Brain and Cognition 55 (2004) 134–147

www.elsevier.com/locate/b&c

Sex differences in early childhood, adolescence, and adulthood

on cognitive tasks that rely on orbital prefrontal cortexq
William H. Overman
Psychology Department, University of North Carolina at Wilmington, 601 South College Road, Wilmington, NC 20403, USA
Accepted 6 June 2003
Available online 8 February 2004

Abstract

Through the use of several tests of cognition we have documented sex differences in young children, adolescents, and adults on
tasks that rely on the integrity of the orbital prefrontal cortex. In children under three years of age, males performed with signif-
icantly fewer errors than did females on tests of object reversals. No significant sex differences were found in older children, despite
the use of a more challenging object reversal task. Sex differences were also found in adolescents and adults on the Iowa Gambling
Task. On this decision-making task, in contrast to males, females appear to be responding to different elements of the task. Dis-
cussion of the implications for these findings is presented.
Ó 2004 Elsevier Inc. All rights reserved.

1. Introduction unique oddity, landmark discriminations, and object

For a number of years, our laboratory has investigated
sex differences on cognitive tasks that are known to rely
on the integrity of the orbital prefrontal cortex. We have
found evidence for ‘‘orbital’’ cognitive sex differences
among the following age groups: children below the age
of three years, adolescents between the ages of 12 and 18,
young adults in their twenties, and older adults over 55
years. We have not found cognitive sex differences in
children between three years of age and puberty. Below
we present these data and discuss their possible meaning.
2. Cognitive sex differences in young children
Our laboratory has tested children on cognitive tasks
that were originally developed for monkeys. These tasks
include object discriminations, delayed matching and
non-matching to sample, single and multiple pair con-
current discriminations, visual preferential viewing, trial
q
This research was supported by National Institute of Child
Health and Human Development Grant RO1 HD 35542-01A1 and by
John D. and Catherine MacArthur Foundation, Research Network on
Psychopathology and Development Grant PR-15001/99.
E-mail address: overmanw@uncw.edu.
reversals (for review, see Overman & Bachevalier, 1999).
Two of these tasks are of particular interest for this
Special Issue because their use has revealed a double
dissociation of cognitive sex differences. The tasks are
object reversal learning and concurrent discrimination. In
monkeys the object reversal task is known to depend
upon the integrity of the orbital prefrontal cortex,
whereas the concurrent discrimination task is known to
depend upon area TE in the inferior temporal lobe (see
Section 2.3). We tested separate groups of children on
these tasks in a modified Wisconsin General Testing
Apparatus, with non-verbal operant procedures, for a
limited number of daily trials, 5 days a week, week after
week, until strict learning criteria were met (for details
see Overman, Bachevalier, Schuhmann, & Ryan, 1996).
First, the tasks are described, and then the results of
testing are reported.
2.1. Object reversal
This task requires the subject, over a number of days,
to learn to discriminate between two stimulus objects
one of which conceals a food reward (e.g., Cheerios or
Froot Loops) and one of which does not. In addition
to the food reinforcer a social reinforcer, (e.g., ‘‘Good

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doi:10.1016/S0278-2626(03)00279-3
 W.H. Overman / Brain and Cognition 55 (2004) 134–147
job!’’) also follows a correct response. After reaching
criterion performance (90% correct responses for two
consecutive test days) the subject learns a second dis-
crimination with two new stimuli. These first two dis-
crimination problems serve as control tasks for general
learning ability. On the first test session after attaining
criterion on the second discrimination problem, the re-
ward contingencies for that problem are reversed with-
out warning. Thus, the previously negative stimulus
becomes positive, and the previous positive stimulus
becomes negative. The subject learns this reversed dis-
crimination to criterion whereupon the reward contin-
gencies are reversed again, and so on, for as many
reversals as the experiment calls for. The subjectÕs score
for this task is the number of errors to criterion for each
reversal and across all reversals. A high number of er-
rors is interpreted as the failure to inhibit a prepotent
response (e.g., the learned response to the initial dis-
crimination) and/or the lack of behavioral flexibility.
2.2. Concurrent discrimination
In this task, on each test day, the subject is consec-
utively presented with a number of object pairs. One
member of each pair conceals food (Cheerio or Froot
Loop) and one does not conceal food. In addition, a
social reinforcer, (e.g., ‘‘Good job!’’) is used as a rein-
forcer. The same series of object pairs is presented once
each test day (24-h retention interval). The positive and
negative objects within each pair and the serial order of
the pairs remain constant across daily sessions, but the
left–right position of the baited and unbaited objects are
randomized daily. Testing continues, day after day, until
the subject attains a criterion of 90% correct choices for
two consecutive test days.
2.3. Sex specificity, neural bases, and the role of gonadal
hormones
There is a convincing double dissociation of perfor-
mance on these two tasks showing that two neural sys-
tems develop at different rates in male and female infant
monkeys. Specifically, infant males perform better than
females on the object reversal task, and infant females
perform better than males on the concurrent discrimi-
nation task. The evidence can be summarized as follows:
(a) Seventy-five day-old male monkeys are superior to
age-matched females on object reversal tasks (Goldman,
Crawford, Stokes, Galkin, & Rosvold, 1974). (b) While
normal infant males out-perform infant females on the
object reversal task, perinatally androgenized infant fe-
males perform as well as normal males and better than
normal females (Clark & Goldman-Rakic, 1989). (c)
Early ablations of orbital prefrontal cortex impair object
reversal performance of normal infant males and an-
drogenized infant females but such lesions do not impair
135
the performance of normal infant females until much
later in life (Clark & Goldman-Rakic, 1989; Goldman,
1971). (d) In contrast, on the concurrent discrimination
task, normal infant females out-perform infant males
(Bachevalier, Hagger, & Bercu, 1989); however, or-
chiectomized infant males perform as well as normal
females and better than normal males. (e) Neonatal
ablations of area TE in the inferior temporal cortex
impair concurrent discrimination performance in nor-
mal females but not in infant males (Bachevalier,
Brickson, Hagger, & Mishkin, 1990). (f) In males, levels
of circulating testosterone are inversely correlated with
performance on the concurrent discrimination task
(Hagger, Bachevalier, & Bercu, 1987).
Taken together, these data strongly indicate that, in
infant males. Perinatal testosterone accelerates the
functional maturation of orbital prefrontal cortex and
slows the functional maturation of area TE in the inferior
temporal lobe.
2.4. Results with children on object reversal and concur-
rent discrimination
We have replicated, in children, the behavioral com-
ponents of the double dissociation found in monkeys. We
tested children in the WGTA using almost exactly the
same testing procedures as used with monkeys, i.e., a
limited number of daily trials for many consecutive days,
food and social reward, no verbal task instructions. The
data can be found in Overman et al., 1996, 1997. The re-
sults were the following: (a) Male and female children
were equal in initial discrimination learning. (b) Males
under the age of 29 months were superior to age-matched
females in reversal learning following a pattern almost
exactly like that found in monkeys. With the addition of
more subjects we have recently discovered that the male
superiority exists through 36 months of age (Overman &
Godin, unpublished data). (c) Females under the age of 36
months were superior to age-matched males in learning a
concurrent discrimination task. (d) There were no sig-
nificant sex differences on either task in older children or
adults. (e) In addition to slower learning relative to males,
about 20% of the females under 29 months showed hyper-
emotional behaviors during reversal training.
As shown in Fig. 1, we found that male and female
children under the age of 30 months display significant
differences in learning object reversals almost exactly as
seen in infant monkeys. Children older than 29 months
and adults showed no sex differences in performance
(lower graph).
Fig. 2 shown the results of testing young monkeys
and children on the concurrent discrimination. In both
groups of youngest subjects, females out-performed
aged-matched males by a factor of nearly two. Older
children and monkeys and adults of both species showed
no sex differences. For infants of both species, the
136 W.H. Overman / Brain and Cognition 55 (2004) 134–147
Fig. 1. (A) Mean errors to criterion for reversals of an object dis-
crimination for 75-day-old male and female monkeys and (B) for two
age groups of male and female children. Vertical bars represent stan-
dard error of the mean (SEM). Top graph is adapted from Clark and
Goldman-Rakic (1989).
direction of the sex differences were distinct from one
learning ability to the other with the majority of males
being superior on object reversal and the majority of
females being superior on concurrent discrimination.
The extremely close similarity of the tasks, behavioral
data, and endocrine functions of infant humans and
monkeys provide inferential evidence that, in both spe-
cies, there is more rapid maturation of orbital prefrontal
circuits in males and more rapid maturation of inferior
temporal circuits in females. Presumably, this phenom-
enon is related to the perinatal testosterone surge that
occurs shortly after birth in males of both species
(Bachevalier et al., 1989; Corbier, Edwards, & Roffi,
1992; Stahl, Gotz, Poppe, Amendt, & Dorner, 1978).
3. A search for cognitive sex differences in older children
The results of the object reversal experiments with
children elicit two important questions: (1) If our infer-
ences are correct and cognitive sex differences in young
children are related to the testosterone surge (that is lim-
ited to the first half of the first year of life), why are the sex
differences found as late as three years of age (Overman &
Godin, unpublished data)? One possibility is that the
perinatal testosterone surge imparts some organizational
Fig. 2. (A) Mean errors to criterion on a 20-pair concurrent discrim-
ination task for young, juvenile, and adult monkeys. (B) Mean errors
to criterion on a 8-pair concurrent discrimination task for male and
female children, 13–35 months of age, 36–54 months of age, and young
adults. Vertical bars represent SEM. Top graph is adapted from
Bachevalier et al. (1989).
effect on brain anatomy and function that persists beyond
the duration of the elevated hormone levels. (2) However,
this possibility raises another question: if there are orga-
nizational changes, why are the cognitive sex differences
not found after three years of age? One possible answer to
the last question, is that beyond three years of age, chil-
dren make very few errors on the object reversal task (see
Fig. 2) and, thus, possible sex differences may be con-
cealed, i.e., there is a floor effect.
In the following experiments, we attempted to un-
cover cognitive sex differences in older children (3–5
years of age), preadolescents (11–12 years of age), and
young adults (college students) by utilizing an object
reversal task that was more challenging (and thus, gen-
erated more errors) than the simple task used above.
This task was a ‘‘partial’’ reversal task.
3.1. Partial reversal task
A total of 369 subjects were tested in this project: 262
preschool children (113 females and 149 males) ranging
in age from 3 to 5 years, 43 adolescent children
 W.H. Overman / Brain and Cognition 55 (2004) 134–147
(19 females and 14 males) ranging in age from 11 to 12
years, and 64 college age adults (14 females and 24
males). Originally we had proposed to test children as
young as two years of age but we quickly discovered that
the computerized reversal task was too demanding in
terms of length of test sessions, response requirements,
and comprehension of the reward system. Consequently,
only children three years of age and above were tested.
3.1.1. Procedure
During testing each child sat in front of a stand-alone
computer screen that was run by a separate lap top
computer in front of the experimenter. This arrange-
ment prevented the subjects from being distracted by the
keyboard. Most children were capable of using a mouse
to point and click on the stimuli, but the program was
designed to teach mouse use to the few inexperienced
children.
3.1.2. Methods
Each child was tested separately in one session that
lasted from 30 to 60 min depending on the complexity of
the reversal task and the ability of the subject. The child
was seated in front of the computer screen and, the ex-
perimenter explained that, in this ‘‘game,’’ stickers and
prizes could be obtained (stickers and prizes were out
of reach but were visible on the right hand side of the
table).
3.1.3. Pretraining: Teaching use of the mouse
On the first trial a 200
200
200 black-outlined trian-
gle appeared on left or right half of the computer screen
while the opposite half screen was black. The experi-
menter said ‘‘watch what happens when I click on the
triangle’’ and then clicked on it. This was followed by a
yellow ‘‘happy face’’ filling most of the screen and a
simultaneous ‘‘tinkle’’ of music. The face lasted 1.5 s,
disappeared and was followed by a 1.5-s message saying
‘‘GET READY.’’ Then the triangle appeared on the
right or left half of the screen. The child was handed the
mouse and asked, ‘‘Can you do what I just did?’’ Eight
such trials occurred with the triangle appearing ran-
domly on the left or right of the screen. On each suc-
cessive trial the darker half of the screen progressively
turned lighter so that on trial 8 the entire screen was
white.
3.1.4. Teaching two-choice responding and the first
discrimination
On the next 8 trials the triangle (S+) appeared on the
left or right half of the screen while a square (S)) was
gradually faded in on successive trials. By the 8th trial,
the screen showed a black outline triangle and an
equally distinctive black outlined square. Next, dis-
crimination training was begun and the subject was re-
quired to respond to the S+ until he/she had correctly
137
responded in two consecutive trials at 87% correct (two
blocks of 13/15 correct responses). During this dis-
crimination and the remaining test trials, the following
reinforcement system was introduced. After each set of 7
correct responses, a 3-s ‘‘colored light show’’ appeared
on the screen rather than the happy face. At each light
show the child was allowed to choose a colorful sticker
and was instructed to move one bead on a colorful
abacus. After a row of 10 beads was moved the child
was allowed to choose a small toy on the table (small
dinosaur, bracelet, animal, yo-yo, etc.). At the end of the
test session, the child was allowed to choose a larger toy
(toy motorcycle, car, necklace, ring, change purse, etc).
Incorrect responses were followed by a 2 s black screen.
3.2. Teaching the second discrimination
After attaining criterion on the triangle–square dis-
crimination, the subject was shown a pentagon (+) vs. a
circle ()). The same reinforcement contingencies were
used as above. Criterion for this discrimination was
again 87% responses for two consecutive blocks of 15
trials.
3.2.1. Reversal learning
After attaining criterion on the pentagon-circle dis-
crimination, without warning, the pentagon became the
negative stimulus and the circle became the positive
stimulus. The percentage of the reversals could be pre
programmed so that the density of reversal was 100, 93,
86, 80%, etc. A 93% reversal consisted of 14 trials in
which the S+ was correct and one (disconfirming) trial
in which S) was correct. The 86 and 80% reversals had 2
and 3 disconfirming trials, respectively, in each block of
15 trails. Table 1 shows trials of full and partial rever-
sals. For any partial reversal, the subject learned to re-
spond to the stimulus that was correct on most of the
trials. Performance criterion was always 87% correct
responses for two consecutive blocks of trials. Correct
responses on disconfirming trials were always counted as
‘‘correct’’ by the computer even though the subject was
not reinforced for that response. After attaining crite-
rion on the first reversal, the reinforcement contingen-
cies switched without warning so that now the pentagon
was again positive and the circle was again negative.
Three such reversals were given to each subject.
4. Results
4.1. Learning in the WGTA compared with learning on
the computer
One hundred and twenty-one children between the
ages of 3 and 5 years were trained on two object dis-
criminations followed by three 100% reversals of the
138 W.H. Overman / Brain and Cognition 55 (2004) 134–147

Table 1
Average number of training sessions and average errors through criterion for discrimination and reversal learning in the Wisconsin General Testing
Apparatus and on a computer
Group # sessions Disc. 1 Disc. 2 Rev. 1 Rev. 2 Rev. 3
WGTA 13.1 6.3 2.1 7.6 2.8 3.4
Computer 1.0 0.6 2.0 7.1 3.5 3.5
There were no disconfirming trials on these reversal tasks.

Table 2
Average errors to criterion on discriminations and reversals for 8 groups of participants
Group Discrimination 1 Discrimination 2 Reversal 1 Reversal 2 Reversal 3
WGTA preschool 6.3 2.1 7.6 2.8 3.4
Computer preschool 0 disco. 0.6 2.0 7.1 3.5 3.5
Computer preschool 1 disco. 0.9 1.8 7.78 6.6 7.0
Computer preschool 2 disco. 1.1 1.1 13.4 11.2 9.3
Computer adoles. 2 disco. 0.03 0.2 19.1 13.6 15.2
Computer adoles. 3 disco. 0.08 0.2 56.7 40.7 29.5
Computer adult 2 disco. 0.2 0.2 10.6 7.9 8.8
Computer adult 3 disco. 0.15 0.0 22.3 16.1 10.1
‘‘Disco.’’ ¼ disconfirming trials.

second discrimination. Thirty-three of these (18 males
and 15 females) were trained in the WGTA at 15 trials a
day, day after day, until attainment of criterion on each
phase of the task as reported earlier (Overman et al.,
1996). Children in this group received food reward
(Froot Loops) and social rewards (‘‘good boy/girl’’) for
each correct response.
As shown in Table 2 the two groups of children who
learned the traditional reversal task (no disconfirming
trials) differed vastly in the amount of time they par-
ticipated in the tasks. Children in the WGTA group
required and average of 13.1 individual test sessions for
completion of the task. Since each test session took
about 20 min each day, each child in the WGTA group
spent a total of approximately 4.4 h with the experi-
menter. In contrast, children in the computer group
were tested in a single session, which was rarely more
than 1 h in duration. Stated another way, children in the
WGTA group experienced distributed practice while
children in the computer group experienced massed
practice.
Despite the difference in practice time, learning per-
formances were remarkably similar (Table 2 and Fig. 3).
The only exception to this was that, because of the fade-
in procedure, children in the computer group learned the
first discrimination significantly faster than those in the
WGTA group, p < :05. There were no other significant
differences between the tasks. The second discrimination
was learned equally fast by both groups (2.1 vs. 2.0,
p > :05). The reversals were learned equally fast in both
groups (average total errors across all reversals 13.8 vs.
14.1, p > :05). Both groups demonstrated the classic

Fig. 3. Mean errors to criterion on ‘‘full’’ (100%) and ‘‘partial’’ object reversal tasks for male and female children, adolescents, and young adults.
Asterisks indicate significant differences between reversal conditions (i.e., number of disconfirming trials).
 W.H. Overman / Brain and Cognition 55 (2004) 134–147
reversal learning pattern with high numbers of errors on
the first reversal followed by decreasing errors thereaf-
ter. The equivalent performance in the WGTA and on
the computer means that results from the more efficient
computerized task can confidently be compared with
results from animals and humans who were tested in the
WGTA.
4.2. Computerized partial reversal learning in children: 1
disconfirming trial
Another group of 79 preschool children (42 males
and 37 females) were tested on two discriminations as
described above and three reversals of the second dis-
crimination; however, the reversal was only a 93% re-
versal (one disconfirming trial randomly occurring
among each block of 15 trials). Criterion requirements
were the same as above: two consecutive blocks of 88%
correct responses at each phase of learning. There were
no differences in learning the first two discriminations
compared with children learning a regular ‘‘full’’ rever-
sal on the computer. This is because the discrimination
tasks were exactly the same in both groups.
Fig. 3 shows the performance of male and female
children on the computer task with one disconfirming
trial. The partial nature of this task (presence of the
disconfirming trial) rendered it significantly more dif-
ficult than the regular reversal task, p < :05. The dif-
ference was about 7 errors for females and 5 errors
for males. Thus, our effort to make a reversal task
more challenging by making it a partial reversal was
successful. However, there were no significant sex dif-
ferences with males learning three reversals in an av-
erage of 19.9 errors and females in an average of 22.9
errors.
4.3. Computerized partial reversal learning in children: 2
disconfirming trials
Another group of 62 preschool children (45 males
and 17 females) were tested on two discriminations as
described above and three reversals of the second dis-
crimination; however, the reversal was only a 87% re-
versal (2 disconfirming trial randomly occurring among
each block of 15 trials). Criterion requirements were two
consecutive blocks of 88% correct responses at each
phase of learning. There were no differences in learning
the first two discriminations compared with children
learning a regular reversal or a reversal with one dis-
confirming trial on the computer.
Fig. 3 shows the total errors on reversals 1, 2, and 3
for male and female on the computer task 2 discon-
firming trials. The additional disconfirming trial ren-
dered the task significantly more difficult than the task
with one disconfirming trial or the task with no dis-
confirming trials (p’s < :05) of 8.1 errors from the pre-
139
vious task. Still, there were no significant differences
between males (35.9 average total errors) and female (29
average total errors) on this task.
4.4. Performance by adolescents: Computer partial
reversal with 2 disconfirming trials
A group of 32 7th grade students (19 boys and 13
girls mean age of 12 years) were administered the com-
puterized partial reversal task with 2 disconfirming tri-
als. As incentives for driving to the university for testing
parents received a $15 coupon for groceries and for
participation children were given a tee-shirt and cou-
pons to local stores.
Fig. 3 shows the total errors across three reversals
for adolescent males and females. There were no sig-
nificant sex differences with or without inclusion of
outlying subjects. One interesting result was that ado-
lescents made significantly more errors on the 2 dis-
confirming-reversal problem than did preschool
children. This was true whether or not the two outlying
adolescent subjects (one male and one female) were
included in the analysis.
Because of the spontaneous comments made by
children and adolescents, we speculate that adolescent
participants differed in that they were trying more than
children to figure out complex (and non-existent) ‘‘pat-
terns’’ for the deliveries of reinforced and non-rein-
forced trials. In other words, the children were better at
probability matching than were adolescents. This is
somewhat similar to reports of animals being better at
probability matching than adult humans (Wolford,
Miller, & Gazzaniga, 2002). Perhaps intellectual devel-
opment allows the older participant to generate erro-
neous pattern strategies of which younger participants
are not yet capable. In addition the adolescent subjects
appeared to be very frustrated by the task and less so
than adults on the same task who made significantly
fewer errors as shown below.
4.5. Adolescents: Computer partial reversal with 3
disconfirming trials
In an effort to make the partial reversal task even
more challenging 11 adolescents (5 males and 6 females)
were given the test with 3 disconfirming trials in every
block of 15 trials. As shown in Fig. 3, this problem was
exceedingly difficult with approximately a 100% increase
in errors as compared to the task with 2 disconfirming
trials. Although the participants completed all three
reversals they became frustrated and upset during the
session which required a minimum of 90 min to com-
pete. Since there was no evidence for sex differences on
this task and since the task was obviously unpleasant for
the participants we stopped testing adolescents with 3
disconfirming trials.
140 W.H. Overman / Brain and Cognition 55 (2004) 134–147
4.6. Adults: Computer partial reversal with 2 and 3
disconfirming trials
Thirty-two adult college students (16 males and 16
females) were testing on partial reversal tasks with 2
disconfirming trials, and an additional 32 adults (8
males and 24 females) were tested with 3 disconfirming
trials. The results are shown in Fig. 3. There were no sex
differences under either condition (p > :05); however, the
3 disconfirming trial condition was significantly more
difficult than the 2 disconfirming trial condition (average
of 48 errors vs. 27 errors). Adults made significantly
fewer errors than did adolescents in both conditions
(p’s < :05)
4.7. Summary of partial reversal testing
It is evident that, for preschool children, computer-
ized discrimination, and reversal learning is comparable
to learning using the WGTA apparatus. There is no
reason to believe that this is not the case for adolescents
and adults. The reversal task was clearly rendered more
difficult for all ages of subjects by adding disconfirming
trials. However, partial reversal manipulation did not
result in significant differences between males or females
at any age. Our failure to find sex differences among
young was puzzling because, as shown below, we were
documenting sex differences among adults on another
task known to be sensitive to orbital prefrontal damage.
5. Cognitive sex differences among adults: The Iowa
Gambling Task
One of the reoccurring themes of this Special Issue
concerns cognitive changes that follow damage to the
orbital prefrontal cortex. One of the most common
changes involves impairments in the quality of personal
decision-making abilities. Patients with orbital damage
demonstrate relative insensitivity to future consequences
even when their decisions are against their personal in-
terest; however, these patients maintain relatively nor-
mal intellectual functioning in other realms (Bechara,
Damasio, Damasio, & Anderson, 1994; see also Bec-
hara, this issue).
5.1. The Iowa Gambling Task
Bechara et al. (1994, & see this issue) developed the
‘‘Iowa Gambling Task’’ (IGT) that detects deficits in
personal decision-making. This task simulates real-life
decision-making in the manner in which it presents an
uncertainty of reinforcers and punishers. The task re-
quires the participant to choose cards from four decks.
Cards in two of the decks, e.g., yellow and blue decks,
are associated with high reward but even higher spo-
radic loss of reward (disadvantageous decks). Cards in
the other two decks, e.g., red and green decks, are as-
sociated with low reward but with even lower sporadic
losses (advantageous decks). Consistent choice of ad-
vantageous cards (red and green) will result in low but
long-term gain, whereas consistent choice of disadvan-
tageous cards (yellow and blue) will result in overall loss
of money. Over the course of 100–200 trials, normal
control participants gradually formulate the strategy of
picking from the low-paying (advantageous) decks,
which result in gain in the long run. In contrast, patients
with damage centered on the ventromedial prefrontal
cortex do the opposite, and select more cards from the
high-paying (disadvantageous) decks, which results in
loss in the long run (Bechara et al., 1994, 1997.) How-
ever, the same patients perform normally on standard
tests of executive function such as the Wisconsin Card
Sorting Test (WCST) Bechara et al. (1994, 1997).
5.2. Orbital prefrontal cortex and decision-making
The role of the orbital PFC in decision-making is not
precisely understood at this time. Several operations
have been proposed. Damasio and his colleagues have
written extensively about the ‘‘somatic markers’’ which
are emotional feelings that gradually come to be con-
nected, by learning, ‘‘to predicted outcomes of certain
scenarios’’ (Damasio, 1994, p. 174). These emotional
‘‘markers,’’ as defined by Skin Conductance Responses
(SCRs), are thought to act as an early warning system
which guides decisional behavior before the subject is
consciously aware of the precise rule. Unlike normal
controls, patients with orbital prefrontal damage do not
generate anticipatory SCRs during the course of playing
the IGT (Bechara et al., 1994). The lack of SCRÕs is
thought to underlie disadvantageous decision-making in
such patients (see Bechara, this volume). It should be
noted that, recently, the somatic marker hypothesis has
come under scrutiny (Tomb, Hauser, Deldin, &
Caramazza, 2002).
In contrast to the somatic marker hypothesis, Rolls
(1996) has suggested that the orbital PFC mediates de-
cision-making by providing action selection mechanisms
with information about the reinforcing properties of
unconditioned and conditioned stimuli. The precise
distinctions and overlaps between these two hypotheses
are not critical for this present research.
5.3. Populations that perform poorly on these decision-
making tasks
The following patient populations have been shown
to perform at sub-normal levels on the IGT (impaired
performance is defined by the continued choice of dis-
advantageous $100 cards): (1) patients with damage
centered on the ventromedial prefrontal cortex (Bechara
 W.H. Overman / Brain and Cognition 55 (2004) 134–147
et al., 1994, 1997); (2) polysubstance abusers (stimulants
or opiods) (Grant, Contoreggi, & London, 2000); (3)
cocaine abusers vs. abstinent cocaine abusers and non-
abusers (Bartzokis, Lu, Beckson, & Rapaport, 2000)
[Patients in these studies performed normally on the
Wisconsin Card Sorting Test]; (4) violent offenders vs.
non-violent offenders (Fishbein, 2000); (5) pathological
gamblers (Cavedini, Riboldi, Keller, DÕAnnucci, &
Bellodi, 2002).
Patients with amygdala damage also perform poorly
on the IGT. However, unlike patients with damage to
the orbital PFC, those with amygdala damage do not
produce SCRs even after receiving a reinforcer or pun-
isher (neither group produced anticipatory SCRs)
(Bechara, Damasio, Damasio, & Lee, 1999). This sug-
gests that the two regions play different roles in decision-
making: amygdala damage has indirect consequence on
attaching affective qualities to stimuli where as the VM
region is not essential for the generation of affective
attributes to (i.e., in response to) emotional stimuli
(Bechara et al., 1999).
5.4. Sex differences in performance on the IGT
Over the past four years we have tested over 700 non-
substance-abusing participants (age range 12–65 years)
on the IGT (plus control tasks) and have discovered the
following (described in detail below). (1) Performance
on the IGT improves with age, from 12 years of age
until adulthood. (2) Gonadal hormone status does not
affect performance in adults. (3) Across age, males
outperform females. (4) Young male adultsÕ perfor-
mance significantly improves when they have prior ex-
perience with another decision-making task (the
California Weather Task). (5) By using a novel error
analysis, we have found that females demonstrate dif-
ferent choices from males due to what we believe are
emotional responses.
5.4.1. Sex differences among adults
In a recent study we tested 191 young and older adults
on the IGT and the California Weather Task (CWT)
(Reavis & Overman, 2001). The latter task is a slowly
acquired probabilistic habit task on which Parkinsons
and Huntingtons patients are significantly impaired
(Knowlton, Mangels, & Squire, 1996). In the California
Weather Task, participants are shown up to four dis-
tinctly patterned cards on a computer screen and grad-
ually learn which combinations of pattern predict ‘‘rain’’
or ‘‘sunshine.’’ A particular card pattern is associated
with the outcome of ‘‘rain’’ 75, 57, 43 or 24% of the time
and thus 24, 43, 57, or 75% of the time with the outcome
of ‘‘sunshine.’’ Participants are exposed to any combi-
nation of cards on each trial, and they gradually learn
which cards and combinations are probabilistically related
to a given outcome. The computer provides written
141
feedback (correct or incorrect) as well as auditory
feedback (high tone for correct and low tone for incor-
rect). Learning difficulty is increased by the inclusion of
prearranged trials that are ‘‘disconfirming’’ (e.g., they
give erroneous feedback about correct or incorrect
choices). Neither disconfirming nor equal-probability
trials are included in scoring.
The participants were divided into 6 groups defined
by age and hormonal status: young males (mean age ¼
19.1 years), older males (mean age ¼ 59.4 years), young
females with low hormones (menstruating, mean age ¼
19.8 years), young females with high hormones (mid-
luteal, mean age ¼ 22.4), older postmenopausal women
either taking estrogen replacement therapy (mean age ¼
54.5 years) or not taking ERT (mean age ¼ 62. 7 years).
Hormonal status was defined via Radio Immuno Assay
of dried blood spots (Shirtcliff, Reavis, Overman, &
Granger, 2001). Each subject was tested individually.
Each subject completed two preliminary questionnaires
and then the two tests of probability learning, the IGT
and the CWT. Half of the participants in each group
took the Iowa Task first and half took the California
Task first. On the IGT, participants were given 150 trials
and every 10th trial they were asked if they knew the
operative rule (that the $50 red and green decks gained
money in the long run and the $100 yellow and blue
decks lost money in the long run).
Results showed no significant task differences be-
tween any of the four hormonal groups of women and
they were collapsed into a single female group. Simi-
larly, there were no task differences between the young
and older men, so they were grouped together. While
there were no sex differences on the California Weather
Task, there were interesting sex differences on the IGT,
as shown in Fig. 4.
Males chose significantly more advantageous cards
($50 cards) than did females on the 2nd and 3rd block of
50 trials (p < :002). By the 3rd block males chose $50
cards on 79% of the trials while females chose them on
68% of the trials. Furthermore, more males stated the
correct rule at some point during the task than did fe-
males (63.5 and 38%, respectively) (p < :009). And
males stated the decision-making rule significantly ear-
lier in training (75th trial on average) than did females
(97th trial on average; p < :009; Reavis & Overman,
2001). It is interesting to note that Kerr and Zelazo (this
issue) reports that 3- to 4-year-old boys out perform
age-matched girls on a child version of a decision-
making task.
5.4.2. Task order effect: Does practice help?
One of the most interesting findings in this study was
an effect of task order. While there was no overall sig-
nificant difference between younger and older males in
choice of $50 cards (70 vs. 64%, respectively), younger
men were clearly superior on the task relative to older
142 W.H. Overman / Brain and Cognition 55 (2004) 134–147
Fig. 4. Mean percent of correct choice ($50 cards) in the IGT by adult
men and women as a function of blocks of 10 trials. Vertical bars
represent SEM.
men and all women when the IGT followed the Cali-
fornia Weather task. Young males significantly improved
in their performance when the IGT was second (64%
choice of $50 cards improved to 75% choice) (p < :05). In
contrast, older males declined in their performance
when the IGT was second (70% choice of $50 cards
declined to 54% choice).
Fatigue can be ruled out as the cause for the older
malesÕ declining order effect on the IGT because they
showed no similar decline on the weather task when it
was second. Rather, it appears that having the probabi-
listic weather task first somehow benefited the younger
men’s performance on the IGT. Thus this improvement
effect in younger men appears to be related to changes in
cognitive processes, not to fatigue, motivation, percep-
tual or motor factors.
These findings generate several questions. Since the
California Weather task involves probabilistic decisions,
one wonders: (1) If practice on such tasks might improve
performance on subsequent and similar tasks? (2) Is this
effect restricted to young adult males or might it occur in
adolescent males or females? (3) Might the effect occur in
adolescent males or females who are substance abusers?
We are currently investigating these questions.
5.4.3. Sex differences among adolescents: Preliminary
results
Until now, the IGT has been given only to adult
populations. We wanted to study the performance of
younger normal subjects on this task to determine pos-
sible developmental trends. We have tested 420 adoles-
cents, 30 males, and 30 females in each school grade 6–
12. In addition, 60 young adults (30 females and 30
males) were tested (Overman et al., 2003; under review).
Each participant was tested individually and each was
given 200 trials of the IGT as in Reavis and Overman
(2001), followed by a computerized WCST, a survey of
risk-taking behavior adapted from the NEO personality
inventory (Costa & McCrae, 1992) and finally an age-
adapted Quantity–Frequency Index of Substance Use
(Polich, Armor, & Braiker, 1981). The latter index
consists of items asking about the participantÕs quantity
and frequency of use of tobacco, alcohol, marijuana,
cocaine, ecstasy, and opioids.
The major results were:
(a) Reports of substance use and risk-taking were not
significantly correlated with performance on either task.
The reason for this was clear from the results of the
surveys: There were few, if any, substance abusers at any
age. We speculate that this is because participation in
the study was voluntary, and those subjects who were
conscientious enough to stay after school and partici-
pate were simply not the type of adolescent who engages
in substance abuse, risk-taking, and poor decision-
making. This study basically provides us with normative
data on the changes in performance across adolescence
on the IGT. (b) In agreement with previous findings,
performance on the WCST was not significantly corre-
lated with sex, reported substance use, or performance
on the IGT (all p’s > 05.) (c) As predicted, across age
there was a significant sex difference in favor of males
(p < :05). This finding is in agreement with previous
results (Reavis & Overman (2001)). (d) As predicted,
performance on the IGT significantly improved with
age (p < :05). A summary of the results is shown in
Fig. 5.
As shown here, performance on the IGT (selection of
‘‘good’’ or advantageous $50 cards) improved with age.
Young participants (e.g., 6th and 7th grade) chose ad-
vantageous ($50) cards less than 60% of the time while
Fig. 5. Mean percent of correct choice ($50 cards) in the IGT across
200 trials as a function of grade. Vertical bars represent SEM.
 W.H. Overman / Brain and Cognition 55 (2004) 134–147
older ones (e.g., 10th, 11th, 12th grades, and adults)
chose $50 cards about 70% of the time. As shown in
Fig. 6, an overall significant sex difference was found in
the ANOVA, but clearly there were no sex differences
among 6th graders, 10th graders or adults. We have no
explanation for the odd appearance of the data points
for 10th graders other than a vagary of sampling.
The lack of sex differences among this sample adults
seemingly conflicts with our finding of a male advantage
in the earlier study (Reavis & Overman, 2001). In that
study, task order was counter balanced and young adult
males who had the IGT second scored significantly
higher than those who had this task first. We speculate
that the previous finding of sex differences may have
been driven by this order effect that so strongly affected
males. This hypothesis was investigated though the use
of a new type of error analysis as presented in the next
section.
5.4.4. New error analysis for the IGT
Heretofore, researchers who have administered the
IGT measured the percentage of advantageous ($50)
cards chosen in blocks of trials (or the number of $50
cards minus the number of $100 cards). In other words,
the only error was considered to be choice of disad-
vantageous ($100) cards. But there are other types of
errors that can be made on this task. In our data set with
normal adolescents and young adults we have devised a
new error analysis that reveals very interesting differ-
ences between males and females.
The new error analysis looks at choice of cards as a
function of the absolute number of minus ‘‘)’’ signs and
plus ‘‘+’’ signs (as well as the absolute net outcome of
the cards). Two of the decks (Yellow $100 and Blue $100
decks) are disadvantageous because the penalties mon-
etarily outweigh the rewards in the long run. Two of the
decks (Red $50 and Green $50 decks) are advantageous
in that the rewards monetarily outweigh the penalties in
Fig. 6. Mean percent of correct choice ($50 cards) in the IGT as a
function of grade and sex. Vertical bars represent SEM.
143
the long run. However, neither the two disadvantageous
decks (Yellow and Blue) nor the two advantageous
decks (Red and Green) are equal in the absolute number
of penalties or ‘‘)’’ (minus) signs. The participant wins
some money (but not necessarily as net win) on every
card, either $50 or $100 and, thus, there is a ‘‘+’’ (plus)
sign on every card in the task. However, losses occur less
often, so only some of the $50 and $100 cards contain a
‘‘)’’ (minus) sign.
Consequently, the cards can be classified as those
which contain only ‘‘+’’ signs vs. those that also contain
a ‘‘)’’ sign. By this definition, both the disadvantageous
and the advantageous decks contain ‘‘+’’ cards and ‘‘)’’
cards. Table 3 shows a summary of each colored deck
with respect to monetary wins and losses.
This table can be summarized as follows: (1) Every
card contains a ‘‘+’’ sign.
(2). Two decks could be considered ‘‘+’’ (plus) decks
because they contain 10 pluses and only one minus per
10 trials; however, one of these is a $50 deck (Red) and
one is a $ 100 deck (Yellow). (3). Two decks could be
considered as ‘‘)’’ (minus) decks (relatively speaking)
because they contain 10 pluses and 5 minuses per 10
trials; however, one is a $50 deck (Green) and one is a
$100 deck (Blue).
In contrast to males, females choose the ‘‘+’’ decks
rather independently of their status of disadvantageous
($100) or advantageous ($50) (Figs. 7A and B).
Results of all females (grade 6 through adults) are
shown in the upper graph, results of all males (grade 6
through adults) are shown in the lower graph. As shown
in Fig. 9, both males and females quickly learn to avoid
the disadvantageous blue deck presumably because it
has five large penalties per 10 trials. Both males and
females quickly learn to choose the advantageous red
deck, presumably because it has only one small penalty
per 10 trials. The sexes differ on choices of the other two
decks. Males learn to choose the advantageous ($50)
Green decks and avoid the disadvantageous ($100) yel-
low deck that has one large loss per 10 trials. In contrast,
females choose the disadvantageous ($100) Yellow card
fairly constantly for the duration of the task.
5.5. Nature of female and male responding
It is possible that females are responding more than
males to the disadvantageous Yellow deck because it
contains 10 ‘‘+’’ and only 1 ‘‘)’’ per 10 trials (exactly as
does the advantageous Red Deck). In other words, they
appear to be guided by the absolute number of pluses
and minuses: they approach pluses ‘‘+’’ and avoid mi-
nuses ‘‘)’’ rather independently of the net outcome.
Males, on the other hand appear to be guided by the net
monetary outcome of the cards rather than simply
number of ‘‘+’’ and ‘‘).’’ Stated another way, males
appear to be more sensitive to the long-term monetary
144 W.H. Overman / Brain and Cognition 55 (2004) 134–147

Table 3
Summary of each colored deck with respect to monetary wins and losses, and number of pluses and minuses
RED (+) Green (+) Yellow ()) Blue ())
Amount won or lost per 10 trials +$250 +$250 )$250 )$250
Number of plus cards per 10 trials 9 5 9 5
Number of minus cards per 10 trials 1 5 1 5
Number of break-even cards per 10 trials 0 3–5 0 0

Fig. 7. Selections in blocks of 50 trials of particular colored cards.

Advantageous cards ($50) are red and green. Disadvantageous cards Fig. 8. Selections of particular colored cards by adult females when (A)
($100) are yellow and blue. (A) Results of all females (grade 6— the IGT was presented first and (B) when the IGT followed the Cali-
adulthood). (B) Results of all males (grade 6—adulthood). fornia Weather task.

outcome while females appear to be more sensitive to
the immediate presence or absence of ‘‘+’’ or ‘‘).’’
During testing we often observed that a female would
verbally respond negatively (e.g., ‘‘oh, no’’) after
choosing an advantageous Green card with the value of
+ $50 and ) $25. Given such reactions and the simple
math involved, the sex difference does not appear to
reflect femalesÕ inability to perform a simple calculation.
Rather, it appears as if the female was reacting emo-
tionally negatively to the presence of the ‘‘)’’ sign. These
data may indicate that, on this task, femalesÕ decision-
making is guided by avoidance of negativity (‘‘)’’ signs)
more than is malesÕ decision-making, which is guided
more by long-term outcome.
Another possibility is that females may be less effi-
cient in reversing associations related to the yellow deck.
In this deck, the first 8 cards were $100 ‘‘wining’’ cards.
The first loss did not occur until the 9th card in that
deck. Perhaps, the females formed strong preferences for
the ‘‘winning’’ yellow cards and were slower to change
to other decks compared to males. This hypothesis
would fit with the slow reversal behavior observed in
infant females (Overman et al., 1996).
Regardless of the cause, it is clear that male and fe-
malesÕ decision-making is being guided by different
factors. We are currently investigating the possible fac-
tors underlying this difference.
5.5.1. What is being learned by males when a probabilistic
task is given prior to the IGT?
Recall that in our previous study, young adult males
showed significantly improved performance (selecting
more advantageous $50 cards) after having the probabi-
listic California Weather Task (Reavis & Overman, 2001).
 W.H. Overman / Brain and Cognition 55 (2004) 134–147
To better understand the nature of this improvement,
we have reanalyzed those data in terms of choice of
color (i.e., plus ‘‘+’’ or minus ‘‘)’’ cards). The results are
shown, first for young adult females who had the IGT
first or second, and next for the young adult males.
Fig. 8 shows the selection of different cards by women
who had the IGT first (upper graph) or second, after the
California Weather Task (lower graph). A comparison
of these graphs reveals that there were little changes
between groups in the avoidance of the disadvantageous
$100 Blue card (with 5 penalties per 10 trials) or in the
preference for the disadvantageous $100 Yellow card
(with one large penalty per 10 trials). When the IGT was
second there appeared to be some increase in the selec-
tion of the advantageous $50 Green card (with 5 minus
signs per 10 trials) whereas preference remained about
the same for the advantageous $50 Red Card (with one
small penalty per 10 trials).
In contrast, as shown in Fig. 9, men showed quite
different card selection when the IGT followed the
probabilistic Weather Task.
The largest change was seen in the decrease of choice
of the disadvantageous $100 Yellow card (with one large
penalty per 10 trials). Aversion was apparent even in the
first block of 50 trials and extremely apparent in the last
block of trial, during which both the disadvantageous
Yellow and Blue cards were rarely chosen. In addition,
Fig. 9. Selections of particular colored cards by adult males when (A)
the IGT was presented first and (B) when the IGT followed the Cali-
fornia Weather task.
145
when the IGT was second in the test series, men selected
significantly more advantageous Red and Green Cards.
It is obvious that ‘‘something’’ was imparted to the
men by having the California Weather task first. We do
not believe that this was an increase in the ability to
perform calculations, because the Weather Task does
not require mathematical calculations.
Two possibilities come to mind: one based on ‘‘cool’’
brain function and one ‘‘hot’’ brain function, to use cur-
rent parlance of frontal lobe function. The ‘‘cool’’ possi-
bility is that the first task imparted some non-emotional
cognitive ability, e.g., probability estimation, that en-
hanced performance on the second task. Given the nature
of the California Weather Task, males may have learned
to judge the long-term probabilities of net gain or loss
associated with the different type of cards. Females did
not appear to benefit in the same manner from having
experience with the California Weather Task, again per-
haps because they were being more strongly guided by the
immediate presence of plus ‘‘+’’ or minuses ‘‘).’’ This
brings us to the possible ‘‘hot’’ or more emotional ex-
planation that may apply differentially to males and fe-
males. The California Weather Task is difficult and
frustrating, due to the presence of disconfirming trials. [It
is also noteworthy to know that all experimenters were
females.] Perhaps the difficulty of the California Weather
Task (in the presence of a female experimenter) aroused
anxiety/emotion in the males. And perhaps such anxiety/
emotion somehow primed the orbital prefrontal system in
such a way as to improve performance in males. This may
not have occurred in females due to the facts that their
experimenter was female, and perhaps less threatening,
and that their affective behavior appeared to be centered
around avoidance of minus ‘‘)’’ cards.
This second speculation is somewhat bolstered by
other reports at this Special Issue. For example, Bec-
hara, this volume, reports that simultaneous stimulation
of the vagus nerve improves performance on the IGT,
presumably by enhancing some emotional state. Seguin
(this issue) reports that anxiety can influence perfor-
mance on the IGT. Blair (this issue) reports that PET
scans of the lateral orbital prefrontal cortex is influenced
by viewing photographs of angry faces.
In summary, it is clear that for males, prior experi-
ence with the California Weather Task somehow en-
hanced performance on subsequent IGT. This may not
be a trivial finding. The final explanation of this phe-
nomenon may significantly expand our understanding
of the orbital prefrontal system.
6. Additional evidence for sex differences in the orbital
prefrontal cortex
In addition to the data cited above, there is accu-
mulating evidence from other methodologies for sex
146 W.H. Overman / Brain and Cognition 55 (2004) 134–147
differences in the functions of orbital prefrontal cortex.
Thus, positron emission tomography (PET) study re-
veals a significant difference in the metabolic rate of
glucose utilization in the orbital prefrontal cortex of 19-
to 32-year-old normal men and women (Andreason,
Zametkin, Guo, Baldwin, & Cohen, 1993). During a
mentally ‘‘idling’’ state, normal adult men have higher
relative metabolism (as shown by PET) than women in
several brain regions including the orbital prefrontal
cortex, and the temporal–limbic system (hippocampus,
amygdala, and temporal lobe) (Gur et al., 1995). In
contrast, women have higher metabolism in the middle
and posterior cingulate gyrus (Gur et al., 1995). Results
from Magnetic Resonance Imaging (MRI) investiga-
tions show that sex differences in the frontal lobe, and,
to a lesser extent, in the temporal lobes, continue into
advanced age. Hemispheric volumes obtained from
MRIÕs of young (18–40 years of age) and older (42–80
years of age) adults reveal a sexually dimorphic, region-
specific, age-related reduction in the volume of frontal
and temporal regions (Cowell et al., 1994). The brain
volume reductions are greater in men than in women.
The authors propose that these region-specific brain
dimorphisms have origins in early neuroendocrinologi-
cal development (Cowell, Allen, Zalatimo, & Dennen-
berg, 1992).
7. Summary and implications
Our lab has documented sex differences in the be-
havior of young children, adolescents, and adults on
tasks that rely on the integrity of the orbital prefrontal
cortex. Others in this Special Issue have at least some
data to support our findings. For the most part, studies
presented in this issue, did not include analyses for sex
differences. Hopefully, these behavioral and imaging
data will be analyzed for sex differences.
Evidence is accumulating that regions of the frontal
cortex, including orbital prefrontal, may be anatomically
and functionally different for males and females. It is
unclear at this time how such sex differences may be
manifest in behaviors other than laboratory tasks. Recent
evidence indicates that performance on the IGT may be
related to several brain areas outside the orbital regions,
e.g., dorsolateral and dorsomedial prefrontal cortices
(Manes et al., 2002). Consequently, the behavioral sex
differences reported in the present data may be related to
differential functioning of one or more of these brain
systems.
Many questions remain: Are such brain differences
related to sex differences in emotional behaviors? Are
they related to sex differences in broader real-life deci-
sion-making processes? Are the sex differences more or
less ‘‘wired’’ at birth or are they conditioned over the life
span? Are sex differences in orbital prefrontal cortex
related to the well-documented numerical difference in
male and females who abuse various substances?
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