Brain and Cognition 55 (2004) 1–10

www.elsevier.com/locate/b&c

Introduction

Development of orbitofrontal function: Current themes

and future directionsq
Keith Happaney,a,* Philip David Zelazo,a,b and Donald T. Stussa,c,d
a
Department of Psychology, University of Toronto, Toronto, Ont., Canada
b
Department of Psychiatry, University of Toronto, Toronto, Ont., Canada
c
Departments of Medicine (Neurology) and Rehabilitation Science, University of Toronto, Toronto, Ont., Canada
d
The Rotman Research Institute, Baycrest Centre for Geriatric Care, Toronto, Ont., Canada M6A 2E1
Accepted 8 January 2004
Available online 5 March 2004

Abstract

In recent years, an exciting thrust in the developmental research literature has been the focus on ‘‘executive’’ functions (EF).
However, the emphasis has been on the more purely cognitive aspects of EF operative in abstract reasoning and problem solving—
aspects associated mainly with dorsolateral frontal regions. Although the literature on adult neuropsychology has seen an emerging
and growing interest in the study of processes more related to orbitofrontal cortex (OFC), including emotion and personality,
research on the development of EF has lagged behind. This special issue of Brain and Cognition is intended to redress this imbalance
by bringing together researchers who are studying the nature and development of orbitofrontal function from a wide variety of
perspectives.
Ó 2004 Elsevier Inc. All rights reserved.

1. Introduction butions to EF, developmental research has lagged be-

In recent years, a great deal of research has been
conducted on the development of executive function
(EF)—the processes required for the conscious control
of thought and action. However, the emphasis has been
on the more cognitive aspects of EF operative in ab-
stract reasoning and problem solving—aspects associ-
ated mainly with dorsolateral frontal regions—rather
than the affective, motivational aspects associated more
with orbitofrontal cortex (OFC) and related medial
regions. DL-PFC and OFC are parts of an interactive
functional system, and they typically work together—
even in a single situation. Although the literature on
adult neuropsychology has seen an emerging and
growing interest in the study of OFC and its contri-
q
The Editors are very grateful for Dana LiebermannÕs help with
the preparation of this Special Issue.
*
Corresponding author. Present address: Department of Psychol-
ogy, Lehman College, City University of New York, 250 Bedford Park
Blvd. West, Bronx, NY 10468, USA..
E-mail address: khappane@lehman.cuny.edu (K. Happaney).
hind.
This special issue of Brain and Cognition is intended to
redress this imbalance by bringing together researchers
who are studying the nature and development of or-
bitofrontal function from a wide variety of perspectives.
Our hope is that the contributions to this issue will help
us to gain a better understanding of the important role
that emotion plays in the typical and atypical develop-
ment of motivated, goal-directed behavior.
2. Orbitofrontal cortex: Structure and function
Prefrontal cortex (PFC) is the part of cerebral cortex
anterior to premotor cortex and the supplementary
motor area (and, according to some authors, excluding
the anterior cingulate gyrus; Fuster, 1989; Stuss &
Benson, 1986). In human beings, it comprises between a
quarter and a third of the entire cortex (Fuster, 1989),
and it is functionally and anatomically heterogeneous.
However, for many purposes, it may suffice to subdivide
it into three broad regions: (1) DL-PFC, (2) medial PFC

0278-2626/$ - see front matter Ó 2004 Elsevier Inc. All rights reserved.
doi:10.1016/j.bandc.2004.01.001
2 K. Happaney et al. / Brain and Cognition 55 (2004) 1–10
(perhaps including the functionally related anterior
cingulate), and (3) OFC, corresponding to the most in-
ferior and ventral parts of PFC (behind the eyes, or
orbits). Both medial PFC and OFC are part of a
frontostriatal circuit that has strong connections to the
amygdala and other parts of the limbic system. Conse-
quently, these regions are anatomically well suited for
the integration of affective and non-affective informa-
tion, and for the regulation of appetitive/motivated re-
sponses. Functionally, these regions are often
considered together, as when researchers focus on effects
of damage to ventromedial PFC (VM-PFC).
Although research on orbitofrontal function in
human patients has only recently begun in earnest,
work with non-human animals has a long history.
Much of this work has relied on two conceptually
related measures, object reversal learning and extinc-
tion. In object reversal learning, animals learn a sim-
ple discrimination between two objects (choice of one
object is rewarded but choice of the other is not).
After this discrimination is learned to a criterion, the
discrimination is reversed (the previously unrewarded
object is rewarded and vice versa). On these tasks,
OFC-lesioned animals fail to switch their responses,
and instead perseverate on the initial discrimination
(e.g., Butter, 1969; Dias, Robbins, & Roberts, 1996;
Iversen & Mishkin, 1970; Jones & Mishkin, 1972).
More recent research has demonstrated that human
patients with acquired OFC damage also reveal defi-
cits in reversal learning, including perseverative re-
sponding to the previously rewarded stimulus (Rolls,
Hornak, Wade, & McGrath, 1994).
Response extinction tasks are similar to reversal
learning tasks in that they also involve a change in the
reinforcement contingencies after a response has been
learned to criterion. In this case, a response is rein-
forced, and then reinforcement is withheld. In such sit-
uations, OFC-ablated non-human primates (e.g., Butter,
Mishkin, & Rosvold, 1963) as well as human patients
with OFC damage (Rolls et al., 1994) display resistance
to extinction, continuing to respond to the non-rein-
forced stimulus. Findings of this sort have led to sug-
gestions that OFC is primarily involved in the
reappraisal of the affective or motivational significance
of stimuli (e.g., Rolls, 1999, 2004). On this view, while
the amygdala is primarily involved in the initial learning
of response-reward contingencies (e.g., Killcross, Rob-
bins, & Everitt, 1997; LeDoux, 1998), reappraisal of
these relations is the province of OFC. Importantly,
deficits in reversal learning and extinction have been
shown in OFC patients who also revealed deficits in
social functioning (Rolls et al., 1994). This finding un-
derscores the fact that reappraisal is likely to play an
important role in social contexts, where one is required
to adapt to rapidly changing contexts (signalled, e.g., by
facial expressions).
3. Affective decision-making and OFC
Interest in OFC and its functions has been boosted
substantially by observations of affective (i.e., moti-
vated) decision-making deficits in patients with damage
to OFC and related (or overlapping) areas, such as VM-
PFC. Affective decision-making deficits found in VM-
PFC patients bear considerable similarity to those noted
in the classic 19th-century case of Phineas Gage (Har-
low, 1848; see Damasio, Grabowski, Frank, Galaburda,
& Damasio, 1994). As is well-known, Gage was a re-
sponsible and affable fellow before an iron tamping rod
was blown through the ventromedial regions of his PFC
in a work-related accident. After the accident, however,
he became irresponsible and impulsive, despite pre-
served general cognitive and motor skills. Contempo-
rary VM-PFC patients, such as EVR, resemble Gage in
manifesting a behavioral profile that has been referred
to as ‘‘acquired sociopathy’’ (e.g., Saver & Damasio,
1991). Although not necessarily violent, these patients
are often grossly insensitive to the consequences of their
behavior—both for themselves and for others. For ex-
ample, they may make disastrous financial decisions,
and have difficulty maintaining personal relationships
(e.g., Dimitrov, Phipps, Zahn, & Grafman, 1999; Rolls
et al., 1994).
In an attempt to create a laboratory analogue of the
‘‘real-life’’ decision-making deficits of VM-PFC pa-
tients, Bechara, Damasio, Damasio, and Anderson
(1994) designed the Iowa Gambling Task. In this task,
participants are presented with four decks of cards that,
when turned, reveal a combination of rewards and losses
(measured in play money). Two decks are advantageous
in the long run: they consistently provide low rewards
on every trial but ultimately have an even lower average
cost. The other two decks are disadvantageous: the re-
wards on every trial are higher than in the advantageous
decks, but the average costs are higher than the average
rewards (so that there is a net cost over trials). Partici-
pants are given an initial sum and told to make as much
money as possible. Bechara et al. (1994) found that both
patients and healthy controls typically preferred the
disadvantageous cards at the outset. Over trials, how-
ever, control subjects were increasingly likely to select
from the advantageous decks, whereas patients were
not. In addition Bechara, Tranel, Damasio, and
Damasio (1996), Bechara, Damasio, Tranel, and
Damasio (1997) found that, over trials, controls but not
VM-PFC patients began to display anticipatory skin
conductance responses (SCRs) that were higher prior to
disadvantageous choices than prior to advantageous
choices.
Damasio (1994) suggested that these anticipatory
SCRs normally serve as ‘‘somatic markers’’ that predict
rewards and punishments and in so doing bias decision-
making away from disadvantageous choices toward
 K. Happaney et al. / Brain and Cognition 55 (2004) 1–10
more advantageous ones. He also suggested that VM-
PFC is essential for the creation and use of these
markers. One might also note, however, that, as with
reversal learning and extinction, success on the Iowa
Gambling Task requires reappraisal of the motivational
significance of stimuli. Although the disadvantageous
choices initially appear profitable, participants must
eventually learn to avoid them. In this regard, the task is
similar to the delayed reward (or delay of gratification)
paradigm, in which animals must choose between a
small immediate reward and a larger delayed reward.
Recent research by Mobini, Chiang, Ho, Bradshaw, and
Szabadi (2002) has shown that OFC-lesioned rats are
more likely than controls to opt for the small immediate
reward.
While the Iowa Gambling Task is perhaps the most
widely used measure of affective decision-making, a
number of other measures have recently been designed,
including measures of guessing with feedback (e.g., El-
liott, Frith, & Dolan, 1997; Rogers et al., 1999a) and
risky decision-making (Rogers et al., 1999b). As sug-
gested by Clark, Cools, and Robbins (2004), RogersÕ
decision-making tasks (also called the Cambridge
Gamble Task; see Clark et al., 2004) may provide a
more selective index of orbitofrontal function than the
Iowa Gambling Task (for comparisons among mea-
sures, see Manes et al., 2002; Monterosso, Ehrman,
Napier, OÕBrien, & Childress, 2001).
4. Development of OFC: Structure and function
Prefrontal cortex exhibits considerable growth during
childhood and into adolescence, as indicated by age-
related changes in myelination (Pfefferbaum et al., 1994;
Yakovlev & Lecours, 1967), interhemispheric connec-
tivity (Thompson et al., 2000), synaptic density (Hut-
tenlocher, 1979, 1990), and metabolic and electrical
activity (e.g., Luciana & Nelson, 1998; Rubia et al.,
2000; Travis, 1998), among other measures. Within
prefrontal cortex, there is some suggestion that OFC
develops earlier than DL-PFC (e.g., Orzhekhovskaya,
1981). The functional implications of these structural
changes are only starting to be understood.
Although historically it was believed that prefrontal
cortex was not functional at all until about 4–7 years of
age (Luria, 1973, p. 87), or even until 12–15 years of age
(Golden, 1981, p. 292), it is now clear that this is in-
correct. Indeed, current evidence indicates that although
important changes continue to occur into adulthood,
prefrontal function first emerges early in development,
probably around the end of the first year of life. For
instance, during the second half of their first year, in-
fants become better able to inhibit responding to an
initial location on the Piagetian A-not-B search task
(Piaget, 1954). Correlations have been found between
3
frontal EEG power (and frontal/parietal EEG coher-
ence) and infantsÕ ability to tolerate longer delays on this
task (Bell & Fox, 1992). Moreover, prefrontal lesions in
monkeys have been shown to impair performance on a
similar task (Diamond & Goldman-Rakic, 1989).
Much of the recent research relevant to understand-
ing the development of prefrontal cortical function has
been conducted under the rubric of EF. It should be
noted that although EF is clearly linked to prefrontal
cortex, and the terms are often used interchangeably, EF
is not synonymous with prefrontal cortical function:
whereas some patients with prefrontal damage do not
show impairments in EF (e.g., Shallice & Burgess, 1991),
some patients with damage outside of prefrontal cortex
do (e.g., Anderson, Damasio, Jones, & Tranel, 1991). In
any event, however, most research on the development
of EF has focussed on the more cognitive aspects of EF;
relatively little is known about the development of af-
fective, motivational contributions to EF. Nonetheless,
some studies have assessed age-related changes using
tasks associated with OFC damage in adult patients and
non-human animals. For example, some studies have
demonstrated age-related decreases in resistance to ex-
tinction during the preschool years, suggestive of the
development of OFC function (e.g., Gladstone, 1969;
Lester, 1966; but see Happaney & Zelazo, 2004, for an
alternative interpretation and findings). Moreover,
Overman, Bachevalier, Schuhmann, and Ryan (1996)
demonstrated age-related improvements in performance
on object reversal in infants and young children. In
addition, these authors found that prior to 30 months of
age, boys performed better than girls (see also Overman,
2004), although a ceiling effect may have obscured sex
differences beyond 30 months. The presence of a sex
difference in object reversal is particularly intriguing in
light of earlier work using this task that suggests both
that object reversal learning develops more slowly in
female monkeys than in male monkeys, and also that
this sex difference is under the control of gonadal hor-
mones (Clark & Goldman-Rakic, 1989; Goldman,
Crawford, Stokes, Galkin, & Rosvold, 1974).
Researchers have also used delayed reward para-
digms in studies with young children. Early studies on
delay of gratification generally failed to find age differ-
ences within the preschool range (e.g., Schwarz, Schr-
ager, & Lyons, 1983), but did find differences within the
school age years (see Mischel, Shoda, & Rodriguez,
1989, for review). However, more recent research has
examined delay of gratification in younger children us-
ing a modified choice task (Thompson, Barresi, &
Moore, 1997). Thompson et al. (1997) found that when
given the option to choose between a small reward now
or a larger reward later (prudence) and between a re-
ward for self now or a reward for self and other later
(altruism), 4- to 5-year-old children demonstrated sig-
nificantly more prudence and altruism than 3-year-old
4 K. Happaney et al. / Brain and Cognition 55 (2004) 1–10
children, and for the 3-year-olds, there was a correlation
between prudence and altruism when age was partialled
out. In addition to replicating these results, Moore,
Barresi, and Thompson (1998) found that prudence and
altruism were related to performance on measures of
theory of mind.
Theory of mind (i.e., making inferences about mental
states when explaining and predicting behavior; As-
tington, 1993; Premack & Woodruff, 1978) has also been
linked to OFC in healthy adults and patients (e.g.,
Stone, Baron-Cohen, & Knight, 1998; Stuss, Gallup, &
Alexander, 2001), and it can be understood as a kind of
affective decision-making. Defined broadly, theory of
mind appears to emerge in infancy—possibly towards
the end of the first year of life, and develops during
childhood, with important change occurring between
about 3 and 5 years. Episodic memory, which has also
been tied to PFC (see Levine, 2004), develops along a
similar trajectory. Borrowing from TulvingÕs (1985)
suggestion that autonoetic (self-knowing) consciousness
is a defining aspect of episodic memory, Perner and
Ruffman (1995) argued that amnesia for early life events
(infantile amnesia) derives from young childrenÕs lack of
self-awareness (see Stuss & Anderson, 2004, for a dis-
cussion of the role of self-awareness in EF). That is,
children below about 4 years of age cannot consciously
re-experience events because they do not yet have the
ability to reflect on their own mental states as such. In
this way, self-awareness, theory of mind, and episodic
memory are intimately connected; they comprise a
constellation of functions for which OFC is likely to be
important.
Finally, changes in affective decision-making have
also been found in preschool-age children using ana-
logues of the Iowa Gambling Task. Kerr and Zelazo
(2004) found that 3-year-olds resembled OFC patients
insofar as they tended to make disadvantageous choices
on a simplified version of the task. In contrast, 4-year-
olds tended to make advantageous choices. Further,
consistent with other research revealing sex differences
on tasks related to OFC (see Overman, 2004), Kerr and
Zelazo demonstrated near significant superior perfor-
mance in 3-year-old boys relative to 3-year-old girls.
Using a more complex version of the task, Garon and
Moore (2004) found that 6-year-oldsÕ chose more ad-
vantageously than younger children (3- and 4-year-
olds). Finally, using the standard Iowa Gambling Task,
Blair, Colledge, and Mitchell (2001) found that age was
inversely related to disadvantageous decisions in 9- to
17-year-olds, suggesting that performance on this task
develops across a wide age range.
4.1. Summary
DL-PFC and OFC are parts of an interactive func-
tional system, and they typically work together—even in
a single situation. Although the literature on adult
neuropsychology has seen an emerging and growing
interest in the study of OFC, research on the develop-
ment of OFC related EF has lagged behind. Most re-
search on EF in children has focused on the relatively
cognitive aspects of EF associated more with DL-PFC
than with OFC. Nonetheless, a number of researchers
have investigated age-related changes using tasks asso-
ciated with OFC damage in adult patients and non-
human animals. For example, researchers have studied
the development of performance on object reversal, ex-
tinction, delay of gratification, theory of mind, and
variants of the Iowa Gambling Task. Results of this
research suggest that like EF in general, the develop-
ment of emotional processes associated with OFC de-
velops across a wide range of ages. This research is
beginning to fill the gap in our understanding of how
orbitofrontal function interacts with other functions of
prefrontal cortex to contribute to the development of
conscious, self-regulated, goal-directed activity.
5. About this issue
This issue is comprised of articles by researchers in
the areas of behavioral neuroscience, neurodevelop-
ment, cognitive and emotional development, neuropsy-
chology, and cognitive neuro-rehabilitation. These
articles are considered together in an effort to stimulate
interdisciplinary dialogue on the important but still
understudied topic of the development of OFC function.
Some of the questions addressed in this issue are: (1)
What does OFC do? (2) How can we best assess or-
bitofrontal function? (3) How does OFC interact with
other brain regions to contribute to EF? (4) How do
OFC and its related functions develop under normal
circumstances? and (5) Can developmental disabilities
be understood in terms of atypical OFC development?
Arguably, these questions are best considered together.
For example, in order to understand the role of atypical
OFC development in various pathologies, it is important
to establish the nature of typical development of this
region at various levels of analysis: neuroanatomical,
neurophysiological, cognitive, and behavioral. At the
same time, the study of atypical development and the
consequences of brain damage provide important in-
formation about typical development and healthy
function. Ultimately, answers to these questions will
shed light on the development of orbitofrontal function,
which clearly lies at the interface between cognition and
emotion.
This issue is divided into three sections. The articles in
Section 1 concern the basic functions of OFC, although
implications for development are discussed. The articles
in Section 2 take a decidedly developmental approach
and examine the development of OFC as indicated by
 K. Happaney et al. / Brain and Cognition 55 (2004) 1–10
lesion studies and neuroimaging. The articles in Section
3 use behavioral indices of affective aspects of EF to
assess age-related changes in OFC function. Whereas
some of the articles in this section focus on typical de-
velopment, others discuss the extent to which various
psychiatric disorders with childhood onset (e.g., obses-
sive compulsive disorder, conduct disorder, and autism)
can be understood in terms of atypical development of
OFC.
5.1. Section 1: OFC function
In the first article, Rolls (2004) reviews research on
OFC functions in human and non-human animals, and
suggests that a critical function of OFC is the reap-
praisal of stimulus–reward relations. In the following
article, Bechara reviews findings from studies using the
Iowa Gambling Task, as well as other decision-making
tasks, and makes a case for the utility of the somatic
marker theory for understanding the role of OFC (and
VM-PFC more generally) in ‘‘real-life’’ decision-mak-
ing. Next, Clark et al. (2004) discuss the conceptual,
anatomical, and neurophysiological (neurotransmitter)
commonalities between decision-making and reversal
learning (two of the most widely studied indices of OFC
function). LevineÕs (2004) article then goes on to discuss
the role of OFC and other frontal regions in episodic
autobiographical memories, using evidence from pa-
tients with frontal lobe damage of various etiologies.
Within his discussion of these findings, Levine suggests a
model for the development of episodic memory. As ep-
isodic memory has been linked (both conceptually and
ontogenetically) to self-concept and social cognition,
this discussion is of particular relevance to any discus-
sion of OFC function.
5.2. Section 2: Developmental approaches to OFC
In the first article in this section, Stuss and Anderson
(2004) suggest that early versus later maturation of OFC
and related frontal structures may be understood in
terms of self-awareness. Namely, they suggest that early
frontal lobe damage leads to global deficits whereas later
damage leads to more selective impairments in self-
awareness and theory of mind. Next, Eslinger, Flaherty-
Craig, and Benton (2004) review cases of early PFC
damage, suggestive of the fact that OFC injury leads to
particularly intractable deficits in social and emotional
functioning. Kolb, Pellis, and Robinson (2004) then
discuss findings that early lesions to DL-PFC (medial
PFC in rodents) in rats leads to the greatest cognitive
deficits whereas damage to OFC is not affected by age of
lesioning. These authors find that the general cognitive
effects of OFC lesions in rats are minimal, but that social
behaviors are drastically affected at all ages. Thus, in
keeping with Eslinger et al.Õs suggestions, damage to
5
OFC appears to have particularly non-remediable ef-
fects. Further, Kolb et al. discuss the differential effects
of drugs of abuse on DL-PFC and OFC, respectively,
and their findings are in keeping with similar decision-
making deficits found between OFC patients and long-
term drug abusers (e.g., see Rogers et al., 1999a).
Finally, in contrast to approaches using non-human
animals and human frontal lobe patients, Segalowitz
and Davies (2004) discuss the utility of assessing the
development of differentiable aspects of EF subserved
by OFC, anterior cingulate cortex (ACC), and DL-PFC
by recording event-related potentials in typically devel-
oping children from 7 to 17 years of age.
5.3. Section 3: Typical and atypical development of
affective and motivational contributions to EF
In the first article in this section, Overman (2004)
reviews findings concerning age-related improvements in
reversal learning in infants and toddlers that mirror
developmental changes in non-human primates. Of
specific emphasis are sex effects favoring boys on re-
versal learning, and men on tests of decision-making.
Kerr and Zelazo (2004) follow this discussion in a report
of their findings using a child appropriate version of the
Iowa Gambling Task. Performance on this task shows
improvements between 3 and 4 years of age, with some
suggestion that, as in reversal learning, boys do better.
Garon and Moore (2004) also discuss findings con-
cerning a slightly more complex version of the Iowa
Gambling Task. Garon and Moore also find evidence of
age-related increases in performance on this task; how-
ever these changes appear later than those observed
using Kerr and ZelazoÕs simpler task. Finally, Happaney
and Zelazo (2004) discuss the potential utility of ex-
tinction tasks in assessing the development of affective
and motivational contributions to EF. These authors
found that between 3 and 6 years of age, there is an
increase in responding during the extinction phase,
which could be explained by an alternative conceptual-
ization of what constitutes adaptive responding on this
task. This study emphasizes that there may be important
differences between participant groups in how the ob-
jectives of such simple tasks are conceptualized.
The remaining articles in this section concern the
relation between atypical OFC development and mal-
adaptive child behaviors as well as developmental psy-
chopathology. In the first article, Seguin (2004) discusses
the utility of considering OFC in an effort to understand
different forms of antisocial behavior in both children
and adults. Next, Blair (2004) suggests that the role of
OFC in antisocial behavior (particularly aggression)
may be limited to reactive aggression (i.e., aggression
deriving from frustration or fear), but argues that in-
strumental aggression (as may be associated with psy-
chopathy) is likely to derive from atypicality in
6 K. Happaney et al. / Brain and Cognition 55 (2004) 1–10
amygdala function. Blair also proposes that socializa-
tion moderates the effect of atypical amygdala function
on the expressions of instrumentally aggressive behav-
ior.
The next two articles have particular implications for
autism and obsessive compulsive disorder (OCD), re-
spectively. Sabbagh (2004) addresses OFC and autism,
both of which have been linked to theory of mind. In his
article, Sabbagh reviews research using ERP, and then
argues that whereas OFC is particularly important for
deciphering othersÕ mental states (particularly emo-
tions), lateral PFC is important for reasoning about
these states. These findings are in keeping with several
studies showing a relation between more cognitive as-
pects of EF and theory of mind in both typically de-
veloping children (e.g., Frye, Zelazo, & Palfai, 1995) and
individuals with autism (e.g., Ozonoff, Pennington, &
Rogers, 1991).
Finally, Evans, Lewis, and Iobst (2004) discuss the
role of OFC in typical age-related changes in repetitive
and ritualistic behavior. These behaviors are character-
istic of OCD, and Evans et al. discuss ways in which
understanding the involvement of OFC in OCD may
help us to understand typical developmental changes in
such behaviors, and vice versa.
6. Common themes
Throughout this issue, a number of themes recur.
Among these are sex differences in OFC growth and
function, and functional specialization within OFC, in-
cluding the lateralization of OFC functions.
6.1. Sexual dimorphism in orbitofrontal development and
function
As reviewed extensively by Overman (2004), evidence
is accumulating that OFC matures earlier in males than
females, potentially as a result of pre-, peri-, and post-
natal testosterone levels. OFC volume in humans is
greater in males than females (Goldstein et al., 2001),
and there is greater dendritic arborization in OFC in
female rats (Kolb et al., 2004). In humans, this sex dif-
ference is evidenced by the superior performance of in-
fants and toddlers on tests of object reversal (Overman
et al., 1996). No sex differences in object reversal have
been found in children beyond 3 years of age, likely due
to ceiling effects by that age (see Overman, 2004).
However, employing a child appropriate version of the
Iowa Gambling Task, Kerr and Zelazo (2004) found
near significant male superiority among 3-year-olds.
While Garon and Moore (2004) failed to find sex dif-
ferences in an alternative childrenÕs version of the Iowa
Gambling Task, their task was more complex than the
one used by Kerr and Zelazo (2004).
Evidence of sex differences favoring males also ex-
tends into adulthood, with men outperforming women
on the Iowa Gambling Task (Reavis & Overman, 2001;
but see Overman, 2004, for qualifications) and pre-
liminary evidence that adolescent boys outperform ad-
olescent girls (Overman, 2004). Further, Overman
(2004) has found that males and females may perform
the Iowa Gambling Task in a different manner, with
females avoiding decks with losses regardless of net
value, and with males more effectively considering the
long-term, net outcome of decisions. As will be dis-
cussed in the next section, we suggest that males and
females might take different approaches to the task in
part because females pay greater attention to local detail
whereas males approach the task in a more holistic
fashion.
Although sex differences have been demonstrated in
some studies, many other studies have simply failed to
consider the effect of sex on performance. Indeed, as
Overman (2004) suggests, it may be worthwhile to re-
analyze data from studies that have employed behav-
ioral indices of OFC function.
6.2. Regional specificity and laterality effects
There is growing evidence for anatomical and func-
tional differentiation within OFC. For instance, as dis-
cussed by Rolls (2004), lesions to the inferior convexity
produce deficits on object reversal whereas damage to
caudal OFC produces resistance to extinction (see also
Rosenkilde, 1979). Blair (2004) also differentiates be-
tween systems that mediate sensitivity to violations of
expectation (as in extinction and response reversal tasks)
and those that mediate social response reversal (as in
responding to expectations of social norms).
As discussed in Evans et al. (2004), medial OFC ap-
pears to be involved in the initial evaluation of the af-
fective or motivational significance of stimuli due to its
more direct connections to limbic and related structures.
Lateral OFC, by contrast, is connected to higher cortical
areas, such as DL-PFC, and has been tied to the reap-
praisal of the emotional significance of stimuli, emotion
regulation, and response suppression and selection, as
assessed in Go-NoGo tasks. Evidence is also mounting
that damage to right (or bilateral) OFC has a greater
effect on social conduct, decision-making, emotional
processing, and other purported OFC functions than
does damage to left OFC (e.g., Manes et al., 2002; Rolls
et al., 1994; Stuss, 1991; Stuss & Alexander, 1999; Stuss
et al., 2001; Tranel, Bechara, & Denburg, 2002). As
discussed by Bechara (2004; see also Tranel et al., 2002),
patients with right VM-PFC damage reveal marked
impairments in everyday functioning as well as on the
Iowa Gambling Task, and these effects are similar to
those revealed in bilateral VM-PFC patients. By con-
trast, patients with left VM-PFC damage are relatively
 K. Happaney et al. / Brain and Cognition 55 (2004) 1–10
unimpaired, suggesting that the reliable impairments
demonstrated by bilateral OFC patients may derive
primarily from right VM-PFC.
There are several possible reasons why right OFC is
so important for these functions. Bechara (2004) sug-
gests that right–left hemispheric asymmetries in OFC
(VM-PFC) function may derive from the differential
involvement of the right and left hemispheres in avoid-
ance (negative affect) and approach (positive affect),
respectively (see also Davidson & Irwin, 1999; David-
son, Jackson, & Kalin, 2000). That is, adaptive decision-
making on the Iowa Gambling Task requires avoidance
of seemingly positive responses (a function for which
right VM-PFC may be particularly well suited). The
right hemisphere has also been implicated in the map-
ping of bodily states and the comprehension of somatic
information (Cohen, Rosen, & Goldstein, 1976; Da-
vidson & Schwartz, 1976), and this too may help to
explain the relative importance of right OFC to every-
day decision-making (Bechara, 2004; Damasio, 1994).
Another possibility is that whereas the right hemi-
sphere is more involved in processing information
globally, the left hemisphere is more involved in pro-
cessing information analytically (see Tucker, 1981, for a
review). If everyday decision-making (as assessed by the
Iowa Gambling Task) is based on hunches abstracted
from feedback on individual trials, then this may explain
why the right hemisphere is so important for this type of
decision-making. Other functions tied to OFC may also
depend on global processing of information, such as
self-reflective awareness, for instance (see Kihlstrom &
Klein, 1994; Stuss & Anderson, 2004). Relatedly, autism
and related disorders may lend speculative support for
relations between global processing deficits and social
and emotional functioning. The link between the syn-
thetic and global processing style of the right hemisphere
and the type of information processing required for
social cognition deserves particular attention (see Ellis &
Gunter, 1999). Indeed, the ability to synthesize indi-
vidual pieces of information into a more holistic picture
has been theorized to underlie many symptoms of au-
tism and AspergerÕs syndrome (Frith & Happe, 1994).
The right hemisphere has also been linked to implicit
information processing, as opposed to the more explicit
and more conscious processing tied to the left hemi-
sphere (Vakil, Galek, Soroker, Ring, & Gross, 1991;
Vakil, Soroker, & Borenstein, 1998). As suggested by
Damasio (1994), considering all relevant factors in-
volved in many personal and immediately social deci-
sions would be too costly in terms of cognitive resources
(see also Bechara, Damasio, Tranel, & Anderson, 1998).
Thus, again, the right hemisphere may be particularly
adept at the processes involved in everyday decisions.
While recent evidence suggests that the Iowa Gambling
Task may be more resource demanding than once
thought (Hinson, Jameson, & Whitney, 2002), these
7
requirements may still be fewer than would be necessary
for more algorithmically based or rational decision-
making. Importantly, the working memory demands of
the Iowa Gambling Task may underlie the fact that
while it serves as a sensitive index of OFC damage, it
may not be a specific index of such damage (see Clark
et al., 2004).
We suggest an even further reason to expect right
hemispheric dominance in OFC function, which draws
on research and theory suggestive of the right hemi-
sphereÕs role in anomaly detection (Ramachandran &
Blakeslee, 1998; Smith, Tays, Dixon, & Bulman-Flem-
ing, 2002). As described by Smith et al. (2002), anomaly
detection involves analyzing information for features
that are inconsistent with memory for that information.
The OFCÕs role in reinterpreting stimuli in light of
changing affective significance (as in the Iowa Gambling
Task, object reversal tasks, and extinction tasks) may
then be understood in terms of anomaly detection, in
that, in these tasks, one must amend initial stimulus–
reward associations on the basis of exposure to associ-
ations at odds with prior experience (i.e., anomalous
ones). Indeed, skills such as detecting faux pas (i.e.,
statements and conduct falling outside of social expec-
tations) have been linked to OFC (Stone et al., 1998),
lending further credence to the link between OFC
(particularly right OFC) and socially valid indices of
anomaly detection.
As alluded to earlier, superior male performance on
tasks tapping OFC function is also consistent with a
right hemispheric account, in that males also show
greater right hemispheric dominance (Diamond, Dow-
ling, & Johnson, 1981; deLacoste, Horvath, & Wood-
ward, 1991) and perform better on tasks tied to the right
hemisphere (e.g., abstract mathematics and spatial vi-
sualization; see Kimura, 1999). Thus, it is possible that
OFC function may mirror other right hemispheric
functions for which male superiority has been observed.
Taken together, then, right OFC may be in a particu-
larly good position, both structurally and functionally,
to support the functions OFC develops to fulfill.
7. Conclusions
Aspects of executive function, such as affective deci-
sion-making and social understanding, undergo marked
changes during childhood, and it is important to un-
derstand the neural mechanisms underlying these
changes. Recent interest in OFC and its functions has
forged a reconsideration of the relations between affect
and cognition (e.g., Damasio, 1994), and these consid-
erations need to be reflected in our understanding of
cognitive, emotional, and social development. It is
hoped that such considerations will allow a mapping of
advances gained from findings concerning adult cognition
8 K. Happaney et al. / Brain and Cognition 55 (2004) 1–10
and emotional and social functioning to both typical
and atypical development in childhood. More generally,
however, any comprehensive understanding of these
functions in adulthood will require an understanding
of their origins and the way in which they change.
Clearly a developmental perspective is crucial to this
latter objective.
References
Anderson, S. W., Damasio, H., Jones, R. D., & Tranel, D. (1991).
Wisconsin card sorting performance as a measure of frontal lobe
damage. Journal of Clinical and Experimental Neuropsychology, 13,
909–922.
Astington, J. W. (1993). The childÕs discovery of the mind. Cambridge,
MA: Harvard University Press.
Bechara, A. (2004). The role of emotion in decision-making: Evidence
from neurological patients with orbitofrontal damage. Brain and
Cognition (Special Issue: Development of Orbitofrontal Function),
55, 30–40.
Bechara, A., Damasio, A., Damasio, H., & Anderson, S. (1994).
Insensitivity to future consequences following damage to human
prefrontal cortex. Cognition, 50, 7–15.
Bechara, A., Damasio, H., Tranel, D., & Anderson, S. W. (1998).
Dissociation of working memory from decision-making within the
human prefrontal cortex. The Journal of Neurosciene, 18, 428–437.
Bechara, A., Damasio, H., Tranel, D., & Damasio, A. (1997). Deciding
advantageously before knowing the advantageous strategy. Sci-
ence, 275, 1293–1295.
Bechara, A., Tranel, D., Damasio, H., & Damasio, A. R. (1996).
Failure to respond autonomically to anticipated future outcomes
following damage to prefrontal cortex. Cerebral Cortex, 6, 215–
225.
Bell, M. A., & Fox, N. A. (1992). The relations between frontal brain
electrical activity and cognitive development during infancy. Child
Development, 63, 1142–1163.
Blair, R. J. R. (2004). The roles of orbital frontal cortex in the
modulation of antisocial behaviour. Brain and Cognition (Special
Issue: Development of Orbitofrontal Function), 55, 198–208.
Blair, R. J. R., Colledge, E., & Mitchell, D. G. V. (2001). Somatic
markers and response reversal: Is there orbitofrontal cortex
dysfunction in boys with psychopathic tendencies. Journal of
Abnormal Child Psychology, 29, 499–511.
Butter, C. (1969). Perseveration in extinction and in discrimination
reversal tasks following selective frontal ablations in macaca
mulatta. Physiology and Behavior, 4, 163–171.
Butter, C. M., Mishkin, M., & Rosvold, H. E. (1963). Conditioning
and extinction of a food-rewarded response after selective abala-
tions of frontal cortex in rhesus monkeys. Experimental Neurology,
7, 65–75.
Clark, L., Cools, R., & Robbins, T. W. (2004). The neuropsychology
of ventral prefrontal cortex: Decision-making and reversal learn-
ing. Brain and Cognition (Special Issue: Development of Orbito-
frontal Function), 55, 41–53.
Clark, A. S., & Goldman-Rakic, P. S. (1989). Gonadal hormones
influence the emergence of cortical function in nonhuman primates.
Behavioral Neuroscience, 103, 1287–1295.
Cohen, H. D., Rosen, R. C., & Goldstein, L. (1976). Electroenceph-
alographic laterality changes during human sexual orgasm. Ar-
chives of Sexual Behavior, 5, 189–199.
Damasio, A. (1994). DescartesÕ error: Emotion, reason, and the human
brain. New York: Grosset/Putnam.
Damasio, H., Grabowski, T., Frank, R., Galaburda, A. M., &
Damasio, A. R. (1994). The return of Phineas Gage: Clues about
the brain from the skull of a famous patient. Science, 264, 1102–
1105.
Davidson, R. J., & Irwin, W. (1999). The functional neuroanatomy of
emotion and affective style. Trends in Cognitive Sciences, 3, 11–21.
Davidson, R. J., Jackson, D. C., & Kalin, N. H. (2000). Emotion,
plasticity, context, and regulation: Perspectives form affective
neuroscience. Psychological Bulletin, 126, 890–909.
Davidson, R. J., & Schwartz, G. E. (1976). The psychobiology of
relaxation and related states: A multi-process theory. In D. I.
Motosky (Ed.), Behavior control and modification of physiological
activity (pp. 342–399). Englewood Cliffs, NJ: Prentice-Hall.
Diamond, M. C., Dowling, G. A., & Johnson, R. E. (1981).
Morphological cerebral cortical asymmetry in male and female
rats. Experimental Neurology, 71, 261–268.
Diamond, A., & Goldman-Rakic, P. S. (1989). Comparison of human
infants and rhesus monkeys on PiagetÕs A-not-B task: Evidence for
dependence on dorsolateral prefrontal cortex. Experimental Brain
Research, 74, 24–40.
Dias, R., Robbins, T. W., & Roberts, A. C. (1996). Dissociation in
prefrontal cortex of affective and attentional shifts. Nature, 380,
69–72.
Dimitrov, M., Phipps, M., Zahn, T. P., & Grafman, J. (1999). A
thoroughly modern Gage. Neurocase, 5, 345–354.
Elliott, R., Frith, C. D., & Dolan, R. J. (1997). Differential neural
response to positive and negative feedback in planning and
guessing tasks. Neuropsychologia, 35, 1395–1404.
Ellis, H. D., & Gunter, H. L. (1999). Asperger syndrome: A simple
matter of white matter? Trends in Cognitive Sciences, 3, 192–200.
Eslinger, P. J., Flaherty-Craig, C. V., & Benton, A. L. (2004).
Cognitive, social and moral development after early prefrontal
cortex damage: Review of available clinical cases. Brain and
Cognition (Special Issue: Development of Orbitofrontal Function),
55, 84–103.
Evans, D. W., Lewis, M. D., & Iobst, E. (2004). The role of the
orbitofrontal cortex in normally developing compulsive-like be-
haviors and obsessive-compulsive disorder. Brain and Cognition
(Special Issue: Development of Orbitofrontal Function), 55, 220–
234.
Frith, U., & Happe, F. (1994). Autism: Beyond theory of mind.
Cognition, 50, 115–132.
Frye, D., Zelazo, P. D., & Palfai, T. (1995). Theory of mind and rule-
based reasoning. Cognitive Development, 10, 483–527.
Fuster, J. (1989). The prefrontal cortex: Anatomy, physiology and
neuropsychology of the frontal lobe (2nd ed.). New York: Raven.
Garon, N., & Moore, C. (2004). Complex decision-making in early
childhood. Brain and Cognition (Special Issue on Development of
Orbitofrontal Function), 55, 158–170.
Gladstone, R. (1969). Age, cognitive control and extinction. Journal of
Experimental Child Psychology, 7, 31–35.
Golden, C. J. (1981). The Luria-Nebraska childrenÕs battery: Theory
and formulation. In G. W. Hynd & J. E. Obrzut (Eds.),
Neuropsychological assessment and the school-aged child (pp. 277–
302). New York: Grune & Stratton.
Goldman, P. S., Crawford, H. T., Stokes, L. P., Galkin, T. W., &
Rosvold, H. E. (1974). Sex-dependent behavioral effects of cerebral
cortical lesions in the developing rhesus monkey. Science, 186, 540–
542.
Goldstein, J. M., Seidman, L. J., Horton, N. J., Makris, N., Kennedy,
D. N., Caviness, V. S., Jr., Faraone, S. V., & Tsuang, M. T. (2001).
Normal sexual dimorphism of the adult human brain assessed by in
vivo magnetic resonance imaging. Cerebral Cortex, 11, 490–497.
Happaney, K., & Zelazo, P. D. (2004). Resistance to extinction as a
measure of orbitofrontal function suitable for children? Brain and
Cognition (Special Issue: Development of Orbitofrontal Function),
55, 171–184.
Harlow, J. M. (1848). Passage of an iron rod through the head. Boston
Medical Surgery Journal, 39, 389–393.
 K. Happaney et al. / Brain and Cognition 55 (2004) 1–10
Hinson, J. M., Jameson, T. L., & Whitney, P. (2002). Somatic markers,
working memory, and decision making. Cognitive, Affective &
Behavioral Neuroscience, 2, 341–353.
Huttenlocher, P. R. (1979). Synaptic density in human frontal cortex:
Developmental changes and effects of aging. Brain Research, 163,
195–205.
Huttenlocher, P. R. (1990). Morphometric study of human cerebral
cortex development. Neuropsychologia, 28, 517–527.
Iversen, S. D., & Mishkin, M. (1970). Perseverative interference in
monkeys following selective lesions of the inferior prefrontal
convexity. Experimental Brain Research, 11, 376–386.
Jones, B., & Mishkin, M. (1972). Limbic lesions and the problem of
stimulus–reinforcement associations. Experimental Neurology, 36,
362–377.
Kerr, A., & Zelazo, P. D. (2004). Development of ‘‘hot’’ executive
function: The childrenÕs gambling task. Brain and Cognition
(Special Issue: Development of Orbitofrontal Function), 55, 148–
157.
Kihlstrom, J. F., & Klein, S. B. (1994). The self as a knowledge
structure. In R. S. Wyer & T. K. Srull (Eds.), Handbook of social
cognition (Vol. 1, pp. 153–208). Hillsdale, NJ: Erlbaum.
Killcross, S., Robbins, T. W., & Everitt, B. J. (1997). Different types of
fear-conditioned behaviour mediated by separate nuclei within
amygdala. Nature, 388, 377–380.
Kimura, D. (1999). Sex and cognition. Cambridge, MA: MIT Press.
Kolb, B., Pellis, S., & Robinson, T. E. (2004). Plasticity and functions
of the orbital frontal cortex. Brain and Cognition (Special Issue:
Development of Orbitofrontal Function), 55, 104–115.
deLacoste, M.-C., Horvath, D. S., & Woodward, D. J. (1991). Possible
sex differences in the developing human fetal brain. Journal of
Clinical & Experimental Neuropsychology, 13, 831–846.
LeDoux, J. (1998). The emotional brain. New York: Weidenfeld &
Nicolson.
Lester, D. (1966). Partial reinforcement effect after a small number of
training trials. Psychological Reports, 19, 970.
Levine, B. (2004). Autobiographical memory and the self in time:
Brain lesion effects, functional neuroanatomy, and lifespan devel-
opment. Brain and Cognition (Special Issue: Development of
Orbitofrontal Function), 55, 54–68.
Luciana, M., & Nelson, C. A. (1998). The functional emergence of
prefrontally-guided working memory systems in four-to-eight-year-
old children. Neuropsychologia, 36, 273–293.
Luria, A. R. (1973). The working brain: An introduction to neuropsy-
chology (B. Haigh, Trans.). New York: Basic Books.
Manes, F., Sahakian, B., Clark, L., Rogers, R., Antoun, N., Aitken,
M., & Robbins, T. (2002). Decision-making processes following
damage to prefrontal cortex. Brain, 125, 624–639.
Mischel, W., Shoda, Y., & Rodriguez, M. L. (1989). Delay of
gratification in children. Science, 244, 933–938.
Mobini, S., Chiang, T. J., Ho, M. Y., Bradshaw, C. M., & Szabadi, E.
(2002). Effects of central 5-hydroxytryptamine depletion on sensi-
tivity to delayed and probabilistic reinforcement. Psychopharma-
cology, 152, 390–397.
Monterosso, J., Ehrman, R., Napier, K. L., OÕBrien, C. P., &
Childress, A. R. (2001). Three decision-making tasks in cocaine-
dependent patients: Do they measure the same construct? Addic-
tion, 96, 1825–1837.
Moore, C., Barresi, J., & Thompson, C. (1998). The cognitive basis of
future oriented prosocial behavior. Social Development, 7, 198–218.
Orzhekhovskaya, N. S. (1981). Fronto-striatal relationships in primate
ontogeny. Neuroscience & Behavioral Physiology, 11, 379–385.
Overman, W. H. (2004). Sex differences in early childhood, adolescence
and adulthood on cognitive tasks that rely on orbital prefrontal
cortex. Brain and Cognition. (Special Issue: Development of
Orbitofrontal Function), 55, 134–147.
Overman, W. H., Bachevalier, J., Schuhmann, E., & Ryan, P. (1996).
Cognitive gender differences in very young children parallel
9
biologically based cognitive gender differences in monkeys. Behav-
ioral Neuroscience, 110, 673–684.
Ozonoff, S., Pennington, B. F., & Rogers, S. J. (1991). Executive
function deficits in high-functioning autistic individuals: Relation-
ship to theory of mind. Journal of Child Psychology and Psychiatry,
32, 1081–1105.
Perner, J., & Ruffman, T. (1995). Episodic memory and autonoetic
consciousness: Developmental evidence and a theory of childhood
amnesia. Journal of Experimental Child Psychology, 59, 516–548.
Pfefferbaum, A., Mathalon, D. H., Sullivan, E. V., Rawles, J. M.,
Zipursky, R. B., & Lim, K. O. (1994). A quantitative magnetic
resonance imaging study of changes in brain morphology from
infancy to late adulthood. Archives of Neurology, 51, 874–887.
Piaget, J. (1954). The construction of reality in the child. New York:
Basic Books.
Premack, D., & Woodruff, G. (1978). Does the chimpanzee have a
theory of mind? Behavioral and Brain Sciences, 1, 515–526.
Ramachandran, V. S., & Blakeslee, S. (1998). Phantoms in the brain:
Probing the mysteries of the human mind. New York: Morrow.
Reavis, R., & Overman, W. H. (2001). Adult sex differences on a
decision-making task previously shown to depend on the orbital
prefrontal cortex. Behavioral Neuroscience, 115, 196–206.
Rogers, R. D., Everitt, B. J., Baldacchino, A., Blackshaw, A. J.,
Swainson, R., Wynne, K., Baker, N. B., Hunter, J., Carthy, T.,
Booker, E., London, M., Deakin, J. F. W., Sahakian, B. J., &
Robbins, T. W. (1999a). Dissociable deficits in the decision-making
cognition of chronic amphetamine abusers, opiate abusers, patients
with focal damage to prefrontal cortex, and tryptophan-depleted
normal volunteers: Evidence for monoaminergic mechanisms.
Neuropsychopharmacology, 20, 322–339.
Rogers, R. D., Owen, A. M., Middleton, H. C., Williams, E. J.,
Pickard, J. D., Sahakian, B. J., & Robbins, T. W. (1999b).
Choosing between small, likely rewards and large, unlikely rewards
activates inferior and orbital prefrontal cortex. Journal of Neuro-
science, 20, 9029–9038.
Rolls, E. T. (1999). The brain and emotion. Oxford: Oxford University
Press.
Rolls, E. T. (2004). The functions of the orbitofrontal cortex. Brain and
Cognition (Special Issue: Development of Orbitofrontal Function),
55, 11–29.
Rolls, E. T., Hornak, J., Wade, D., & McGrath, J. (1994). Emotion-
related learning in patients with social and emotional changes
associated with frontal lobe damage. Journal of Neurology,
Neurosurgery & Psychiatry, 57, 1518–1524.
Rosenkilde, C. E. (1979). Functional heterogeneity of the prefrontal
cortex in the monkey: A review. Behavioural and Neural Biology,
25, 301–345.
Rubia, K., Overmeyer, S., Taylor, E., Brammer, M., Williams, S. C.,
Simmons, A., Andrew, C., & Bullmore, E. T. (2000).
Functional frontalisation with age: Mapping neurodevelopmental
trajectories with fMRI. Neuroscience & Biobehavioral Reviews, 24,
13–19.
Sabbagh, M. (2004). Frontal contributions to recognizing and
reasoning about mental states: Implications for autism. Brain and
Cognition (Special Issue: Development of Orbitofrontal Function),
55, 209–219.
Saver, J. L., & Damasio, A. R. (1991). Preserved access and processing
of social knowledge in a patient with acquired sociopathy
due to ventromedial frontal damage. Neuropsychologia, 29, 1241–
1249.
Schwarz, J. C., Schrager, J. B., & Lyons, A. E. (1983). Delay of
gratification by preschoolers: Evidence for the validity of the choice
paradigm. Child Development, 54, 620–625.
Segalowitz, S. J., & Davies, P. L. (2004). Charting the maturation of
the frontal lobe: An electrophysiological strategy. Brain and
Cognition (Special Issue: Development of Orbitofrontal Function),
55, 116–133.
10 K. Happaney et al. / Brain and Cognition 55 (2004) 1–10
Seguin, J. R. (2004). Neurocognitive elements of antisocial behaviour:
Relevance of an orbitofrontal cortex account. Brain and Cognition
(Special Issue: Development of Orbitofrontal Function), 55,
185–197.
Shallice, T., & Burgess, P. W. (1991). Deficits in strategy
application following frontal lobe damage in man. Brain, 111,
727–741.
Smith, S. D., Tays, W. J., Dixon, M. J., & Bulman-Fleming, M. B.
(2002). The right hemisphere as an anomaly detector: Evidence
from visual perception. Brain and Cognition, 48, 574–579.
Stone, V. E., Baron-Cohen, S., & Knight, R. T. (1998). Frontal lobe
contributions to theory of mind. Journal of Cognitive Neuroscience,
10, 640–656.
Stuss, D. T. (1991). Self, awareness, and the frontal lobes: A
neuropsychological perspective. In J. Strauss & G. R. Goethals
(Eds.), The self: Interdisciplinary approaches (pp. 255–278). New
York: Springer.
Stuss, D. T., & Alexander, M. P. (1999). Affectively burnt in: A
propose drole of the right frontal lobe. In E. Tulving (Ed.),
Memory, consciousness and the brain: The Tallinn conference (pp.
215–227). Philadelphia: Psychology Press.
Stuss, D. T., & Anderson, V. (2004). The frontal lobes and theory of
mind: Developmental concepts from adult focal lesion research.
Brain and Cognition (Special Issue: Development of Orbitofrontal
Function), 55, 69–83.
Stuss, D. T., & Benson, D. F. (1986). The frontal lobes. New York:
Raven Press.
Stuss, D. T., Gallup, G. G., & Alexander, M. P. (2001). The
frontal lobes are necessary for ‘‘theory of mind’’. Brain, 124,
279–286.
Thompson, C., Barresi, J., & Moore, C. (1997). The development of
future-oriented prudence and altruism in preschoolers. Cognitive
Development, 12, 199–212.
Thompson, P. M, Giedd, J. N., Woods, R. P., MacDonald, D., Evans,
A. C., & Toga, A. W. (2000). Growth patterns in the developing
brain detected by using continuum mechanical tensor maps.
Nature, 404, 190–193.
Tranel, D., Bechara, A., & Denburg, N. L. (2002). Asymmetric
functional roles of right and left ventromedial prefrontal cortices in
social conduct, decision-making, and emotional processing. Cor-
tex, 38, 589–612.
Travis, F. (1998). Cortical and cognitive development in 4th, 8th and
12th grade students: The contribution of speed of processing and
executive functioning to cognitive development. Biological Psy-
chology, 48, 37–56.
Tucker, D. M. (1981). Lateral brain function, emotion, and concep-
tualization. Psychological Bulletin, 89, 19–46.
Tulving, E. (1985). Memory and consciousness. Canadian Psychology,
26, 1–12.
Vakil, E., Galek, S., Soroker, N., Ring, H., & Gross, Y. (1991).
Differential effect of right and left hemishpheric lesions on two
memory tasks: Free recall and frequency judgement. Neuropsych-
ologia, 29, 981–992.
Vakil, E., Soroker, N., & Borenstein, E. (1998). The effect of right and
left hemispheric lesions on effortful and automatic memory tasks.
Laterality, 3, 143–159.
Yakovlev, P. I., & Lecours, A. R. (1967). The myelogenetic cycles of
regional maturation of the brain. In A. Minkowski (Ed.), Regional
development of the brain in early life (pp. 3–70). Oxford, England:
Blackwell.