i

Training Attention through Intensive Meditation:

Improvements in Sustained Performance and Response Inhibition

By

KATHERINE ANNE MACLEAN
A.B. (Dartmouth College) 2003
M.A. (University of California, Davis) 2006

DISSERTATION

Submitted in partial satisfaction of the requirements for the degree of

DOCTOR OF PHILOSOPHY

in

Psychology

in the

OFFICE OF GRADUATE STUDIES

of the

UNIVERSITY OF CALIFORNIA

DAVIS

Approved:

Dr. George R. Mangun

Dr. Clifford D. Saron

Dr. Steven J. Luck

Dr. Phillip R. Shaver

Committee in Charge

2009




UMI Number: 3379639






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ii









Training Attention through Intensive Meditation:
Improvements in Sustained Performance and Response Inhibition


DISSERTATION

KATHERINE ANNE MACLEAN

2009







iii

Acknowledgments
Many people have guided me along my journey through graduate school. The
completion of this dissertation would not have been possible without their thoughtful
contributions, inspiring ideas, and friendship. I would like to start by thanking my
advisors, Ron Mangun and Clifford Saron, for allowing me the freedom to pursue my
ideas and challenging me to strive for excellence. Ron You have supported me
wholeheartedly from the very beginning, and I will always look up to you as an example
of how to be a successful scientist while also enjoying life. Cliff You are truly a fearless
leader and have shown me that the wildest dreams are attainable, through hard work and
a little mystical charm. I am lucky to have had both of you as mentors. I would also like
to thank the members of my dissertation committee: Phillip Shaver, for bravely
confronting errors in writing style and grammar and diligently passing on your wisdom to
me at every opportunity; and Steven Luck, for always being available at a moments
notice to provide thoughtful feedback and suggestions. I would also like to thank other
faculty members in the Department of Psychology who have contributed to my research
and provided valuable guidance: Ewa Wojciulik, David Whitney, Petr Janata, Robert
Post, and Debra Long. Finally, I would like to thank my undergraduate advisor, Yale
Cohen, for exercising the right balance of skepticism and encouragement that enabled me
to pursue my craziest career goals.
I arrived at UC Davis just as a courageous, once-in-a-lifetime project was about to
begin. The Shamatha Project was a large-scale, collaborative research endeavor aimed at
understanding the longitudinal effects of intensive meditation training on behavioral,
psychological, and neural functioning. This is just the sort of project that is usually
iv

impossible to undertake as a graduate student. And yet, with the support of my committee
and other faculty members, I successfully developed a dissertation that would be
embedded within the Shamatha Project. The findings presented here reflect the tireless
work of many collaborators, and I extend my deepest gratitude to all of them for making
this dissertation possible. First, I would like to thank the team of researchers who endured
the stress of living in close (quiet) quarters while collecting data for 12 hours a day for
many months: Stephen Aichele, Anthony Zanesco, David Bridwell, Tonya Jacobs, and
Brandon King. I would also like to thank Baljinder Sahdra, for a truly pleasant and
rewarding collaborative writing experience; Erika Rosenberg, for thoughtful
contributions and compassionate support; and Emilio Ferrer, for invaluable statistical
advice and insights. Finally, I would like to thank Alan Wallace, meditation teacher
extraordinaire, for introducing me to meditation and inspiring me to push the envelope in
my science career. Working on the Shamatha Project has taught me that great science is
produced when creative, intelligent, hard-working people come together to bravely
attempt to achieve what others deem impossible.
I would like to thank my laboratory mates, past and present: Mita Puri, Santani Teng,
Evelijne Bekker, Sean Fannon, Bong Walsh, Joy Geng, Risa Sawaki, Sharon Corina,
Jesse Bengson, Andre Bastos, and Ali Mazaheri. Thank you also to all of the faculty
members and researchers at the Center for Mind and Brain for helping create a collegial,
productive, and fun atmosphere to work in. Also, a huge thank you to the administrative
and support staff at the Center for Mind and Brain: Noelle Blalock, Carmina Caselli,
Patricia Schuler, Jeremy Phillips, and Chris Brick. This dissertation literally would not
have been completed without your help.
v

I would not have made it through graduate school without the love and support of my
friends and family. Thank you to Erin Sullivan and Jason Haberman, who have been with
me every step of the way since we arrived in Davis as first-year psychology graduate
students. Thank you to my parents, Julie and Richard, for raising me to always
confidently and passionately pursue my dreams; to my sister, Rebecca, for inspiring me
to be competitive and strong; and to my brother, Edward, for always making me smile.
Finally, thank you to my partner, John, for showing unconditional love and amazing
patience, for following me to the ends of the earth (or, at least to a remote meditation
retreat center in the mountains of Colorado), for listening to me rant about data and
theories and new ideas, and above all, for being a great human being. John You are the
love of my life, and I am lucky to have found someone to share my dreams.












vi

Abstract
The ability to focus attention underlies success on many everyday tasks, but voluntary
attention cannot be sustained indefinitely. In the laboratory, perceptual sensitivity
declines with increasing time on task, a phenomenon called the vigilance decrement. The
aim of this dissertation was to demonstrate that vigilance that is, performance over
extended periods of time - can be improved in healthy adults.

The experiments in Chapter 1 describe the effects of attentional cues on performance in
two related vigilance tasks: a sustained attention task that required responses to rare
targets occurring in a sequence of non-targets, and a sustained response inhibition task
that required responses to the frequent non-targets and withholding of responses to the
rare targets. In the sustained attention task, sudden-onset cues presented immediately
before each stimulus improved overall perceptual sensitivity while predictable timing
cues attenuated the decline in perceptual sensitivity over time. Attentional cues did not
improve response inhibition performance. Finally, performance on the most challenging
versions of the sustained attention and response inhibition tasks did not improve with
repeated task practice, indicating the suitability of these tasks as outcome measures of the
effects of attention training.

The experiments in Chapters 3 and 4 describe the effects of intensive meditation training
on vigilance. Training (~5 hours/day for 3 months) consisted of a meditation practice that
entailed learning to regulate and control voluntary attention by sustaining attention
selectively on a chosen stimulus (e.g., the breath). Participants were randomly assigned
vii

either to receive training first (N = 30) or to serve as wait-list controls and receive
training second (N = 30). Training produced improvements in visual discrimination
which led to increases in perceptual sensitivity and reductions in the vigilance decrement
during the sustained attention task. Training also produced increases in response
inhibition accuracy without concomitant slowing of reaction time. Finally, improvements
in discrimination and response inhibition were maintained several months after the
completion of formal training, indicating enduring changes in behavior.
















viii

Table of Contents

Chapter 1 Introduction 1

Chapter 2 Benefits of Attentional Cues during Vigilance:
A Comparison of Sustained Attention and
Response Inhibition 13

Chapter 3 Improvements in Perception and Sustained Attention
with Intensive Meditation Training 60

Chapter 4 Training-related Changes in Sustained Response
Inhibition: The Influence of Age and Response Speed
on Improvements in Accuracy 85

Chapter 5 Conclusion 110




1











CHAPTER 1

INTRODUCTION










2

The ability to control the focus of attention while resisting distraction is a central
feature of adaptive human behavior. Attention can be voluntarily guided in line with
personal goals, prior knowledge, or explicit instructions to improve the accuracy and
efficiency of behavior (referred to as endogenous or voluntary attention). Attention can
also be automatically drawn to stimuli and actions because of momentary salience or
well-established habits (referred to as exogenous attention). Attentional control involves
exerting voluntary, goal-directed control over which stimuli, thoughts, emotions, and
actions are selected despite automatic or habitual tendencies (see reviews in Fan &
Posner, 2004; Posner & Petersen, 1990).
Although attention can be successfully maintained over relatively short time intervals
(e.g., the 1-second interval between a cue and a subsequent stimulus; Posner, 1980;
Posner & Cohen, 1984), voluntary attention cannot be sustained indefinitely. Decades of
research on sustained attention has consistently demonstrated declines in accuracy with
increasing time on task, a phenomenon called the vigilance decrement (see Parasuraman,
1986 for review). Although traditional sustained attention tasks required observers to
respond to rare targets, recent research has shown that performance also declines on tasks
that require response inhibition (i.e., responding to frequent non-targets and withholding
responses to rare targets; Grier et al., 2003; Helton et al., 2005). Both sustained target
discrimination and sustained response inhibition place high demands on the attentional
control system to maintain performance over extended periods. The overarching goal of
this dissertation is to investigate how vigilance - that is, performance over extended
periods of time - can be improved in healthy adults.
3

In this chapter, I will first provide an overview of the major theories of vigilance and
outline the task factors that have been shown to modulate the vigilance decrement. Next,
I will discuss results from previous training studies and introduce meditation training as a
technique for improving sustained voluntary attention and attentional control. At the end
of the chapter, I will introduce the series of experiments that were carried out to
investigate (1) whether attentional cues could improve vigilance performance in
untrained adults and (2) whether improvements in vigilance could be achieved with
intensive meditation training.
Theories of Vigilance
Most vigilance tasks are variations of the original Mackworth Clock Test
(Mackworth, 1948) used to test vigilance performance in radar operators. In the Clock
Test, observers monitored a pointer moving in single-step increments across a blank
clock face for periods up to 2 hours and were instructed to indicate when they detected a
rare double jump in the movement of the pointer. Current vigilance tasks share several
core design elements of the Clock Test in that they require subjects to monitor the
sequential display of neutral, non-target stimulus events over a long period of time and
initiate responses to rare target events occurring unpredictably. Typically, performance
declines over time, with the greatest decrement in correct detections occurring within the
first 15-20 minutes (Davies & Tune, 1969). Of particular interest are those tasks that
produce a decline in perceptual sensitivity (d' or A'), indicating that subjects are actually
detecting fewer targets over time, not simply adopting a more conservative response
criterion (D. M. Green & Swets, 1966).
4

Since Mackworths (1948) initial investigation, a multitude of experiments have
contributed to a better understanding of why humans are not very good at maintaining
accurate performance for long periods of time. Several theoretical models have been
proposed to explain the classic vigilance decrement, including the arousal model
(Frankmann & Adams, 1962), which views the decrement as a consequence of reductions
in general alertness over time, and the mindlessness model (Manly, Robertson, Galloway,
& Hawkins, 1999; Robertson, Manly, Andrade, Baddeley, & Yiend, 1997), which views
the decrement as the result of failures of a supervisory attention system to appropriately
direct awareness toward a relatively boring, undemanding task. Experimental findings
seem to challenge both of these theories and instead support an attentional resource
model (Davies & Parasuraman, 1982) as the best explanation of performance decrements.
Recent studies have used self-report measures in combination with vigilance tasks to
test key predictions of these competing theories and to more fully characterize sustained
attention as effortful and resource-demanding (Warm, Parasuraman, & Matthews, 2008).
For example, adding irrelevant stimuli to reduce task monotony (i.e., increase overall
arousal) does not improve vigilance performance (Smit, Eling, & Coenen, 2004b), in
contradiction to the arousal theory of vigilance. With respect to the mindlessness theory,
studies have shown that observers find vigilance tasks to be effortful rather than
undemanding (Grier et al., 2003; Szalma et al., 2004), that performance is influenced by
implicit stimulus patterns during the task (i.e., observers do not withdraw awareness from
the task; Helton et al., 2005), and that self-reported mind wandering (e.g., task-unrelated
thought) is not related to the vigilance decrement (Helton & Warm, 2008).
5

In addition, brain imaging studies offer direct evidence for the models assumption
that vigilance relies on attentional resources. Early studies suggested reductions in
cerebral blood flow in right frontal cortex with increasing time on task (Paus et al., 1997).
More recently, Hitchcock and colleagues (2003) showed that these reductions were
directly related to the extent of the vigilance decrement, and that the change in cerebral
blood flow was specific to task factors that had previously been interpreted as increasing
resource demands (e.g., difficult target discrimination). Furthermore, recent
neuroimaging and electrophysiological investigations of sustained attention suggest a link
between pre-stimulus patterns of brain activity and subsequent attention failures. For
example, reductions in brain activity in attentional control regions (prefrontal and parietal
cortex) predict errors during sustained attention (Weissman, Roberts, Visscher, &
Woldorff, 2006). Changes in brain activity are also observed in regions that receive
projections from attentional control areas. Specifically, increases in the amplitude of the
ongoing electroencephalograph in specific frequency bands related to visual and motor
processing predict lapses in attention during sustained response inhibition (Mazaheri,
Nieuwenhuis, van Dijk, & Jensen, in press). Together these findings indicate that ongoing
brain activity in attentional control regions and in regions that control stimulus and
response processing underlies successful sustained performance.
Factors Affecting Sustained Performance
A combination of experimental (e.g., Parasuraman, 1979) and meta-analytic
approaches (e.g., See, Howe, Warm, & Dember, 1995) have been used to identify the key
factors that increase resource demands and thus promote declines in perceptual sensitivity
during vigilance. Event rate (i.e., the rate of stimulus events that must be inspected to
6

detect the presence of the rare target event) has been judged to be one of the prepotent
parameters that affects vigilance performance, with high event rate (> 24 events per
minute, Parasuraman & Davies, 1977) increasing the resource demands of the task and
thereby increasing the perceptual sensitivity decrement (Davies & Parasuraman, 1982;
Parasuraman & Davies, 1977; See et al., 1995; Warm & Jerison, 1984). Perceptual
sensitivity also declines more reliably in tasks in which the target discrimination process
loads memory (Parasuraman, 1979). For example, when the currently visible stimulus in
a series of successive presentations is compared to a standard, non-target held in memory
(e.g., detecting a short line target occurring in a sequence of long line non-targets),
observers perform more poorly over time compared to performance on tasks that require
a comparative judgment based only on the currently visible stimuli (e.g., detecting a line
length difference between two lines presented at the same time). Finally, vigilance
decrements are large and reliable when target discrimination is difficult. Parasuraman and
Mouloua (1987) demonstrated that targets that are initially perceptually difficult to
discriminate from non-targets become more difficult to detect over time than targets that
are easy to discriminate, regardless of working memory load. Even tasks with numeric
targets, which do not consistently produce decrements in healthy subjects because they
are easy to discriminate, promote declines in perceptual sensitivity if the targets are
highly degraded and thus rendered more difficult to discriminate from non-target
numbers (Nuechterlein, Parasuraman, & Jiang, 1983).
In addition to event rate, target discriminability, and working memory load, other
factors contribute to the overall resource demand. In a meta-analysis of vigilance studies,
See and colleagues (1995) found that several task factors, such as sensory modality,
7

signal regularity, and spatial uncertainty did not independently predict the extent of the
vigilance decrement but were correlated with the overall level of sensitivity during
vigilance. Overall sensitivity in turn significantly predicted the extent of the vigilance
decrement: When overall sensitivity is low, vigilance decrements are large. Thus, these
authors proposed that the average level of sensitivity might well be viewed as an
index of total task demand brought about by the combination of task parameters (p.
243). Together, these findings demonstrate that the vigilance decrement is modulated by
the resource demands associated with task performance.
Finally, studies that have directly compared performance on perceptually demanding
tasks with both response initiation and inhibition requirements have found similar
decrements in accurate performance over time, as well as similar estimates of stress and
effort between the two types of responses (Grier et al., 2003; Helton et al., 2005). These
findings indicate that specific response requirements are not major determinants of the
vigilance decrement. Rather, both sustained target discrimination and sustained response
inhibition place high demands on resources needed for attentional and behavioral control.
Training Sustained Attention
In many perceptual, cognitive, and motor domains, skill learning is often limited to
the training task. That is, individuals show improved task performance with repetitive
practice but show no improvements on related tasks that have not been practiced but
presumably recruit similar skills. There has been growing research interest in identifying
training procedures that lead to generalizable improvements in cognition (see review in
C. S. Green & Bavelier, 2008). For example, computer-based training programs have
been shown to improve working memory capacity and non-verbal intelligence (Jaeggi,
8

Buschkuehl, Jonides, & Perrig, 2008; Olesen, Westerberg, & Klingberg, 2004). Likewise,
more naturalistic training regimens, such as playing action video games (C. S. Green &
Bavelier, 2003, 2006) or practicing a musical instrument (Graziano, Peterson, & Shaw,
1999) can improve performance on untrained tasks, suggesting the transfer of learned
skills.
Most studies that have attempted to improve vigilance in healthy individuals have
examined the effects of repetitive task practice or performance feedback (see review in
Davies & Parasuraman, 1982). Although moderate practice (1- 4 hours) of perceptually
demanding vigilance tasks does not improve performance (e.g., Nuechterlein et al., 1983;
Williams, 1986), extensive task practice (e.g., 20 sessions of a 30-minute vigilance task)
has been shown to increase overall accuracy and reduce the vigilance decrement in young
and elderly adults (Parasuraman & Giambra, 1991). But even this extensive practice did
not completely eliminate the vigilance decrement in all individuals who underwent
training. Moreover, task-specific improvements following repetitive practice do not
inform us about the possibility for more general improvements in vigilance. A recent
study suggests that learning to self-regulate arousal or alertness (as measured by electrical
skin conductance) can improve overall accuracy during a sustained response inhibition
task (Self Alert Training, O'Connell et al., 2008). However, participants in this study
made few errors and did not show performance decrements over time before training.
Thus, it is unclear whether training could improve performance over extended periods of
time on demanding vigilance tasks.
In contrast to the relative paucity of psychological research on attention training,
experts in the Buddhist contemplative tradition have long recognized the possibility of
9

improving attention through mental training. Historical accounts (see translations in
(Buddhaghosa, 1979; Tsong-kha-pa, 2002) describe Shamatha as a class of meditation
practices specifically designed to improve sustained attention. Shamatha involves the
repeated practice of maintaining voluntary attention on a chosen stimulus or event, such
as the tactile sensations of the breath (Wallace, 1999). During training, practitioners aim
to develop attentional stability (maintaining attentional focus on the chosen stimulus) and
perceptual vividness (enhancing the detail with which the attended stimulus is perceived).
While attention is sustained in this manner, introspection is used to recognize when
attention has wandered and guide attention back to the chosen stimulus. In this way,
practitioners develop general (i.e., non-stimulus-specific) skill in regulating and
controlling voluntary attention. In particular, attentional control related to response
inhibition is cultivated as practitioners learn to resist the automatic tendency to attend to
and engage with passing thoughts or momentary emotions. Historical texts note that
response inhibition is not used to suppress normal thoughts and emotions from arising,
but rather to refrain from (mentally) pushing away undesirable thoughts and emotions or
clinging to desirable ones (Lingpa, referenced in Wallace, 2006); (Padmasambhava,
1997). Thus, the practice entails using self-monitoring to control attention and behavior.
This metacognitive or meta-attentive aspect of training may underlie the transfer of these
skills to other domains and daily life activities (Wallace & Shapiro, 2006).
Overview of Experiments
In the following chapters, I present results from a series of experiments that
investigated how attentional cues (Chapter 2) and meditation training (Chapters 3 and 4)
modulate performance over extended periods of time. In Chapter 2, I describe three
10

behavioral experiments that were designed to examine the benefits of exogenous and
endogenous attentional cues during vigilance. In the first experiment, participants were
presented with sudden-onset luminance changes that were designed to capture attention
exogenously and thus improve perception. We tested the effect of exogenous cues on
performance on two related vigilance tasks that differed in response requirements: The
sustained attention task required responses to rare (10%) short lines (targets) while the
response inhibition task required responses to frequent (90%) long lines (non-targets) and
withholding of responses to the short target lines. In both tasks, line stimuli were
presented at a constant rate (interstimulus interval [ISI] = 1.85 seconds). The results
demonstrated that exogenous cues reduced the vigilance decrement in the sustained
attention task but not in the response inhibition task. In a second experiment, I explored
the influence of predictable timing on performance in the sustained attention task by
varying the ISI. I reasoned that the constant ISI in Experiment 1 provided information
about when to pay attention, thus acting as an endogenous attention cue. The results
showed that exogenous cues improved overall accuracy when the ISI was variable, but
did not reduce the vigilance decrement. This result suggested a strong interaction
between exogenous and endogenous cues during sustained attention. In addition, a direct
comparison of the results from Experiment 1 (constant ISI) and Experiment 2 (variable
ISI) showed that predictable timing independently reduced the vigilance decrement.
Finally, in a third experiment, I explored the stability of task performance across repeated
testing sessions using the versions of the sustained attention and response inhibition tasks
that produced the largest vigilance decrements in Experiment 1. The results from this
experiment showed that performance did not improve in either task with simple practice
11

and thus indicated that these tasks were appropriate to use as outcome measures in the
subsequent training studies.
Next, I investigated the influence of intensive meditation training on performance of
the sustained attention task (Chapter 3) and the response inhibition task (Chapter 4). Each
chapter presents the results from two studies: The first study was a between-groups
comparison of participants who underwent three months of intensive meditation training
(retreat participants) and participants who underwent testing at the same time points
(wait-list control participants). The second study was a within-subjects assessment of the
wait-list control participants when they subsequently underwent the same meditation
training approximately three months after the end of the first training period.
Performance was measured at three testing points during each training period (pre, mid,
and post) using the versions of the sustained attention and response inhibition tasks that
reliably produced significant performance decrements in untrained participants (Chapter
1, Experiment 3).
In Chapter 3, I present data demonstrating training-related improvements in target
perception that led to improved performance on the sustained attention task. In the first
study, target parameters were set according to individual threshold at each assessment.
Because retreat participants showed improvements in threshold, more difficult targets
were presented as training progressed. Thus, there were no observed improvements in
vigilance. In the second study, target parameters were held constant for each individual,
and increases in average sensitivity and reductions in the vigilance decrement were
observed. The results suggest that meditation training leads to improved perception which
12

reduces the resource demand associated with difficult target discrimination and thus
improves vigilance.
In Chapter 4, I present data showing training-related improvements in accuracy on the
response inhibition task. Participants in both groups showed increases in overall response
inhibition accuracy during their respective training period. I also investigated the
influence of response speed, age, and previous meditation experience on response
inhibition accuracy. Although older participants and participants with more previous
meditation experience had slower reaction times than younger participants and
participants with less previous meditation experience, respectively, reaction times did not
change with training and did not influence the observed changes in accuracy. Controlling
for demographic factors and reaction time also did not change the results. Thus, these
findings suggest replicable and robust training-related improvements in response
inhibition accuracy.
Finally, performance on a subset of the tasks was assessed at a follow-up that was
conducted approximately five months after the completion of training. Improvements in
discrimination (Chapter 3) and response inhibition accuracy (Chapter 4) were maintained
at the follow-up assessment, indicating long-term stability in training-related behavioral
changes. Moreover, discrimination performance was correlated with the amount of
continued daily meditation, suggesting a link between the maintenance of training-related
improvements and regular but less intensive meditation practice.



13










CHAPTER 2

BENEFITS OF ATTENTIONAL CUES DURING VIGILANCE:
A COMPARISON OF SUSTAINED ATTENTION AND RESPONSE INHIBITION









14

Goal-driven attention is referred to as top-down or endogenous attention, whereas
stimulus-driven attention is referred to as bottom-up or exogenous attention, being driven
by external events (e.g., Posner & Cohen, 1984). Allocating attention over short time
periods can be referred to as phasic orienting (Posner, 1980), while maintaining attention
over longer time periods is referred to as sustained attention, or vigilance (Parasuraman,
1986). The interactions among these varieties of attention come into to play in our
momentary experience as our goals and intentions compete with the attractions of the
environment over milliseconds, seconds and minutes (for review see Ruz & Lupianez,
2002). The primary focus of this initial set of experiments was to investigate the effects
of endogenous and exogenous attention during vigilance.
Endogenous attention can be engaged explicitly to select one thing over another (e.g.,
attend to one location and ignore others, Posner, 1980), as well as to follow a particular
instructional set or general rule (e.g., attend to and report the color of the presented word,
Stroop, 1935). Endogenous attention can be also guided by spatiotemporal regularities
that are learned implicitly over time (Large & Jones, 1999). In contrast to both explicit
and implicit forms of endogenous attention, exogenous attention refers to effects that are
generated externally by the physical properties of stimuli (e.g., brightness, color, shape).
Exogenous attention allows novel or salient information (e.g., a sudden change in
luminance, Jonides & Yantis, 1988) to transiently interrupt goal-directed behavior. The
dynamic and flexible interplay between endogenous and exogenous attention has been
well-studied in relation to visuospatial selective attention over short time periods (Folk,
Leber, & Egeth, 2002; Folk, Remington, & Johnston, 1992), and this interaction
continues to be characterized in electrophysiological and brain imaging studies (e.g.,
15

Hopfinger & West, 2006; Serences et al., 2005) that seek to delineate functionally the
brain networks supporting these attention systems. Current theories posit that endogenous
and exogenous attention systems interact and compete over short time periods to guide
behavior (Corbetta, Patel, & Shulman, 2008; Corbetta & Shulman, 2002), with the
outcome of that interaction resulting in real-time prioritization of attentional focus and
stimulus processing based on both endogenous and exogenous factors (Fecteau & Munoz,
2006; Gottlieb, 2007).
However, less is known about how exogenous and endogenous attention systems
interact over longer periods of time. Among the major theories of vigilance, the resource
model (Davies & Parasuraman, 1982) proposes that the drop-off in performance accuracy
over time the vigilance decrement - is a result of the exhaustion of information
processing resources that are not replenished over time. In support of this model, task
factors that increase processing demands also increase the vigilance decrement. For
example, vigilance decrements are large and reliable when stimulus events occur at a
high rate (~ 30 events/minute), when targets are difficult to discriminate from non-
targets, and when the task loads working memory (Parasuraman, 1979; Parasuraman &
Mouloua, 1987; See et al., 1995). Moreover, vigilance is deemed to be effortful and
stressful (Grier et al., 2003; Szalma et al., 2004; Warm et al., 2008). More recently,
researchers have demonstrated that the vigilance decrement is not specific to tasks that
require rare responses in that accuracy also declines over time in tasks that require
response inhibition (Grier et al., 2003; Helton et al., 2005). Sustained response inhibition
tasks combine the effort associated with normal vigilance, with the challenge of
inhibiting impulsive behavioral responses (Helton, 2009; Helton, Kern, & Walker, 2009).
16

In general, most vigilance research has focused on describing the task factors, such as
stimulus and response features, that contribute to worse performance over time (see
Chapter 1).
Assuming that attentional resources are limited and vigilance requires attentional
effort, it is important to determine whether endogenous and/or exogenous attention
manipulations might improve vigilance performance, and how these effects are related to
the response requirements of the task. Although benefits from endogenous and exogenous
attention cues during vigilance have been demonstrated, the interaction between
exogenous attention and endogenous attention during vigilance remains unclear,
primarily owing to the fact that no investigators have examined effects of exogenous and
endogenous cuing in the same study. Thus, this was the purpose of the current study. I
directly investigated the interaction between endogenous and exogenous attention during
vigilance in order to better understand how manipulations of both forms of attention
would affect the performance decrement. Specifically, I investigated (1) whether simple,
sudden-onset visual events could improve sustained performance (exogenous attention
manipulation), (2) whether benefits due to sudden onsets would interact with fluctuations
in attention as a function of the predictability of stimulus timing (endogenous attention
manipulation), and (3) whether performance on the most resource-demanding versions of
the vigilance tasks would change with simple task practice.
Perceptual Benefits of Endogenous Spatial Attention
Many studies outside of the vigilance domain have investigated the extent to which
endogenous attention improves performance under conditions of low sensitivity (e.g., low
contrast, short stimulus duration). In the current study, I interpret internally represented
17

forms of attention, either explicit (voluntary) or implicit, as aspects of endogenous
attention, although I will discuss the behavioral and neural effects of each in turn. In
designs that manipulate attention explicitly on a trial-by-trial basis, voluntary spatial
attention improves the accuracy and speed with which stimuli are detected (e.g., Posner,
1980). Voluntary attention can also be deployed to varying degrees, with associated
perceptual sensitivity benefits commensurate with the amount of attention (e.g., 100%
versus 75%) to a given spatial location (Mangun & Hillyard, 1990). Event-related
potential (ERP) studies in humans have shown that voluntary attention improves
sensitivity by enhancing the gain of attended as compared to unattended stimulus
representations at early levels of visual processing. For example, many studies have
found amplitude enhancements of early latency visual evoked responses to stimuli that
occur in the location of voluntary attention (Mangun & Hillyard, 1991; Martinez, Di
Russo, Anllo-Vento, & Hillyard, 2001; Van Voorhis & Hillyard, 1977). Moreover,
behavioral and neurophysiological enhancements occur at the fovea with focused versus
distributed attention (Miniussi, Rao, & Nobre, 2002), demonstrating that directed
voluntary attention can enhance processing even when spatial resolution is already high.
In light of the benefits of endogenous attention, it is perhaps not surprising that
performance on sustained attention tasks suffers as voluntary attention wanes over time.
Endogenous attention can also be guided by regularities or patterns that are learned
implicitly over time, such as the probability of a target item appearing in a particular
spatial location (Chun & Jiang, 1998; Geng & Behrmann, 2005; Hoffmann & Kunde,
1999) or temporal patterns that predict when task-relevant stimuli will occur (Martin et
al., 2005; Olson & Chun, 2001). Like explicit manipulations of attention, implicit
18

manipulations of endogenous attention have been shown to be strong predictors of
behavioral improvements. For example, Geng and Behrmann (2005) showed that spatial
probability robustly influenced attention, with behavioral improvements (faster reaction
times, better accuracy) occurring at likely versus unlikely locations. Moreover, they
showed that implicit cueing through spatial probability had just as strong an influence on
attention as explicit endogenous cues and was effective in reducing distraction during
task performance.
Perceptual Benefits of Exogenous Spatial Attention
Exogenous selective attention also improves performance under conditions of low
sensitivity. Behaviorally, exogenous attention improves detection of, and facilitates
responses to, salient sudden-onset stimuli (Jonides & Yantis, 1988; Posner & Cohen,
1984). In addition, sudden-onset cues that occur within ~200 milliseconds of an
upcoming stimulus have been shown to enhance apparent contrast (Carrasco, Ling, &
Read, 2004; Carrasco, Penpeci-Talgar, & Eckstein, 2000; Ling & Carrasco, 2006) and
increase spatial resolution (Carrasco, Williams, & Yeshurun, 2002a; Yeshurun &
Carrasco, 1999), thus improving perceptual sensitivity to stimuli that occur in the location
of the cue. Although most studies of exogenous attention have examined peripheral
cueing, sudden onsets have been shown to capture attention at the fovea as well. For
example, Coull, Frith, Buchel, and Nobre (2000) demonstrated that targets that occurred
unexpectedly at fixation while subjects prepared to attend to a later point in time (but still
at fixation) captured attention in a stimulus-driven manner that was similar to exogenous
capture in the periphery. In supporting of the exogenous influence of these stimuli, visual
cortex activity to the unexpected targets (as measured using functional magnetic
19

resonance imaging, fMRI) was preferentially enhanced, supporting the idea that
exogenous attention results in sensitivity and gain enhancements in visual processing,
even at the fovea.
Interactions between Endogenous and Exogenous Attention
In cases of highly focused voluntary attention, observers are usually able to resist
distraction from salient, abrupt onsets outside the focus of attention (Theeuwes, 1991;
Yantis & Jonides, 1990). However, several lines of research indicate that endogenous and
exogenous visuospatial attention interact, in that the consequences of sudden-onset
salient events (exogenous attention) can be modulated by the current goal state
(endogenous attention). Although these combined influences may sometimes lead to
performance impairments, such as when you are distracted from paying attention to a
certain location in space by a salient interruption in the periphery (van der Lubbe &
Postma, 2005), exogenous orienting is not truly automatic (for review see Santangelo &
Spence, 2008). Instead, the total impact of a sudden-onset event on endogenous attention
has been shown to depend on how much the event shares in common with the target of
voluntary attention (e.g., the target and the sudden-onset distracter are the same color), a
mechanism labeled contingent attentional capture (Folk et al., 2002). Relevant to the
present investigation, such sudden onsets can lead to performance enhancements, such as
when endogenous and exogenous attention cues converge on the same location. In a
study of directed spatial attention, Hopfinger and West (2006) showed that brief,
exogenous cues that occurred in the location of voluntary attention (e.g., left or right
visual field) amplified the behavioral benefits of endogenous attention: Subjects were
faster to respond to an upcoming target when an exogenous cue occurred in the location
20

where they were already paying voluntary attention. Overall, these findings demonstrate
that endogenous and exogenous attention systems together optimize both behavior and
the distribution of neural resources.
Sustained Interactions between Endogenous and Exogenous Attention
Robertson and colleagues (Robertson, Mattingley, Rorden, & Driver, 1998) examined
the relationship between vigilance and spatial attention, and demonstrated that
improvements in sustained attention can lead to better spatial attention (e.g., less severe
symptoms in patients with hemispatial neglect after sustained attention training). But the
reverse - that spatial attention benefits may lead to better vigilance performance - has
received less consideration. Exogenous and endogenous cues may improve vigilance in
different ways. For example, endogenous cues may help the observer conserve attentional
resources over time. Previous studies using voluntary attention cues, such as the
presentation of explicit warning cues (Hitchcock, Dember, Warm, Moroney, & See,
1999; Hitchcock et al., 2003) have shown better vigilance when cues reliably (80%)
predict the occurrence of an upcoming target. That is, observers can use the cues to direct
their attention over short intervals of time (i.e., the cue indicates that a target will occur in
the next few seconds) rather than maintain attention continuously over long time
intervals. Manipulations of implicit endogenous attention also appear to be strong
predictors of better sustained attention. Observers benefit from temporal regularities in
the structure of stimulus presentation, showing better performance over time when
stimuli occur at a constant, rhythmic rate (Scerbo, Warm, Doettling, Parasuraman, &
Fisk, 1987) or when targets occur at constant intervals (e.g., 1 target every 30 seconds,
Helton et al., 2005).
21

The deleterious effects of insufficient resources over time during a vigilance task
might also be offset by manipulating exogenous attention, such as by inserting attention-
grabbing, salient stimuli to transiently prime resources. In this vein, previous research on
the effects of exogenous stimulation during vigilance has shown promising effects for
procedures that capitalize on the effects alerting cues (O'Connell et al., 2008; Robertson,
Tegner, Tham, Lo, & Nimmo-Smith, 1995). However, in these studies, the benefits to
sustained performance were the result of intensive training with loud auditory cues that
primarily affected overall physiological arousal (i.e., increases in skin conductance;
O'Connell et al., 2008). It is therefore still an open question as to whether the kind of
simple, sudden-onset visual events, such as those used in classic cueing studies of
exogenous attention over short time periods (e.g., Posner & Cohen, 1984), can change the
trajectory of sustained attention performance.
The benefits of exogenous attention due to sudden-onsets during vigilance tasks that
require response inhibition are not as straightforward. Abundant evidence suggests that
exogenous attention boosts stimulus perception regardless of the particular response
mapping (e.g., Hopfinger & Mangun, 1998). However, response inhibition tasks place
the additional demand of controlling impulsive motor responses (Helton et al., 2009).
Thus, improved target perception due to exogenous attention may not be enough to
counteract the efficient but occasionally error-prone response routine of responding to
most stimuli, especially without strong, endogenous attentional control to inhibit
responses to rare targets (Robertson et al., 1997).


22

Overview
The overarching goal of this set of experiments was to examine how vigilance is
affected by exogenous and endogenous attention cues that I hypothesized would reduce
resource demands. Although the primary aim was to determine whether such cues
benefited performance, the eventual aim was to use this information to design maximally
challenging vigilance tasks that could be used in training studies. In the first experiment, I
investigated how exogenous attention affects performance on vigilance tasks requiring a
demanding perceptual discrimination. I compared performance on two display versions
that differed in features designed to capture exogenous attention. I examined the
influence of display version during a vigilance task that required responses to rare targets
(hereafter referred to as the sustained attention task) and a vigilance task that required
withholding of responses to rare targets (hereafter referred to as the response inhibition
task). I hypothesized that sudden onsets would improve sensitivity by drawing attention
exogenously to the task, and thus attenuate the usual vigilance decrement that occurs
when observers fail to maintain high levels of voluntary attention. However, I anticipated
that this benefit due to exogenous cueing might not improve performance in the response
inhibition task due to the competing and detrimental influence of a habitual motor
routine. In a second experiment, I varied interstimulus timing to determine whether the
benefit due to sudden onsets observed in Experiment 1 depended on precise and
predictable temporal information about when to allocate endogenous attention. In the
third experiment, I assessed performance across a two-week interval on the most difficult
versions of the sustained attention and response inhibition tasks to confirm that vigilance
decrements would not be reduced with simple repeated testing. To anticipate the results, I
23

will mention that exogenous attention cues (Experiment 1) improved performance on the
sustained attention task but not on the response inhibition task. Predictable timing also
improved performance during sustained attention (Experiment 2). Relevant to choosing
outcome measures to examine training effects, vigilance did not improve with simple
practice on either the sustained attention or the response inhibition tasks (Experiment 3).
Experiment 1
The purpose of this experiment was to examine how sudden-onset exogenous
attention cues affected performance on perceptually demanding vigilance tasks. One of
the necessary objectives was to create target stimuli that were equally demanding across
individuals. Vigilance researchers have often noted striking individual differences in
baseline performance (Davies & Tune, 1969; Parasuraman, 1986), which poses serious
problems for comparing performance over time across individuals. Simply put, a target
that is initially difficult for one person to discriminate might be easy for another person to
discriminate, and hence these two people will experience very different resource demands
in maintaining endogenous attention for a long period of time. While some studies have
set target parameters based on average group discrimination performance (Parasuraman
& Mouloua, 1987), vigilance tasks are not commonly based on individually determined
target parameters. The few exceptions have included adaptive threshold procedures to
change a target stimulus parameter (e.g., brightness) throughout the task, in effect making
the targets easier to detect in order to compensate for performance decline (Bakan, 1955;
Berger & Mahneke, 1954; Frome, MacLeod, Buck, & Williams, 1981; Wiener, 1973). In
contrast, my approach was to determine target parameters for each participant using an
individual discriminability threshold procedure, and then present that threshold-level
24

target throughout the duration of the subsequent sustained attention task. This novel
design ensures equivalent attentional and perceptual demands across individuals, which
makes the task highly sensitive to fluctuations of attention over time. In this first
experiment, I implemented threshold-based targets and compared perceptual sensitivity
over time on two versions of the vigilance tasks that differed in display features that were
designed to capture exogenous attention. I predicted that perceptual sensitivity would
decline less in the sustained attention task that displayed brief, sudden-onset cues
immediately before a potential target stimulus due to the influence of exogenous
attention. However, I predicted that the influence of exogenous attention might not
benefit performance when response inhibition was required.
Methods
Participants. Forty volunteers (mean age = 20, range = 18 - 34; 29 females) from the
University of California, Davis gave informed consent and participated in exchange for
course credit. All participants had normal or corrected-to-normal vision and were free
from neurological insult. Two participants could not complete the practice and were
excluded from further participation. Additionally, two participants were unable to
complete the response inhibition task and were excluded. Analyses of performance over
time did not include those participants who could not perform the task to criterion
initially. Four participants performed below the initial accuracy cut-off (at least 50%
correct detections) during the first 4 min of at least one version of the assigned task (3
performed the response inhibition task) and were excluded from all analyses. Data from
the remaining 32 participants are presented here.
25

Stimuli. For all tasks, participants were seated comfortably in a dark, sound-
attenuating booth, with a chin rest ensuring a viewing distance of 57 cm. Presentation
software (Neurobehavioral Systems, http://www.neurobs.com) was used to control
display of all stimuli on a 53-cm CRT monitor (Eizo Flexscan F980) at a refresh rate of
60 Hz. Responses were made using the top right key on a ProPad game pad response
device. Participants were assigned to one of two successive-presentation (Parasuraman,
1979) vigilance tasks: a sustained attention task (N = 16) that required button presses to
rare targets or a response inhibition task (N = 16) that required withholding button
presses to rare targets. Uninterrupted task duration was 32 minutes. During both tasks,
single lines (light grey; 1.31 cd/m
2
) were presented at the center of the screen against a
black background (0.02 cd/m
2
) while participants fixated a small red square (see Figure
2-1). The fixation square was continuously visible at the center of the screen and
participants were instructed to maintain their gaze on the square throughout the task.
Stimuli were presented at a high event rate (30 events/minute; stimulus duration = 150
ms, interstimulus interval [ISI] = 1850 ms) that is considered to be resource demanding
and has been shown previously to promote declines in perceptual sensitivity over time
(Parasuraman, 1979). Long non-target lines (4.5) were presented 90% of the time and
short target lines (M = 3.4, range = 3.2- 4.1) were presented 10% of the time. Line
width (0.035) did not differ between short and long lines. Two target lines occurred
pseudo-randomly every 20 trials, with the restriction that only 2 short lines could occur
consecutively at any point in the task. Instructions emphasized accuracy in making
simple button responses with the right index finger to the target short lines (sustained
attention task) or the non-target long lines (response inhibition task). Each participant
26

performed two versions (order counterbalanced) of their assigned task that differed
according to display features: The stable version displayed a mask pattern (same
brightness as line stimuli) during the entire interstimulus interval, and the transient
version displayed the same mask pattern for 100 ms before and after each line stimulus,
with a blank intervening screen (see Figure 2-1). The sudden onset of the mask before the
line in the transient display was designed as an exogenous attention cue, in line with
previous findings showing a sudden change in luminance contrast to be one of the
strongest forms of exogenous attentional cueing (Steinman, Steinman, & Lehmkuhle,
1997). The presentation of the mask following the line was included to equate visibility
between the transient and stable displays (i.e., to prevent a persisting after-image in the
transient display). The mask was composed of many short lines positioned throughout a
5.0 x 1.0 space; each short line was 0.07 wide and ranged in height from 0.28 to
0.45. The mask pattern was never present during line stimulus presentation. The
Michelson contrast ratio between the line/mask stimuli and the black background was
97% (Coren, Ward, & Enns, 1999).
Threshold procedure. The length of the short target line was set to each participants
discrimination threshold using a self-paced 3 down/1 up staircase procedure that
determined an 80% accuracy threshold (see Leek, 2001). Display characteristics and
event durations were matched to the stable and transient versions of the subsequent
vigilance task. During each trial of the threshold procedure, a warning signal
(enlargement of the red fixation square) preceded two sequential lines at fixation and
participants judged the length difference between the two lines, indicating same or
different length with the top left (left index finger) and top right (right index finger) game
27








Figure 2-1. Stimuli and timing for the transient display version of the vigilance task.
Sequence of events is shown for one non-target trial (long line stimulus; 90% of stimuli)
and one target trial (short line stimulus; 10% of stimuli). Participants made responses to
the short lines during the sustained attention task and to the long lines during the response
inhibition task. The transient mask appeared before and after every line (both targets and
non-targets) as depicted. In the stable display version, the mask was presented
continuously throughout the entire 1850 ms inter-event interval (e.g., in place of the sole
fixation square shown in the figure). Stimuli are shown in white for illustrative clarity,
but were presented in light grey during the tasks; the fixation square was presented in red
in Experiment 1 and 3, and yellow in Experiment 2. In Experiment 2, the inter-event
interval varied randomly across trials (M = 1850, range = 1350 - 2350 ms). The fixation
square was continuously visible at the center of the screen during all tasks.





28

pad buttons, respectively. Each trial was composed of (a) a long and short line (order
randomized), (b) two short lines, or (c) two long lines. Participants made an untimed
response and then received sound feedback (a ding sound for correct, a whoosh sound for
incorrect) indicating response accuracy. The length of the short line varied throughout the
task in ~0.07 steps (the first short line subtended 3.6; possible range = 3.2- 4.4), with
increments in line length occurring after 3 successive correct responses and decrements in
line length occurring after each incorrect response. The task terminated after eight length
reversals (i.e., a switch from an increment in line length to a decrement in line length).
The length of the long line was the same as in the main task and never varied.
Overview of testing procedures. Participants were introduced to the task through brief
verbal instructions and practice of the threshold procedure (20 trials with the short line
length held constant at a length that would be easily detectable by all participants).
Participants then completed the threshold procedure (~10 minutes) followed by practice
on the assigned vigilance task with the target line length set to their threshold (5 minutes).
An extra 5 minutes of practice was allowed if participants correctly detected fewer than
75% of targets. (Note: One participant received extra practice and subsequently
performed at 87% accuracy during the first block of the task, well within the range of
normal [< 1 SD] group performance.) Participants then performed the first version (e.g.,
stable display) of the assigned vigilance task (sustained attention or response inhibition)
for 32 minutes (960 trials) without sound feedback or breaks. This sequence was repeated
for the second version (e.g., transient display) after a 15-minute break. Order of display
version (stable or transient first) was counterbalanced across participants.
29

Overview of data analysis. Hit and false alarm rates were calculated over 4
contiguous blocks (240 trials each) for each version of the vigilance task. To allow a
comparison between tasks with opposite response requirements, hits were defined as
correct detections of targets (short lines), whether that corresponded to a button press
(sustained attention task) or a correct inhibition (response inhibition task). Likewise, false
alarms were defined as failures to detect non-targets (i.e., responding to a long line as if it
were a short line), corresponding to incorrect button presses to non-targets in the
sustained attention task and accidental inhibitions to non-targets in the response
inhibition task. The non-parametric index of perceptual sensitivity, referred to as A', was
calculated from the hit and false alarm rates (Stanislaw & Todorov, 1999) and served as
the main dependent measure of performance over time. The non-parametric index of
response bias, ''
D
(See, Warm, Dember, & Howe, 1997) was also calculated and used to
assess changes in response tendencies over time. Finally, correct RTs were examined;
RTs were trimmed to remove the top and bottom 5% outliers on a cell-by-cell basis for
each individual.
Separate analyses were conducted for the sustained attention task and for the response
inhibition task. Multivariate analysis of variance (Maxwell & Delaney, 2004, pp. 671-
675, 682-751), which makes no assumption about compound sphericity, was used to
assess all within- and between-subjects effects, and Bonferroni correction was used to
control the family-wise error rate for all post-hoc analyses (e.g., u .05/2 when testing
two simple main effects to follow-up a significant interaction).


30

Results
Threshold. The threshold procedure successfully produced a ~.80 hit rate during the
first block (240 trials) of the sustained attention task (M = .77, SD = .14) and the response
inhibition task (M = .78, SD = .17). Difference thresholds the visual angle difference
between the standard long line and the participants threshold-level short line were
analyzed in separate mixed-model ANOVA with the within-subjects factor of display
(stable or transient) and the between-subjects factor of order (stable first or transient
first). The only significant source of variance in each analysis was an interaction between
display and order (sustained attention: F (1, 14) = 18.29, p < .001; response inhibition: F
(1, 14) = 27.98, p < .001), which revealed that thresholds were lower (i.e., smaller
difference in visual angle between the two lines) on the threshold task that was performed
second regardless of display version (see Table 2-1). The results suggest that participants
perceptual thresholds improved with exposure to the tasks in general, but did not differ
according to display version.
Order effects on performance. Measures of performance over time were subjected to
separate mixed-model ANOVAs with within-subjects factors of time (4 blocks) and
display version, and between-subjects factor of order of display version. It was first
important to determine if task order affected performance. In terms of accuracy (A')
during the sustained attention task, the omnibus ANOVA showed no main effects of
order, but a significant interaction between order and display version (F (1, 14) = 6.57, p
= .022). Overall accuracy was higher in the task that was performed first, regardless of
display version (see Table 2-1), although order did not modulate performance over time.
31

The same interaction was just significant in the ANOVA of response inhibition (F (1, 14)
= 4.54, p = .05). Due to this effect, order was retained as a between-subject factor in
all subsequent analyses of accuracy. Order did not influence reaction times (all p values >
.08) and was excluded from RT analyses (Note: Including order in the analyses of RT did
not change the reported results).

Table 2-1
Order Effects on Threshold and Performance
________________________________________________________________________

Difference Threshold
a
Overall Perceptual Sensitivity (A')

______________________________________________________

Task Order First Task Second Task First Task Second Task
________________________________________________________________________

Stable First 1.08 0.84 .946 .916


Transient First 0.99 0.71 .919 .880
________________________________________________________________________

a
Visual angle difference between the long line and the 80%-threshold-level short line obtained through a 3
down/1-up staircase procedure. Values represent the average across sustained attention and response
inhibition tasks.


Perceptual sensitivity. I examined changes in perceptual sensitivity (A') over time for
each task. In the sustained attention task, mean A' values during the initial block did not
differ between display versions (M = .940 for transient and .937 for stable, t < 1),
indicating that the threshold procedure successfully matched initial performance.
Perceptual sensitivity declined over time in both versions, although the main effect of
time in the omnibus ANOVA was a non-significant trend (Ms blocks 1 - 4 = .939, .934,
.917, .907, F (3, 12) = 3.13, p = .065). Importantly, the decline in A' over time was
32

qualified by a significant interaction between time and display version (F (3, 12) = 5.35,
p = .015). Examination of the performance trajectories over time revealed that there was
comparatively less performance decline in the transient version as compared to the stable
version, in line with our prediction (see Figure 2-2). A test of the simple main effect of
time within each display separately indicated that the performance decline was significant
in the stable version (F (3, 12) = 4.87, p = .019), but not in the transient version (F (3, 12)
= 1.27, p = .32). No other sources of variance were significant in the analysis of A'. A
similar model of response bias revealed non-significant effects of time (F (3, 12) = .386,
p = .76), display (F (1, 14) = 1.02, p = .32) and the interaction between display and time
(F (3, 12) = 1.09, p = .38). Thus, changes in response bias did not accompany the
observed changes in perceptual sensitivity.
In the response inhibition task, mean A' values during the initial block did not differ
between display versions (M = .946 for transient and .931 for stable, t (15) = 1.22, p =
.23), indicating that the threshold task successfully matched initial performance.
Perceptual sensitivity declined over time in both display versions, as indicated by a main
effect of time in the omnibus ANOVA (Ms blocks 1 - 4 = .939, .917, .905, .887, F (3, 12)
= 5.74, p = .011). There were no other significant sources of variance (ps > .24).
However, an examination of the performance trajectories revealed that the performance
decrement was numerically worse on the transient display, the exact opposite pattern as
in the sustained attention task (see Figure 2-2). Finally, the model of response bias
revealed no significant sources of variance (all ps > .26).


33

a.

b.


Figure 2-2. Changes in perceptual sensitivity (A') as a function of time on task in
Experiment 1. A' plotted over four contiguous 8-minute blocks (240 trials each) for each
display version (stable and transient) during (a) the sustained attention task (N = 16) and
(b) the response inhibition task (N = 16).
0.88
0.9
0.92
0.94
0.96
1 2 3 4
A
'
Block (8 min)
Transient
Stable
0.86
0.88
0.90
0.92
0.94
0.96
1 2 3 4
A
'
Block (8 min)
Transient
Stable
34

Reaction time. Although participants were encouraged to respond accurately rather
than quickly, I wanted to determine whether changes in perceptual sensitivity were
accompanied by changes in RT (e.g., a speed-accuracy trade-off). In the sustained
attention task, mean correct RTs (correct detections of targets) were submitted to a
repeated-measures ANOVA with within-subjects factors of time (blocks 1-4) and display
version. A main effect of time revealed that RTs significantly increased over time (Ms =
691 msec, 745 msec, 748 msec, 745 msec, F (3, 13) = 9.23, p = .002), but RTs did not
differ according to display (F (1, 15) = .009, p = .92). In addition, the interaction between
time and display version was not significant (F (3, 13) = 2.37, p = .11). These results
confirm that RTs did not become differentially faster in the stable display, which is the
display version that showed the most decline in A' (i.e., there was no speed-accuracy
trade-off). In the response inhibition task, mean correct RTs (correct responses to non-
targets) were submitted to a repeated-measures ANOVA with within-subjects factors of
time (blocks 1-4) and display version. A significant main effect of time revealed that RTs
significantly decreased over time (Ms = 556 msec, 537 msec, 523 msec, 506 msec, F (3,
13) = 5.68, p = .01). In addition, an effect of display revealed that RTs were significantly
slower in the transient display (M = 559 msec for transient and 502 msec for stable, F (1,
15) = 6.00, p = .027). The interaction between display and time was not significant (p >
.45). These results suggest that the decrease in accuracy over the course of the task was
accompanied by a speeding of reaction time, regardless of display version. Thus, there
appeared to be a speed-accuracy trade-off in the response inhibition task that did not exist
in the sustained attention task. This finding is consistent with a recent report that
35

characterizes sustained response inhibition tasks as measures of impulsive responding
(Helton et al., 2009).
Discussion
The effects of exogenous attention and response requirements on vigilance were
examined. Using threshold-based targets, I found that perceptual sensitivity declined
predictably over time, which was mitigated by the influence of exogenous attention
during performance of the sustained attention task. The perceptual benefit due to
exogenous attention did not improve performance over time when response inhibition
was required, indicating that this type of responding demands a high level of voluntary
attentional control. Taken together, the results extend previous findings suggesting that
errors in sustained attention tasks are a consequence of the limits of effortful, voluntary
attention when resource demands are high (Grier et al., 2003; Helton et al., 2005).
In the sustained attention task, the typical decline in performance over time was
attenuated when display features of the transient task presumably facilitated exogenous
capture of attention and thereby enhanced perception. When the positive effects of
exogenous attention were removed in the stable display, the decline in perceptual
sensitivity significantly increased. I interpret the improvement in perceptual sensitivity to
be a beneficial consequence of sudden-onset cues exogenously attracting attentional
resources to the display. In addition, there were concomitant changes in target response
speed over time. Because accuracy was strongly emphasized over speed in both
experiments, I interpret the modulation of target responses as further support that sudden
onsets made the perceptual decision easier (Parasuraman, 1986, p. 14). In line with this
interpretation, Parasuraman and Davies (1976) examined changes in response latencies in
36

a vigilance task that emphasized accuracy over speed in making different button
responses to both targets and non-targets. They found that while correct detection and
false alarm response times (i.e., positive response latency) increased over time,
response times associated with correct rejections and errors to non-targets (i.e., negative
response latency) decreased or remained stable over time. The increase in RT associated
with positive responses was taken as a marker of the relative strength of evidence
supporting a positive (target) v. negative (non-target) decision. That is, as observers
attempt to be as accurate as possible, correct discrimination of targets is faster in the case
where better perception supports the target v. non-target decision. In the present study,
sudden-onsets led to both better perception and faster responses to targets over time on
task.
The present study demonstrated that resource demands can be reduced by enhancing
perception through simple sudden-onset cues. Importantly, sudden onsets occurred on
every trial, and they presumably increased sensitivity on both target and non-target trials.
In a sustained attention task with rare targets, maintaining a reliable percept of the non-
target stimulus is important for accurate identification of the target when it does occur.
Indeed, when the percept of the non-target does not have to be held in working memory,
such as in simultaneous-presentation tasks (e.g., deciding whether two lines presented
simultaneously are different in length), sensitivity does not reliably decline over time
(Parasuraman, 1979). Furthermore, Caggiano and Parasuraman (2004) demonstrated that
simultaneous performance of a spatial working memory task interfered with the
discrimination of a rare target defined by its spatial location (i.e., eccentricity). They
interpreted the increased perceptual sensitivity decrement under the condition of a
37

competing working memory task to reflect the consequence of worse working memory
representations of both the standard, non-target position and the infrequent target
position. In line with the critical role of memory load in vigilance (Parasuraman, 1979;
Parasuraman & Davies, 1977), the improvement in perception with exogenous attention
in the present study could help maintain an accurate working memory representation of
the non-target line, thus making target discrimination easier. This finding is important
since it demonstrates for the first time that sustained discrimination depends not only on
the availability of voluntary attention resources, but is also a result of an interaction
between voluntary attentional control and the exogenous features of the task.
In the response inhibition task, the presumed perceptual benefit following exogenous
cues did not attenuate the decline in performance: There was not a significant difference
in the vigilance decrement between transient and stable display versions. Although
exogenous attention to the sudden-onset stimuli in the transient display may have
improved target detection during the response inhibition task as it did during the
sustained attention task, this perceptual enhancement was not enough to offset waning
voluntary attention and inhibitory control. To the contrary, features of the transient
display may have actually hurt performance during response inhibition. In the transient
display, the sudden on/offsets are recurrent and rhythmic visual events that may have
served to synchronize the simple motor response with each stimulus (for a review, see
Repp, 2005). Indeed, non-target responses became faster over time while accuracy
decline, indicating a speed-accuracy trade-off. This mode of responding is consistent with
synchronous, habitual motor responding. Furthermore, overall RTs were significantly
slower in the transient display, which may reflect a strategic attempt to resist accidental
38

inhibition errors by making slower, more careful responses to non-targets. Together,
these data suggest that sustained response inhibition places high resource demands on the
attentional control system to maintain voluntary attention while also counteracting
habitual motor responses to non-targets when presented with a target (Robertson et al.,
1997).
Experiment 2
In Experiment 1, I found that sudden-onset cues improved performance over time
during sustained attention. The primary purpose of Experiment 2 was to examine the
impact of predictable timing cues on the perceptual benefits due to sudden-onset cues. I
introduced a variable ISI in Experiment 2 to prevent subjects from taking precisely timed
and efficient breaks in attention, thus increasing the overall resource demand. This
manipulation allowed the assessment of the independent effects of sudden onsets and
predictable timing on sustained performance.
The secondary purpose of Experiment 2 was to clarify the nature of changes in
threshold and overall accuracy observed in the second task in Experiment 1.
Improvements in perceptual threshold on the second task suggested either short-term
perceptual changes (i.e., true changes in threshold due to stimulus exposure) or practice
effects (i.e., improved performance on the threshold procedure itself rather than true
changes in threshold). In Experiment 2, I designed a more ecologically valid threshold
procedure that would be less susceptible to procedure-specific practice effects. To test for
any changes in threshold due to stimulus exposure, threshold was re-measured
immediately following completion of the sustained attention task. In Experiment 1, I also
observed that overall accuracy was lower in the second task performed. This pattern
39

suggested a possible interaction between threshold changes and changes in accuracy. If
improvements in threshold were mainly driven by procedure-specific practice effects,
then thresholds would be artificially lower on average, leading to a more difficult target
during the sustained attention task and lower overall accuracy. At the group level, this
could be expressed as accuracy going down as thresholds get better, as we observed. The
decrease in accuracy could alternatively reflect fatigue; however, the 15-minute break
between task versions should have restored performance to normal levels (e.g.,
Mackworth, 1948). The between-subjects manipulation of display in Experiment 2
directly addressed practice effects and fatigue issues related to threshold and accuracy
during sustained attention.
Methods
Participants. Forty volunteers (mean age = 20, range = 18 34, 26 females) from the
University of California, Davis gave informed consent and participated in exchange for
course credit. All participants had normal or corrected-to-normal vision and were free
from neurological insult.
Stimuli. Task and stimulus parameters (including visual angle, contrast ratio and
average event rate) for the sustained attention tasks were the same as in Experiment 1,
except for a few important modifications. First, a variable ISI was included representing a
range of 500 ms above and below the interval used in Experiment 1 (range = 1350 2350
ms). Second, debriefing in Experiment 1 revealed that some participants attempted to
assess line length by comparing the currently presented line with the background mask
parts. To prevent this strategy, each of the lines forming the mask was vertically
repositioned on every presentation. The change in position of each mask part (0.07 x
40

0.28) was randomly chosen within a range of -0.14 to +0.14, with negative values
indicating a downward shift and positive values indicating an upward shift. The subtle
shift was not immediately noticeable, but prevented the comparison strategy if a
participant attempted it. Finally, the type of display (stable or transient) was manipulated
between subjects (n = 20 in each task version) to avoid possible fatigue or order effects
on performance. As in Experiment 1, instructions emphasized accuracy in making simple
responses with the left mouse button (right index finger) to the target short lines. Stimuli
were presented on a 21-inch CRT monitor (Viewsonic AccuSync 120) at a refresh rate of
60 Hz. Line and mask stimuli were presented in light grey (3.25 cd/m
2
) on a black
background (0.05 cd/m
2
; Michelson contrast ratio = 97%).
Threshold procedure. The length of the short target line was set to each participants
80% discrimination threshold at each assessment, which was determined using a variant
of Parameter Estimation through Sequential Testing (PEST; Taylor & Creelman, 1967).
PEST is an adaptive thresholding procedure that dynamically adjusts the amount of
change (step size) between testing levels based on current task performance. This design
ensures that testing levels quickly converge on a threshold value. PEST is more efficient,
accurate, and flexible than fixed thresholding methods such as the staircase procedure
used in Experiment 1.
In PEST, one specifies a testing level for the first target stimulus, and testing levels of
subsequent target stimuli are determined using a Wald sequential likelihood-ratio test
(Wald, 1947). The Wald test indicates whether the percent of correct responses at a given
testing level is within a range determined by the threshold probability (P
t
) and the
deviation limit of the sequential test (W) chosen by the experimenter at the beginning of
41

the procedure. If the current testing level is above or below the range determined by the
Wald test, it is decreased or increased appropriately by a specified step size. The initial
testing level (e.g., the length of the short line), as well as P
t
, W and the starting step size,
are all pre-determined by the experimenter at the beginning of the procedure. Subsequent
step sizes (e.g., halving the step size with a reversal in direction) are determined as
described by Taylor and Creelman (1967, p. 783). The PEST procedure terminates when
a minimum step size value is obtained. The resulting threshold estimate is equal to the
testing level that would have next been presented.
According to Taylor and Creelman (1967), the starting values of W, P
t
and minimum
step size influence the outcome of the procedure. Specifically, the starting value of W
primarily affects the power of the test, such that small values of W yield quick but not
very powerful decisions, while large values of W give, after many trials, decisions of
great power (p. 783). The starting values of P
t
and the minimum step size affect the
efficiency and the precision of the test estimate, respectively (pp. 783-784). In the current
study, I chose starting values of .75 for the threshold probability (P
t
), .85 for W, and a
minimum step size of 1 pixel (.035).
Overview of testing procedures. Participants followed the same sequence of tasks as
in Experiment 1. The display characteristics, event durations, and response requirements
of the PEST threshold procedure exactly matched those of the corresponding sustained
attention task, except that the target short line occurred 33% of the time (cf. 10% target
probability during the sustained attention task). Participants received sound feedback
indicating (a) correct detections (ding sound), (b) target misses (whoosh sound), and (c)
incorrect responses to non-targets (whoosh sound). Next, participants perform the
42

sustained attention task without breaks or sound feedback. After completing the sustained
attention task, participants performed the threshold task again to assess short-term
changes in threshold.
Results
Threshold. The PEST threshold procedure successfully produced an ~.80 hit rate
during the first block of the corresponding sustained attention task (M = .81, SD = .13).
Difference thresholds were subjected to a mixed-model ANOVA with a between-subjects
factor of display (stable or transient) and a within-subjects factor of exposure (pre or
post). No significant differences were found between pre and post thresholds (M = 0.98
for pre and 0.96 for post; F (1, 38) = .196, p = .66). No other sources of variance were
significant (range of p values = .18 - .34). These results demonstrate that low-level
perception, as measured in a threshold task with similar structure to the actual sustained
attention task, did not change after continuous performance.
Performance over time. Perceptual sensitivity (A') over time was assessed with a
mixed-model ANOVA with a between-subjects factor of display and a within-subjects
factor of time (blocks 1- 4). A main effect of time demonstrated that perceptual
sensitivity declined steadily over time for both display versions (Ms = .944, .910, .887,
.872, F (3, 36) = 37.7, p < .0001). In addition, a main effect of display (F (1, 38) = 4.74, p
= .036) revealed that overall performance was better in the transient display. However,
the interaction between display and time was not significant (F (3, 36) = .547, p = .65)
(see Figure 2-3). Thus, the data show that sudden-onset cues improved overall sensitivity,
but did not prevent a decline in perceptual sensitivity with unpredictable timing.

43



Figure 2-3. Changes in perceptual sensitivity (A') as a function of time on task in
Experiment 2. A' plotted over four contiguous 8-minute blocks (240 trials each) for each
display version (stable and transient) of the sustained attention task.

A similar model of response bias (''
D
) revealed a main effect of time (Ms = .69, .78,
.90, .83, F (3, 36) = 5.78, p = .002), indicating that subjects were less likely to respond as
time passed. The non-significant interaction between display and time (F (3, 36) = 1.09, p
= .36) indicated that this increase in response bias over time was not different between
the two displays. Although there was no difference in response bias over time between
the two display versions, a significant main effect of display (F (1, 38) = 4.11, p = .049)
indicated that subjects exhibited a more liberal overall response bias (i.e., were more
likely to respond) in the transient (M = .70) than in the stable display version (M = .90).
0.84
0.86
0.88
0.9
0.92
0.94
0.96
0.98
1 2 3 4
A
'
Block (8 min)
Transient
Stable
44

Finally, trimmed mean correct RTs were submitted to a mixed-model ANOVA with
between-subject factor of display and within-subjects factor of time (blocks 1-4). In line
with Experiment 1, reaction times significantly increased over time (Ms = 683, 739, 737,
780 ms, F (3, 36) = 16.87, p < .0001). In addition, a significant interaction between time
and display revealed that the pattern of RT over time differed according to display (F (3,
36) = 3.06, p = .04). Although subjects slowed down over time with both displays,
subjects were able to maintain faster responses to targets for a longer portion of the task
in the transient display condition (transient display: Ms = 665 msec, 699 msec, 701 msec,
778 msec; stable display: Ms = 701 msec, 799 msec, 773 msec, 783 msec). No other
sources of variance were statistically significant (main effect of display: F (1, 38) = 2.12,
p = .15). These results replicate the pattern of increasing target RT over time revealed in
Experiment 1, but also suggest that reaction time performance suffered more in the stable
version with unpredictable timing.
Effects of event timing on performance over time. To assess the independent effect of
event timing on perceptual sensitivity over time, we compared performance between
participants who completed the transient display version of the sustained attention task
with a constant ISI (Experiment 1, N =16) and those who completed the transient display
version with a variable ISI (Experiment 2, N = 20). A mixed-model ANOVA with a
between-subjects factor of ISI (constant or variable) and a within-subjects factor of time
(blocks 1- 4) revealed a significant main effect of time (F (3, 32) = 13.3, p < .0001) that
was moderated by a significant interaction between time and ISI (F (3, 32) = 5.87, p =
.003; see Figure 2-4). The main effect of ISI was not significant (F < 1). The data show
45

that performance over time was better with predictable interstimulus timing, controlling
for the effect of sudden-onset attention cues.
Figure 2-4. Changes in perceptual sensitivity (A') as a function of time on task in
Experiments 1 and 2 for the transient display version of the sustained attention task.
A' plotted over four contiguous 8-minute blocks (240 trials each) for constant
(Experiment 1; N = 16) and variable (Experiment 2; N = 20) ISI conditions.

Discussion
The results from Experiment 2 revealed that the perceptual benefit of exogenous cues
depended on the endogenous modulation of attention by predictable timing. When event
timing was variable, exogenous cues did not prevent a decline in perceptual sensitivity
over time. However, exogenous cues did lead to an increase in overall perceptual
sensitivity and a decrease in response times to targets in the transient display. These
effects support the conclusion that exogenous cues captured attention and improved
0.86
0.88
0.9
0.92
0.94
0.96
0.98
1 2 3 4
A
'
Block (8 min)
Experiment 1 - Constant ISI
Experiment 2 - Variable ISI
46

performance, even in the face of variable timing, but not to the same extent as when
exogenous cues coincided with maximal endogenous attention guided by the implicit
endogenous cue of predictable timing (Experiment 1). In addition, predictable timing was
a strong independent predictor of successful performance over time. The results from the
comparison of performance over time on the transient tasks in Experiment 1 and 2
demonstrate that in tasks equated for individual difficulty and initial accuracy,
performance declined significantly more over time with variable stimulus timing
(Experiment 2). The independent benefit due to predictable timing confirms previous
reports of better sustained attention with rhythmic stimulus presentation (Scerbo et al.,
1987).
Predictable timing likely reduced resource demands by providing information about
when to pay maximal attention. Studies of attention in both the visual (Martin et al.,
2005; Olson & Chun, 2001) and the auditory domain (Jones, Moynihan, MacKenzie, &
Puente, 2002) have shown that endogenous attention is modulated according to the
underlying temporal structure of stimulus events, leading to more accurate performance
with predictable timing. This kind of anticipatory attending to specific moments in time
has been described as an important mechanism for synchronizing attentional resources
with critical external events in a way than can flexibly improve behavior (Large & Jones,
1999). Moreover, the modulation of endogenous attention in this manner is an efficient
way to conserve attentional resources so that maximal attention coincides primarily with
task-relevant events. The current replication of the effects of implicit timing cues on
sustained performance points to a critical role for this kind of attentional cueing in
preserving limited resources over long periods of time.
47

Temporal predictability can modulate endogenous attention without an explicit
intention on the part of the observer to voluntarily direct attention differently across time
(Martin et al., 2005). However, it is unclear whether the regular underlying temporal
structure in the current study also manipulated voluntary attention explicitly. Previous
studies of the effects of both spatial and temporal probability on attention have shown
large variability in how much observers notice and volitionally use the underlying
expectancies to direct attention during the task (Geng & Behrmann, 2005). The constant
ISI in Experiment 1 (1.85 seconds) provided observers with a relatively long and
predictable period during which attention was not required. Since voluntary attention can
be generated toward a particular location in space in ~300 ms after the onset of an
instructional cue (Muller & Rabbitt, 1989; Posner & Cohen, 1984), observers could
afford to take a break in attention after presentation of the previous line and then
explicitly redirect attention toward the end of the ISI to prepare for the next line. Indeed,
studies of temporal attention within the range of the ISI used in the present study have
shown that observers can accurately direct voluntary selective attention to specific
moments in time based on instructional cues (Coull, 2004; Coull et al., 2000; Coull &
Nobre, 1998). In terms of temporal complexity, the present task was relatively simple and
straightforward. Thus, some participants likely became aware of the regular temporal
pattern and used it to take voluntary time outs during the interstimulus interval and then
direct maximal attention to the point in time of the next stimulus. Although we can
confidently conclude that temporal regularity modulated endogenous attention implicitly,
it is possible that temporal regularity also modulated endogenous attention explicitly.
48

Given that observers could afford to take an attention time-out, why would they?
Although voluntary attention can be maintained in a given location for many seconds
(e.g., Ling & Carrasco, 2006; Silver, Ress, & Heeger, 2006), there is evidence that
maintaining attention continuously even for relatively short periods of time (~5 minutes)
is not always successful, resulting in attentional lapses along with behavioral and neural
processing consequences (Weissman et al., 2006). In this vein, there is mounting
evidence for a phenomenological basis of mind-wandering, or breaks in attention, that
occur spontaneously during performance of a central task. Behavioral studies using self-
reports, thought probes, and RT measures (for a review see Smallwood & Schooler,
2006) and neuroimaging studies of non-task related brain activity (Mason et al., 2007;
Smallwood, Beach, Schooler, & Handy, 2007) have demonstrated that observers often
experience disruptions in focused attention while performing sustained attention tasks,
such as visual monitoring or reading. Thus, voluntary attention is not usually maintained
at a maximum level but rather appears to wax and wane over long periods of time.
Predictable timing information about when to pay attention can be used to efficiently
deploy maximal attention only during presentation of a potential target, and this helps a
person avoid the consequences of unintentional mind wandering.
Experiment 3
The primary purpose of Experiment 3 was to use a within-subjects design to assess
the effect of repeated practice on vigilance performance. Previous studies have found that
vigilance does not improve with moderate practice (e.g., Nuechterlein et al., 1983),
although vigilance decrements can be reduced with extensive practice (e.g., > 20 practice
sessions of a 30-minute vigilance task; Parasuraman & Giambra, 1991). However, such
49

repeated exposure to visual stimuli can produce a limited (i.e., stimulus-specific) form of
perceptual expertise referred to as perceptual learning (Seitz & Watanabe, 2005). In this
way, repetitive stimulus exposure during training might lead to incidental improvements
in baseline perception, which could influence vigilance performance. In order to use these
tasks as outcome measures of training-related improvement, it was critical to characterize
how repeated task practice influenced both baseline thresholds and vigilance.
I reasoned that performance on the most resource-demanding versions of the
vigilance tasks would be robust to simple practice effects and would also be most
sensitive to real training-related change. Because exogenous attention aided performance
in the transient display version of the sustained attention task, I chose the stable version
as the most demanding sustained attention task. While sudden-onset display features in
the transient display version did not significantly affect accuracy during the response
inhibition task, the transient version certainly produced a larger numeric decrement
compared to the stable version; thus, I chose the transient version as the most demanding
response inhibition task. I predicted that performance would decline significantly in both
the sustained attention and response inhibition tasks, and that it would not improve with
repeated practice.
Methods
Participants. Seventeen volunteers (mean age = 25, range = 20 32; 8 females) gave
informed consent and were paid $10/hour for participation. Three participants performed
below the initial accuracy cut-off (at least 50% correct detections) during the first 4 min
of at least one of the tasks and were excluded from all analyses. Additionally, three
50

participants did not complete all testing sessions and were excluded. Data from the
remaining 11 participants are presented here.
Stimuli. At each of two assessments, each participant performed the stable version of
the sustained attention task and the transient version of the response inhibition task. Task
and stimulus parameters (including visual angle) for these tasks were the same as in
Experiment 1, except for a few modifications. First, each participant was pseudo-
randomly assigned a non-variable ISI for each task (M = 1880 ms), with assigned ISIs
representing a range of 300 ms above and below the ISI used in Experiment 1 (range =
1550 2150 ms). Second, instructions emphasized both speed and accuracy. Stimuli
were presented on an LCD monitor with 2ms black/white transition times at a refresh rate
of 60 Hz (Viewsonic VX-922). Line and mask stimuli were presented in light grey (40.29
cd/m
2
) on a black background (.35 cd/m
2
). (Note: Neither task order nor ISI influenced
performance and were excluded from all analyses.)
Overview of testing procedures. Participants visited the lab for four testing sessions:
one session on each of two consecutive days for the first assessment and the same format
repeated two weeks later for the second assessment. The order of tasks (e.g., sustained
attention task on day 1 followed by response inhibition task on day 2) and time of day
were counterbalanced across participants and then held constant for each participant at
the second assessment. On the first day of testing, participants followed the same
sequence of tasks as in Experiments 1 and 2 (e.g., brief practice followed by the threshold
procedure and then the sustained attention task). We used the PEST procedure from
Experiment 2 to determine 75% threshold-level target line lengths at each assessment.
(Note: We increased the difficulty to 75% to make the target discrimination challenging
51

across repeated testing sessions.) Additionally, so that I could assess short-term changes
in perceptual threshold, participants performed the threshold procedure again after
completing the vigilance task. This sequence was repeated for the second task (e.g.,
response inhibition task) on the second day.
Analysis of vigilance performance. Initial data processing (e.g., calculating perceptual
sensitivity for each block of trials) followed the same approach as in Experiment 1.
However, instead of analyzing vigilance performance using repeated measures ANOVA,
I analyzed performance using hierarchical linear regression. Hierarchical linear models
produce estimates of group performance (as in standard linear regression) as well as
individual deviations from group estimates. This modeling approach is well-suited for
analyzing nested data (i.e., individuals testing repeatedly over time) (see details in
Raudenbush & Bryk, 2002).
Analyses were conducted in SAS 9.1 using the proc mixed function (Littell, Miliken,
Stoup, & Wolfinger, 1996; Singer, 1998). Full maximum likelihood estimation was
implemented and the error covariance structure was modeled as unstructured. Separate
analyses were conducted for the sustained attention task and for the response inhibition
task. Each model of perceptual sensitivity included the fixed (i.e., group-level) effects of
time (the slope of A' across blocks) and assessment (test vs. retest), and the interaction
between time and assessment. Random effects on the intercept (i.e., A' during the first
block) were included to allow for individual differences in initial performance. The
following model was implemented for each task:
A' =
0

1
(time)
2
(assessment)
3
(time*assessment) + e
With random eIIects on
0

52

In this model, the
0
estimate represents initial A' at the beginning oI the task, and the
1

estimate represents the slope oI A' (i.e., the vigilance decrement). The
2
estimate
represents change in overall A' due to repeated testing, and the
3
estimate represents the
change in the vigilance decrement due to repeated testing.
Results
Threshold. The PEST threshold procedure successfully produced an ~.75 hit rate
during the first block (120 trials) of the corresponding vigilance task (M = .76, SD = .15).
Difference thresholds were subjected to two separate repeated measures ANOVAs with
within-subjects factors of assessment (test vs. retest) and exposure (pre-task vs. post-
task). For the sustained attention task, I found no significant effects of assessment (M =
.69 at test and .70 at retest, F (1, 10) = .37, p = .55) or exposure (M = .69 for pre-task
and .60 for post-task, F (1, 10) = 3.31, p = .099). The interaction was also not significant
(p > .36). Likewise, in the response inhibition task, I found no significant effects of
assessment (M = .86 at test and .88 at retest, F (1, 10) = .32, p = .58) or exposure (M =
.88 for pre-task and .97 for post-task, F (1, 10) = 3.33, p = .098). The interaction was
also not significant (p > .39). These findings confirm that thresholds did not improve
significantly after either short-term exposure (pre-task to post-task) or repeated practice
of the vigilance tasks.
Perceptual sensitivity. Perceptual sensitivity (A') was assessed in two separate
hierarchical linear models as a function of time (blocks 1 - 8) and assessment (test vs.
retest) and their interaction. In the sustained attention task, the effect of time was
signiIicant ( -.006, p = .028) demonstrating that perceptual sensitivity declined over
time. Examination of the performance trajectories showed that performance was very
53

similar at the two testing points (see Figure 2-5). Neither the effect of assessment nor the
interaction between assessment and time was significant (p values > .20), confirming that
performance did not change with repeated testing.
In the response inhibition task, the eIIect oI time was signiIicant ( -.009, p = .005)
demonstrating that perceptual sensitivity declined over time as in the sustained attention
task. Again, the performance trajectories at test and retest were very similar (see Figure
2-5). Neither the effect of assessment nor the interaction between assessment and time
was significant (p values > .89), confirming that performance did not change with
repeated testing.
Reaction time. In contrast to the instructions in Experiment 1, instructions in
Experiment 3 emphasized speed and accuracy equally. To assess the effect of this
instruction variation, trimmed mean correct RTs were submitted to two separate
hierarchical linear regression analyses (similar to the analyses of perceptual sensitivity).
In the sustained attention task, reaction times to correctly detected targets significantly
increased over time ( 15.39, p = .004), replicating the findings from Experiments 1
and 2. Neither the effect of assessment nor the interaction between assessment and time
was significant (p values > .60), indicating that response speed did not change with
repeated testing.
In the response inhibition task, reaction times to non-targets significantly decreased
over time ( -5.14, p = .03) as in Experiment 1. In addition, a significant effect of
assessment revealed that reaction times were significantly faster at the second assessment
(M 474 msec at test and 440 msec at retest, -30.51, p = .034). The interaction
between time and assessment was not significant (p = .74). These results suggest that
54

a.

b.


Figure 2-5. Changes in perceptual sensitivity (A') as a function of time on task in
Experiment 3. A' plotted over four contiguous 4-minute blocks (120 trials each) for (a)
the sustained attention task (N =11) and (b) the response inhibition task (N = 11). The
two testing points (test and retest) were separated by a 2-week interval.

0.82
0.84
0.86
0.88
0.9
0.92
0.94
0.96
1 2 3 4 5 6 7 8
A
'
Block (4 min)
Test
Retest
0.82
0.84
0.86
0.88
0.9
0.92
0.94
0.96
1 2 3 4 5 6 7 8
A
'
Block (4 min)
Test
Retest
55


overall response speed decreased at retest, possibly as a result of participants becoming
more familiar with the response requirements of the task. Importantly, controlling for
changes in reaction time did not change the reported results for perceptual sensitivity.
In addition, overall reaction times were ~ 50 msec faster in Experiment 3 than in
Experiment 1. These results suggest that instructions that emphasize both speed and
accuracy (Experiment 3) motivate faster responding overall compared to instructions that
emphasize only accuracy (Experiment 1). To further investigate this result, I examined
whether reaction time correlated with accuracy. Although there was not a significant
correlation at the first testing point (r = .29, p = .38), worse accuracy was significantly
correlated with faster reaction times at the second testing point (r = .61, p = .048). This
finding is consistent with previous reports of speed-accuracy trade-offs during sustained
response inhibition (Helton, 2009), and it indicates that one should carefully examine the
correlations between speed and accuracy across repeated testing sessions.
Discussion
The results demonstrate that perceptual sensitivity declined significantly over time in
the most resource-demanding versions of the sustained attention and response inhibition
tasks. Importantly, performance within the same individuals did not improve with
repeated testing. However, reaction times sped up on the response inhibition task at
retest, and thus it became important to track changes in both accuracy and reaction time
in subsequent training studies. Finally, perceptual thresholds did not change over either a
short period of time (30 minutes on task) or a longer period of time (at the end of the
second testing session), suggesting that baseline perception did not improve with simple
stimulus exposure. Together, the results from Experiment 3 demonstrate that threshold
56

and vigilance performance was stable across repeated testing sessions, making these tasks
appropriate indices of training-related improvements.
Summary
I examined changes in perceptual sensitivity over time the vigilance decrement as
a function of various experimental manipulations that we hypothesized would modulate
resource demands. In all tasks, I employed parameters previously shown to increase
resource demands and promote declines in perceptual sensitivity, including a high event
rate and successive presentation. In addition, I experimentally manipulated resource
demands in four different ways: (1) by individualizing target discrimination difficulty
using threshold-based targets; (2) by manipulating response requirements (response
initiation vs. response inhibition); (3) by manipulating display features to vary the degree
of exogenous attention drawn to the task; and (4) by manipulating the predictability of
stimulus timing, which provided information about when to allocate endogenous
attention. Using threshold-based targets, I found that sensitivity declined reliably over
time in tasks that required response initiation to rare targets (sustained attention task) and
in tasks that required withholding of responses to rare targets (response inhibition task).
In the sustained attention task, I found that the vigilance decrement was mitigated by the
influence of sudden-onset exogenous attention cues and by predictable stimulus-timing
information. In the response inhibition task, exogenous cues did not help performance.
Indeed, response inhibition appeared to increase demands on attentional resources
independently such that cues were not beneficial in preventing performance decline. In
sum, the best sustained performance was achieved (1) when response inhibition was not
required; (2) when predictable timing information guided the efficient deployment of
57

endogenous attention over time; and (3) when exogenous sudden-onset cues attracted
attention to the task to increase overall sensitivity. These data extend previous findings
that have shown that errors in vigilance tasks are a consequence of the limits of effortful,
voluntary attention given high resource demands (Grier et al., 2003; Helton et al., 2005;
Helton & Warm, 2008; Smit et al., 2004b; Szalma et al., 2004). Here, I have shown that
attentional cues can improve performance by reducing resource demands.
Together, these findings support the idea that successful vigilance performance is a
result of a dynamic interaction between endogenous attentional control and exogenous
events that capture attention, in line with theories concerning the neurobiological basis of
attention (Corbetta et al., 2008; Corbetta & Shulman, 2002). I interpret the effects of
exogenous and endogenous attention in light of evidence supporting a resource model of
vigilance. Specifically, exogenous cues independently improved overall sensitivity by
transiently increasing attentional resources, but they were arguably not effective in
preventing a decline in vigilance (Experiment 2). On the other hand, implicit endogenous
cues independently improved sustained performance (Experiment 1 vs. 2), in line with
previous reports of better vigilance with rhythmic, regular presentation of stimuli (Scerbo
et al., 1987). If the major limiting factor during vigilance is the overall resource demand
imposed on the endogenous attention system, then it appears that manipulations of
endogenous attention that result in the conservation of resources over time are more
effective at preventing the vigilance decrement than exogenous cues that only transiently
prime resources. The combined benefit of exogenous and endogenous cues was clearly
evident in that the best performance over time was achieved when exogenous cueing
coincided with maximal endogenous attention (Experiment 1, transient display).
58

Controlling for individual differences in low-level perception is critical in any study
of vigilance. My novel approach of setting target parameters according to individual
threshold using the PEST procedure (Experiments 2 and 3) proved to be an effective way
to create vigilance tasks that were equally challenging, both attentionally and
perceptually, for all individuals. Because the PEST method can flexibly converge on any
chosen threshold level (within an appropriate range), this approach allows an assessment
of sustained attention at varying levels of baseline performance. In effect, standardizing
baseline sensitivity helps control the total demand of the task (the amount of effort
required for task completion). In a meta-analysis of the sensitivity decrement in 42
vigilance studies, See and colleagues (1995) suggested that total task demand helps
explain the variation in the vigilance decrement not directly attributable to other critical
design factors. The present threshold-calibration approach is a promising method for
equating overall task difficulty according to individual perceptual ability and thus
equating the extent of the vigilance decrement across individuals. Indeed, although
thresholds did not change with moderate exposure to the line stimuli in Experiment 3 (~2
hours of total stimulus exposure between the two tasks and two assessments), it is
possible that more extensive stimulus exposure could lead to perceptual learning.
Alternatively, training interventions might influence baseline perception. Thus, setting
target parameters according to threshold at each testing point is an effective way to
control the difficulty of target discrimination with repeated testing.
Finally, repeated testing did not improve vigilance performance on the most resource-
demanding versions of the sustained attention and response inhibition tasks. This finding
is consistent with previous reports showing no improvements in vigilance with moderate
59

practice (Nuechterlein et al., 1983; Williams, 1986). These results indicate that simple
practice is not sufficient to reduce the information-processing demands associated with
perceptually challenging vigilance tasks. In other words, the vigilance decrement
observed in nave participants is not due to lack of familiarity with overall task demands
or with specific task features such as target probability and task duration. The stability of
performance in the response inhibition task further indicates that normal performance
decrements are not due to lack of practice with the particular motor requirements of this
kind of vigilance task. The level of performance stability I observed across repeated
testing suggests that these tests are robust to simple practice effects and will likely be
sensitive to real training-related changes in voluntary attention.













60










CHAPTER 3

IMPROVEMENTS IN PERCEPTION AND SUSTAINED
ATTENTION WITH INTENSIVE MEDITATION TRAINING









61

The ability to focus attention while resisting distraction is critical for adaptive, goal-
directed behavior. Voluntary attention guided by goals, previous knowledge, or explicit
instructions can be directed to spatial locations (e.g., Posner, 1980) and moments in
time (e.g., Correa, Lupianez, Madrid, & Tudela, 2006) to improve behavioral accuracy
and efficiency. However, voluntary attention is limited and cannot be sustained
indefinitely. Research on sustained attention consistently demonstrates declines in target-
discrimination accuracy (perceptual sensitivity) with increasing time on task, a
phenomenon called the vigilance decrement (see Parasuraman, 1986, for review). Despite
this evidence, few attempts have been made to improve healthy adults sustained
attention with training. Studies have demonstrated improvements on vigilance tasks
following repetitive practice (e.g., Parasuraman & Giambra, 1991), but no study has yet
identified a general training regimen for reducing the vigilance decrement. Similarly,
training-related improvements in other domains are often limited to the training task.
However, some forms of training, such as practice on action video games, can improve
performance on untrained attention tasks, indicating the transfer of learned skills to new
situations (see review in C. S. Green & Bavelier, 2008). The present study investigated
improvements in voluntary sustained attention with meditation training.
The attentional resource model attributes the vigilance decrement to exhaustion of
limited information-processing resources (Davies & Parasuraman, 1982; Warm et al.,
2008). Specifically, factors known to increase information-processing demands, such as
perceptually difficult targets (Nuechterlein et al., 1983) or working memory load
(Parasuraman, 1979) cause rapid and reliable decrements in perceptual sensitivity. When
the total task demand (the combined resource demands of multiple aspects of a task) is
62

high, vigilance decrements are large (See et al., 1995). Attentional cues can mitigate the
impact of resource demands during vigilance. Hitchcock and colleagues (2003) found
that both accuracy and cerebral blood flow (a measure of resource consumption) declined
less over time when a warning cue reliably (80%) predicted the occurrence of an
upcoming target. Similarly, other kinds of attentional cues have been shown to improve
vigilance (see MacLean, Aichele, Bridwell, Mangun et al., 2009 for review). Less is
known about how training might affect vigilance.
Historical accounts from the Buddhist contemplative tradition describe meditation
practices (Shamatha) that are designed to improve sustained attention (see translations in
Buddhaghosa, 1979; Tsong-kha-pa, 2002). Shamatha practitioners learn to stabilize
attention on a chosen stimulus, such as the tactile sensation of breathing, and enhance the
perceived detail of the stimulus in order to cultivate non-task-specific skill in regulating
and controlling voluntary attention (Wallace, 1999). Introspection is used to monitor the
quality of attention, recognize when attention has wandered, and guide attention back to
the chosen stimulus. This metacognitive or meta-attentive aspect of training may support
the transfer of meditation skills to other domains (Wallace & Shapiro, 2006) and
improvements in perception and attention that are reported in daily life (Wallace, 1999, p.
185).
The present study tests this claim empirically. Findings from longitudinal research on
meditation training in non-experts (1-3 months of full-time daily practice) suggest
improvements on laboratory tests of temporal attention (Slagter et al., 2007; Slagter,
Lutz, Greischar, Nieuwenhuis, & Davidson, 2009) and attentional alerting (Jha,
Krompinger, & Baime, 2007, one-month group). Cross-sectional studies have shown that
63

long-term meditation practitioners show superior sustained attention compared to
meditation-nave controls (Brefczynski-Lewis, Lutz, Schaefer, Levinson, & Davidson,
2007; Valentine & Sweet, 1999). However, longitudinal improvements in sustained
attention with meditation training have not been demonstrated.
I examined the effects of Shamatha training on 60 participants who were divided into
two groups by random assignment: a group of individuals who would enter training first
(N = 30; Retreat 1) and a group of individuals who would serve as wait-list controls for
the first retreat, then attend a second retreat (N = 30; Retreat 2). Retreat participants lived
in a remote mountain setting (Shambhala Mountain Center, Red Feather Lakes,
Colorado) for three months and received instruction from B. A. Wallace in Shamatha, as
well as complementary meditation practices that use imagery and concentration to
develop compassion and kindness toward others (see Wallace, 2006). Participants
attended two daily group meditation and discussion sessions, practiced in solitude (M = 5
hours/day of Shamatha, 45 min/day of complementary practices), and had weekly
individual meetings with the instructor.
Outcomes were assessed at three points during each retreat - before the start of the
retreat (pre), halfway through the retreat (mid), and at the end of the retreat (post) - using
a test of sustained visual attention that produced significant decrements in perceptual
sensitivity before training. I predicted that meditation training would reduce this
decrement. I also predicted that improvements in vigilance would be related to the
resource demands of the task. Specifically, training-related improvements in baseline
perception could improve vigilance by reducing the resource demands associated with
discriminating difficult targets (Nuechterlein et al., 1983; Parasuraman & Mouloua,
64

1987). To characterize the relationship between perception, resource demand, and
vigilance, I analyzed: (1) discrimination (visual threshold on a task that did not require
sustained attention), (2) average sensitivity during sustained attention (an index of total
resource demand; See et al., 1995), and (3) vigilance (decrement in perceptual sensitivity
over time).
Method
Participants
Volunteers were recruited using magazine and online advertisements. Participants
were selected who were (i) between 21 and 70 years old; (ii) willing to be randomly
assigned to either retreat; (iii) familiar with intensive meditation practice (at least three 5-
day retreats); and (iv) willing to abstain from recreational drugs, tobacco and alcohol one
month prior to, and during the study. Selected participants had normal or corrected-to-
normal vision and hearing, and no known neurological or Axis I psychiatric impairments
(M.I.N.I. screen, Sheehan et al., 1998). After selection (~50% inclusion rate), participants
were assigned to either the first retreat group or the wait-list control group through
stratified (age, sex, handedness, ethnicity, meditation experience) random assignment.
Retreat and wait-list control participants were matched on demographic factors and
psychological characteristics (N = 60, see Table 3-1).
1
Three months after the end of
Retreat 1 wait-list control participants began training and underwent assessment in
Retreat 2 (N = 29).
Due to technical problems, performance data were missing for one assessment for
two participants. Analyses of average sensitivity and vigilance included only participants
with complete data (Retreat 1: N = 59; Retreat 2: N = 27).
65

Table 3-1
Group Matching on Demographic Information and Self-report Questionnaires

Retreat Wait-list Control All t value
______________________________________________________________________________________

Age
a
49 (23 69) 46 (22 65) 48 0.79
Sex 14 M, 16 F 14 M, 16 F 28 M, 32 F
Handedness
b
29 R, 1 L 28 R, 2 L 57 R, 3 L
Education
c
4.9 (3 - 6) 5.2 (1 - 6) 5.07 1.09
Non-verbal IQ
d
10.8 (4.0 17.0) 11.2 (6.0 17.0) 11.0 0.65

Meditation Experience
Retreats
e
13 (2 50) 16 (2 100) 14.3 0.67
Daily meditation
f
56 (8.5 180) 54 (12.8 155) 55 0.18
Lifetime hours
g
2800 (250 9500) 2644 (200 15,000) 2716 0.16

BFI
h

Openness 5.5 (3.8 7.0) 5.6 (4.0 7.0) 5.57 0.45
Conscientiousness 5.2 (2.4 6.8) 5.0 (2.3 6.9) 5.1 0.73
Neuroticism 3.2 (1.2 5.2) 3.2 (1.7 5.1) 3.25 0.04
Extroversion 4.2 (2.7 6.7) 4.3 (2.6 6.5) 4.27 0.67
Agreeableness 5.2 (3.3 7.0) 5.3 (3.5 6.8) 5.25 0.25
STAI
i
1.7 (1.0 2.3) 1.8 (1.0 2.7) 1.77 1.06
CES-D
j
1.6 (1.0 4.9) 1.8 (1.0 2.9) 1.71 0.96
Wellbeing
k
5.6 (4.1 6.6) 5.4 (4.5 6.6) 5.54 0.37
_____________________________________________________________________________________

Note. Descriptive statistics (mean value with range in parentheses) of demographic factors reported for
participants who completed all assessments during the first retreat (n = 30 in each group) except as noted.
Scores on all questionnaire measures are based on a 7-point Likert-type scale. Scores at pre assessment
were available for 30 retreat and 28 wait-list control participants for the BFI and STAI, and for 28
participants in each group for CES-D. There were no significant differences between groups on any
demographic or self-report measure at pre assessment (p > .05)

a
Age in years at the beginning of participation in the study.
b
Hand dominance as assessed by the Edinburgh Inventory (Oldfield, 1971).
c
Scale of educational achievement (1 = less than high school; 2 = high school degree; 3 = some college; 4
= college degree; 5 = some graduate; 6 = graduate degree).
d
Total correct on a timed version of half of the full Ravens Progressive Matrices (Raven, Court, & Raven,
1988); second half administered at post assessment.
e
Total number of meditation retreats lasting at least 5 consecutive days. Reports available for 28 wait-list
control and 30 retreat participants.
f
Average daily minutes of formal meditation practice. Reports available for 28 wait-list control and 25
retreat participants.
g
Total lifetime hours of formal meditation practice. Reports available for 30 wait-list control and 29 retreat
participants.
h
Big Five Inventory, 44 items (John, Donahue, & Kentle, 1991).
i
State and Trait Anxiety Inventory (Spielberger, Gorsuch, Lushene, Vagg, & Jacobs, 1983).
j
Center for Epidemiologic Studies Depression Scale (Radloff, 1977).
k
Psychological Wellbeing Scale (Ryff, 1989).




66

All study and task details were approved by the University of California, Davis
Institutional Review Board. Participants gave informed consent and were debriefed after
training. Participants paid for room and board during the retreat (~$5300) but were paid
$20/hr during data collection. Travel expenses were covered for wait-list controls, who
were flown to the retreat center to be tested during Retreat 1.
Measures
On-site testing. Testing sessions were conducted in two laboratories in the building
where participants lived and meditated. Each included a sound-attenuated, darkened
testing room and an adjacent control room. Presentation software
(http://www.neurobs.com) was used to deliver stimuli on an LCD monitor (Viewsonic
VX-922). At each of the three assessments in Retreat 1, wait-list controls arrived three
days (range = 65-75 hours) before testing for acclimatization.
2

At each assessment, participants completed the threshold procedure (~10 min)
followed by the sustained attention task (32 min). In both tasks, participants saw a single
vertical line appear at the center of the screen; the line could be either long (frequent non-
target) or short (rare target). Instructions emphasized speed and accuracy in making
responses with the left mouse button to the short lines (see Figure 3-1).
In the threshold procedure the length of the short target line varied according to a
PEST algorithm (Taylor & Creelman, 1967), which determined the short line length that
participants could correctly detect at a given level of accuracy (e.g., 75%). Participants
received auditory feedback indicating correct detections (ding), target misses (whoosh)
and incorrect responses to non-targets (whoosh). We defined discrimination as the visual
angle difference between the threshold-level short line and the long line, with smaller
67






Figure 3-1. Stimuli and timing for the threshold procedure and sustained attention
task. Single lines (light grey; 40.29 cd/m
2
) were presented at the center of the screen
against a black background (.35 cd/m
2
) while participants fixated a small yellow square
(shown in white) at a viewing distance of 57 cm. In the threshold procedure, long non-
target lines (4.82) were presented 70% of the time and short target lines (possible range
= 2.76 - 4.78) were presented 30% of the time. In the sustained attention task, target
frequency was reduced to 10% of stimuli. A mask was presented whenever a line was not
on the screen, during the variable inter-stimulus interval (ISI = 1550 - 2150 ms).
Instructions emphasized speed and accuracy in responding to the short target lines.






68

threshold values representing better discrimination. In Retreat 1, all participants
performed the threshold procedure again after completing the sustained attention task to
assess short-term changes.
In Retreat 1, the length of the short target line was set to each participants threshold
at each assessment, an approach I have used previously (MacLean, Aichele, Bridwell,
Mangun et al., 2009). Threshold was set to 85% at pre-assessment of Retreat 1 and to
75% at all other assessments. Thresholds accordingly decreased for all participants from
pre to mid assessment. Because testing level was not the same across all assessments, I
conducted separate analyses of group differences in outcome measures at each
assessment in Retreat 1.
Follow-up testing. Participants completed two follow-up assessments of
discrimination after the completion of each retreat (approximately 5 and 15 months after
training), which were conducted via laptop computers sent to each participants home.
Participants received instructions to set viewing distance and ambient lighting. The
threshold procedure at follow-up exactly matched the laboratory procedure.
Analysis
The non-parametric index of perceptual sensitivity, A' (Stanislaw & Todorov, 1999),
was calculated from hit and false alarm rates for each of 8 contiguous 120-trial blocks
during the sustained attention task. The decline in A' was largest during the first half of
the task, and thus we defined improvements in vigilance as positive changes in the slope
of A' during the first 4 blocks. I analyzed changes in slope using hierarchical linear
models, which produced estimates of fixed effects at the group level (similar to standard
regression models) as well as estimates of random effects at the individual level (i.e.,
69

variability around fixed-effects estimates). Analyses of vigilance were conducted in SAS
9.1 (see details in Littell et al., 1996; Singer, 1998).

Results
Retreat 1
Discrimination. Group differences in threshold at each assessment were tested using
analysis of variance (ANOVA; SPSS 17.0). Each ANOVA included group (retreat vs.
control) as the between-subjects factor and task exposure (threshold tested before vs.
after the sustained attention task) as the within-subjects factor. At the pre assessment I
found no differences between groups (F (1, 58) = .32, p = .57) indicating that the groups
were matched on discrimination ability at the beginning of training. I found significantly
lower thresholds in the retreat group at mid (F (1, 58) = 4.80, p = .032) and post (F (1,
58) = 5.80, p = .019) indicating training-specific improvements in discrimination (see
Figure 3-2).
I found no significant effects of task exposure or the interaction of task exposure and
group at any assessment (p > .41 at pre; p >.08 at mid; p >.31 at post), indicating no
short-term change in threshold tested before versus after the sustained attention task (M =
0.79 vs. 0.78). Therefore, participants did not repeat the threshold procedure after the
sustained attention task in Retreat 2.
Average sensitivity. I found no significant group differences in average A' (across all
8 blocks) at pre (M = .95 for both groups, p = .61), mid (M = .90 for both groups, p = .83)
or post (M = .89 for retreat and .90 for control, p = .22), indicating that the groups were
matched on total task demand.

70

a.

b.

*p < .05 **p < .01

Figure 3-2. Improvements in discrimination in Retreat 1 and Retreat 2. (a)
Thresholds plotted for each assessment in Retreat 1 (N = 30 in each group) and the first
follow-up (N = 29 in each group). Retreat participants exhibited significantly better
discrimination (lower thresholds) than wait-list control participants at mid, post, and
follow-up. (b) Thresholds plotted for each assessment in Retreat 2 (N = 29) and the first
follow-up (N = 27). Participants showed significant improvements in discrimination from
pre to mid assessment, and these improvements were maintained at the post and first
follow-up assessments. There were no significant changes in discrimination during non-
training periods.
0
0.2
0.4
0.6
0.8
1
1.2
PRE MID POST FOLLOW-UP
D
i
f
f
e
r
e
n
c
e

T
h
r
e
s
h
o
l
d

(
d
e
g
r
e
e
s

o
f

v
i
s
u
a
l

a
n
g
l
e
)
Assessment
Control Retreat
0
0.2
0.4
0.6
0.8
1
PRE MID POST FOLLOW-UP
D
i
f
f
e
r
e
n
c
e

T
h
r
e
s
h
o
l
d
(
d
e
g
r
e
e
s

o
f

v
i
s
u
a
l

a
n
g
l
e
)
Assessment
* * *
** *
**
71

Vigilance. At each assessment, perceptual sensitivity declined linearly over time, and
the mean vigilance decrement did not appear to differ between retreat and control groups
at any assessment (see Figure 3-3). Using hierarchical linear regression, changes in
vigilance were modeled as a function of the fixed effects of block and group, and their
interaction. Random effects on the intercept were included in the models to allow for
individual differences in initial performance (A' during the first block). At all
assessments, the best-fitting model of performance over time confirmed significant
declines in A' across blocks for all participants, regardless of group membership (p <
.0001; see Table 3-2). Thus, contrary to my prediction, vigilance did not improve with
training.
Influence of discrimination on vigilance. There were no differences in vigilance
between retreat and control groups when task demand (average A') was equated by
experimentally adjusting target length according to individual threshold at each
assessment. However, I also explored how threshold, task demand, and vigilance were
related at the individual level. Before training, lower average sensitivity correlated with
poorer vigilance (r = .45, p < .0001) underscoring the relationship between task demand
and vigilance. As anticipated, there was no correlation between threshold and average
sensitivity or vigilance before training (ps > .27). Next, I correlated change scores in
each pair of the three performance indices. I calculated change on each behavioral
measure by regressing post score on pre score and retaining the standardized residual for
each individual as the index of change from pre to post. I then calculated partial
correlation coefficients between pairs of change scores, controlling for the third change
score. Consistent with the results at pre assessment, decreases in average sensitivity
72

a.

b.

c.

Figure 3-3. Vigilance performance in Retreat 1. Data plotted for participants with complete data at
all assessments (N = 59). Average sensitivity (A') declined significantly during the first 4 blocks (16
minutes) of the sustained attention task at (a) pre, (b) mid, and (c) post assessments. Retreat and control
participants did not differ in the extent of the vigilance decrement at any assessment.
0.88
0.90
0.92
0.94
0.96
0.98
1.00
1 2 3 4
A
'
Block (4 min)
Control
Retreat
0.88
0.90
0.92
0.94
0.96
0.98
1.00
1 2 3 4
A
'
Block (4 min)
Control
Retreat
0.88
0.90
0.92
0.94
0.96
0.98
1.00
1 2 3 4
A
'
Block (4 min)
Control
Retreat
73


Table 3-2
Parameter Estimates from Models of Vigilance in Retreat 1

Model Parameter Estimate Test Statistic
a
BIC
b
_________________________________________________________________________________


Pre
Fixed effects
0
intercept .972 212 ***

1
slope -.006 3.94 ***
-902
Random effects o
2
0
- intercept .001 4.06 ***

o
2
e
residual variance .001 9.41 ***

Mid
Fixed effects
0
intercept .939 142***

1
slope -.012 5.37***
-759
Random effects o
2
0
intercept .001 4.39***

o
2
e
residual variance .001 9.41 ***

Post
Fixed effects
0
intercept .936 149 ***

1
slope -.013 6.12 ***
-765
Random effects o
2
0
intercept .001 4.20 ***

o
2
e
residual variance .001 9.41 ***

__________________________________________________________________________________

* p < .05 ** p < .01 *** p < .001

Note. Full maximum likelihood estimates reported for the best-fitting models of change in perceptual
sensitivity during sustained performance (N = 59 at all assessments). Slope estimates refer to the amount of
decrease in sensitivity per block (4 blocks). At each assessment, slope was centered to the first block of the
sustained attention task (block 1 = 0). In all cases, the simpler models (shown) were better fits than models
that included group and interaction effects (not shown; BIC = -898 for pre, -751 for mid and -759 for post).

a
Test statistics are reported as t values for fixed effects estimates and as z values for random effects
estimates.
b
Bayesian information criterion. Smaller (more negative) values indicate a better model fit.


correlated with decreases in vigilance (pr = .27, p = .036). In addition, decreases in
average sensitivity correlated with improvements in discrimination (pr = .53, p < .0001)
indicating increases in task demand for participants with improvement in threshold. Thus,
adjusting line length during training created systematic differences in task demand
between individuals with the most improvement in threshold (the retreat group) and
74

individuals with the least improvement in threshold (the control group). These results
suggest that although retreat participants may have actually improved in their ability to
sustain attention, the setting of line length based on threshold at each assessment
precluded the observation of such improvements as reflected in measured vigilance.
Retreat 2
The aim in Retreat 2 was to examine how vigilance changed with training when target
parameters were held constant. I fixed the length of the target line for each individual to
the threshold value achieved at the beginning of training, so that task demand would not
increase systematically as training progressed and discrimination improved. Participants
completed the threshold procedure before the sustained attention task at each assessment
and were unaware of the design modification.
Discrimination. Repeated-measures ANOVA was used to test within-subjects change
in threshold across the five assessments that had been tested at the 75% threshold level.
The effect of assessment was significant (F (4, 25) = 4.05, p = .011) indicating
improvement in discrimination over time.

Comparisons between pairs of assessments
confirmed that discrimination improved significantly from pre to mid assessment during
Retreat 2 (F (1, 28) = 8.27, p = .008) with no additional improvement from mid to post (F
(1, 28) = .45, p = .50) (see Figure 3-2). No significant changes in discrimination were
observed during the non-training period (including Retreat 1 mid and post, and Retreat 2
pre; all ps > .21). These findings replicate the training-specific changes in discrimination
observed in Retreat 1.
3

Average sensitivity. A repeated-measures ANOVA of within-subjects change in
average sensitivity revealed a significant effect of assessment (F (4, 23) = 3.26, p = .03).

75

Sensitivity improved significantly from pre to mid during Retreat 2 (F (1, 26) = 8.38, p =
.008) with no additional improvement from mid to post (F (1, 26) = .004, p = .95; see
Figure 3-3). No significant changes in sensitivity were observed between non-training
assessments (all ps

> .83). The observed increases in average sensitivity during training
suggest reductions in total task demand due to improved discrimination.
Vigilance. As can be seen in Figure 3-4, perceptual sensitivity declined over time at
each assessment during the non-training period (mid and post assessments of Retreat 1,
and pre assessment of Retreat 2). However, the performance decrement was reduced at
mid and post assessments of the training period (i.e., the slope of the decrement was
flatter, or less negative, at mid and post assessments of Retreat 2). Using hierarchical
linear models, I modeled changes in vigilance as a function of the fixed effects of block
and assessment, and their interaction. As in Retreat 1, random effects on the intercept
were included in the models. The model of vigilance during the non-training period
revealed a significant effect of block ( -.009, p = .008) but no significant effects of
assessment or the interaction between assessment and block (p > .32; see Table 3-3),
confirming that the vigilance decrement did not change with simple task practice before
training. I then modeled within-subjects change in vigilance during the training period,
with the prediction that training would reduce the vigilance decrement. The model
revealed a significant effect of block ( -.013, p < .0001) and a non-significant effect of
assessment (p = .94). A significant interaction between block and assessment ( .005, p
= .012) indicated that vigilance improved with training, consistent with my prediction.



76

a.

b.


*p < .05 **p < .01

Figure 3-4. Improvements in average sensitivity and vigilance in Retreat 2. Data
plotted for participants with complete data at all assessments (N = 27). The non-training
period of Retreat 1 is designated by R1 and the training period of Retreat 2 is
designated by R2. (a) Average sensitivity (A') across 8 blocks (32 minutes) of the
sustained attention task at each assessment. Participants showed significant increases in
average sensitivity during the training period. (b) A' trajectories plotted across four
contiguous 4-min blocks at each assessment (120 trials/block; 16 min total duration).
Participants showed significant reductions in the vigilance decrement (i.e., less negative
slope) across the first half of the sustained attention task during the training period.
0.88
0.9
0.92
0.94
0.96
0.98
1
Mid - R1 Post - R1 Pre - R2 Mid - R2 Post - R2
A
'
0.88
0.9
0.92
0.94
0.96
0.98
1
1 2 3 4
A
'
Block (4 min)
Mid - R1 Post - R1 Pre - R2 Mid - R2 Post - R2
**
*
77

Table 3-3
Parameter Estimates from Models of Vigilance in Retreat 2


Model Parameter Estimate Test Statistic
a
BIC
b
_______________________________________________________________________________
Non-training
Fixed effects
0
intercept .935 111 ***

1
slope -.009 2.67 **

2
assessment .005 0.91

3
slope x assessment -.003 1.00
-1001
Random effects o
2
0
intercept .001 2.80 **

o
2
e
residual variance .002 12.2 ***
_______________________________________________________________________________
Training

Baseline Model
Fixed effects
0
intercept .944 119 ***

1
slope -.013 4.81***

2
assessment .001 0.07

3
slope x assessment .005 2.52 *
-1165
Random effects o
2
0
intercept .001 3.32 **

o
2
e
residual variance .001 12.2 ***

Threshold Model
Fixed effects
0
intercept .944 122 ***

1
slope -.013 4.82 ***

2
assessment .001 0.10

3
threshold change .001 0.04

4
slope x .005 2.51*
assessment

5
threshold change x -.008 2.06 *
assessment

5
slope x -.003 1.34
threshold change

6
slope x
threshold change x .004 1.74
assessment
-1158
Random effects o
2
0
intercept .001 3.30 ***

o
2
e
residual variance .001 12.2 ***
________________________________________________________________________________
* p < .05 ** p < .01 *** p < .001

Note. Full maximum likelihood estimates reported for models of change in perceptual sensitivity across
blocks (slope) and assessments (N = 27 participants in all models). Slope was centered to the first block of
the sustained attention task (block 1 = 0) and assessment was centered to the first assessment in the model.
Change in threshold was calculated as the standardized residual of the regression of post threshold on pre
threshold, and was centered to zero residual change.

a
Test statistics are reported as t values for fixed effects estimates and as z values for random effects
estimates.
b
Bayesian information criterion. Smaller (more negative) values indicate a better model fit.
78

Follow-up
Follow-up 1. The stability of discrimination improvements was examined at two
follow-up assessments, the first of which was conducted approximately five months after
each retreat. In Retreat 1, I compared performance of Retreat 1 participants at the first
follow-up (N = 29) with wait-list control performance at the beginning of Retreat 2 (N =
29). Retreat 1 participants thresholds were significantly lower than control participants
thresholds (F (1, 56) = 4.58, p = .037) confirming the maintenance of discrimination
improvements after completion of training. In Retreat 2, participants exhibited
significantly lower thresholds at their first follow-up (N = 27) compared to the beginning
of their training (F (1, 26) = 16.9, p < .0001). These results demonstrate enduring
changes in discrimination after training (Figure 3-2).
Next, I explored whether the maintenance of discrimination improvements might be
related to the amount of post-retreat daily meditation by combining participants from
both retreats (N = 54 with complete meditation reports and threshold data). T tests
confirmed that there were no significant differences between retreat groups in daily
meditation after retreat (129 min/day vs. 112 min/day, t (52) = 0.41, p = .68) or follow-up
thresholds (0.56 vs. 0.55, t (52) = 0.20, p = .84). Amount of daily meditation
significantly predicted thresholds at the first follow-up (r = -.36, p = .007) indicating a
correlation between the stability of training-induced discrimination improvement and the
maintenance of regular, but less intensive, meditation (see Figure 3-5).
4



79

Figure 3-5. Daily meditation after retreat predicts discrimination 5 months after
completion of training. Data shown for participants with complete meditation report (N
= 54). Thresholds (visual angle of short line) at the first follow-up plotted as a function of
average minutes of daily practice. Amount of daily meditation after retreat significantly
predicted follow-up threshold.
Follow-up 2. The second follow-up assessment was conducted approximately 15
months after each retreat. I defined performance stability as no significant difference in
discrimination between the first and second follow-up assessments. Paired t tests
confirmed that thresholds were not different between the first and second follow-ups for
either Retreat 1 (N = 23; 0.55 vs. 0.51, t (22) = .93, p = .36) or Retreat 2 (N = 22; 0.55
vs. 0.56, t (22) = .34, p = .73). I also explored whether discrimination performance at the
second follow-up was related to the amount of daily meditation since the first follow-up
(N = 40 with complete meditation reports and threshold data). Amount of daily
80

meditation (M = 113 min/day) did not significantly predict thresholds at the second
follow-up (r = -.15, p = .36). Thresholds were also not significantly related to amount of
change in daily meditation from the first to second follow-up (M = -7 min/day, range = -
210 to +150 min/day; r = -.26, p = .10). Together, the results from both follow-up
assessments suggest that regular meditation may be more critical for maintaining and
consolidating behavioral improvements during the first few months following training.
Although thresholds at the second follow-up were not directly related to meditation, the
stability of discrimination performance more than one year after training nevertheless
suggests long-lasting changes in perception.
Discussion
In a longitudinal wait-list controlled study of long-term daily meditation training, I
found improved vigilance performance in healthy adults. I also found improvements in
visual discrimination, which is consistent with a previous report of increases in visual
sensitivity with intensive meditation training (Brown, Forte, & Dysart, 1984). The results
suggest that mental training of attention on non-visual perceptions (e.g., sensations of
breathing, mental events) generalized to improved visual perception of task-relevant
stimuli.
Previous vigilance research has described task factors that increase information-
processing resource demands, leading to poorer performance over time (Warm et al.,
2008). The present findings suggest that training-related improvements in perception can
decrease resource demands and thus lead to better vigilance. The variation in Retreat 1
versus Retreat 2 illustrates the important relationship between task features, resource
demand, and vigilance. When target length was adjusted according to individual
81

threshold in Retreat 1, I observed systematic increases in task demands (reductions in A')
as threshold improved with training and no changes in observed vigilance. However,
when target length was held constant in Retreat 2, improvements in target discrimination
ability led to reductions in task demands (increases in A') and therefore to improved
vigilance. I interpret these results as follows: Training-related improvements in
perception reduced the resources required to discriminate an unchanging target, which in
turn increased the resources available for sustaining voluntary attention. This implies that
a limited central resource is tapped by both increased perceptual and attentional demands.
Line discrimination performance relies on several stages of information processing
that could be affected by training, from early-stage perceptual processing to later-stage
decision and response execution. Because the dependent measures used in the present
study took into account both hit and false alarm rates, improvements in threshold and
perceptual sensitivity likely reflect improvements in perceptual encoding (Macmillan &
Creelman, 2005). This interpretation is supported by electrophysiological studies that
have linked improvements in perceptual sensitivity to early-stage perceptual processing
(e.g., Luck et al., 1994). Critically, I found no significant group differences or changes in
target reaction time (median correct RT) or response bias (
D
) (See et al., 1997) in
either retreat (all ps > .15), suggesting a decision/response-stage account of the results
can be largely ruled out.
However, improvements in vigilance remained significant after controlling for
changes in threshold (see Table 3-3), suggesting that improvements in perception could
not fully explain the results. In the line discrimination task, an accurate working memory
representation of the non-target supports the perceptual identification of the rare target
82

when it occurs. Using stimuli similar to those in the present study, Parasuraman and
Mouloua (1987) demonstrated that both perceptual manipulations (changing the length
difference between the target and non-target lines) and working memory manipulations
(requiring a comparison between the currently visible target and a non-target line held in
memory) modulated decrements in perceptual sensitivity. During meditation,
practitioners receive extensive experience in attending to the representations of sensory
experience and observing how these representations change moment-to-moment. Thus, a
core aspect of meditation training involves maintaining and accessing information in
working memory from decaying sensory traces. It is therefore possible that training-
related improvements in the precision of visual working memory representations (see
Zhang & Luck, 2008) contributed to observed changes in vigilance.
Interpretations of previous meditation findings have been complicated by possible
alternate explanations, such as pre-existing group differences (in cross-sectional designs)
and self-selection bias (when assignment to training is not random). Although the use of a
longitudinal, wait-list controlled design in the present study eliminated these particular
confounds, factors other than meditation may still have contributed to the present results.
First, features of the training environment, such as the secluded retreat setting and regular
interactions with a committed teacher could have influenced performance, but the
persistence of discrimination improvements after completion of formal training indicates
that retreat-specific factors were not necessary for superior discrimination. Moreover, the
significant correlation between daily meditation and thresholds at the first follow-up
suggests that discrimination ability was directly related to meditation per se and not to
incidental factors. Second, changes in personal motivation during training could have
83

played a role in improved vigilance (e.g., Tomporowski & Tinsley, 1996). Before data
collection, all participants had committed $5300 and were interested in meditation, the
projects scientific goals, and the instructors teachings. Given strongly motivated retreat
and control participants, and the stability of control group data during non-training
assessments, it is unlikely that motivation differentially contributed to results between
groups. In future training studies it will be important to empirically assess the
contributions of motivation and environment, for example by comparing meditation and
active control treatments that are dispensed by the same teacher in the same environment.
The present results add to the growing body of evidence that meditation training can
improve aspects of attention (Lutz, Slagter, Dunne, & Davidson, 2008) while specifically
suggesting that the enhanced sustained attention ability that has been linked to long-term
meditation practice (Brefczynski-Lewis et al., 2007; Valentine & Sweet, 1999) likely
reflects plasticity in the adult brain. The present findings also add to reports of training-
induced improvements in other core cognitive processes, such as working memory
capacity and non-verbal intelligence (Jaeggi et al., 2008; Olesen et al., 2004). Together,
these findings raise the possibility of general improvements in mental function that can
benefit daily activities.
Footnotes
Footnote 1: Age, sex, non-verbal IQ, and previous meditation were not significant
predictors of training-related changes in discrimination, average sensitivity or vigilance
in either retreat (all p values > .16).

84

Footnote 2: Deficits in perception and vigilance were unlikely to occur at an altitude of
2500 m (Virues-Ortega, Buela-Casal, Garrido, & Alcazar, 2004). However, the 3-day
period was sufficient for any such deficits to resolve (Tripathi, Apte, & Mukandan,
2005).

Footnote 3: Improvements in discrimination remained significant after controlling for
visual acuity (M = 20/20, Titmus T2a vision screener; Retreat 1: group difference at post,
p = .009; Retreat 2: effect of assessment, p = .018).

Footnote 4: A similar analysis showed that daily meditation during retreat (M = 368
min/day, SD = 88 min/day) did not significantly predict threshold, average sensitivity or
vigilance at post (p > .51), suggesting that average meditation throughout the reported
range was effective for producing training-related changes in performance.











85










CHAPTER 4

TRAINING-RELATED CHANGES IN SUSTAINED RESPONSE INHIBITION:
THE INFLUENCE OF AGE AND RESPONSE SPEED ON
IMPROVEMENTS IN ACCURACY









86

In their tripartite model of attention, Posner and Petersen (1990) proposed a
functionally distinct executive component of attention that prioritizes and selects among
competing stimuli and actions (see also Fan & Posner, 2004). Executive attention is
involved in exerting voluntary, goal-directed control over which stimuli, thoughts,
emotions, and actions are selected despite automatic or habitual tendencies. The ability to
inhibit habitual or impulsive responses can be tested in the laboratory using tasks that
require quickly responding to frequently occurring stimuli (e.g., 90% probability) and
withholding responses to infrequently occurring stimuli (e.g., 10% probability). Despite
the seemingly simple nature of these response inhibition tasks, errors in responding to the
infrequent stimulus are common. Response inhibition tasks place high processing
demands on executive attention, requiring a person to monitor his or her performance,
counteract the habitual motor response when the low-frequency stimulus appears, and
make behavioral adjustments when errors occur (Carter, Botvinick, & Cohen, 1999;
Miller & Cohen, 2001; Ridderinkhof, van den Wildenberg, Segalowitz, & Carter, 2004).
Moreover, response inhibition errors in the laboratory predict attentional lapses and
action slips in everyday life (Robertson et al., 1997), suggesting that response inhibition
tasks tap general attentional control skills.
Although healthy individuals are often unable to prevent errors during response
inhibition, strategic behavioral adjustments can improve performance (see review in
Ridderinkhof et al., 2004). Specifically, speed-accuracy trade-offs have been observed
during response inhibition tasks, such that responding more slowly to the frequent
stimulus improves overall response inhibition accuracy (Helton et al., 2009). Similarly,
studies have found that speeding of responses prior to the infrequent stimulus increases
87

the likelihood of accidental response inhibition errors (Manly et al., 1999; Robertson et
al., 1997). Emphasizing speed over accuracy can also improve performance in tasks that
require more indirect forms of response inhibition, such as tasks that present conflicting
stimulus-response mappings (e.g., trial-to-trial adjustments in speed during performance
of the Stroop task; Kerns et al., 2004). These findings indicate that individuals can
improve response inhibition performance by adopting a more careful or cautious response
strategy to avoid accidental errors. However, it is unknown whether individuals can
improve response inhibition accuracy through training, possibly without sacrificing
response efficiency.
In the present study, I investigated whether training in Shamatha meditation could
improve response inhibition. During Shamatha meditation individuals cultivate
attentional and behavioral control skills that may transfer to other domains and tasks (see
Chapter 1). In Chapter 3, I observed training-related improvements in perception and
sustained voluntary attention that map onto the primary goals of the practice, including
learning to stabilize attention on the chosen stimulus for long periods of time and
enhancing the perceived detail of the attended stimulus. However, successfully
maintaining voluntary attention on the chosen stimulus requires learning to regulate and
control attention. During meditation, distractions inevitably arise that automatically draw
attention away from the chosen stimulus. In other domains, such as vision-related
attention, executive control plays a key role in guiding attention back to the chosen
stimulus following a salient distraction, thus bringing the focus of attention in line with
current goals (Corbetta & Shulman, 2002). In this way, meditation training likely
88

produces improvements in attentional control that may transfer to other domains and
tasks that also require attentional control, such as response inhibition.
Response inhibition is also cultivated more directly during meditation training.
Response inhibition is involved at all levels of meditation practice, from resisting
habitual behavioral movements while meditating (e.g., not scratching an itch), to resisting
the desire to follow interesting trains of spontaneous thought, to refraining from engaging
with momentary emotions. Learning to recognize and regulate (but not suppress) ones
automatic tendencies to attend to and engage with thoughts and emotions that arise is
deemed a significant step in achieving attentional stability and progressing in the practice
(Wallace & Shapiro, 2006). In addition, classical meditation manuals emphasize
refraining from usual activities and habitual tendencies during formal training (e.g.,
Asanga, 1950; Bodhi, 1995, 2000; Shantideva, 2008). For example, practitioners may
limit normal behaviors such as speaking and reading when not engaged directly in
meditation. Thus, many aspects of formal meditative training cultivate response
inhibition and behavioral control.
There is empirical evidence that meditation can improve executive attention. Jha,
Krompinger, and Baime (2007) recently reported that experienced meditators, compared
to meditation-nave controls, had superior executive control as indexed by less
interference from irrelevant distracting stimuli (on the executive component of the
Attention Network Task; Fan, McCandliss, Sommer, Raz, & Posner, 2002). However, it
is unclear whether this result reflects meditation training per se or other pre-existing
differences between regular meditators and non-meditators.
89

The aim of the present study was to assess longitudinal improvements in response
inhibition as a consequence of Shamatha training. Previous studies (Grier et al., 2003;
Helton et al., 2005) have shown that sustained response inhibition tasks combine the
effort associated with normal vigilance (see review in Warm et al., 2008) and the
challenge of inhibiting impulsive behavioral responses (Helton, 2009; Helton et al.,
2009). Thus, I developed a 32-minute response inhibition task to measure sustained
behavioral control (see Chapter 1).
Method
Study design, participant details, and stimuli were the same as in Chapter 3, except as
noted.
Measures
At each assessment, participants first completed a threshold procedure (duration = 10
min) used to calibrate task difficulty for each individual (see Chapter 2). Stimulus timing
and task details were the same as in the threshold procedure for the sustained attention
task (see Figure 3-1) except that a blank screen was presented during the interstimulus
interval (see Figure 2-1). Instructions required making responses with the left mouse
button to frequent long lines (70% of stimuli) while inhibiting responses to rare short
lines (30% of stimuli). Participants received sound feedback indicating (a) correct
inhibitions of responses to short lines, (b) accidental responses to short lines, and (c)
missed responses to long lines. The procedure determined the length of the short line
required for each participant to perform at 85% overall accuracy. Immediately after the
threshold procedure, participants performed the sustained response inhibition task
(duration = 32 minutes) with the short line set to his or her individual threshold. The
90

response inhibition task exactly matched the threshold procedure except that it did not
include sound feedback and the short line occurred less frequently (10% of stimuli).
Analysis
Response inhibition performance can be quantified as overall error rate, or the rate of
accidental responses to the rare target. However, error rate does not take into account the
tendency to respond (or not respond) during the task (i.e., response bias; see Macmillan
& Creelman, 2005). For example, a participant could achieve many correct inhibitions
using a strategy of responding to fewer trials overall. Thus, I chose a dependent measure
of accuracy that captured how well a participant could correctly inhibit responses to short
lines while also correctly responding to long lines. Specifically, I calculated the non-
parametric index of perceptual sensitivity, A', using correct and incorrect inhibition rates
(see formula in Stanislaw & Todorov, 1999, in applying this formula, I defined correct
inhibitions as hits, and incorrect inhibitions as false alarms). Because I was interested in
how accuracy changed over time during task performance, I calculated A' for each of
eight contiguous blocks of trials (120 trials per block, 4 minutes each) within the
response inhibition task at each assessment. Thus, the dependent measures of response
inhibition task accuracy were (1) average A' and (2) the slope of A' across blocks. I
analyzed changes in response inhibition accuracy using hierarchical linear regression;
analyses were conducted in SAS 9.1 (Singer, 1998).
Some participants unexpectedly did not show large declines in accuracy at pre
assessment. To keep task demands high for all individuals across multiple sessions, I
increased the overall difficulty level of the task by setting target threshold at 75%
accuracy at all subsequent assessments in both retreats. To compare Retreat 1 pre-
91

assessment performance (set at 85% threshold level) to all other assessments (set at 75%
threshold level), I mathematically adjusted A' for each individual at each block at pre-
assessment to compute performance at 75% (adjusted A' = (original A' x .75) / .85).
Assessment Schedule
Before assignment to groups (~4 months prior to the beginning of Retreat 1), all
participants completed a pre-assignment assessment to confirm that the eventual groups
would be matched on performance before training. This testing was conducted via laptop
computers (Dell 600D) sent to each participants home. Participants received detailed
instructions for setting viewing distance and controlling ambient light and sound. The
task procedures completed at pre-assignment exactly matched the procedures completed
at all other assessments.
Participants completed three on-site testing sessions during each retreat (pre, mid, and
post) that were conducted in two laboratories located in the same building where
participants lived and practiced meditation. At each of the three assessments in Retreat 1,
wait-list control participants arrived three days before testing for acclimatization. The
response inhibition task was performed on the second day of lab testing at each
assessment. (Note: The sustained attention task in Chapter 3 was performed on the first
day of lab testing at each assessment.)
Finally, participants completed a follow-up assessment approximately five months
after the completion of each retreat. Testing was similar to the pre-assignment assessment
and was again conducted via laptop computers (Dell 600D for Retreat 1, IBM T-40 for
Retreat 2) sent to each participants home.

92

Results
Pre-assignment
Threshold. A between-subjects analysis of variance (ANOVA) revealed no significant
differences in threshold between retreat and wait-list control groups (M = 1.24 vs. 1.27,
F (1, 58) = .02, p = .89) confirming that the groups did not differ on RIT threshold before
training.
Response inhibition task. An examination of the observed performance trajectories
revealed that response inhibition accuracy (A') declined over time for retreat and control
groups (see Figure 4-1). Using hierarchical linear regression, response inhibition
performance was modeled as a function of the fixed effects of block (1 - 8) and group
(retreat and control), and the interaction of block and group. Random effects on the
intercept were included to allow for individual differences in initial performance (A
during the first block). This model revealed a significant effect of block ( = -.009, p <
.001), confirming that performance declined over time before training. The effect of
group was not significant ( = -.013, p = .59), nor was the interaction between block and
group ( = .002, p = .50), confirming that the groups were matched on performance
before training.
Retreat 1
Participants were removed from analyses who were outliers (3 SD below the grand
mean) in either accuracy at pre assessment (M = .80, SD = .06) or change in accuracy (M
= .07, SD = .07) on the RIT. One Retreat 1 participant was an outlier on change in
accuracy and one control participant was an outlier on accuracy at pre assessment. Thus,
the analyses in Retreat 1 included 58 individuals.
93


Figure 4-1. Response inhibition performance before assignment to groups. Response
inhibition accuracy (A') plotted as a function of time on task (eight 4-minute blocks) for
retreat and control groups (N = 60) at the pre-assignment assessment. Performance
declined significantly across blocks, and the groups did not differ in response inhibition
accuracy.

Threshold. Analysis of variance (ANOVA) was used to test the between-subjects
effect of group (retreat v. control) and the within-subjects effect of assessment (pre-, mid-
, and post-retreat) on threshold. A main effect of assessment (F (2, 55) = 48.5, p < .0001)
revealed that thresholds decreased (i.e., improved) across assessments in both groups
(Retreat: M = 1.0 at pre-retreat, .71 at mid-retreat, and .65 at post-retreat; Control: M =
1.2 at pre-retreat, .84 at mid-retreat, and .74 at post-retreat). Although thresholds were
generally lower in the retreat group at all assessments, the main effect of group was only
a trend (F (1, 56) = 3.31, p = .07). Moreover, the interaction between group and
assessment was not significant (p = .81). These findings are consistent with practice
effects and not with training-related change (cf., training-specific improvements in target
discrimination threshold, Chapter 3). However, because I set target line length according
0.80
0.85
0.90
0.95
1.00
1 2 3 4 5 6 7 8
A
c
c
u
r
a
c
y

(
A
'
)
Block (4 min)
Control Retreat
94

to threshold at each assessment, individual differences in threshold change may have
influenced response inhibition performance. Thus, I statistically controlled for individual
differences in threshold and change in threshold in all models of response inhibition
performance reported next.
Response inhibition task. Similar to performance at the pre-assignment assessment,
response inhibition accuracy declined over time at pre assessment of Retreat 1 (see
Figure 4-2). An examination of the performance trajectories revealed increases in
accuracy from pre to mid assessment for both retreat and control groups. Although the
groups showed similar increases in accuracy during the first half of the task (i.e., blocks 1
4), the retreat group showed larger increases in accuracy toward the end of the task (i.e.,
blocks 6 8) compared to the control group (see Figure 4-2). These observations
suggested training-specific improvements in response inhibition performance.
To analyze changes in response inhibition accuracy, I first tested a model that
included the fixed effects of block, assessment, group and threshold. Random effects on
the intercept were included to allow for individual differences in initial performance. This
model revealed a significant effect of block ( = -.009, p < .0001), a significant effect of
threshold ( = .034, p = .001), and a non-significant effect of group (p = .79).
Importantly, the effect of assessment was significant ( = .043, p < .0001), indicating
increases in overall accuracy. Next, I added the interaction between group and assessment
to the model to test the prediction that increases in accuracy would be greater for retreat
participants. This model also included the interaction between threshold and assessment
to control for individual differences in threshold change. Consistent with my prediction,

95

a.

b.

Figure 4-2. Response inhibition performance during Retreat 1. Response inhibition
accuracy (A') plotted as a function of time on task (eight 4-minute blocks) for (a) retreat
and (b) control groups (n = 29 participants in each group). Both groups showed increases
in overall response inhibition accuracy across testing points (pre, mid, and post).
Increases in accuracy were significantly greater for the retreat group.

0.70
0.75
0.80
0.85
0.90
0.95
1.00
1 2 3 4 5 6 7 8
A
c
c
u
r
a
c
y

(
A
'
)
Block (4 min)
Pre Mid Post
0.70
0.75
0.80
0.85
0.90
0.95
1.00
1 2 3 4 5 6 7 8
A
c
c
u
r
a
c
y

(
A
'
)
Block (4 min)
96

the interaction between group and assessment was significant ( = .012, p = .013).
Moreover, this model fit the data better than the baseline model (see Table 4-1). Finally,
the addition of 2-way and 3-way interaction effects (e.g., the interaction between block
and assessment) did not improve the model fit. Thus, the findings demonstrate training-
related increases in average response inhibition accuracy.

Table 4-1
Parameter Estimates from Models of Response Inhibition in Retreat 1

Model Parameter Estimate Test Statistic
a
BIC
b
_______________________________________________________________________________
Baseline
Fixed effects
0
intercept .842 76.2 ***

1
slope -.009 10.9 ***

2
group .003 0.26

3
threshold -.034 3.23 **

4
assessment .043 12.8 ***

-3055
Random effects o
2
0
intercept .002 4.90 ***

o
2
e
residual variance .006 25.8 ***
_______________________________________________________________________________
Group Differences

Fixed effects
0
intercept .851 74.7***

1
slope -.009 11.0 ***

2
group -.006 0.39

3
threshold -.010 0.90

4
assessment .041 9.76 ***

5
assessment x -.056 6.53 ***
threshold

6
assessment x .012 2.48 *
group
-3090
Random effects o
2
0
intercept .002 4.88 ***

o
2
e
residual variance .005 25.8 ***
________________________________________________________________________________
* p < .05 ** p < .01 *** p < .001
Note. Full maximum likelihood estimates reported for models of response inhibition accuracy (A') as a
function of block (slope), assessment (pre, mid and post), group (retreat and control) and threshold (N = 58
participants in all models). Slope was centered to the first block; assessment was centered to pre assessment
of Retreat 1; and threshold was centered to the grand mean. Retreat group membership was coded as 1 and
control group membership as 0.

a
Test statistics are reported as t values for fixed effects estimates and as z values for random effects
estimates.
b
Bayesian information criterion. Smaller (more negative) values indicate a better model fit.

97

Retreat 2
Threshold. In the model of response inhibition performance in Retreat 1, individual
differences in threshold improvement influenced response inhibition accuracy (p < .0001;
see Table 4-1). In Retreat 2, I addressed this issue by fixing the length of the target line
for each participant to the threshold achieved at the beginning of training. A repeated-
measures ANOVA across four assessments (Retreat 1 post, Retreat 2 pre, Retreat 2 mid,
and Retreat 2 post) confirmed that thresholds did not change significantly after the end of
Retreat 1 (F (3, 26) = 1.06, p = .38). Thus, changes in response inhibition accuracy in
Retreat 2 could be attributed to the direct effect of training and not to the indirect
influence of threshold.
Response inhibition task. As in Retreat 1, I modeled response inhibition performance
using hierarchical linear regression. I tested separate models of (1) the non-training
period (including mid and post assessments of Retreat 1, and pre assessment of Retreat 2)
and (2) the training period (including pre, mid, and post assessments of Retreat 2). Each
model included the fixed effects of block and assessment, and their interaction, as well as
random effects on the intercept. As can be seen in Figure 4-3, response inhibition
accuracy declined similarly over time at each assessment during the non-training period.
The model of performance revealed a significant effect of block ( = -.013, p < .0001)
and non-significant effects of assessment and the interaction of block and assessment (p >
.44), confirming that performance did not change with simple task practice before
training. During the training period, retreat participants showed increases in overall
accuracy from pre to mid assessments, as well as a flatter slope over time (see Figure 4-
3). The model of the training period revealed a significant effect of block ( = -.015, p <
98





Figure 4-3. Improvements in response inhibition during Retreat 2. Data plotted for
participants with complete data at all assessments (N = 29). The non-training period of
Retreat 1 is designated by R1 and the training period of Retreat 2 is designated by
R2. Response inhibition accuracy (A') plotted across 8 blocks (32 minutes) of the
response inhibition task at each assessment. Participants showed significant increases in
average response inhibition accuracy and better sustained performance (i.e., less negative
slope) during the training period.

.0001), a significant effect of assessment ( = .011, p = .02), and a significant interaction
between assessment and block ( = .003, p = .02). The significant effect of assessment
confirmed the result from Retreat 1 of improvements in average response inhibition
accuracy with training. In addition, the significant interaction between block and
assessment demonstrated that training led to reductions of the vigilance decrement when
task difficulty and target parameters were held constant.
0.8
0.85
0.9
0.95
1
1 2 3 4 5 6 7 8
A
c
c
u
r
a
c
y

(
A
'
)
Block (4 min)
Mid - R1 Post - R1 Pre - R2 Mid - R2 Post - R2
99

Follow-up
I observed significant training-related improvements in overall response inhibition
accuracy in both retreats, and an examination of the observed performance trajectories
confirmed that increases in accuracy were evident throughout the 32-min task (see Figure
4-4 for combined data from both retreats). I explored whether these improvements in
accuracy were maintained after completion of training by combining participants from

Figure 4-4. Improvements in response inhibition are maintained after the
completion of training. Response inhibition accuracy (A') plotted as a function of time
on task (eight 4-minute blocks) for each of four testing points. Data shown for
participants of both retreats during their respective training periods (N = 59), and at a
follow-up assessment (N = 54) conducted approximately five months after the end of
each retreat. Both retreat groups showed significant improvements in overall accuracy
with training that endured after the completion of formal training.
0.80
0.85
0.90
0.95
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both retreats with complete data at follow-up (n = 27 for each retreat). A repeated-
measures ANOVA revealed significantly higher accuracy at follow-up compared to pre
assessment (F (1, 53) = 42.38, p < .001). Follow-up t tests confirmed that this result was
independently significant for Retreat 1 participants (M = .87 at follow-up and .80 at pre;
t(26) = 5.80, p < .001) and Retreat 2 participants (M = .91 at follow-up and .87 at pre;
t(26) = 3.52, p = .002). For both groups, accuracy at follow-up was nearly identical to
accuracy at post (M = .89 at both assessments), indicating long-term stability of training-
related improvements (see Figure 4-4).
Influence of Reaction Time on Accuracy
Reaction time and accuracy. I combined participants from both retreats (N = 59) to
examine the relationship between reaction time (median correct reaction time to non-
targets) and response inhibition accuracy. Reaction time and accuracy were not
significantly correlated before training (r = .11, p = .39), and there were no significant
changes in reaction time from pre to post (482 msec vs. 478 msec; t(58) = .50, p = .61).
However, slower reaction times were correlated with higher accuracy at post (r = .38, p =
.003). This result suggests a possible speed-accuracy trade-off, which would be consistent
with the relationship between response inhibition errors and the speed of non-target
responses reported in other studies (e.g., Helton et al., 2009).
I further explored this finding by examining the influence of demographic variables
on reaction time and accuracy. To anticipate the following results, both age and previous
meditation were significantly related to reaction time. Importantly, controlling for
reaction time, age, and previous meditation experience in the hierarchical regression
models of response inhibition accuracy did not change the reported results in either
101

retreat (Retreat 1: interaction between group and assessment, p = .004; Retreat 2: main
effect of assessment, p = .016, interaction between assessment and block, p = .018). Thus,
training-related improvements in accuracy (Retreats 1 and 2) and slope (Retreat 2) were
robust to the influence of these variables.
Reaction time and age. Due to the large age range tested and the possibility for an
interaction between age and response speed (see Caserta et al., 2009), I investigated
whether age was related to changes in either reaction time or accuracy with training. At
pre assessment, age was not correlated with accuracy (r = .12, p = .36) but was
significantly correlated with reaction time (r = .52, p < .001). The findings at post
assessment were similar: Age was not correlated with accuracy (r = .12, p = .34) but was
significantly correlated with reaction time (r = .46, p < .001). These results indicate that
older participants responded more slowly than younger participants.
Improvements in accuracy in younger and older participants. To explore the relations
between age, reaction time, and accuracy, I compared performance in younger (n = 30, M
= 36 years, range = 22 - 50 years) and older (n = 29, M = 59 years, range = 51 - 69 years)
participants. Although older participants had numerically higher accuracy than younger
participants at all assessments, accuracy did not differ significantly between the two
groups (p > .21). However, older participants were significantly slower than younger
participants both before (t (57) = 4.05, p < .001) and after training (t (57) = 3.92, p <
.001; see Figure 4-5). Next, I examined training-related changes separately in the two age
groups using repeated-measures ANOVA across three assessments (pre, mid and post).
The younger group showed significant improvements in overall accuracy (F (2, 28) =
23.70, p < .001) and no change in reaction time with training (F (2, 28) = .23, p = .79).
102

Similarly, the older group showed significant improvements in accuracy (F (2, 27) =
17.63, p < .001) and no change in reaction time (F (2, 27) = 1.11, p = .34). These results
confirm that both younger and older participants improved in response inhibition
accuracy with training (see Figure 4-5). Importantly, training-related improvements in
accuracy were not accompanied by changes in reaction time in either group. However,
the data suggest that older participants may have relied more on slowing of non-target
responses to achieve accurate response inhibition at all assessments.
Influence of Previous Meditation Experience on Performance
I found that older participants were significantly slower than younger participants at
all time points. Although this result could reflect changes in response speed that occur
during the normal ageing process, I also wanted to explore the influence of other
demographic factors on this result. Specifically, older participants entered training with
more previous meditation experience (on average) compared to younger participants.
Thus, presumed age-related differences in response speed could instead reflect the
influence of long-term meditation practice.
Previous meditation and performance before training. Previous meditation
experience (N = 58 with complete data, M = 2572 total hours) was correlated with both
reaction time (r = .44, p < .001) and age (r = .52, p < .001) at pre assessment.
Furthermore, the correlation between previous meditation and reaction time held up after
controlling for the effect of age (pr = .30, p = .02). This result indicates that participants
with more meditation experience performed more slowly on the response inhibition task.
Previous meditation was not related to accuracy (r = .11, p = .39).

103

a.

b.

*p < .05 **p < .01

Figure 4-5. Age effects on accuracy and reaction time. (a) Response inhibition
accuracy (A') and (b) non-target reaction time (RT) plotted as a function of assessment
for younger (n = 30, M = 36 years) and older (n = 29, M = 59 years) participants. Both
age groups showed significant increases in A', but no changes in RT with training. Older
participants were significantly slower than younger participants at all assessments, but the
age groups did not differ in response inhibition accuracy.

0.76
0.8
0.84
0.88
0.92
0.96
1
PRE MID POST
A
c
c
u
r
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(
A
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Younger Older
300
400
500
600
700
PRE MID POST
N
o
n
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t
a
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g
e
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r
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Younger Older
** ** **
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Previous meditation and training-related changes in accuracy. I explored the
influence of previous meditation experience on training outcomes by examining training-
related changes in response inhibition accuracy separately in participants with less
experience (N = 29, M = 933 hours, range = 200 1700 hours) and participants with
more experience (N = 29, M = 4211 hours, range = 1800 15,000 hours). Repeated-
measures ANOVA revealed that accuracy increased across assessments (pre, mid, and
post) both for participants with less experience (F (2, 27) = 29.42, p < .001) and for
participants with more experience (F (2, 27) = 19.36, p < .001). This result suggests that
training-related improvements in response inhibition accuracy were not influenced by
previous meditation experience.
Discussion
In a longitudinal wait-list controlled study of meditation training, I found replicable
improvements in accuracy during sustained response inhibition. Improvements in
accuracy were greatest toward the end of the 32-minute response inhibition task (see
Figures 4-2a and 4-3), indicating that participants were better able to maintain accurate
performance over extended periods of time after training.
These results suggest training-related improvements in executive control, specifically
the voluntary control of impulsive responses. However, improvements in response
inhibition accuracy could also reflect training-related changes in perceptual
discrimination. Previous vigilance studies have reported increases in accuracy and
reductions in the vigilance decrement when targets are easy to discriminate from non-
targets (e.g., Parasuraman & Mouloua, 1987); however, the ease of target discrimination
does not improve performance when response inhibition is required. For example,
105

response inhibition errors are common in vigilance tasks that present perceptually
unambiguous (e.g., alphanumeric) stimuli (e.g., Conners, 1995). In addition, participants
commit response inhibition errors despite being perceptually aware of the rare target
(e.g., the 'Oops' phenomenon; see Robertson et al., 1997). These findings suggest that
improvements in response inhibition accuracy were not likely due to improvements in
perception. Indeed, controlling for changes in baseline target discrimination (i.e., the
measure of threshold reported in Chapter 3) in the analyses of response inhibition
performance did not change the reported results in either retreat, suggesting that
improvements in response inhibition accuracy were not primarily due to changes in
perceptual discrimination. However, it will be important in ongoing analyses to
characterize changes in both perceptual and response-related brain activity that are
correlated with behavioral improvements in response inhibition accuracy (see Chapter 5).
There is growing interest in both research and clinical fields in the possibility for
flexible changes in cognitive ability throughout the life span. In particular, researchers
have attempted to develop cognitive training programs (e.g., Mahncke, Connor et al.,
2006) for ameliorating age-related declines in perception, attention, and cognition (see
reviews in C. S. Green & Bavelier, 2008; Mahncke, Bronstone, & Merzenich, 2006). The
present finding that meditation training improves response inhibition accuracy in older as
well as younger participants is promising (see Figure 4-2). Healthy ageing is
characterized by a natural loss of volume and white matter structural integrity,
particularly in prefrontal cortex, which may be associated with cognitive decline (see
Caserta et al., 2009 for review). Evidence from structural brain studies of long-term
meditators suggests that meditation may reverse this pattern. Lazar and colleagues (2005)
106

demonstrated increased cortical thickness in long-term meditators compared to
meditation-nave controls. In particular, the group differences in prefrontal cortical
thickness were most pronounced in older participants, suggesting that meditation may
attenuate normal age-related deterioration in frontal brain structures. In addition, a recent
study demonstrated that changes in gray matter volume with long-term meditation are
also evident in other areas of frontal cortex implicated in emotion regulation and response
inhibition (Luders, Toga, Lepore, & Gaser, 2009). Although such cross-sectional studies
cannot definitively link meditation training with changes in brain structure, the results
suggest an intriguing connection between the improvements in behavioral response
inhibition observed in older adults in the present study and possible structural and/or
functional changes prefrontal cortex. To test this hypothesis, further research is needed
that tracks longitudinal changes in both behavioral measures of executive control and
changes in brain structure and function in the same participants.
In addition to response inhibition accuracy, I also explored changes in reaction time
with training. Researchers have observed speed-accuracy trade-offs during response
inhibition tasks such that faster responding is linked to a greater likelihood of accidental
errors and slower responding enables better inhibition (Garavan, Hester, Murphy,
Fassbender, & Kelly, 2006; Helton, 2009; Helton et al., 2009; Manly et al., 1999).
Consistent with these findings, I observed a significant correlation between increases in
accuracy and increases in reaction time at post assessment. However, reaction times did
not change with training, suggesting that improvements in accuracy could not be
explained by strategic changes in response speed. I did observe that reaction time was
related to age: Older participants were slower than younger participants at all testing
107

points. This result may reflect normal age-related declines in speed of information
processing (Caserta et al., 2009) or strategic changes in response speed to enable better
inhibitory control. Importantly, older participants did not differ from younger participants
in accuracy before training, nor in amount of improvement in accuracy with training. It is
unclear why meditation training improved accuracy in older participants but did not
affect response speed. Considering the possible relationship between meditation and age-
related changes in prefrontal brain structure discussed above, it will be necessary in
future research to explore how training-related changes in response inhibition accuracy
and response speed map differently onto daily life functioning in older adults.
Slower reaction times were also independently correlated with more previous
meditation experience after controlling for the influence of age. Participants entered the
study with experience in various kinds of meditation practice, and thus it is not possible
to attribute slowing of reaction time to any particular type of training. In addition, slower
reaction times may not be related to meditation per se but rather to life experiences
and/or characteristics specific to individuals who have chosen to dedicate much of their
life to regular meditation practice (see C. S. Green & Bavelier, 2008, for a general
discussion of this topic). In contrast, previous meditation experience was not related to
response inhibition accuracy before training. Thus, the observed changes in accuracy
after Shamatha training are likely mediated by a different mechanism than possible
changes in response speed with long-term meditation.
Executive attention and response inhibition are core dimensions of adaptive, goal-
directed behavior (Miller & Cohen, 2001), and training-related improvements on the
response inhibition task may translate into beneficial changes in daily life. This
108

possibility is supported by previous research showing a relation between response
inhibition performance in the laboratory and attentional and behavioral mistakes in daily
life (Robertson et al., 1997). Motivated by the prediction that response inhibition
performance might be related to positive changes outside the laboratory, my colleagues
and I explored the influence of training-related improvements in response inhibition on
socio-emotional functioning (Sahdra et al., under review). We operationalized self-
reported adaptive functioning (AF) as a single latent factor underlying self-report
measures of anxious and avoidant attachment, mindfulness, ego resilience, empathy, the
five major personality traits, difficulties in emotion regulation, depression, anxiety, and
psychological well-being (all assessed by a battery of questionnaires that participants
completed before and after training). Participants in both retreats showed significant
improvements in AF with training. Longitudinal dynamic models with data combined
across retreats showed that improvement in response inhibition accuracy positively
influenced the change in AF over time. This finding is consistent with a key claim in the
Buddhist literature that enhanced inhibitory control is an important precursor of positive
subjective changes experienced following meditation training. Importantly, these results
suggest that the observed improvements in behavioral response inhibition contribute to
improvements in mental, emotional and psychological functioning.
To conclude, improvements in response inhibition accuracy were observed in two
training studies, suggesting a strong link between Shamatha meditation training and
improvements in executive attention and behavioral control. Moreover, improvements in
response inhibition accuracy endured several months after the completion of formal
training. Although the demographic factors of age and previous meditation experience
109

influenced aspects of task performance (reaction time), improvements in accuracy were
not limited to a particular subset of participants (i.e., all results remained significant after
controlling for individual differences in reaction time, age and previous meditation
experience). Future studies with larger sample sizes will be required to fully characterize
these improvements in individuals of different ages and with varying amounts of previous
experience with meditation. Nevertheless, these findings suggest that flexible changes in
executive attention and behavioral control are possible throughout the life span.
















110











CHAPTER 5

CONCLUSION










111

The overarching aim of the experiments contained in this dissertation was to
demonstrate that vigilance can be improved in healthy adults. In this chapter, I will
provide an overview of the main results and discuss the neural mechanisms that may
underlie the observed training-related improvements in sustained performance.
The series of experiments in Chapter 1 explored the effects of attentional cues on
sustained attention and sustained response inhibition. In the sustained attention task,
exogenous attention cues (sudden-onset luminance changes) increased overall perceptual
sensitivity, while endogenous attention cues (predictable stimulus timing) reduced the
perceptual sensitivity decrement. These results suggest that (1) overall resource demands
can be reduced by manipulations that transiently improve target perception (exogenous
cues), and (2) limited resources can be conserved by using implicit timing information
(endogenous cues) to pay maximum attention at certain moments in time. The best
performance was achieved in the version of the sustained attention task that combined
both types of cues, demonstrating a strong interaction between exogenous and
endogenous attention during vigilance. On the other hand, improvements in perception
due to exogenous cues did not facilitate better sustained response inhibition. This result
suggests that inhibiting impulsive behavioral responses for extended periods of time is
more resource-demanding than sustained target discrimination. Finally, performance on
the most resource-demanding versions of the sustained attention and response inhibition
tasks did not change with repeated testing, indicating that the detrimental effects of high
resource demands are not reduced by increased familiarity with task stimuli or more
practice with executing correct responses.
112

The series of experiments in Chapters 3 and 4 showed that intensive mental training
of voluntary attention through meditation improved performance on the most resource-
demanding vigilance tasks. Training led to increases in overall perceptual sensitivity and
reductions of the vigilance decrement during sustained attention (Chapter 3). These
improvements were specifically linked to training-induced changes in visual perception,
indicating that sustained attention can be improved by reducing the resource demand
associated with difficult target discrimination. Training also improved accuracy on a
vigilance task that required response inhibition (Chapter 4). Improvements in response
inhibition accuracy were not accompanied by adjustments in reaction time (i.e., there was
no speed-accuracy trade-off), indicating that participants were able to inhibit habitual
responses without sacrificing speed.
The observed behavioral improvements are consistent with aspects of meditation
training that entail learning to (1) stabilize attention on a chosen stimulus (sustained
attention), (2) enhance the perceived detail of that stimulus (perceptual discrimination),
and (3) regulate and control attention and behavior (response inhibition). Now it will be
important to elucidate the underlying neural mechanisms of meditation-related
improvements in each of these behavioral domains. Such investigations will also
contribute to a better understanding of the brain resources required for successful
vigilance. During the training studies presented in Chapters 3 and 4,
electroencephalograph (EEG) activity was recorded continuously while participants
performed the behavioral tasks. In the following I will present preliminary EEG results
and discuss how changes in brain activity may support training-related changes in
behavior.
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Perceptual Processing and Stimulus-Evoked Neural Activity
Behavioral studies of visual attention have shown that endogenous and exogenous
attention cues improve perceptual sensitivity by enhancing the quality of stimulus
representations. For example, attention enhances apparent contrast (Carrasco et al., 2004;
Carrasco et al., 2000; Ling & Carrasco, 2006) and increases spatial resolution (Carrasco,
Williams, & Yeshurun, 2002b; Yeshurun & Carrasco, 1999), particularly when a target is
difficult to discriminate. Changes in stimulus-evoked brain electrical activity (event-
related potentials [ERPs]) have been used to describe time-related modulations of brain
activity that underlie such attention-related improvements in perception. In general, ERP
studies of visual-spatial selective attention have shown that representations of attended
visual stimuli are enhanced relative to ignored stimuli at early stages of visual processing
(i.e., within the first 200 milliseconds after stimulus presentation). Specifically, voluntary
attention increases the amplitude of early-latency visual ERPs (P1 and N1) to the
attended stimulus (Mangun & Hillyard, 1991; Martinez et al., 2001; Van Voorhis &
Hillyard, 1977). Moreover, attention-related changes in perceptual sensitivity have been
linked to early-stage perceptual processing rather than later-stage decision processing
(Luck et al., 1994). Enhancements in the amplitude of early-latency components also
occur with focused (versus distributed) attention at the fovea (Miniussi et al., 2002),
demonstrating that directed attention can enhance perceptual processing when spatial
resolution is high.
In addition to early perceptual processing, attention also modulates post-perceptual
target processing. For example, the P300 (also referred to as the P3) is a positive-going
ERP that occurs between 300 and 600 ms after the correct detection of a task-relevant
114

target. The amplitude of the P3 varies according to the amount of attentional resources
devoted to the task, with greater attentional resources resulting in larger amplitude. The
P3 is also modulated by various task manipulations such as target frequency (larger for
infrequent targets), stimulus intensity (smaller for degraded targets), and uncertainty
about target/non-target categories (smaller amplitude with greater uncertainty) (see
Soltani & Knight, 2000 for a review).
Findings from ERP studies of vigilance support the general conclusion that changes
in behavioral performance during vigilance are related to changes in post-perceptual
target processing (see review in Parasuraman, Warm, & See, 1998). For example, Davies
and Parasuraman (1977) found reductions in both early- and late-latency visual ERPs
with increasing time on task; however, only changes in the relatively late components,
including the P3 were related to the behavioral performance decrement. Decreases in P3
amplitude during vigilance could reflect reductions in attentional resources or increasing
uncertainty associated with target identification. However, there is unfortunately little
evidence that demonstrates that changes in stimulus-evoked activity are necessary to
produce the vigilance decrement.
Linking changes in stimulus-evoked brain activity with training-related changes in
sustained attention will advance our understanding of the processing resources that are
required for successful vigilance. Although the studies mentioned above suggest that the
normal vigilance decrement is related to changes in post-perceptual target processing,
training-related improvements in target perception (i.e., perceptual sensitivity during
vigilance) may instead reflect changes in early-stage perceptual processing. Preliminary
analyses I have conducted indicate training-related increases in the amplitude of the N1
115

component following correctly detected targets (vs. non-targets) (MacLean, Aichele,
Bridwell, Jacobs et al., 2009). The N1 component reflects a discrimination process that is
applied within the focus of attention (Vogel & Luck, 2000). Thus, increases in N1
amplitude with training indicate improvements in perceptually discriminating targets
from non-targets. The focus of my ongoing analyses is to describe how changes in N1
amplitude relate to the observed behavioral improvements in baseline threshold, overall
perceptual sensitivity, and sustained performance.
Error-related Processing during Response Inhibition
In the preceding section I discussed how changes in stimulus-locked neural activity
may underlie improvements in target perception during sustained attention. However,
improvements in response inhibition accuracy are not likely mediated by the same
mechanism. Rather, because successful response inhibition requires controlling impulsive
motor responses and because accidental errors occur even after training, improvements in
response inhibition accuracy may be related to changes in error- and response-related
processing. Functional neuroimaging studies have shown that errors in response
inhibition result in recruitment of the anterior cingulate cortex (ACC), which is involved
in monitoring for conflict (e.g., the conflict between a habitual response and a task-
relevant response) and signaling the need for increased attentional control (Carter et al.,
1998; MacDonald, Cohen, Stenger, & Carter, 2000). Several lines of evidence (Dehaene,
1994; van Veen & Carter, 2002a, 2002b) point to the ACC as the source of the error-
related negativity (ERN), a negative deflection in the ERP that peaks between 50 and 150
milliseconds after a subject makes an erroneous response (Falkenstein, Hohnsbein,
Hoormann, & Blanke, 1991). Training-related improvements in response inhibition
116

accuracy may depend on being able to effectively recover from accidental errors by
making attentional and behavioral adjustments. Specifically, enhancements in the
amplitude or changes in the latency of the ERN could play a role in signaling increases in
attentional control after an error has been committed, thus preventing subsequent errors
and improving overall accuracy.
Individuals are usually aware of response inhibition errors, as evidenced by the
common utterance of Oops! (or other expressions of frustration) following an
accidental response to the infrequent stimulus (Robertson et al., 1997). While the ERN
does not appear to reflect conscious error processing, the error positivity (Pe) that occurs
at a later latency (between 200 and 400 milliseconds after an erroneous response) is
modulated by error awareness (Endrass, Franke, & Kathman, 2005; Endrass, Reuter, &
Kathmann, 2007; Nieuwenhuis, Ridderinkhof, Blom, Band, & Kok, 2001; O'Connell et
al., 2007). Changes in the amplitude or latency of the Pe component may thus provide a
window into changes in the subjective awareness of errors following training. In ongoing
analyses I am examining how training-related changes in subconscious (ERN) and
conscious (Pe) error processing contribute to improvements in response inhibition
accuracy.
Modulations of Ongoing EEG
As discussed above, the evoked neural response to discrete events reveals much about
how attention enhances perceptual processing and how errors are processed in the brain.
At the same time, the ongoing EEG also reflects changes in attention during task
performance. The EEG is composed of several frequency bands, some of which show
consistent modulations in oscillatory power (i.e., amplitude) in correspondence with
117

space- and feature-based attention, attentional load/effort, and conscious perception (see
review in Womelsdorf & Fries, 2007). A rapidly growing body of evidence suggests that
modulations in the amplitude of the alpha band (8-14 Hz) over occipital cortex relate to
visual-spatial attention. First, reductions in pre-stimulus alpha are spatially specific and
reflect the location of attention (i.e., reductions in alpha are observed contralateral to the
attended location in space; Kelly, Lalor, Reilly, & Foxe, 2006; Thut, Nietzel, Brandt, &
Pascual-Leone, 2006; Worden, Foxe, Wang, & Simpson, 2000; Yamagishi, Goda, Callan,
Anderson, & Kawato, 2005). Second, decreases in alpha amplitude predict increases in
target detection accuracy (Ergenoglu et al., 2004; Hanslmayr et al., 2007; Hanslmayr et
al., 2005; Linkenkaer-Hansen, Nikulin, Palva, Ilmoniemi, & Palva, 2004; Thut et al.,
2006; van Dijk, Schoffelen, Oostenveld, & Jensen, 2008). Together, these findings
suggest that lower levels of occipital alpha reflect the positive top-down influence of
voluntary attention on visual information processing.
Changes in ongoing EEG during vigilance are generally characterized by a shift
toward slower-frequency bands with increasing time on task (see reviews in Oken,
Salinsky, & Elsas, 2006; Parasuraman et al., 1998). For example, increases in the theta
band (4-8 Hz) have been reported during auditory (Paus et al., 1997) and visual vigilance
tasks (Smit, Eling, & Coenen, 2004a). However, despite the strong link between alpha
activity and visual attention/perception, there is little evidence that the vigilance
decrement is related to changes in alpha activity. One study of the effects of sleep
deprivation on vigilance showed that increases in alpha reliably predicted subsequent
target misses during a 40-minute vigilance task (Kaida, Akerstedt, Kecklund, Nilsson, &
Axelsson, 2007). However, because sleepiness was explicitly manipulated in this study, it
118

is unclear whether these findings reflect changes in overall arousal or visual attention.
More promising are results from a recent study showing that increases in pre-stimulus
alpha predicted subsequent response inhibition errors (Mazaheri et al., in press).
Specifically, errors were related to increases in alpha oscillations (~10 Hz) over occipital
and sensorimotor scalp regions. This finding suggests that the state of the brain before the
occurrence of a stimulus is critical for successful target discrimination (i.e., visual
processing) and subsequent behavior (i.e., sensorimotor processing).
Changes in ongoing brain activity have been hypothesized to underlie meditation-
related changes in perception and attention. However, a recent longitudinal study of
meditation training failed to find changes in pre-stimulus EEG patterns related to
successful attentional performance (Slagter et al., 2009). In contrast, my preliminary
analyses indicate reliable training-related reductions in pre-stimulus alpha (8 14 Hz)
during sustained attention and response inhibition (MacLean, Aichele, Bridwell, Jacobs
et al., 2009). Moreover, training-related reductions in pre-stimulus alpha over occipital
cortex predicted the amount of improvement in vigilance (i.e., reduction in the slope of
performance over time) in both tasks. These results indicate a strong link between pre-
stimulus alpha activity and successful performance over extended periods of time. The
aim of my ongoing analyses is to relate changes in pre-stimulus EEG with behavioral
indices of target discrimination and response speed, as well as to explore how changes in
pre-stimulus EEG relate to changes in stimulus- and response-locked ERPs.
Conclusions
The results from this dissertation demonstrate improvements in perception, sustained
attention, and response inhibition after three months of intensive meditation training. In
119

addition, the preliminary analyses discussed above indicate neurophysiological changes
that are associated with improved behavioral performance. These findings add to the
growing body of evidence that meditation training can improve behavioral performance
(see review in Lutz, Slagter et al., 2008) and lead to enduring changes in brain function
(Brefczynski-Lewis et al., 2007; Lutz, Brefczynski-Lewis, Johnstone, & Davidson, 2008;
Lutz, Greischar, Rawlings, Ricard, & Davidson, 2004; Slagter et al., 2009). Indeed, the
persistence of improvements in discrimination and response inhibition after the
completion of formal training points to the potential for long-lasting behavioral changes
with regular, but less intensive meditation practice. Finally, my ongoing collaborative
work shows that improvements in response inhibition during the formal training period
predicted positive changes in adaptive socio-emotional function (Sahdra et al., under
review), suggesting that the ability to control impulsive behavioral responses in the
laboratory contributes to self-reported psychological wellbeing. It will be important in
future research to continue to explore how training-related changes in behavior and brain
function relate to the more lofty goals of meditation practice, such as learning to regulate
ones emotions and achieving enhanced psychological wellbeing (Wallace & Shapiro,
2006).
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