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Abstract
Here, we update our 1990 Annual Review of Neuroscience article, The
Attention System of the Human Brain. The framework presented in
the original article has helped to integrate behavioral, systems, cellular,
and molecular approaches to common problems in attention research.
Our framework has been both elaborated and expanded in subsequent
years. Research on orienting and executive functions has supported the
addition of new networks of brain regions. Developmental studies have
shown important changes in control systems between infancy and childhood. In some cases, evidence has supported the role of specic genetic
variations, often in conjunction with experience, that account for some
of the individual differences in the efciency of attentional networks.
The ndings have led to increased understanding of aspects of pathology and to some new interventions.
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Contents
INTRODUCTION . . . . . . . . . . . . . . . . . .
THE ORIGINAL NETWORKS . . . . .
Alerting . . . . . . . . . . . . . . . . . . . . . . . . . . .
Orienting . . . . . . . . . . . . . . . . . . . . . . . . . .
Executive . . . . . . . . . . . . . . . . . . . . . . . . . .
ELABORATIONS OF THE
FRAMEWORK . . . . . . . . . . . . . . . . . . .
Alerting . . . . . . . . . . . . . . . . . . . . . . . . . . .
Orienting . . . . . . . . . . . . . . . . . . . . . . . . . .
Executive Control . . . . . . . . . . . . . . . . .
EXTENDING THE
FRAMEWORK . . . . . . . . . . . . . . . . . . .
Self-Regulation . . . . . . . . . . . . . . . . . . . .
Differences in Network Efciency . .
Training . . . . . . . . . . . . . . . . . . . . . . . . . .
Evolution . . . . . . . . . . . . . . . . . . . . . . . . . .
FUTURE . . . . . . . . . . . . . . . . . . . . . . . . . . . .
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INTRODUCTION
Twenty years ago, when neuroimaging was in
its infancy, we summarized the current state
of knowledge on attention in the 1990 volume
of the Annual Review of Neuroscience (Posner &
Petersen 1990). At that time, most available evidence was from behavioral studies of normal
adults or patients with varying forms of brain injury. However, the ability to image brain activity with positron emission tomography seemed
to hold great promise for the physiological analysis of mental processes, including attention. In
our review, we were able to integrate ndings of
the initial imaging studies. We never imagined
that the growth of cognitive neuroscience over
the subsequent 20 years would make it possible
to revisit our analysis, with 4,0006,000 imaging papers on attention or cognitive control and
nearly 3,500 citations of our original review.
The original review suggested three basic
concepts about the attention system. The rst
is that the attention system is anatomically separate from processing systems, which handle incoming stimuli, make decisions, and produce
outputs. We emphasized the sources of the attentional inuences, not the many processing
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Alerting
The concept of arousal goes back to the classic work of Moruzzi & Magoun (1949) on the
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role of the brain stem reticular system in maintaining alertness (Figure 1, for macaque brain).
As more became known of the neuromodulatory systems of the brain stem and thalamus,
it was necessary to qualify the general concept
of arousal into more differentiated components.
Within cognitive psychology, a major emphasis
has been on producing and maintaining optimal
vigilance and performance during tasks; this is
the sense of alertness that we discussed in our
1990 article.
One approach to the study of alerting is to
use a warning signal prior to a target event to
produce a phasic change in alertness. The warning cue leads to replacing the resting state with
a new state that involves preparation for detecting and responding to an expected signal. If a
speeded response is required to the target, reaction time improves following a warning. This
improvement is not due to the buildup of more
accurate information about the target, which
is not changed by the warning signal, but the
warning signal does change the speed of orienting attention and thus responding to the signal.
Several other methods have been used to
study tonic alertness. These include changes
over the course of the day (circadian rhythm).
Reaction times are usually longer in the early
morning and decline over the course of the day
only to rise again during the night and peak in
the early morning (Posner 1975). These measures reect other diurnal changes such as body
temperature and cortisol secretion. A long established approach to tonic alertness is to use
a long and usually rather boring task to measure sustained vigilance. Some of these tasks
have grown out of the job of radar operators looking for near-threshold changes over
long periods of time. Vigilance tasks rely heavily on mechanisms of the right cerebral cortex
(Posner & Petersen 1990). Both classical lesion
data and more recent imaging data conrm that
tonic alertness is heavily lateralized to the right
hemisphere.
Alerting
Locus coeruleus:
norepinephrine
Figure 1
The locus coeruleus projections of the alerting system shown on a macaque
brain. The diffuse connections interact with other, more strongly localized
systems. The alerting system also includes regions of the frontal and parietal
cortices (not shown). Reproduced from Aston-Jones & Cohen (2005).
Orienting
Executive
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b
a
Executive control
dACC/msFC
Orienting
mCC
Precuneus
IPS/SPL
Precuneus
aPFC
FEF
Thalamus
Thalamus
IPS
dFC
dlPFC
IPL
aPFC
VFC
(IFg/MFg)
TPJ
(IPL/STG)
dFC
dlPFC
aPFC
al/fO
al/fO
Cingulo-opercular system:
task set maintenance
Figure 2
(a) The dorsal and ventral orienting networks (after Corbetta & Shulman 2002). The dorsal attention network (light green) consists of
frontal eye elds (FEF) and the intraparietal sulcus/superior parietal lobe (IPS/SPL). The ventral attention network (teal ) consists of
regions in the temporoparietal junction (TPJ) and the ventral frontal cortex (VFC). (b) Two networks of the executive control system.
The circled region indicates the original member of the executive control system from Posner & Petersen (1990). The remaining
regions come from the elaboration of the original cingulo-opercular system (black) and the addition of the frontoparietal system ( yellow)
(adapted from Dosenbach et al. 2007). (c) Resting-state correlation reecting separate control systems. The gure illustrates three views
of the brain (left, dorsal view; middle, tilted lateral view; right, medial view). These separable resting networks are consistent with the
distinctions based on functional criteria exhibited in panels a and b: dorsal attention ( green), ventral attention (teal ), cingulo-opercular
(black), frontoparietal ( yellow) (adapted from Power et al. 2011).
Petersen
Posner
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ELABORATIONS OF THE
FRAMEWORK
The intervening 20 years since our original
article have produced a surprising amount of
support for the basic outlines of the framework
described above. There has also been a signicant amount of elaboration or evolution of the
ideas during that timeframe. The next three
sections review some of the studies deepening
or expanding our understanding of the original
networks.
Alerting
Our understanding of the physiology and
pharmacology underlying the alerting system
has changed signicantly. For example, strong
evidence relates the neuromodulator norepinephrine (NE) to the alerting system. A warning signal is accompanied by activity in the locus coeruleus, the source of NE (Aston-Jones
& Cohen 2005). Warning-signal effects can be
blocked by drugs such as guanfacine and clonodine, which decrease NE release (Marrocco &
Davidson 1998). Drugs that increase NE release can also enhance the warning-signal effect. The NE pathway includes major nodes in
the frontal cortex and in parietal areas relating
to the dorsal but not the ventral visual pathways
(Morrison & Foote 1986).
To examine the specicity of these effects
to the warning signal, researchers used a cued
detection task with humans, monkeys, and
rats (Beane & Marrocco 2004, Marrocco &
Davidson 1998) to separate information about
where a target will occur (orienting) from when
it will occur (alerting). To accomplish this, one
of four cue conditions was presented prior to
a target for a rapid response. By subtracting a
double cue condition, where the participant is
informed of when a target will occur but not
where, from a no cue condition, they receive
a specic measure of the warning inuence
of the signal. When the cue that indicates
the targets location is subtracted from an
alerting cue, the difference represents effects of
orienting. Results of drug studies with humans
and monkeys show that NE release inuences
alerting effects, whereas drugs inuencing
the neuromodulator acetylcholine (Ach) affect
orienting but not alerting. Studies have shown
that individual differences in alerting and
orienting are largely uncorrelated (Fan et al.
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Orienting
In a series of imaging experiments using cuing methodology in combination with
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Executive Control
Our third original network has been elaborated considerably. As noted above, our original
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start +
+ stop
Sustained
fixate
Correct
trials
Cue
Errors
Cue
Task-related (adjusted)
Trial-related
0.4
0.4
Error
Correct
0.3
0.3
0.2
Transient
taskinitiation
signal
0.1
0.2
% change
% change
fixate
0.1
0.1
0.1
1
11
21
31
41
Figure 3
Executive control signals. The top panel shows three putative executive control signals: a task initiation signal in yellow, a taskmaintenance signal in red, and activity related to correct (black) and error (blue) trials (adapted from Dosenbach et al. 2006). Regions
showing differences in error versus correct trials are considered to be computing or receiving performance feedback. The bottom gure
shows activity in the left anterior insula during a task that contains all the putative signals (plus a transient transition signal at the end of
the block of trials). MR, magnetic resonance.
stable background maintenance for task performance as a whole. The frontoparietal system,
in contrast, showing mostly start-cue signals, is
thought to relate to task switching and initiation
and to adjustments within trials in real time.
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The addition of two separate orienting networks and two separate executive networks
raises the possibility that additional control networks will be elaborated in the future. However, for several reasons, we do not expect the
number of control networks to be much larger
than the number described here. The study
of many complex systems, from ecosystems to
protein-protein interactions, seems to indicate
that these systems follow a rule of hand and
have approximately ve controlling variables
(ranging from three to seven) (Gunderson &
Holling 2002). For example, the maintenance
of upright balanced posture appears to be controlled by at least three separate systems: vision, the vestibular system, and kinesthetic joint
sensors. These systems act relatively independently and have different spatial and temporal
characteristics. From this perspective, the presence of ve relatively separate attention networks appears reasonable. A second argument
in favor of this view is an empirical one. In
a recent large-scale study of resting state networks (Power et al. 2011), with effectively all
the brain represented, all the cortical networks,
found by the more piecemeal approaches described above, are present.
Self-Regulation
The ability to control our thoughts, feelings,
and behavior in developmental psychology
is called self-regulation; with adults it is
often called self-control (see sidebar on Will,
Self-Regulation, and Self-Control for further
denitions). Neuroimaging presents strong
evidence that conict tasks such as the Stroop
effect activate common areas of the anterior
cingulate gyrus: the dorsal portion for more
strictly cognitive tasks and the ventral area for
emotion-related tasks (Botvinick et al. 2001,
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(incongruent) direction as the target. Cues presented prior to the target provide information
on where or when the target will occur. There
are strong individual differences in each attention network and there are surprisingly low correlations between these network scores (Fan
et al. 2002), although the networks interact in
more complex tasks and in everyday life (Fan
et al. 2009).
Normal functions including attention are
undoubtedly inuenced by many genes in
complex interaction with epigenetic and environmental factors. Most studies have involved
various pathologies and have not centered on
common human functions; hence relatively
little is known about the full range of genes
involved in attention networks. One strategy
would be to use emerging genomic and
epigenomic technologies to carry out studies
of large cohorts using various attention tasks as
phenotypes to determine genes that relate to
performance differences. A more limited
approach, based on what is known about
attention networks, takes advantage of the
association between different neuromodulators
and attention networks to examine specic
genetic alleles (e.g., related to dopamine)
to examine individual performance on the
appropriate network (see Green et al. 2008 for
review). As one example, the ANT has been
used to examine individual differences in the
efciency of executive attention. A number
of polymorphisms in dopamine and serotonin
genes have been associated specically with
the scores on executive attention (Green et al.
2008). This work is still just getting started,
and reports are conicting. One reason for the
conict may be that genetic variations are also
inuenced by environmental factors.
Genetic modulation by environmental factors is perhaps clearest for the dopamine 4 receptor gene (DRD4), which has been associated
with the executive network in adult imaging
studies (Fan et al. 2003b). Data at 1820 months
showed that quality of parenting interacted
with the 7 repeat allele of the DRD4 gene
to inuence the temperamental dimensions of
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Training
Because parenting and other cultural factors
interact with genes to inuence behavior, it
should be possible to develop specic training
methods that can be used to inuence underlying brain networks. Two forms of training
methods have been used in the literature. One
involves practice of a particular attention network. Several such attention training studies
have shown improved executive attention function and produced changes in attention-related
brain areas (Klingberg 2011, Rueda et al. 2005).
The practice of a form of meditation has been
used to change the brain state in a way that
improves attention, reduces stress, and also improves functional connectivity between the anterior cingulate and the striatum (Tang et al.
2007, 2009).
Evolution
The ACC is a phylogenetically old area of the
brain. Comparative anatomical studies point
to important differences in the evolution of
cingulate connectivity between nonhuman
FUTURE
It has been exciting for us to see the expansion of work on networks of attention over the
past 20 years. We now have the opportunity to
go from genes to cells, networks, and behavior
and to examine how these relationships change
from infancy to old age. In development, the
number of active control systems increases and
their inuence changes.
Although much has been learned, many
questions remain unanswered. We are hopeful
that the study of attention will continue to
provide greater understanding of how control
develops typically and in pathology (Posner
2012a, Posner et al. 2011) and will provide
promising leads for translating basic research
into interventions to aid children and families.
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DISCLOSURE STATEMENT
The authors are not aware of any afliations, memberships, funding, or nancial holdings that
might be perceived as affecting the objectivity of this review.
ACKNOWLEDGMENTS
This article was supported in part by grant HD060563 to Georgia State University subcontracted
to the University of Oregon. Prof. Mary K. Rothbart made important contributions to the research
and writing of this review. This article was also supported by NIH grants NS32797 and 61144
and the McDonnell Foundation.
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Annual Review of
Neuroscience
Volume 35, 2012
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p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 309
What Is the Brain-Cancer Connection?
Lei Cao and Matthew J. During p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 331
The Role of Organizers in Patterning the Nervous System
Clemens Kiecker and Andrew Lumsden p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 347
The Complement System: An Unexpected Role in Synaptic Pruning
During Development and Disease
Alexander H. Stephan, Ben A. Barres, and Beth Stevens p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 369
Brain Plasticity Through the Life Span: Learning to Learn and Action
Video Games
Daphne Bavelier, C. Shawn Green, Alexandre Pouget, and Paul Schrater p p p p p p p p p p p p p 391
The Pathophysiology of Fragile X (and What It Teaches Us about
Synapses)
Asha L. Bhakar, Gul
Dolen, and Mark F. Bear p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 417
Central and Peripheral Circadian Clocks in Mammals
Jennifer A. Mohawk, Carla B. Green, and Joseph S. Takahashi p p p p p p p p p p p p p p p p p p p p p p p p 445
Decision-Related Activity in Sensory Neurons: Correlations Among
Neurons and with Behavior
Hendrikje Nienborg, Marlene R. Cohen, and Bruce G. Cumming p p p p p p p p p p p p p p p p p p p p p p 463
Compressed Sensing, Sparsity, and Dimensionality in Neuronal
Information Processing and Data Analysis
Surya Ganguli and Haim Sompolinsky p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 485
The Auditory Hair Cell Ribbon Synapse: From Assembly to Function
Saaid Saeddine, Aziz El-Amraoui, and Christine Petit p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 509
Multiple Functions of Endocannabinoid Signaling in the Brain
Istvan Katona and Tamas F. Freund p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 529
Circuits for Skilled Reaching and Grasping
Bror Alstermark and Tadashi Isa p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 559
Indexes
Cumulative Index of Contributing Authors, Volumes 2635 p p p p p p p p p p p p p p p p p p p p p p p p p p p 579
Cumulative Index of Chapter Titles, Volumes 2635 p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p p 583
Errata
An online log of corrections to Annual Review of Neuroscience articles may be found at
http://neuro.annualreviews.org/
viii
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