), and so we omitted it from the task battery.
Thus, our EF battery was composed of the three
remaining tasks: grass–snow, card sort, and less is
more. This battery also showed moderate internal
consistency (a = .554), but was not significantly
associated with children’s age, r(28) = .113, p = .559.
The results for each of the behavioral tasks from
the final RTM and EF batteries are presented in
Table 1, along with statistical comparisons of the
groups who were included in the final EEG analyses
versus those who were excluded. Apparent from
these analyses was that the included and excluded
groups differed significantly on only one task—that
is, children in the EEG excluded group performed
better on the knowledge access task than did children
who were included in the final sample. The overall
parity between the included and excluded groups
was most powerfully demonstrated in the compari-
son of children’s performance on the battery aggre-
gates, which were nearly identical across groups.
These findings provide some confidence that there
was nothing particularly special about the included
versus excluded group, and thus strengthen our abil-
ity to generalize from our findings.
For the final sample, children’s performance
on the RTM battery was highly correlated with
their performance on the EF battery, r(27) = .575,
p = .001, and this relation was unaffected by statisti-
cally controlling for age, r(26) = .590, p = .001.
sLORETA Analyses
Our main goal in these analyses was to deter-
mine whether individual differences in regional

Table 1
Children’s Performance on the Theory-of-Mind (ToM) and Executive Functioning (EF) Batteries

Included (n = 29) Excluded (n = 49) Test of difference

ToM tasksa
Knowledge access 72.4% 94.0% v2(1) = 8.27, p = .004
False belief: contents 37.9% 52.0% v2(1) = 1.51, p = .219
False belief: location 72.4% 66.0% v2(1) = 0.36, p = .547
Appearance–reality 69.0% 66.0% v2(1) = 0.07, p = .785
Battery (mean total of 4) 2.55 (1.30) 2.76 (1.11) t(76) = 0.734, p = .465
EF tasksb
Grass–snow 78.22% (27.67) 78.68% (25.53) t(76) = 0.07, p = .944
Card sort 72.41% (42.79) 72.67% (42.16) t(76) = 0.02, p = .984
Less is more 79.96% (27.11) 77.62% (25.71) t(76) = 0.38, p = .705
Battery (mean of 3, with SD) 79.91% (22.09) 75.84% (22.89) t(76) = 0.768, p = .445

a
Figures in this category represent percent passing each task. bFigures in this category represent mean percent trials correct (with
standard deviation) on each task.

current-source estimations predicted children’s
performance on the RTM battery while controlling
for children’s age and performance on the EF bat-
tery. Preliminary inspection of the sLORETA data
showed that activation values were positively
skewed, and so were submitted to a natural log (ln)
transformation prior to the focal analyses. All anal-
yses were conducted using purpose-designed
scripts that called functions from the general and
statistics toolboxes of MATLAB (The MathWorks,
Natick, MA).
Whole Brain Analyses
Our analytical approach was to perform voxel-
wise partial correlations measuring the relationship
between sLORETA activation values and children’s
RTM performance while statistically controlling
children’s age and EF performance. To establish a
significance criterion for the whole brain analyses,
we conducted a permutation test in which 2,000
random permutations of the behavioral data were
submitted to the partial correlation analyses with
the sLORETA activation values. Results of this
showed that voxel-wise tests with a relatively leni-
ent p-value criterion (p < .01), combined with a
cluster-size criterion of 20 contiguous voxels was
associated with an acceptable family-wise alpha
level (p < .05).
Using these criteria, we identified five regions
at which current-source density was positively asso-
ciated with performance on the RTM tasks (see
Table 2). The strongest associations were located
within the dMPFC and the right TPJ (rTPJ) (see
Figure 1). Three additional clusters also met the
significance criteria (see Figure 2). One cluster was
located in motor and premotor cortex, and the
correlation with RTM performance was strongest in
the precentral gyrus. A second cluster was within
Table 2
Brain Regions in Which sLORETA Current Density Estimates Predicted Theory-of-Mind Performance
Region (MNI coordinates
of max. r) L⁄R Included BAs
dMPFC (5, 55, 40) L⁄R 8⁄9
rTPJ (55, )55, 30) R 40
Precentral gyrus (50, )15, 45) R 3⁄4⁄6
Cuneus (20, )95, 15) R 17 ⁄ 18 ⁄ 19
Inferior temporal (50, )70, )5) R 19 ⁄ 37
Note. sLORETA = standardized low-resolution electromagnetic tomography; BA = Brodmann area; dMPFC = dorsal medial prefrontal
cortex; rTPJ = right temporal–parietal juncture.
**p < .01.
Neurodevelopment and Theory of Mind 1153
middle occipital regions, and the correlation
with RTM was maximal at the cuneus. The third clus-
ter included posterior inferior temporal regions and
fusiform gyrus, but the correlation with RTM was
maximal at the posterior inferior temporal gyrus.
To further characterize these effects, we averaged
across the significant voxels in each region to create
one aggregate measure of activity in each region for
each subject. Each of these aggregates were strongly
correlated with RTM reasoning, with age and EF
performance statistically controlled (see Table 2).
Furthermore, the aggregates showed were largely
intercorrelated—only dMPFC and rTPJ did not
show a significant correlation with one another (see
Table 3). Because of the high degree of intercorrela-
tion among the aggregates, we conducted a stepwise
regression to determine the most parsimonious
model of the relation between regional current den-
sity and RTM performance. For this analysis, age
and EF were entered in the first block, and the aggre-
gate current density estimations were added step-
wise in the second block. Age and EF performance
each significantly predicted RTM reasoning in the
first block, F(2, 26) = 11.825, p < .001, Radj2 = .436.
The stepwise analysis of current-source density
aggregates showed that adding aggregates from the
dMPFC and rTPJ made a significant unique contri-
bution, DR2 = .235, Fchange(2, 24) = 10.315, p = .001.
The final model (see Table 4) including age, EF per-
formance and dMPFC and rTPJ aggregates
explained 67.1% of the variance in RTM reasoning,
F(4, 24) = 15.306, p < .001. The additional regions
(e.g., cuneus, inferior temporal, and precentral) were
excluded from the model in the stepwise analysis
because they did not explain unique variance in
RTM reasoning, thereby suggesting that the contri-
bution of those other regions could be largely
accounted for by their associations with either
dMPFC or rTPJ.
Max. rpartial Agg. rpartial
k df = 25 df = 25
44 .585** .580**
35 .552** .537**
30 .526** .521**
33 .525** .529**
28 .511** .513**
1154 Sabbagh, Bowman, Evraire, and Ito

(a)

(b)

Figure 1. Thresholded statistical map and scatterplot of partial correlations measuring the relation between RTM reasoning and
sLORETA estimates of current density at (a) dMPFC and (b) rTPJ. Only significant voxels (p < .01) are shown (in white). Voxels in all
figures are shown projected onto a template structural MRI to illustrate their neuroanatomical locations.
Note. RTM = representational theory of mind; sLORETA = standardized low-resolution electromagnetic tomography; dMPFC = dorsal
medial prefrontal cortex; rTPJ = right temporal–parietal juncture; MRI = magnetic resonance imaging.

(a) (b) (c)

Figure 2. Additional regions that contained clusters of voxels showing significant relations between sLORETA estimates of current
density and RTM reasoning. Only significant voxels (p < .01) are shown (in white).
Note. sLORETA = standardized low-resolution electromagnetic tomography; RTM = representational theory of mind.

children’s theory-of-mind reasoning. Thus, we con-

Dorsal Medial Prefrontal Cortex and Right Temporal–
Parietal Juncture
Of the regions identified in the whole-brain anal-
yses, the dMPFC and rTPJ have been most consis-
tently implicated in several adult and school-aged
ducted further analyses to determine whether the
regions we identified were essentially homologous
with those shown in previous research. We began
by defining regions of interest in the dMPFC and
rTPJ as 2-cm-diameter sphere drawn around the
 Neurodevelopment and Theory of Mind 1155

Table 3 voxel region in the rTPJ. The reason for the differ-
Partial Correlations Between Cluster Aggregates Identified in Whole- ence in the number of voxels included in each region
Brain Analysis (Controlling for Age and EF) is that the rTPJ has less cortical gray matter than the
Variable Inferior Temp. dMPFC rTPJ Precentral
dMPFC region. We then statistically compared the
clusters we identified with these regions of interest.
Cuneus .905 .418 .757 .596 The results of this analysis are summarized in Fig-
Inferior Temp. .389 .779 .662 ure 3. Descriptively speaking, 22 ⁄ 32 (68.8%) voxels
dMPFC .366 .549 in the dMPFC and 12 ⁄ 21 (57.1%) voxels included in
rTPJ .746 the region of interest overlapped with the activation
Note. EF = executive functioning; dMPFC = dorsal medial
cluster we identified in the whole-brain analysis. In
prefrontal cortex; rTPJ = right temporal–parietal juncture. the dMPFC, our cluster extended to slightly more
Table 4
inferior regions. For the rTPJ, our cluster included
Regression Coefficients for Final Model Predicting RTM Development voxels that were slightly superior and posterior to
From Age, Executive Functioning, and Current-Source Density the region of interest, and did not extend to more
inferior regions. Although we did not achieve com-
Variable Standarized b t value p value plete overlap, permutation tests (2,000 iterations)
showed that this degree of overlap is an extremely
Age .321 2.903 .008
rare result by chance in both the dMPFC and the
Executive functioning .574 4.994 <.001
dMPFC aggregate .317 2.707 .012
rTPJ (both p £ .0005). This suggests that the dMPFC
rTPJ aggregate .292 2.402 .024 and rTPJ regions in which current-source density
was positively correlated with RTM performance
Note. RTM = representational theory of mind; dMPFC = dorsal were likely homologous with those that have been
medial prefrontal cortex; rTPJ = right temporal–parietal juncture.
shown to be active during RTM reasoning in adults.
average reported MNI coordinates for each area
showing peak activation in recent theory-of-mind
studies (dMPFC: 9, 54, 36, rounded to 10, 55, 35 to
Discussion
match our resolution 5 mm3; rTPJ: 56, )54, 19,
rounded to 55, )55, 20). This procedure resulted in a Our main finding was that individual differences in
32-voxel region of interest for dMPFC, and a 21- estimated current density attributed to the dMPFC

(a)

(b)

Figure 3. Overlap between observed results and predicted regions of interest in (a) dMPFC and (b) rTPJ.
Note. dMPFC = dorsal medial prefrontal cortex; rTPJ = right temporal–parietal juncture.
1156 Sabbagh, Bowman, Evraire, and Ito
and rTPJ were positively associated with preschool
children’s performance on a battery of RTM tasks. In
addition to these regions, we found some evidence
that individual differences in postcentral gyrus, cun-
eus, and posterior inferior temporal regions also
positively predicted RTM performance. We will dis-
cuss each of these findings in turn and discuss some
implications and limitations of this study.
Dorsal Medial Prefrontal Cortex
The region of dMPFC that we found was criti-
cally associated with RTM performance in children
is similar to the regions that have been associated
with RTM reasoning in functional magnetic reso-
nance imaging with adults and school-aged chil-
dren. From the outset, we questioned whether any
possible contributions of dMPFC to RTM develop-
ment could be accounted for by the role that
dMPFC might play in negotiating response conflict.
On this question, we showed that current density
attributable to dMPFC was related to RTM perfor-
mance, even when controlling for performance on
an EF battery that was also strongly correlated with
RTM performance. Furthermore, other research
from our laboratory has provided evidence that
when the partial correlation analyses are conducted
to assess the relations between current-source den-
sity and EF (controlling for age and RTM), the
results show that current-source density at dMPFC
are not associated with EF performance; instead, EF
is associated with individual differences in current-
source density attributed to cingulate (anterior and
posterior) and ventral-medial prefrontal cortex
(Sabbagh, Bowman, Evraire, & Ito, 2009). These
findings suggest that increased current density
attributable to this region of dMPFC is associated
with the emergence of explicit RTM reasoning in
young children, independent of the contributions
that these increases might also make to response-
conflict EF skills.
It is important to clarify that we do not wish to
downplay the role of EF in preschoolers’ explicit
RTM reasoning abilities. Indeed, our own findings
showed that EF was associated with RTM reason-
ing, and that this association survived the regres-
sion analysis in which the EF-RTM relation was
assessed while statistically controlling for age, and
current-source densities in dMPFC and rTPJ.
Clearly, then, EF (and presumably its associated
neural substrate) makes some independent contri-
bution to RTM performance in the preschool years.
Our finding here helps to clarify the uniqueness of
this contribution by highlighting that it is not
directly through a common relation with a particu-
lar region of dMPFC. Likewise, these findings clar-
ify that the role that the dMPFC plays in RTM
reasoning cannot easily be attributed to the role
that this region may play in response-conflict EF,
even in preschoolers.
Our finding that the dMPFC plays a role in
RTM reasoning that is independent of its possible
association with EF is consistent with a number of
studies suggesting that the neural substrates of
RTM reasoning are dissociable from those associ-
ated with response-conflict EF, even in closely
matched tasks (Kain & Perner, 2005; Saxe, Schulz,
& Jiang, 2006). Further, they are consistent with
some recent developmental data suggesting that
the timetable of EF and RTM can be dissociated in
development. For instance, Sabbagh et al. (2006)
recently showed that Chinese preschoolers’ EF
skills were more advanced than those in a sample
of North American preschoolers; however, the two
groups showed no differences in their RTM devel-
opment (see also Oh & Lewis, 2008). Likewise,
there are a number of reports now suggesting that
RTM skills might vary across groups that are gen-
erally equated for their EF abilities (e.g., Liu et al.,
2008). These cross-cultural dissociations between
the developmental trajectories of EF and RTM may
be attributable to the fact that the two skills may
have partly dissociable neural substrates during
the preschool years.
Yet, more research is required to better under-
stand what the dMPFC does for explicit RTM
development. Two different hypotheses about the
role that dMPFC might play in RTM reasoning
have emerged from recent reviews of the adult
neuroimaging literature. Amodio and Frith (2006)
suggest that the region of dMPFC that predicted
RTM reasoning in the current study (what they
call the anterior rostral medial frontal cortex) per-
forms the computations associated with complex
metarepresentational, and possibly emotional rea-
soning. This suggestion is based upon the fact
that dMPFC activations are observed as adults
perform false belief tasks, and in tasks in which
participants make probabilistic judgments about
others’ likely beliefs and intentions (e.g., judg-
ments of others’ subjective experience of pain and
emotion, economic games). Saxe (2006), in con-
trast, summarizes the available data by suggesting
a more limited role for the dMPFC. Specifically,
Saxe argues that the dMPFC is important for
representing triadic attentional relations (i.e., rep-
resenting the relations between self, other, and
an object), but finds no conclusive evidence to

suggest that the dMPFC generates or represents
the propositional contents of mental states them-
selves. A clear direction for future research is to
adapt paradigms like the one we used here to
better understand the neural bases of preschool-
ers’ theory of mind development, based upon
insights gleaned from the burgeoning adult
literature.
Right Temporal–Parietal Juncture
Our findings showed that individual differences
in current density estimations localized to rTPJ
predicted preschoolers’ performance on the RTM
battery. These findings confirm that the region that
appears to be most consistently and specifically
associated with RTM reasoning in adults is also
implicated in explicit RTM development. Saxe and
colleagues (Saxe, 2006; Saxe & Powell, 2006) have
presented evidence to suggest that rTPJ performs
the highly specific task of reasoning about repre-
sentational mental states. Our findings are certainly
consistent with the notion that rTPJ plays a critical
role in RTM development; though, our study was
not designed with the aim of licensing fine-grained
inferences about this important region.
More generally, the hypothesis of a domain-spe-
cific role for rTPJ in RTM reasoning has not been
explored in the developmental literature. However,
a recent challenge has emerged from work in visual
neuroscience noting that the region of rTPJ that is
involved in RTM reasoning is isomorphic with
regions that are critical for coordinating shifts of
visual attention (Mitchell, 2008). When tested in the
same subject, the same regions that are associated
with attention shifting (in an ostensibly nonsocial
context) tasks show considerable overlap with those
recruited for RTM reasoning. These findings raise
the intriguing possibility that the association of
rTPJ with RTM may be attributable, at least in part,
to a domain-general rather than domain-specific
skill. Were an ontogenetic relation between atten-
tion shifting and RTM performance to exist, we
suggest that an approach like the one we adopted
here would be fruitful in clarifying the functional
relation between rTPJ and RTM reasoning in the
preschool years.
Additional Regions
We found evidence that variations in current
density attributed to the cuneus, inferior temporal
lobe or fusiform gyrus, and precentral gyrus were
also associated with RTM development. Each of
Neurodevelopment and Theory of Mind 1157
these areas has been linked with performance in
social cognitive tasks, if not RTM reasoning more
specifically. We will address briefly the possible
contributions of each region to RTM development
in turn.
Precentral Gyrus
Across several studies, the precentral gyrus has
been implicated in tasks that require distinguishing
one’s own from another’s visual perspective (Aich-
horn, Perner, Kronbichler, Staffen, & Ladurner,
2006; Ruby & Decety, 2001). Although the concep-
tual links between perspective taking and theory of
mind are clear in that both would seem to require
reasoning about other’s psychological states (Flav-
ell, 1999), there is relatively scant empirical evi-
dence to suggest that tasks tapping the two
constructs are related in development. Nonetheless,
it does seem likely that the cognitive operations
associated with suppressing one’s own perspective
are critical for performance on RTM tasks in chil-
dren (Birch & Bloom, 2007), and even in adults
under circumstances in which one’s own perspec-
tive is particularly salient (Keysar, Lin, & Barr,
2003).
Cuneus
Several studies of social-cognitive skills related
to theory of mind have noted cuneus activation in
at least some of their statistical comparisons (e.g.,
Vogeley et al., 2001). Even more interesting with
respect to the present findings is that Kobayashi
et al. (2007) found that activation in the cuneus
distinguished theory-of-mind from non-theory-of-
mind reasoning in children, but not in adults. These
preliminary findings, together with ours suggest
that the cuneus may make an especially important
contribution to RTM development.
With respect to characterizing the contribution of
the cuneus to RTM development, it can be noted
that the cuneus is most reliably engaged during
tasks that require reasoning about self knowledge
(Aichhorn et al., 2006). One possibility, then, is that
the relation we observed between RTM perfor-
mance and current density variations in cuneus
may reflect the extent to which RTM performance
in children relies on reflecting upon one’s own
knowledge states. The cuneus is also recruited in
tasks that involve mental imagery (Kosslyn et al.,
1999). Thus, a second possibility is that cuneus
function is important to RTM reasoning because
RTM tasks may require children to reimagine the
1158 Sabbagh, Bowman, Evraire, and Ito
scenario to answer correctly the question. In some
respects, this latter explanation may be preferred
because it potentially accounts for why the cuneus
is implicated in RTM reasoning in children (but
less reliably so in adults), for whom using mental
imagery might provide some necessary assistance
in recalling the foregoing events.
Inferior Temporal Lobe
In adults, participation in the now common
‘‘intentional attribution’’ task reliably elicits activity
in the inferior temporal lobe extending into the
fusiform gyrus (Brunet, Sarfati, Hardy-Bayle, &
Decety, 2000; Vollm et al., 2006). In children, inten-
tion attribution is typically a precursor of explicit
RTM reasoning, and is present quite early in devel-
opment (Woodward, 2003). These precursors are
ontogenetically associated with later RTM develop-
ment. For instance, Wellman and colleagues (Well-
man, Lopez-Duran, LaBounty, & Hamilton, 2007;
Wellman, Phillips, Dunphy-Lelil, & LaLonde, 2004)
have shown that performance in a looking time
study designed to measure 12-month olds’ under-
standing of the link between emotion and intention
was associated with those same children’s theory-
of-mind development during the preschool years.
Although the mechanism underlying this develop-
mental relation is currently unclear, the fact that a
developmental relation exists raises the possibility
that the development of inferior temporal regions
associated with intention attribution may play some
crucial role in the development of the regions that
are more specifically related to later RTM advances
(i.e., rTPJ, dMPFC).
Summary
In general, these additional regions have been
associated with cognitive abilities that might be
either considered necessary for solving RTM tasks
(i.e., mental imagery and suppressing first-
person perspective), or conceptual precursors to
RTM reasoning (i.e., intentional understanding).
One possibility, then is that the computations associ-
ated with these regions may play a particularly
important role in RTM development per se. Preli-
minary support for this hypothesis comes from our
regression analyses in which we found that the con-
tributions of these additional regions were mediated
by their relations with either rTPJ or dMPFC.
Although there are many possible explanations for
this mediation effect, one is that the development of
these regions somehow contributes to the functional
maturation of dMPFC and rTPJ, which in turn make
up the primary neural substrate for RTM reasoning.
Longitudinal research may prove fruitful in further
understanding the role that these additional regions
play in the development of the neural systems
critical for RTM development.
Implications
From the outset, we took as our main goal estab-
lishing the neurodevelopmental correlates of RTM
reasoning in preschoolers. Of particular interest
was whether regions that are known to play a role
in RTM reasoning in adults would also be impli-
cated in children. On this question, our evidence
was clear. Individual differences in current density
estimates attributable to dMPFC and rTPJ were cor-
related with RTM skills in preschoolers. Thus, the
link between these brain areas and RTM develop-
ment is present in children just as these skills are
explicitly emerging. Further, insofar as variations in
current density estimates reflect the ongoing func-
tional maturation of these associated neural
regions, our findings provide some evidence that
the functional maturation of dMPFC and rTPJ is
associated with RTM development.
This finding lays the foundation for the explora-
tion of two interrelated questions in future
research. The first concerns whether these func-
tional specializations are a cause or a consequence
of RTM development. Some researchers have artic-
ulated the hypothesis that humans have evolved a
specific neurocognitive system that is dedicated to
performing the computations that are associated
with constructing and reasoning about representa-
tions of others’ mental states (Leslie, Friedman, &
German, 2004). Although far from providing con-
clusive evidence for this hypothesis, the current
findings are consistent with the possibility that the
dMPFC and rTPJ are constituents of that specially
evolved system, and that functional maturation of
this system must reach a threshold level to enable
explicit RTM reasoning. An alternative hypothesis
is that the increased coherence in dMPFC and rTPJ
reflect the outcome of a developmental process.
Across domains, the neural systems associated with
particular computations become reorganized and
more circumscribed with expertise (Casey, Giedd,
& Thomas, 2000). Thus, the relations we observed
may be attributable to children’s RTM performance
and neural coherence each reflecting increasing
expertise with explicit RTM reasoning. Although
we expect that it will be challenging, longitudinal
research that takes an approach similar to that

presented here may provide crucial insight into the
causal relations among regional neural develop-
ments and RTM reasoning.
It is important to note that the functional matura-
tion of any particular neural region is likely to be
affected by both endogenous and experiential fac-
tors. There are reasons to think that both might
play a particularly important role in establishing
the functional maturation of dMPFC and rTPJ.
With respect to endogenous factors, the develop-
ment of the frontal lobes has been associated with
maturational changes in dopaminergic functioning
(Diamond, 2002). Insofar as there is some endo-
genous mechanism underlying the developmental
trajectory of dopaminergic expression, it seems
possible that this mechanism would in turn set
constraints on the functional maturation of the
neural systems associated with RTM reasoning. The
presence of these kinds of endogenous neurodevel-
opmental constraints on RTM-related cortical devel-
opment might help explain why, as mentioned at
the outset of the article, the developmental trajectory
of RTM looks so similar across cultures, and is
specifically impaired in the case of autism.
With respect to experiential factors, there is now
substantial evidence to suggest that several experi-
ential variables, including parent–child conversa-
tions about mental states (Ruffman, Slade, &
Crowe, 2002), family size (Lewis, Freeman, Kyriaki-
dou, Maridaki-Kassotaki, & Berridge, 1996), and
socioeconomic status (Pears & Moses, 2003) affect
the timetable of RTM development. More dramati-
cally, the development of RTM is greatly protracted
in children with highly atypical experiences owing
to being born either blind (Peterson, Peterson, &
Webb, 2000) or born deaf into non-native signing
families (Woolfe, Want, & Siegal, 2002). An impor-
tant question, then, concerns whether and how
these experiential factors might affect the matura-
tion of the neural systems we identified as associ-
ated with RTM development. Understanding these
fundamentally developmental processes will pro-
vide deep insights into how endogenous neuromat-
urational constraints might interact with experience
to render the time course and trajectory of RTM
development in the preschool years.
Limitations
There are several limitations of this study. First,
the sLORETA estimates are based upon volume
conductance parameters (how electrical activity is
conducted through the scalp, skull, and cerebrospi-
nal fluid) that were developed from research on
Neurodevelopment and Theory of Mind 1159
adults. There is some research to suggest that adult
skulls are somewhat thicker than children’s skulls,
though the age-related trends are modest and not
as pronounced as individual differences within age
groups (Simms & Neely, 1989). It is unclear as to
how these minor differences in the conductance
parameters might have affected our intracerebral
current density estimates. Thus, some caution is
warranted in interpreting the present findings.
Second, there are two factors that may have
reduced the statistical power of our tests. One fac-
tor stems from the fact that we developed the trans-
formation parameters for the sLORETA based upon
average electrode locations because we did not
have efficient technology for gathering electrode
coordinates from participants individually. Relying
on average sensor locations, though common in
much EEG–ERP research, may have added noise to
the group localization analyses and prevented us
from detecting the involvement of additional areas.
Another factor that may have reduced the statistical
power of our tests detecting the involvement of
additional neural regions was the relatively limited
range scores possible on our RTM measure.
Although neither of these factors call into question
any of the findings we report, both leave open the
possibility that other regions might have been
detected, including regions that have been shown
in the prior literature—such as the left temporal–
parietal juncture and the precuneus or posterior
cingulate—with more sensitive dependent mea-
sures.
Finally, it was unfortunate that our inexperience
with pediatric EEG collection led to our excluding
so many participants at the outset of our study.
Although statistical analyses of the behavioral mea-
sures showed that there was no reason to believe
that the participants we excluded were substan-
tially different from the participants retained in the
final sample, our high exclusion rate does raise
some concern about the generality of these find-
ings. Future work that replicates and extends these
findings will be valuable in assuaging these
concerns.
Summary
We used regional current density estimates of
preschoolers’ resting EEG alpha to show that indi-
vidual differences in the functional maturation of
the dMPFC and rTPJ are associated with preschool-
ers’ performance on a battery of RTM tasks. Statisti-
cal analyses showed that these relations could not
be accounted for by their relations with children’s
1160 Sabbagh, Bowman, Evraire, and Ito
age, or their relations with EF skills. In addition,
we found that the cuneus, posterior inferior tempo-
ral lobe, and precentral gyrus were associated with
RTM performance; however, statistical analyses
suggested that their influence was mediated
through their common associations with dMPFC
and rTPJ. Taken together, these findings suggest
that the functional maturation of the dMPFC and
rTPJ may be a critical neural correlate of preschool-
ers’ RTM development. Future work is necessary to
determine both the specific contributions of these
regions to RTM development and the endogenous
and experiential factors that affect their functional
maturation.
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