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ANRV296-PS58-01 ARI 17 November 2006 1:10

Research on Attention
Networks as a Model for the
Integration of Psychological
Science
Annu. Rev. Psychol. 2007.58:1-23. Downloaded from arjournals.annualreviews.org

Michael I. Posner and Mary K. Rothbart1

by Ball State University on 01/04/09. For personal use only.

Psychology Department, University of Oregon, Eugene, Oregon 97403-1291;

email: [email protected], [email protected]

Annu. Rev. Psychol. 2007. 58:1–23 Key Words

First published online as a Review in attention, candidate genes, orienting, neural networks,
Advance on October 9, 2006
temperament
The Annual Review of Psychology is online at
http://psych.annualreviews.org Abstract
This article’s doi: As Titchener pointed out more than one hundred years ago, at-
10.1146/annurev.psych.58.110405.085516
tention is at the center of the psychological enterprise. Attention
Copyright  c 2007 by Annual Reviews. research investigates how voluntary control and subjective experi-
All rights reserved
ence arise from and regulate our behavior. In recent years, attention
0066-4308/07/0110-0001$20.00 has been one of the fastest growing of all fields within cognitive psy-
1
We appreciate the invitation of the editors chology and cognitive neuroscience. This review examines attention
of the Annual Review of Psychology to submit a as characterized by linking common neural networks with individual
prefatory essay to this year’s volume. We
differences in their efficient utilization. The development of atten-
have taken the opportunity to propose a
unified basis for psychological science based tional networks is partly specified by genes, but is also open to specific
upon an effort to combine experimental and experiences through the actions of caregivers and the culture. We
differential approaches to the field. This
believe that the connection between neural networks, genes, and
article is an improved and expanded version
of an earlier one along these lines (Posner & socialization provides a common approach to all aspects of human
Rothbart 2004), and its developmental cognition and emotion. Pursuit of this approach can provide a basis
aspects are further expanded in a book
for psychology that unifies social, cultural, differential, experimen-
(Posner & Rothbart 2007).
tal, and physiological areas, and allows normal development to serve
as a baseline for understanding various forms of pathology. D.O.
Hebb proposed this approach 50 years ago in his volume Organiza-
tion of Behavior and continued with introductory textbooks that dealt
with all of the topics of psychology in a common framework. Use of
a common network approach to psychological science may allow a
foundation for predicting and understanding human behavior in its
varied forms.

1
ANRV296-PS58-01 ARI 17 November 2006 1:10

At the time Hebb wrote his monograph,

Contents relatively little was known about how the
structure and organization of the central ner-
INTRODUCTION . . . . . . . . . . . . . . . . . 2
vous system contribute to the functions ob-
HEBB’S NETWORK
served in psychological studies. This led Hebb
APPROACH . . . . . . . . . . . . . . . . . . . . 4
to talk in terms of the conceptual nervous
New Tools . . . . . . . . . . . . . . . . . . . . . . . 5
system, that is, ideas about its structure that
IMAGING ATTENTION
might be imagined or inferred from psycho-
NETWORKS . . . . . . . . . . . . . . . . . . . 6
logical studies. We suggest that the meth-
Simulating Attention Networks . . . 8
ods available to Hebb, mostly animal research
INDIVIDUAL EFFICIENCY . . . . . . . 9
and behavioral human experiments, were not
NETWORK DEVELOPMENT . . . . 11
sufficient to provide empirical methods for
Socialization and Culture . . . . . . . . . 13
linking his conceptual nervous system to real
PATHOLOGY OF ATTENTION
Annu. Rev. Psychol. 2007.58:1-23. Downloaded from arjournals.annualreviews.org

events in the human brain. This methodology

NETWORKS . . . . . . . . . . . . . . . . . . . 15
has now been provided by neuroimaging. Al-
Plasticity . . . . . . . . . . . . . . . . . . . . . . . . . 16
though Hebb also recognized the importance
INTEGRATION OF
by Ball State University on 01/04/09. For personal use only.

of studying individual differences in intelli-

PSYCHOLOGICAL SCIENCE . . 17
gence and affect, at that time there were no
Generality . . . . . . . . . . . . . . . . . . . . . . . 17
methods for exploring the specific genes that
are an important source of these differences.
The human genome project has provided new
INTRODUCTION methods for exploring this issue.
Can psychology be a unified science, or must In Hebb’s time, the idea of a network (cell
it remain fragmented and subject to transient assembly or phase sequence) was a rather
research interests rather than cumulative de- vague verbal abstraction that did not allow
velopment? Is psychology teachable or must for models that could produce specific pre-
Attention: the
regulating of various psychology textbooks remain encyclopedias dictions. As a result of the rapid changes in
brain networks by of often unrelated findings? In this article, cellular biology (Bullock et al. 2005) and in
attentional networks we examine research on attention to suggest the mathematics of multilevel networks, this
involved in ways of looking at psychological questions and too has changed (Rumelhart & McClelland
maintaining the alert
findings that might lead to hopeful answers to 1986). Although early versions of these net-
state, orienting, or
regulation of conflict these questions. We offer a model based on works were inspired by simple versions of
Hebbian psychology that reaches out from neurons as all-or-none elements, more recent
Neural networks: a
number of brain biological roots to tackle questions of emo- versions (O’Reilly & Munakata 2000) have be-
areas that when tion and thought as well as behavior. We, with gun to use the details of neuroanatomy and
orchestrated carry many of our colleagues, see different levels cellular structure as provided by imaging and
out a psychological of analysis in psychology as informing each cellular studies to develop networks that take
function
other, with each level of equal scientific valid- more realistic advantage of the structure of
ity. Bridging these levels can allow a higher the human brain. Hebb’s basic idea, together
level of understanding and prediction. with the new methodological tools and new
We argue that D.O. Hebb (see Figure 1), disciplines (e.g., cognitive, affective, and so-
beginning with his monograph in 1949 and cial neuroscience), all based on network views,
continuing through a series of introductory give abundant evidence of the value of em-
textbooks (1958, 1966), has convincingly pre- ploying the converging operations strategy
sented the basis for such integration. This in- advocated by Sternberg (Sternberg 2004).
tegration lies in understanding how genes and It is important to recognize the need for in-
experience shape neural networks underlying tegrating cognitive, affective, and social neu-
human thoughts, feelings, and actions. roscience with psychology because many of

2 Posner · Rothbart
ANRV296-PS58-01 ARI 17 November 2006 1:10
Annu. Rev. Psychol. 2007.58:1-23. Downloaded from arjournals.annualreviews.org
by Ball State University on 01/04/09. For personal use only.

Figure 1
Photograph of Professor Donald O. Hebb.

www.annualreviews.org • Research on Attentional Networks as a Model 3

ANRV296-PS58-01 ARI 17 November 2006 1:10

the theoretical questions that need to be ad- rons and between neurons and other brain
dressed by neuroimaging and genetic studies cells (Bullock et al. 2005). These develop-
are exactly those that a century of psychol- ments have reduced the gap between networks
ogists have explored. The neuroscience ap- revealed in imaging studies and the complex
proach provides crucial constraints for psy- intracellular activity that underlies them. In
chological theories, but it also benefits when particular, they show that learning may reflect
a closer connection is made with the psycho- the activity of interactions at many time scales
logical level of analysis. and may be modified by aspects of the organ-
ism’s overall state.
Hebb also introduced the concept of the
HEBB’S NETWORK APPROACH phase sequence involved in the coordination
In 1949, D.O. Hebb published his epic work, of multiple cell assemblies. He recognized the
The Organization of Behavior. His book was importance of the temporal correspondence
Annu. Rev. Psychol. 2007.58:1-23. Downloaded from arjournals.annualreviews.org

immediately recognized as providing the po- between assemblies. In recent years these
tential for an integrated psychology. One ideas have been supported by studies show-
reviewer (Attneave 1950, p. 633) wrote: ing that synchronization of brain areas may be
by Ball State University on 01/04/09. For personal use only.

critical to detecting stimuli (Womelsdorf et al.

I believe The Organization of Behavior to be 2006) and for transfer of information between
the most important contribution to psycho- remote areas (Nikolaev et al. 2001). In later
logical theory in recent years. Unlike those years, Hebb (1958) developed an introductory
of his contemporaries who are less interested psychology textbook that integrated much of
in psychology than in some restricted as- psychology using the framework provided by
pect thereof to which their principles con- his 1949 theory. He applied his network the-
fine them, Hebb has made a noteworthy at- ory to heredity, learning and memory, motiva-
tempt to take the experimentally determined tion, perception, thought, and development.
facts of behavior, as they are, and account for In later editions, he extended his approach
them in terms of events within the central to emotions in their social contexts, individ-
nervous system. ual differences in intelligence, and abnormal
psychology.
The most important basic idea that Hebb Despite his efforts and those of his fol-
presented was the cell assembly theory out- lowers at McGill and elsewhere, Hebb’s work
lined in chapters 4 and 5 of his book (Goddard was unsuccessful in providing a fully integra-
1980, Harris 2005). Hebb argued that ev- tive psychology, and many still seek to de-
ery psychological event, sensation, expecta- velop such an integration (Sternberg 2004,
tion, emotion, or thought is represented by Sternberg & Grigorenko 2001). One of the
the flow of activity in a set of interconnected major problems with Hebb’s framework was
neurons. Learning occurs by a change in that it left no clear empirical path for the ac-
synaptic strength when a synapse conducts quisition of new knowledge about how hu-
excitation at the same time the postsynap- man brain networks develop, how they dif-
tic neuron discharges. This provided a basis fer among individuals, why they break down,
for the modification of synapses and showed and how they can be restored to functioning.
how neural networks might be organized un- Most of the research by Hebb and his asso-
der the influence of specific experiences. The ciates was performed on nonhuman animals,
Hebb synapse plays a central role in modern while most of psychology concerns human be-
neuroscience (see Kolb 2003, Milner 2003, havior, brain, and mind.
Sejnowski 2003). There are important new In 1955, Hebb argued for the utility of a
developments in the study of synapses and in conceptual nervous system inferred from psy-
the discovery of other influences among neu- chological studies of motivation. This idea

4 Posner · Rothbart
ANRV296-PS58-01 ARI 17 November 2006 1:10

remains of key importance. Despite our make human brain networks accessible for de-
growing understanding of the function and tailed physiological study.
structure of the nervous system, psycholog- Some have thought that the influence of
PET: positron
ical knowledge needs to be used to pro- imaging has been merely to tell us where in emission
pose hypotheses about how the central ner- the brain things have happened (Utall 2001). tomography
vous system (CNS) works to support feelings, Certainly many, perhaps even most, imaging fMRI: functional
thoughts, and behaviors. We argue that only studies have been concerned with anatomical magnetic resonance
within an integrated field of psychology can issues. As Figure 2 illustrates, several func- imaging
knowledge at all levels be used to develop and tions of attention have been shown to involve Temperament:
constrain these hypotheses. specific anatomical areas that carry out impor- relatively enduring
tant functions. However, imaging also probes biological
characteristics of
neural networks that underlie all aspects of
New Tools individuals that
human thought, feelings, and behavior. The include both reactive
Annu. Rev. Psychol. 2007.58:1-23. Downloaded from arjournals.annualreviews.org

Several major late-twentieth century events full significance of imaging for (a) viewing and self-regulatory
give improved prospects for an integration brain networks, (b) examining their compu- (attentional) aspects
of psychological science around the ideas in- tation in real time, (c) exploring how they are
by Ball State University on 01/04/09. For personal use only.

troduced by Hebb. Cell assemblies and phase assembled in development, and (d ) their plas-
sequences are names for aspects of neural net- ticity following physical damage or training is
works. Now, thanks to work on the compu- a common theme in current research that is
tational properties of neural networks (i.e., just beginning to reach its potential.
Rumelhart & McClelland 1986), we are in a A third development, the mapping of the
much better position to develop detailed the- human genome (Venter et al. 2001), offers the
ories integrating information from physiolog- potential for an increased understanding of
ical, cognitive, and behavioral studies. the physical basis for individual differences,
In addition, new neuroimaging methods including individual differences in tempera-
now allow us to examine neuronal activity ment and personality. Many genes exhibit a
in terms of localized changes in blood flow number of relatively high-frequency variants
or metabolism in positron emission tomogra- (polymorphisms) that can code for different
phy (PET) or changes in blood oxygenation physical configurations. These in turn can al-
by functional magnetic resonance imaging ter the efficiency of a network. For exam-
(fMRI). By using tracers that bind to different ple, different types of genes (alleles) form-
transmitters, PET can also be used to examine ing dopamine receptors can lead to different
transmitter density. By measuring electrical efficiency in binding to dopamine and thus
(electroencephalogram, or EEG) and mag- differences in underlying neural networks. In
netic (magnetoencephalogram) signals out- a number of cases, it has been possible to relate
side the skull, the time course of activation these genetic differences to individual per-
of different brain areas localized by fMRI can formance in tasks involving the network (see
be measured (Dale et al. 2000). Pathways of Fossella & Posner 2004 for a review).
activation can also be imaged by use of dif- Genetics-based research also provides an im-
fusion tensor imaging, a form of MRI that portant approach to the development of char-
traces white matter tracts. In addition to the acteristics of neural networks common to all
study of naturally occurring lesions, interrupt- members of the species.
ing information flow by transcranial magnetic We recognize that not all topics of psy-
stimulation (TMS) can produce temporary chology have been sufficiently explored to il-
functional lesions of pathways (see Toga & lustrate this framework and that limitations
Mazziotta 1996 for a review of these and other in our knowledge prevent us from explor-
methods). These methods provide a toolkit ing all of the areas where these develop-
that can be used either alone or together to ments can be applied. However, attention and

www.annualreviews.org • Research on Attentional Networks as a Model 5

ANRV296-PS58-01 ARI 17 November 2006 1:10
Annu. Rev. Psychol. 2007.58:1-23. Downloaded from arjournals.annualreviews.org
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Figure 2
Anatomy of three attentional networks: alerting, orienting, and executive attention (from Posner &
Rothbart 2007).

temperament, areas of our study, can be used tive experiments, behavioral observations, and
to consider how a large number of psycho- self-report as important elements of psycho-
logical questions can be explored using the logical science. We hope to illustrate in this ar-
neural network framework outlined by Hebb. ticle how important such psychological meth-
Attention serves as a basic set of mechanisms ods are and how they can be integrated within
that underlie our awareness of the world and a brain network framework, as Hebb (1955)
the voluntary regulation of our thoughts and illustrated in his Textbook of Psychology.
feelings. The methods used to understand at-
tentional networks in terms of anatomy, indi-
vidual differences, development, and plastic- IMAGING ATTENTION
ity can be applied readily to explore networks NETWORKS
related to other aspects of human behavior. Functional neuroimaging has allowed many
Any approach based on neural networks cognitive tasks to be analyzed in terms of the
raises the issue of crude reductionism. Many brain areas they activate, and studies of atten-
agree that all human behavior must ulti- tion have been among those most often ex-
mately be traceable to brain activity, but cor- amined (Corbetta & Shulman 2002, Driver
rectly argue for the importance of cogni- et al. 2004, Posner & Fan 2007). Imaging

6 Posner · Rothbart
ANRV296-PS58-01 ARI 17 November 2006 1:10

data have supported the presence of three net-

Table 1 A summary of the anatomy and chemical modulators
works related to different aspects of attention
involved in the alerting, orienting, and executive attention
(Fan et al. 2005). These networks carry out
networks.
the functions of alerting, orienting, and exec-
Function Structures Modulator
utive attention (Posner & Fan 2007). A sum-
Orient Superior parietal Acetylcholine
mary of the anatomy and chemical modula-
Temporal parietal junction
tors involved in the three networks is shown in
Frontal eye fields
Table 1. Superior colliculus
Alerting is defined as achieving and main- Alert Locus coeruleus Norepinephrine
taining a state of high sensitivity to incom- Right frontal
ing stimuli; orienting is the selection of in- Parietal cortex
formation from sensory input; and executive Executive attention Anterior cingulate Dopamine
attention involves mechanisms for monitor- Lateral ventral
Annu. Rev. Psychol. 2007.58:1-23. Downloaded from arjournals.annualreviews.org

ing and resolving conflict among thoughts, Prefrontal

feelings, and responses. The alerting system Basal ganglia
has been associated with thalamic as well as
by Ball State University on 01/04/09. For personal use only.

frontal and parietal regions of the cortex (Fan

et al. 2005). A particularly effective way to vary to the lateral intraparietal area (LIP) in
alertness has been to use warning signals prior monkeys, which is involved in the production
to targets. The influence of warning signals of eye movements (Andersen 1989). When Orienting: the
interaction of a brain
on the level of alertness is thought to be due a target occurs at an uncued location and
network with sensory
to modulation of neural activity by the neu- attention has to be disengaged and moved to a systems designed to
rotransmitter norepinephrine (Marrocco & new location, there is activity in the temporal improve the selected
Davidson 1998). parietal junction (Corbetta & Shulman 2002). signal
Orienting involves aligning attention with Lesions of the temporal parietal junction
a source of sensory signals. This may be overt, lobe and superior temporal lobe have been
as when eye movements accompany move- consistently related to difficulties in orienting
ments of attention, or may occur covertly, (Karnath et al. 2001).
without any eye movement. The orienting Executive control of attention is often
system for visual events has been associated studied by tasks that involve conflict, such as
with posterior brain areas, including the various versions of the Stroop task. In the
superior parietal lobe and temporal parietal Stroop task, subjects must respond to the
junction, and in addition, the frontal eye fields color of ink (e.g., red) while ignoring the color
(Corbetta & Shulman 2002). Orienting can word name (e.g., blue) (Bush et al. 2000). Re-
be manipulated by presenting a cue indicating solving conflict in the Stroop task activates
where in space a target is likely to occur, midline frontal areas (anterior cingulate) and
thereby directing attention to the cued loca- lateral prefrontal cortex (Botvinick et al. 2001,
tion (Posner 1980). It is possible to determine Fan et al. 2005). There is also evidence for
the anatomy influenced by the cue separately the activation of this network in tasks involv-
from that influenced by the target by using ing conflict between a central target and sur-
MRI to trace changes in the blood that specif- rounding flankers that may be congruent or
ically follow the cue. This method is called incongruent with the target (Botvinick et al.
event-related functional magnetic resonance 2001, Fan et al. 2005).
imaging, and its use has suggested that the Recently, the role of the anterior cingulate
superior parietal lobe is associated with in modulating sensory input has been demon-
orienting following the presentation of a cue strated experimentally by showing enhanced
(Corbetta & Shulman 2002). The superior connectivity between the anterior cingulate
parietal lobe in humans is closely related cortex and the sensory modality to which the

www.annualreviews.org • Research on Attentional Networks as a Model 7

ANRV296-PS58-01 ARI 17 November 2006 1:10

person is asked to attend (Crottaz-Herbette & condition) that informs the participant either
Menon 2006). This finding supports the gen- that a target will occur soon or where it will
eral idea that anterior cingulate cortex activity occur, or both. The target always occurs either
ANT: attention
network test regulates other brain areas, at least for sen- above or below fixation, and consists of a cen-
sory areas. Experimental tasks may also pro- tral arrow, surrounded by flanking arrows that
RT: reaction time
vide a means of fractionating the contribu- can point either in the same direction (con-
ACT-R: adaptive
tions of different areas within the executive gruent) or in the opposite direction (incon-
control of
thought-rational attention network (MacDonald et al. 2000). In gruent). Subtracting RTs for congruent from
accord with the findings in recent neuroimag- incongruent target trials provides a measure of
ing studies (Beauregard et al. 2001, Ochsner conflict resolution and assesses the efficiency
et al. 2001), we have argued that the exec- of the executive attention network. Subtract-
utive attention network is involved in self- ing RTs obtained in the double-cue condition
regulation of positive and negative affect as that provides information on when but not
Annu. Rev. Psychol. 2007.58:1-23. Downloaded from arjournals.annualreviews.org

well as a wide variety of cognitive tasks under- where the target will occur from RTs in the
lying intelligence (Duncan et al. 2000). This no-cue condition gives a measure of alerting
idea suggests an important role for attention due to the presence of a warning signal. Sub-
by Ball State University on 01/04/09. For personal use only.

in moderating the activity of sensory, cogni- tracting RTs to targets at the cued location
tive, and emotional systems. (spatial cue condition) from trials using a cen-
tral cue gives a measure of orienting, since the
spatial cue, but not the central cue, provides
Simulating Attention Networks valid information about where a target will
Quantification has had a high value in psycho- occur.
logical research. An advantage of the network The attention network task has now been
approach is that it lends itself to the devel- simulated in the framework of the adaptive
opment of precise computer models that al- control of thought-rational (ACT-R) theory
low both summarization of many findings in (Wang et al. 2004). The ANT task is di-
the field and prediction of new findings. Cur- vided into subroutines. Cue processing in-
rently, symbolic models, such as rule-based volves switching of attention to the cued loca-
systems (Newell 1990), appear to be a good tion. Target processing involves detection of
way to capture data from reaction time and the direction of the arrow in the center and in-
other psychological findings. In these models, volves an attention switch and response initia-
cognitive functions are represented as chains tion. Each of these operations is implemented
of production rules and can be identified with by a chain of production rules. The operations
the mental operations postulated by cogni- are similar to those discussed in most psycho-
tive studies. On the other hand, subsymbolic logical studies and localized in neurological
models, such as connectionist models (e.g., studies. For example, the switching of atten-
O’Reilly & Munakata 2000), permit a more tion based on the peripheral cue or target is
biologically realistic implementation of the thought to be implemented by the temporal
operations and closer links to imaging studies. parietal junction (Corbetta & Shulman 2002).
We have developed the attention network A connectionist simulation of the ANT
test (ANT) to examine individual differences (H. Wang & J. Fan, manuscript in prepara-
in the efficiency of the brain networks of alert- tion) is based upon a local error-driven and
ing, orienting, and executive attention dis- associative, biologically realistic algorithm
cussed above (Fan et al. 2002, Rueda et al. (LEABRA) (O’Reilly & Munakata 2000). The
2004). The ANT uses differences in reaction subroutines of ACT-R are now replaced by
time (RT) between conditions to measure the specific connections between hypothesized
efficiency of each network. Each trial begins neurons. These neurons are somewhat real-
with a cue (or a blank interval, in the no-cue istic, and are designed to represent the known

8 Posner · Rothbart
ANRV296-PS58-01 ARI 17 November 2006 1:10

properties of specific brain areas. Thus, the The ability to measure differences in at-
orienting network can be designed to reflect tention among adults raises the question of
the known properties of the frontal eye fields, the degree to which attention is heritable. To
Candidate genes:
superior parietal lobe, and temporal parietal explore this issue, the ANT was used to assess genes that may be
junction, and they can be connected within attention in monozygotic and dizygotic same- involved in the
the simulation. As shown in Figure 3, sim- sex twins (Fan et al. 2001). Strong heritabil- development of
ulations do a reasonable job of fitting with ity was found for the executive network, some utilization of
attentional networks
the known ANT data, although improvement heritability for the orienting network, and no
can be expected in the future. The symbolic apparent heritability for the alerting network.
model makes contact with the mental opera- These data support a search for genes in exec-
tions related to imaging, whereas the connec- utive attention and in orienting of attention.
tionist framework allows a strong treatment We then used the association of the ex-
of the underlying biology. Together they illus- ecutive network with the neuromodulator
Annu. Rev. Psychol. 2007.58:1-23. Downloaded from arjournals.annualreviews.org

trate how network views provide a computa- dopamine (see Table 1) as a way of search-
tional means for summarizing many findings ing for candidate genes that might relate to
within psychology, allowing novel predictions the efficiency of the networks (Fossella et al.
by Ball State University on 01/04/09. For personal use only.

reflecting properties of the network. 2002). To do this, 200 persons performed

the ANT and were genotyped to examine
frequent polymorphisms in genes related to
INDIVIDUAL EFFICIENCY dopamine. We found significant association
Psychology is often divided into two ap- of the dopamine 4 receptor and monoamine
proaches that are almost completely separate oxidase A genes. We then conducted a neu-
in the literature (but see Gardner 1983 for an roimaging experiment in which persons with
effort at integration). The discussion above different alleles of these two genes were com-
focused on general features of the human pared while they performed the ANT (Fan
mind, such as the ability to attend. Another et al. 2003). Groups with different alleles
approach deals with differences among indi- of these genes showed differences in perfor-
viduals. These differences may involve cogni- mance on the ANT and also produced sig-
tion, as in the measurement of intelligence, nificantly different activations in the anterior
or they may involve temperamental differ- cingulate, a major node of the executive atten-
ences, many of which relate to energetic fac- tion network.
tors such as the expression and control of the Recent studies have confirmed and ex-
emotions. Almost all studies of attention have tended these observations to other dopamine
been concerned either with the general abili- genes and to the orienting network. In two dif-
ties involved or with effects of brain injury or ferent studies employing other conflict tasks,
pathology on attention. However, it is clear the catecholamine-O-methyltransferase gene
that normal individuals differ in their ability to was linked to the mental operations re-
attend to sensory events, and it is even clearer lated to resolving conflict (Blasi et al. 2005,
that they differ in their ability to concentrate Diamond et al. 2004). Different alleles of
for long periods on internal trains of thought. cholinergic genes were also related to perfor-
We used the attention network task to ex- mance on orienting tasks such as visual search
amine the individual efficiency of the alerting, (Parasuraman et al. 2005), thus confirming the
orienting, and executive networks (Fan et al. link between orienting and the neuromodula-
2002). In a sample of 40 normal persons, each tor acetylcholine (see Table 1).
of these scores was reliable over repeated pre- Hebb (1955) thought that most of the
sentations. In addition, we found no correla- networks involved in higher functions were
tion among the orienting, alerting, and exec- shaped primarily through experience. We
utive scores. now know that there is a great deal in common

www.annualreviews.org • Research on Attentional Networks as a Model 9

ANRV296-PS58-01 ARI 17 November 2006 1:10

ALERT ORIENT CONFLICT

EXPERIMENTAL RESULTS 47 51 84

ACT-R SIMULATION 55 45 86

LEABRA SIMULATION 70 33 67
Annu. Rev. Psychol. 2007.58:1-23. Downloaded from arjournals.annualreviews.org
by Ball State University on 01/04/09. For personal use only.

1. Fixation &
Output Cue
Executive Control Expectation
Network (2)

Orienting Network Object Pathway 2. Cue or Stimulus

Stimulus? State Switching
(2)

Alerting Network Primary Visual 3. Cue

Cortex Processing
(7)

Next Trial?

4. Stimulus
Visual Input Expectation
(1)

5. Stimulus
Processing
(18)

6. Response
(6)

Figure 3
Experimental results compared with simulations of attention networks from ACT-R (Wang et al. 2004)
and from a connectionist model (H. Wang & J. Fan, manuscript in preparation). The network scores are
in milliseconds.

10 Posner · Rothbart
ANRV296-PS58-01 ARI 17 November 2006 1:10

among humans in the anatomy of these high- tress can be traced to early infancy (Harman
level networks, and thus that they must have et al. 1997). Early in life, most regulation de-
a basis within the human genome. It seems pends on the caregiver providing ways to con-
likely that the same genes that are related to trol infant reactions. In our study, we first in-
individual differences in attention are also im- duced a mild level of distress, and then showed
portant in the development of the attentional how attentional orienting calms that distress
networks that are common among people. while the infant remains engaged with the ob-
Some of these networks are also common to ject. When the orienting is broken, distress re-
nonhuman animals. By examining these net- turns to the level present prior to the introduc-
works in animals, it should be possible to test tion of the object. It is likely that the distress
these assumptions further and to understand remains present and is held by networks in the
the role of genes in shaping networks. amygdala. Parents often use manipulation of
Of importance for this effort is the de- the infant’s orienting to control distress, and
Annu. Rev. Psychol. 2007.58:1-23. Downloaded from arjournals.annualreviews.org

velopment of methods to manipulate relevant the infant also exhibits coping behaviors in-
genes in specific anatomical locations that are volving orienting to form the basis for one
important nodes of a particular network. Usu- aspect of early self-regulation (Rothbart et al.
by Ball State University on 01/04/09. For personal use only.

ally genes are expressed at multiple locations 1992).

so that changes (e.g., gene knockouts) are not Developmental changes in executive atten-
specific to one location. However, using sub- tion found during the third year of life are cor-
tractive genomics, a method is currently be- related with parent reports of temperamental
ing developed to do this (Dumas et al. 2005). effortful control (Gerardi-Caulton 2000). Ef-
We believe that this kind of genetic analysis of fortful control is a broad variable identified in
network development will become a very pro- temperament research; it includes the ability
ductive link between genes and both normal to inhibit a dominant response in order to pro-
and pathological psychological function. duce subdominant response and to detect and
correct errors (Rothbart & Rueda 2005). Be-
cause children of this age do not read, the loca-
NETWORK DEVELOPMENT tion and identity rather than the word mean-
Every parent of more than one child recog- ing and ink color served as the dimensions in
nizes that from birth, infants are individuals a spatial conflict task. In one study (Gerardi-
with their own distinct characteristics and dis- Caulton 2000), children sat in front of two re-
positions. These include reactive traits such as sponse keys, one located to the child’s left and
emotionality, activity level, and orienting to one to the right. Each key displayed a picture,
sensory events, and regulatory traits like at- and on every trial, a picture identical to one
tention focusing and shifting and inhibitory of the pair appeared on either the left or right
control. We believe that these early devel- side of the screen. Children were rewarded
oping temperamental differences reflect the for responding to the identity of the stimu-
maturation of particular neural networks. We lus, regardless of its spatial compatibility with
have studied individual differences in atten- the matching response key. Reduced accuracy
tion and related these differences to emotional and slowed reaction times for spatially incom-
and behavioral control (Rothbart & Rueda patible trials relative to spatially compatible
2005, Ruff & Rothbart 1996). trials reflected the effort required to resist
A major advantage of viewing attention as the dominant response and to resolve conflict
an organ system is to trace the ability of chil- between these two competing dimensions.
dren and adults to regulate their thoughts and Performance on this task produces a clear in-
feelings. Over the early years of life, the regu- terference effect in adults and activates the
lation of emotion is a major issue of develop- anterior cingulate (Fan et al. 2003). Children
ment. The ability of attention to control dis- 24 months of age tended to perseverate on a

www.annualreviews.org • Research on Attentional Networks as a Model 11

ANRV296-PS58-01 ARI 17 November 2006 1:10

Table 2
Annu. Rev. Psychol. 2007.58:1-23. Downloaded from arjournals.annualreviews.org
by Ball State University on 01/04/09. For personal use only.

single response, whereas 36-month-old chil- they showed similar independence among the
dren performed at high accuracy levels, but three networks. Children had much larger
like adults, responded more slowly and with scores for conflict and alerting, suggesting
reduced accuracy to incompatible trials. that they have trouble in resolving conflict
We have traced the development of ex- and in maintaining the alert state when not
ecutive attention into the preschool and pri- warned of the new target. Rather surprisingly,
mary school periods (Rueda et al. 2004) by the ability to resolve conflict in the flanker
using a version of the ANT adapted for chil- task, as measured by the ANT, remains about
dren (Table 2). In some respects, results were the same from age eight to adulthood (see
remarkably similar to those found for adults Table 2).
using the adult version of the task. Reaction There is considerable evidence that the
times for the children were much longer, but executive attention network is of great

12 Posner · Rothbart
ANRV296-PS58-01 ARI 17 November 2006 1:10

importance in the acquisition of school sub- 1994). To display empathy toward others re-
jects such as literacy (McCandliss et al. 2003) quires that we interpret their signals of distress
and in a wide variety of other subjects that or pleasure. Imaging work shows that sad faces
draw upon general intelligence (Duncan et al. activate the amygdala. As sadness increases,
2000). It has been widely believed by psychol- this activation is accompanied by activity in
ogists that training involves only specific do- the anterior cingulate as part of the attention
mains, and that more general training of the network (Blair et al. 1999). It seems likely that
mind, for example, by formal disciplines like the cingulate activity represents the basis for
mathematics or Latin, does not generalize be- our attention to the distress of others.
yond the specific domain trained (Thorndike Developmental studies find two routes
1903, Simon 1969). However, attention may to successful socialization. A strongly reac-
be an exception to this idea. Attention involves tive amygdala would provide the signals of
specific brain mechanisms, as we have seen, distress that would easily allow empathic feel-
Annu. Rev. Psychol. 2007.58:1-23. Downloaded from arjournals.annualreviews.org

but its function is to influence the operation ings toward others and a hesitancy to per-
of other brain networks (Posner & Rothbart form behaviors that might cause harm re-
2007). Anatomically, the network involving lated to fear. These children are relatively
by Ball State University on 01/04/09. For personal use only.

resolution of conflict overlaps with brain ar- easy to socialize. In the absence of this
eas related to general intelligence (Duncan form of control, development of the cingu-
et al. 2000). Training of attention either ex- late would allow appropriate attention to the
plicitly or implicitly is sometimes a part of the signals provided by amygdala activity. Con-
school curriculum (Posner & Rothbart 2007), sistent with its influence on empathy, effort-
but additional studies are needed to deter- ful control also appears to play a role in the
mine exactly how and when attention training development of conscience. The internaliza-
can best be accomplished and its long-lasting tion of moral principles appears to be facili-
importance. tated in fearful preschool-aged children, espe-
cially when their mothers use gentle discipline
(Kochanska 1995). In addition, internalized
Socialization and Culture conscience is facilitated for children high in
Cognitive measures of conflict resolution effortful control (Kochanska et al. 1996). Two
have been linked to aspects of children’s self- separable control systems, one reactive (fear)
control in naturalistic settings. Children who and one self-regulative (effortful control),
are relatively less affected by spatial conflict appear to regulate the development of
also received higher parental ratings of tem- conscience.
peramental effortful control and higher scores Individual differences in effortful control
on laboratory measures of inhibitory control are also related to some aspects of metacog-
(Gerardi-Caulton 2000). Questionnaires have nitive knowledge, such as theory of mind,
shown that the effortful control factor, defined which is the knowledge that people’s behavior
in terms of scales measuring attentional focus- is guided by their beliefs, desires, and other
ing, inhibitory control, low intensity pleasure, mental states (Carlson & Moses 2001). Tasks
and perceptual sensitivity (Rothbart & Rueda that require the inhibition of a prepotent re-
2005), is inversely related to negative affect. sponse are related to theory of mind tasks
This is in keeping with the notion that at- even when other factors, such as age, intelli-
tentional skill may help attenuate negative af- gence, and working memory, are factored out
fect while also serving to constrain impulsive (Carlson & Moses 2001). The mechanisms of
approach tendencies. self-regulation and of theory of mind share
Empathy is strongly related to effortful a similar developmental time course, with
control, with children high in effortful con- advances in both areas between the ages of
trol showing greater empathy (Rothbart et al. 2 and 5.

www.annualreviews.org • Research on Attentional Networks as a Model 13

ANRV296-PS58-01 ARI 17 November 2006 1:10

Efforts to determine the neural network the mental connection between the situation
involved in theory of mind tasks reveal some and its component reactions. In the eastern
of the reasons for the common developmen- tradition, this is done partly through medita-
tal time course in self-regulation. Theory of tion, when ideas can be brought up in a con-
mind tasks activate a network that includes text of calmness and safety, and partly through
areas of the anterior cingulate that are also in- changing the view of the self so that situations
volved in self-regulation, as well as temporal will become less threatening. Links between
lobe areas (Gallagher & Frith 2003). These thoughts and emotions or action tendencies
anatomical links provide further support for are also weakened. Western therapy similarly
efforts to integrate psychological topics at a works through the client’s patterns of reac-
network level. tion, attempting to rework previously consoli-
Emotion, thought, and behavior form a dated patterns and to provide new frameworks
cluster of temporally associated processes in for meaning. From a developmental view, one
Annu. Rev. Psychol. 2007.58:1-23. Downloaded from arjournals.annualreviews.org

specific situations as experienced by the child. would attempt to give the child the kinds of
Single and repeated life experiences can thus experiences that will form favorable and non-
shape connections between elicited emotion, injurious mental habits.
Clusters of reactions are found within the
by Ball State University on 01/04/09. For personal use only.

conceptual understanding of events, and use

of coping strategies to deal with these events. young child’s temperament. Later clusters of
Several theorists have made contributions to thought, emotion, and behavior will also be
this approach (e.g., Epstein 1998, Mischel & based on individual differences in personal-
Ayduk 2004), but the overall framework is in ity, including emotions, expectancies, beliefs,
keeping with the idea of Hebbian learning values, goals, self-evaluations, appraisals, and
through network activation. Mischel and his thoughts about the situation, self, and/or oth-
colleagues have recently developed a cogni- ers. These clusters will be influenced by tem-
tive affective personality theory, making use of peramental predispositions, but they will also
cognitive affective units (CAUs) seen to oper- be influenced by socialization and experience.
ate within a connectionist network (Mischel Coping and the application of effortful con-
& Ayduk 2004). In their model, CAUs are trol operate when “the subjective meaning
variables encoding the features of situations, of the situation, including its self-relevance
which include environmental features as well and personal importance, are appraised. The
as self-initiated thoughts. appraisal itself activates a cascade of other
When they are repeatedly exercised, tem- cognitive-affective representations within the
porally linked clusters of thoughts, emotions, system—expectations and beliefs, affective re-
and action tendencies to a particular situation actions, values and goals” (Mischel & Ayduk
can become highly likely to reoccur and dif- 2004, p. 105).
ficult to change. Some of the processes may Different coping strategies follow and may
be conscious and others unconscious; in most be consolidated or rejected in the future, de-
cases, the thoughts about affectively signifi- pending in part on their consequences. To de-
cant material will be self-referent. Research velop this idea, Mischel & Ayduk (2004) give
on the distinction between conscious and un- the example of individual differences in rejec-
conscious processes has shown that special tion sensitivity. When persons are particularly
networks are active only when items are con- prepared to perceive rejection from others, at-
scious (Dehaene et al. 2003). Studies of self- tention is likely to be narrowly focused on this
reference have also suggested activation of possibility, and it has been demonstrated that
specific networks of neural areas (Gusnard rejection by a social group can influence ar-
et al. 2001). eas of the anterior cingulate related to exec-
In applying this approach to the control of utive attention and pain (Eisenberger et al.
distress, one basic question is how to weaken 2003). Defensive behaviors such as anger or

14 Posner · Rothbart
ANRV296-PS58-01 ARI 17 November 2006 1:10

preventative rejection of the other may serve for example, a child may pursue a positive view
to fend off feelings of rejection. These strate- of the self even when achievements result from
gies in turn can also provide further support following goals and values that are not socially
for the idea that the self is unworthy of pos- acceptable.
itive social relationships. Different levels of
generality of these clusters are possible. Re-
jection sensitivity, for example, might extend PATHOLOGY OF ATTENTION
to a wide range of human relationships, but NETWORKS
the sensitivity may also be more specific, so Much of modern psychology is involved in
that only rejection by the child’s peers, but the diagnosis and treatment of mental illness
not by adults, is sensitized. The reaction may or disturbance, and the network framework
in fact be limited to a single person in a sin- may be an ideal one for incorporating ideas
gle kind of situation, in which case a particular related to clinical remediation. An excellent
Annu. Rev. Psychol. 2007.58:1-23. Downloaded from arjournals.annualreviews.org

person, but not other persons, elicits rejection example of this approach has been in the re-
sensitivity. cent studies of Mayberg (2003) on clinical in-
Socialization in western cultures strongly terventions for depression based upon a neural
by Ball State University on 01/04/09. For personal use only.

emphasizes the individual, promoting the pur- network model. Treatments involved drugs or
suit of individual security, satisfaction of indi- cognitive behavioral therapy and both forms
vidual desires, and achievement of a positive of therapy were about equally effective, based
self-concept. In other cultures, the shaping of on the percentage of persons showing im-
the child’s experienced world can be quite dif- provement. Imaging data, however, indicated
ferent. Mascolo et al. (2003) suggest, for ex- that the two therapies involved very differ-
ample, that the biological systems on which ent brain networks. The drugs remediated a
pride and shame are based are the same across largely subcortical network of brain areas that
cultures, yet can be shaped in quite differ- might be difficult to control voluntarily. The
ent directions. In the United States, pride re- cognitive-behavioral therapy affected cortical
actions develop as parents and others praise networks, including areas involved in atten-
the child’s accomplishments; shame reactions tion that would be more easily subject to vol-
occur when there is self-referent failure. In untary control. These findings show how im-
China, however, parents downplay or criti- portant network approaches are likely to be in
cize children’s performance while other adults evaluating the outcome of clinical trials.
praise it, leading to more moderate prideful A similar story may emerge in the study
affect. Shame, on the other hand, is directly of dyslexia. Many forms of dyslexia involve a
encouraged by parents and others when chil- difficulty in phonological processing that can
dren do not fulfill their obligations to family. be remediated by an intervention targeting
The biological equipment or temperament the ability to convert visual letters to sound
is thus similar across cultures, but the men- (McCandliss et al. 2003). Following remedi-
tal habits and representation of self created ation, there is normal activation of a brain
as a result of the child’s actions varies from region at the boundary between the tem-
culture to culture (Ahadi et al. 1993). By the poral and parietal lobes related to phonol-
time a child is a well-socialized member of ogy. However, although these students can
the society, biologically based responses will now decode words, they do not show flu-
have been shaped into a set of values, goals, ent reading. This may require development
and representations of the self and others. of the visual word form system, which in-
These representations specify what is good volves an extrastriate visual region (fusiform
and bad for the person. Even for the child gyrus) quite distinct from the phonological
who is not well socialized, cultural socializa- areas (McCandliss et al. 2003). It is likely
tion may have an effect. In the United States, that time spent reading is one way to develop

www.annualreviews.org • Research on Attentional Networks as a Model 15

ANRV296-PS58-01 ARI 17 November 2006 1:10

Table 3 Disorders that have been number of disorders of children and adults
related to attentional networks (from (Rothbart & Bates 2006). Executive atten-
Rothbart & Posner 2006) tion represents a neurodevelopmental process
Alerting in children and adolescents, the alteration of
Normal aging which could affect the propensity for the de-
Attention deficit disorder velopment of a number of disorders.
Orienting It is thus important to link efforts at reme-
Autism diation to the training of underlying brain net-
Executive control works. A central aspect of the executive atten-
Alzheimer’s tion network is the ability to deal with conflict.
Borderline personality disorder We used this feature to design a set of exer-
Schizophrenia cises for children adapted from efforts to train
22Q11 deletion syndrome monkeys to perform tasks during space mis-
Annu. Rev. Psychol. 2007.58:1-23. Downloaded from arjournals.annualreviews.org

sions (Rumbaugh & Washburn 1995), which

this brain area, but it may also be possible resulted in training monkeys to resolve con-
to create special training exercises that target flict in a Stroop-like task (Washburn 1994).
the area, as has been done for phonological
by Ball State University on 01/04/09. For personal use only.

Our exercises began by training the child

intervention. to control the movement of a cat on a com-
The possibility that aspects of brain net- puter screen by using a joystick and to predict
works involved in depression and dyslexia where an object would move on the screen,
might be remediated by therapies based on given its initial trajectory. Other exercises em-
training illustrates the close connection be- phasized the use of working memory to retain
tween therapeutic interventions designed to information for a matching-to-sample task
correct deficits and education designed to im- and the resolution of conflict.
prove the performance of people in general. We have tested the efficacy of a very brief
From the perspective of improving neural net- five days of attention training with groups of
works through specific training, therapy and 4- and 6-year-old children (Rueda et al. 2005).
education can represent similar approaches to The children were brought to the laboratory
improving network efficiency. for seven days for sessions lasting approxi-
The ANT has been applied to a number mately 40 minutes, conducted over a two-
of forms of pathology in adults, the aging, to three-week period. The first and last days
adolescents, and children. Table 3 presents involved assessment of the effects of train-
classification of a number of disorders that ing by use of the ANT, a general test of in-
have been related to specific nodes of at- telligence (the K-BIT; Kaufman & Kaufman
tentional network; a review by Rothbart & 1990), and a temperament scale (the Chil-
Posner (2006) provides a fuller account. dren’s Behavior Questionnaire or CBQ;
Rothbart et al. 2001). During administra-
tion of the ANT, we recorded 128 chan-
Plasticity nels of EEG to observe the amplitude and
Executive attention as measured by the ANT time course of activation of brain areas associ-
and other conflict-related tasks provides a ba- ated with executive attention in adult studies
sis for the ability of children to regulate their (Rueda et al. 2005).
behavior through the use of effortful control. Five days is of course a minimal amount
Executive attention has a well-defined neu- of training to influence the development
roanatomy, and much is known about the role of networks that develop over many years.
of genes in modulating its efficiency. Diffi- Nonetheless, we found a general improve-
culties in effortful control provide the basis ment in intelligence in the experimental
for problems in child socialization and in a group as measured by the K-BIT. This was

16 Posner · Rothbart
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Mea dicata

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Anigrum

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