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 FOCUS ON NEUROPSYCHOLOGY
RESEARCH

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liability is assumed for incidental or consequential damages in connection with or arising out of information
contained herein. This digital document is sold with the clear understanding that the publisher is not engaged in
rendering legal, medical or any other professional services.
FOCUS ON NEUROPSYCHOLOGY
RESEARCH

JOSHUA R. DUPRI
EDITOR

Nova Science Publishers, Inc.

New York
Copyright © 2006 by Nova Science Publishers, Inc.

All rights reserved. No part of this book may be reproduced, stored in a retrieval system or
transmitted in any form or by any means: electronic, electrostatic, magnetic, tape, mechanical
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liability is assumed for incidental or consequential damages in connection with or arising out of
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AMERICAN BAR ASSOCIATION AND A COMMITTEE OF PUBLISHERS.

Library of Congress Cataloging-in-Publication Data

Focus on neuropsychology research / Joshua R. Dupri (editor).
p. cm.
Includes bibliographical references and index.
ISBN: 978-1-60876-231-6 (E-Book)
1. Neuropsychology. I. Dupri, Joshua R.
[DNLM: 1. Neuropsychology. 2. Mental Processes--physiology. 3. Brain--physiology. 4. Models,
Neurological. WL 103.5 F652 2006]
QP360.F63 2006
612.8'2--dc22 2005030795

Published by Nova Science Publishers, Inc. New York

 CONTENTS

Preface vii
Chapter 1 Neural Substrates of Executive Functions: Insights from Functional
Neuroimaging 1
Robert M. Roth, John J. Randolph,
Nancy S. Koven, and Peter K. Isquith
Chapter 2 ERP Signs of Frontal and Occipital Processing
of Visual Targets and Distractors within and
without the Channel of Spatial Attention 37
Alberto Zani and Alice Mado Proverbio
Chapter 3 Differential Contributions of Viewer-Centred and
Environment-Centred Representations in Unilateral Neglect 89
Maria Luisa Rusconi, Raffaella Ricci and Francesca Morganti
Chapter 4 Neuropsychological Parameters Affecting the Academic
Aptitude Test (AAT) Achievement at the End of High School
in 1996 and their Impact on Job Status in 2002:
A Multifactorial Approach in a Follow-Up Study 103
Daniza M. Ivanovic, Hernán T. Pérez, Boris P. Leiva,
Nora S. Díaz, Bárbara D. Leyton, Atilio Aldo F. Almagià,
María Soledad C. Urrutia, Cristián G. Larraín, Paulina E. Olave,
Nélida B. Inzunza and Rodolfo M. Ivanovic
Chapter 5 Reliability and Magnitude of Laterality Effects in a Dichotic
Listening Task: Influence of the Testing Procedure 141
Daniel Voyer, Vanessa G. Boudreau and Aileen M. Russell
Chapter 6 Neuropsychology and Pseudo-Memories 163
Maarten J. V. Peters, Marko Jelicic and Harald Merckelbach
Chapter 7 The Beneficial Effects of Polyphenols in Age-Related Neurological
Disorders 185
Bastianetto Stéphane and Rémi Quirion
vi Joshua R. Dupri

Chapter 8 Current Issues in Response Bias During Neuropsychological

Assessment: Incomplete Effort to Malingering 199
Thomas M. Dunn
Index 219
 PREFACE

Neuropsychology is the study of brain-behavior relationships and examines such domains

of cognitive functioning as memory, attention, visual-perceptual abilities, language and
intellectual function. It is strongly scientific in its approach and shares an information
processing view of the mind with cognitive psychology and cognitive science. It is one of the
most eclectic of the psychological disciplines, overlapping at times with areas such as
neuroscience, philosophy (particularly philosophy of mind), neurology, psychiatry and
computer science (particularly by making use of artificial neural networks).
Executive function has been conceptualized as a set of interrelated cognitive processes
that are essential for regulation of cognition, behavior, and emotion. Hypothesized component
processes include cognitive flexibility, decision making, inhibitory control, planning and
organization, self-monitoring, as well as working memory. Historically, the neurobiological
basis of executive functions in humans was largely examined through the administration of
performance-based neuropsychological tests to patients with acquired focal brain lesions.
Such studies led to the widely held belief that the integrity of the frontal lobes is central to
executive functions. However, while lesion studies have been highly informative, they suffer
from a number of methodological weaknesses including the inability to investigate the
broader neural circuitry that contributes to executive functions. Over the past twenty five
years functional neuroimaging techniques, in particular positron emission tomography (PET)
and functional magnetic resonance imaging (fMRI), have been increasingly applied in the
effort to elucidate the neural circuitry subserving executive functions. In chapter I, the authors
review functional neuroimaging research addressing the neural substrates of executive
functions in healthy adults, and discuss applications of neuroimaging techniques to the
identification of the neural correlates of executive dysfunction in selected psychiatric and
neurological illnesses.
Available neurophysiological and neuroimaging studies support the view that attentional
processing is carried out by means of the reciprocal communications between distributed
neural networks of brain areas including frontal and prefrontal cortex, as well as the occipital-
temporal-parietal cortices. In chapter II, ERPs were recorded using a 32 channels whole-head
montage while volunteers performed conjoined selection tasks of spatial location and spatial
frequency. Target stimuli and distractors were presented within and without attended
locations. The data indicate that during spatially directed selective processing, the visual
system, although overall increasing processing activation for all stimuli falling within the
attended channel of attention, treats the relevant and irrelevant sources of information
viii Joshua R. Dupri

differently. They also suggest that this occurs not only within the relevant channel of
attention, but without it too. Indeed, the present findings suggest that, while enhancing
processing of frequency-relevant gratings with respect to frequency-irrelevant ones since the
earliest sensory level, as reflected by C1, at the attended location, the system seems also
actively to suppress the processing of the latter stimuli with respect to the former ones at the
neglected location, already starting at early sensory level, as reflected by P1. As for the
prefrontal cortex, ERPs data and their topography suggested that distinct sub-regions of the
latter might serve distinct attention functions. In fact, while dorsolateral regions would control
the orienting of attention toward relevant space locations, the left-hemisphere inferior
posterior prefrontal cortex (PFC) may suppress processing of distractors within an attended
channel. Conversely, the dorsal frontal-polar PFC might bust, although in different degrees,
the post-perceptual/semantic processing of relevant information by posterior brain areas
independent of location relevance.
Unilateral neglect refers to the failure, due to unilateral brain damage, in exploring
portion of space contralateral to the side of lesion. It is a largely shared opinion that the deficit
could be described with reference to specific spatial co-ordinate systems. Where neglect is
referring to the mid-sagittal plane of the patient’s body, it has been defined viewer–centred
(or egocentric) and within it a variety of components have been distinguished, such as
retinotopic-, head-, limb- and trunk-centred. Whereas, were patients show failures in
representing stimuli within the extra-personal domain, such as the surrounding ‘environment’
and the hand-reach visual space, disorder has been specifically defined “allocentric” neglect.
Several studies have compared the specific contribution of viewer and allocentric frames of
reference on neglect. Some evidence suggests a major contribution of the environment-
centred system on neglect. Other data support a viewer-centred predominance when the
environmental frame is derived exclusively from gravitational information without using
visual cues. In chapter III, the authors wished to learn how neglect allocation is affected by
the differential intervention of viewer, environment and array centered spatial representations.
Fifteen left-neglect patients were asked to search for visually presented targets. They
performed the searching either in an upright position or lying on their right side, in a position
orthogonal to the environment. This latter condition was meant to dissociate viewer-centered
and environment-centered frames. The visual display, within which stimuli were presented,
could be either aligned with the patient’s body or rotated of 90° counterclockwise. Finally,
they analyzed the effects of stimulus content on neglect severity. Numbers, letters, and
drawings of objects or animals were used as stimuli. Patients’ spatial neglect was found to be
allocated mainly with reference to the viewer’s body and, to a lesser degree, to the
environment. Moreover, the amount of neglect centered on the environment was found to be
enhanced by the alignment of the display with its vertical. The authors interpreted these data
as an evidence that environment-centered neglect may be partially due to a mental rotation of
viewer-based representations. This phenomenon would very likely be mediated by
environmental visual cues as well as the visual display within which patients are searching for
targets. Regarding stimulus material, patients manifested more severe neglect with figures,
improved with letters and did even better with numbers. This pattern of results is in line with
Weintraub and Mesulam’s (1988) finding of a less severe neglect with ‘verbal’ than with ‘non
verbal’ stimuli.
The aim of chapter IV was to determine the impact of neuropsychological parameters on
the academic aptitude test (AAT) achievement at the end of high school in 1996 when they
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 Preface ix

should be graduating from high school and on job status carried out six years later during
2002. From a representative sample of 1817 Chilean school-age children (mean age 18.0 ±
0.9 y) graduating from high school in 1996 in Chile’s Metropolitan Region, 96 were selected
with high (> 120 WAIS-R) and low IQ (< 100 WAIS-R) (1:1), from the high and low socio-
economic strata (SES) (1:1) and of both sexes (1:1). AAT scores for university admission
were obtained for 84 school-age children from University of Chile records and were divided
into two groups: high AAT (≥ median (Md)= score 631) and low AAT (< Md). IQ was
determined by the Wechsler Intelligence Scale for Adults (WAIS-R) and the Raven
Progressive Matrices Test in the school-age children and their parents. Scholastic
achievement (SA) was measured applying the standard Spanish language and mathematics
tests. SES was evaluated using Graffar's modified method. Nutritional status was assessed
through anthropometric measurements of weight and height to establish the body mass index
(BMI) according to Garrow; head circumference (HC) was compared with Tanner, Nellhaus,
Roche et al. and Ivanovic et al. tables and was expressed as Z score (Z-HC); body
composition parameters such as arm circumference-for-age, triceps skinfold-for-age, arm
muscle area-for-age and arm fat area-for-age were calculated using data from Frisancho.
Brain morphology was determined by magnetic resonance imaging (MRI). Job status was
expressed as: (1) jobless, (2) workers without further schooling, (3) students at institutes and
(4) students at universities. Statistical analysis included correlation and logistic regressions
using the Statistical Analysis System (SAS). Results showed that students with high AAT
score presented IQ, parental IQ, SA, brain volume (BV), Z-HC, maternal schooling, house-
hold head occupation and quality of housing significantly higher than their peers that
achieved the lowest AAT scores of whom 19% had suffered severe undernutrition in the first
year of life (Fisher p< 0.0054). However, logistic regression revealed that student IQ is the
best predictor of AAT score and the odds ratio value (1.252) implies that when the IQ score
increases by one point, the probability to obtain a high AAT score increases in 25.2%.
Students at universities presented AAT, child and maternal IQ, SA, BV, birth weight and
birth height, Z-HC and socio-economic conditions significantly higher than their peers from
the other three job statuses; however, AAT score at the end of high school was the best
predictor of the job status six years later (odds ratio value= 1.025) which indicates that when
AAT score increases by one point the probability for university admission and for university
graduation increases 2.5%. In a multifactorial approach, these results point out the importance
of neuropsychological parameters on children’s achievement for university admission and
future jobs.
In chapter V, two experiments investigated the reliability of dichotic listening tasks under
two conditions of administration: on tape or on a computer. Stimuli in both experiments were
words (bower, dower, power, and tower) pronounced with an emotional tone of anger,
happiness, neutrality, or sadness. Experiment 1 required participants to detect a target word in
each dichotic pair. Accordingly, 40 right-handed participants were randomly assigned to a
condition in which the dichotic word recognition task was completed twice either in a taped
version or directly on a computer. Experiment 2 involved the detection of a target emotion. A
different group of 40 right-handed participants completed the dichotic emotion recognition
task twice either on tape or on-computer. In Experiment 1, results showed a large right ear
advantage of similar magnitude and test-retest reliability on both versions of the task. In
Experiment 2, the laterality effect also had a similar reliability in both versions. However, the
magnitude of the observed left ear advantage (LEA) varied as a function of procedure.
x Joshua R. Dupri

Specifically, on computer, the LEA was significant only in females. Implications of these
findings for the use of computer tasks to measure functional lateralization in research and
applied settings are discussed. Factors responsible for variability in the results obtained in
Experiment 2 are also discussed.
Although the power of memory is evident in various daily life experiences (e.g., personal
history, knowledge of facts and concepts, and learning of complex skills), memory also has
its fallible side. Memory experiences are encoded as separate pieces of a puzzle. At retrieval,
this puzzle has to be reconstructed. This reconstruction makes our memory susceptible to
distortions. Inspired by ongoing discussions about recovered memories, researchers have
studied personality and social factors that make people vulnerable to memory distortions,
including pseudo-memories. Recent studies highlight the role of higher cognitive functions in
reconstructing our memories. Chapter VI aims to give an overview of the current state of
affairs linking executive functions to pseudo-memories. Evidence from aging, lesion, and
imaging studies (Positron Emission Tomography and functional Magnetic Resonance
Imaging) will be discussed. Studies conducted in our laboratory also suggest that suboptimal
inhibition, monitoring and working memory functions contribute to the development of
pseudo-memories. Explaining and identifying the neural basis of pseudo-memories can be
regarded as a promising and new domain within neuropsychology.
Polyphenols have received particular attention because of their possible beneficial health
effects in age-related neurological disorders. In support of this hypothesis, epidemiological
studies reported a lower incidence of stroke, Parkinson's disease and dementia in populations
that consume beverages or food (i.e. red wine, green tea, fruits, vegetables) enriched in
polyphenolic compounds. These findings concur with animal and in vitro studies indicating
that polyphenols derived from either beverages, fruits (e.g. catechins, resveratrol) or plant
extracts (i.e. blueberry, Ginkgo biloba and tea) displayed neuroprotective abilities. For
example, the authors’ studies and those obtained by other groups indicated that various
polyphenols derived from green tea and red wine protected cultured neuronal cells against
toxicity induced by free radicals and beta-amyloid (Aß) peptides, whose accumulation likely
play a deleterious role in age-related neurological disorders. These effects involved their well-
known antioxidant activities, but also their abilities to directly interact with Aß and to
modulate intracellular effectors and genes associated with cell death/survival. Chapter VII
overviews epidemiologic and pre-clinical studies that support the role of polyphenols in the
beneficial effects of diet in human.
The purpose of chapter VIII is to review the recent literature regarding the assessment of
biased responding, especially in the context of malingering. It makes the case that biased
responding is multifaceted and that clinicians need to be aware of a variety of issues
pertaining to effort. The different types of malingering are discussed, as well as the particular
situations when patients are likely to feign symptoms. The more common commercially-
available tests to assess effort are reviewed, with a discussion of their strengths and
weaknesses. Other means of assessing response bias are mentioned, including the use of
traditional neuropsychological assessments, as well as the issue of coached malingerers.
Finally, specific suggestions for the practicing neuropsychologist are given.
Preface xi
In: Focus on Neuropsychology Research ISBN 1-59454-779-3
Editor: Joshua R. Dupri, pp. 1-36 © 2006 Nova Science Publishers, Inc.

Chapter 1

NEURAL SUBSTRATES OF EXECUTIVE FUNCTIONS:

INSIGHTS FROM FUNCTIONAL NEUROIMAGING

Robert M. Roth,* John J. Randolph,

Nancy S. Koven, and Peter K. Isquith
Neuropsychology Program and Brain Imaging Laboratory
Department of Psychiatry; Dartmouth Medical School

ABSTRACT
Executive function has been conceptualized as a set of interrelated cognitive processes
that are essential for regulation of cognition, behavior, and emotion. Hypothesized
component processes include cognitive flexibility, decision making, inhibitory control,
planning and organization, self-monitoring, as well as working memory. Historically, the
neurobiological basis of executive functions in humans was largely examined through the
administration of performance-based neuropsychological tests to patients with acquired
focal brain lesions. Such studies led to the widely held belief that the integrity of the
frontal lobes is central to executive functions. However, while lesion studies have been
highly informative, they suffer from a number of methodological weaknesses including
the inability to investigate the broader neural circuitry that contributes to executive
functions. Over the past twenty five years functional neuroimaging techniques, in
particular positron emission tomography (PET) and functional magnetic resonance
imaging (fMRI), have been increasingly applied in the effort to elucidate the neural
circuitry subserving executive functions. In the present chapter, we review functional
neuroimaging research addressing the neural substrates of executive functions in healthy
adults, and discuss applications of neuroimaging techniques to the identification of the
neural correlates of executive dysfunction in selected psychiatric and neurological
illnesses.

*
Corresponding author: Robert M. Roth, Ph.D., Brain Imaging Laboratory, Dartmouth Medical School/DHMC, 1
Medical center Drive, Lebanon, NH, USA 03756-0001. Phone: 603-650-5824; Fax: 603-650-5842; E-Mail:
[email protected]
2 Robert M. Roth, John J. Randolph, Nancy S. Koven, and Peter K. Isquith

INTRODUCTION

The Executive Function Construct

Executive function may be conceptualized as a collection of interrelated higher level or

supervisory cognitive processes involved in the selection, initiation, execution and monitoring
of complex domain specific cognitive processes and motor responses, as well as aspects of
emotional processing. Stated more broadly, executive functions are concerned with the self-
regulation of cognition, behavior and emotion [1-3]. Executive functions have also been
conceptualized as exerting regulatory control over the basic, domain-specific
neuropsychological functions (e.g., language, visuospatial functions, memory, emotional
experience, motor skills) in the service of reaching intended goals.
Executive function is generally considered an umbrella construct that subsumes a
collection of related yet at least partially distinct processes that provide for intentional, goal-
directed problem solving, as well as organization and direction of cognitive activity,
emotional response, and overt behavior. However, the operational definition of executive
function and the specific cognitive processes subsumed under this construct have varied
somewhat among authors [4, 5, 6]. Theoretical concepts or models such as the supervisory
attention system [7], the central executive [8], or managerial knowledge units [9] have been
put forth in an effort to explain the structure and role of executive functions, as well as the
relationship between executive and other cognitive functions. Processes commonly regarded
as executive functions include the ability to initiate behaviors, inhibit competing actions or
stimuli, select relevant goals, plan and organize means to solve complex problems, shift
problem-solving strategies flexibly when necessary, regulate emotions, and monitor and
evaluate behavior [3, 6, 10, 11]. Working memory capacity, whereby information is actively
held “online” so that it may be manipulated and transformed in the service of planning and
guiding cognition and behavior, is also described as a key aspect of executive function [11,
12].

Developmental Course

The development and maturation of executive functions is considerably more prolonged

than that of other cognitive functions such as language [13-18], and differences in the
developmental course of specific executive functions may also be observed [19]. This
extended developmental course parallels the prolonged pattern of neurodevelopment of the
frontal lobes including growth in synaptic connections, myelination, and bioelectric signal
coherence [20-22]. The development of executive functions such as the self-regulation of
emotion and behavior begins in infancy [23] and continues into the preschool period [24-27]
through adolescence and early adulthood [28-30]. Decline in the use of executive functions in
later adulthood and old age is also observed, though change tends to be modest and is not
found for all executive functions nor for all healthy individuals [31-34]. As with many other
dimensions of psychological and neuropsychological functioning, the decline of executive
functions varies across individuals in terms of the age of onset, rate of decline, and actual
proficiency level at any given age. Several variables have been associated with age-associated
 Neural Substrates of Executive Functions: Insights from Functional Neuroimaging 3

decline in executive functions such as changes in the structural integrity of white matter
pathways, neurodegenerative changes in frontal lobe grey matter, as well as genetic factors
[35-38].

Historical Perspective on the Neuroanatomical Basis

Executive functions were thought to be subserved primarily, if not solely, by the frontal
lobe since at least the 19th century when Phineas Gage demonstrated dramatic changes in self-
regulatory function after a dynamite tamping rod was propelled through his frontal lobe [39].
Further, more substantial evidence of executive deficits associated with frontal lobe damage
came about in the 1940’s and 1950’s in the context of war related brain injury [e.g., 40].
However, a wealth of evidence has accumulated indicating that while damage to the frontal
lobes can indeed result in significant executive dysfunction [41-43], these complex, higher order
processes are not solely a product of frontal activity. Executive dysfunction may also be seen in
cases of injury or pathology to non-frontal cortical as well as subcortical regions [e.g., 44, 45,
46]. Thus, the historical view of executive functions as subserved solely by the frontal lobes is an
oversimplification of the complex organization of the brain, leading some authors to caution
against attributing executive deficits to frontal lobe pathology except at a hypothesis generating
level [47]. Nevertheless, an understanding of the frontal region of the brain is important in any
discussion of the executive functions. The frontal lobes are richly and reciprocally interconnected
through numerous neuroanatomical pathways with other cortical and subcortical regions of the
brain such as the limbic (motivational/mnemonic) system, the reticular activating (arousal)
system, the posterior association cortex (perceptual/cognitive processes and knowledge base),
and the motor (action) regions [11, 48-51]. This broad connectivity underlies the regulatory
control that the frontal brain systems exert over the posterior cortical and subcortical systems.

FUNCTIONAL NEUROIMAGING OF EXECUTIVE FUNCTIONS

Brain-behavior relationships underlying executive functions remain elusive given the
challenges inherent in traditional neuropsychological methods. Recent advances in
technology, specifically functional neuroimaging, are beginning to provide a more detailed
picture of these relationships. In the present chapter we provide a brief description of the
functional neuroimaging techniques most commonly employed to study executive functions.
We then selectively review functional neuroimaging studies investigating the neural
substrates of executive functions in healthy humans. Specifically, we discuss cognitive
flexibility, decision making, planning, self-monitoring, response inhibition, organization, and
working memory. We then illustrate the utility of functional neuroimaging for studying
executive dysfunction in selected psychiatric and neurological disorders.
4 Robert M. Roth, John J. Randolph, Nancy S. Koven, and Peter K. Isquith

Some Functional Neuroimaging Basics

By far the most commonly employed functional neuroimaging technologies for studying
the neural substrates of executive functions are positron emission tomography (PET) and
functional magnetic resonance imaging (fMRI). Briefly, PET is a scanning technique that
involves visualizing brain structure and function through the detection of small amounts of
radioactively labeled compounds that are injected (e.g., water or glucose) or inhaled (e.g.,
oxygen) shortly before or during scans/tasks. As the radioactive atoms in the compound decay
they release small positively charged particles called positrons. The positrons subsequently
collide with negatively charged electrons resulting in the creation of a pair of particles of light
called photons that are picked up by detectors in the scanner. Brain regions where the labeled
compound has accumulated, and thus it is believed where activated related to a given task was
greatest, are identified through computer programs designed to analyze the information and to
generate three-dimensional, cross-sectional images that represent the biological activity where
the compound has accumulated. PET is increasingly used in combination with computerized
tomography and structural magnetic resonance imaging in order to provide more accurate
localization of activation.
fMRI is based on the observation that hemodynamic activity is closely related to neural
activity [52-54]. About four to six seconds after an electrical burst of neural activity can be
detected in a region, a hemodynamic response occurs as active neurons use up oxygen from
oxygen-rich blood that is infused into the region. fMRI takes advantage of the different
magnetic properties of oxygenated and deoxygenated hemoglobin, each of which gives off a
slightly different signal. The MRI scanner detects this slight difference, which is known as the
blood oxygen level dependent (BOLD) contrast, and the information is used to generate three-
dimensional images.
The two most commonly employed task designs in functional neuroimaging are referred
to as blocked and event-related designs. Blocked designs involve the presentation of different
task conditions in separate trains of stimuli called epochs. Conditions of interest may involve
one or more cognitive, sensory or motor tasks. These conditions are then compared to control
conditions in order to facilitate identification of brain regions specifically related to the task
condition of interest. Control tasks are usually quite similar to the condition of interest, but
exclude demands on the process of interest. For example, an imaging study of response
inhibition may use a go/nogo task that has a condition in which 20% of stimuli are nogo and
80% of stimuli are go (i.e., inhibit condition), while another condition may involve only the
presentation of only go stimuli without any demand for inhibition (i.e., control condition).
Activation related to inhibition is found by subtracting activation to the 100% go condition
from that found for the 20% nogo condition. In event-related designs, stimuli of interest are
presented in random or pseudorandom order within the same epoch. Using the go/nogo
example above, an event-related design could present the nogo and go stimuli in a random
order within the same train of stimuli (e.g., go - go - nogo - go - nogo), and thus a direct
comparison can be made between activations to the nogo and go stimuli.
Event-related designs have several advantages over blocked designs, such as allowing for
greater control over the potential impact of fluctuations in arousal, attention, and mood over
the course of an experiment. However, under some circumstances, blocked designs can be
more powerful in detecting effects of interest. Several excellent texts are available for readers
 Neural Substrates of Executive Functions: Insights from Functional Neuroimaging 5

interested in more detailed explanations of these neuroimaging methods and their limitations,
as well as data analytic techniques [55-57].

Cognitive Flexibility

The ability to think flexibly (also referred to as set shifting) is considered an integral
component of normal executive functioning. Deficits in cognitive flexibility are often seen in
patients with frontal lobe damage, especially when the dorsolateral cortex is involved. Such
patients are generally slower to switch between two tasks or concepts, even when they are
able to perform each of the two tasks accurately, and have difficulty benefiting from feedback
indicating that their current pattern of responding is no longer correct [58].
Trail making tests are often administered to assess cognitive flexibility, and require the
respondent to alternate cognitively and motorically between sets of numbers and letters. To
examine brain regions activated during this kind of task-switching, Moll and colleagues used
a verbal adaptation of a traditional paper-and-pencil trail-making test in an fMRI paradigm,
reporting critical roles for the dorsolateral and medial frontal cortex, as well as the
intraparietal sulcus [59]. Gurd et al. reported prominent posterior parietal fMRI activation
when participants were required to produce triads of words from three semantic categories
[60]. This pattern of activation underscores the role of widespread networks involved in set-
shifting that extend beyond the frontal lobes and require coordination with posterior areas.
Another popular test of executive functioning, the Wisconsin Card Sorting Test (WCST),
has also been examined using neuroimaging techniques to uncover frontal lobe activation
correlates. Early PET and fMRI studies reported dorsolateral activation during WCST
performance [61, 62], specifically bilateral inferior frontal sulci [63]. However, without
further investigation, it is unclear whether this activation represents set-shifting, working
memory, error recognition, or some combination of these executive processes, as the WCST
is multidimensional in its task demands. To address this, Konishi and colleagues used event-
related fMRI to compare patterns of brain activation during a set-shifting task condition of the
WCST to those found during a working memory condition [64]. Results showed that
activation associated with set-shifting overlapped in the same region of the inferior frontal
sulci in which activation was found to be associated with working memory, suggesting that
set-shifting and working memory act in concert in the same areas of the frontal cortex. A
recent study that further decomposed elements of set-shifting found that right frontal regions
were responsive to receipt of negative feedback during the WCST administration whereas
analogous left frontal regions were activated during updating of behavior [65]. Given this
evidence for hemispheric specialization within the set-shifting domain, it appears that
interhemispheric transfer within frontal regions is the key for successful completion of this
task. Monchi et al. also reported dissociations between regions activated to different aspects
of the WCST [66]. They found that the mid-dorsolateral frontal cortex was activated when
participants received either positive or negative feedback likely related to working memory
demands; a cortical-basal ganglia circuit involving the mid-ventrolateral frontal cortex,
caudate nucleus, and thalamus increased activity only for negative feedback, related to the
need to shift response set; and posterior frontal cortex activity to both responding on the task
and feedback, interpreted as suggesting a role in the association of specific stimuli to actions.
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