research on attention networks as a model for the...

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

Research on AttentionNetworks as a Model for theIntegration of PsychologicalScienceMichael I. Posner and Mary K. Rothbart1

Psychology Department, University of Oregon, Eugene, Oregon 97403-1291;email: [email protected], [email protected]

Annu. Rev. Psychol. 2007. 58:1–23

First published online as a Review inAdvance on October 9, 2006

The Annual Review of Psychology is online athttp://psych.annualreviews.org

This article’s doi:10.1146/annurev.psych.58.110405.085516

Copyright c© 2007 by Annual Reviews.All rights reserved

0066-4308/07/0110-0001$20.00

1We appreciate the invitation of the editorsof the Annual Review of Psychology to submit aprefatory essay to this year’s volume. Wehave taken the opportunity to propose aunified basis for psychological science basedupon an effort to combine experimental anddifferential approaches to the field. Thisarticle is an improved and expanded versionof an earlier one along these lines (Posner &Rothbart 2004), and its developmentalaspects are further expanded in a book(Posner & Rothbart 2007).

Key Words

attention, candidate genes, orienting, neural networks,temperament

AbstractAs Titchener pointed out more than one hundred years ago, at-tention is at the center of the psychological enterprise. Attentionresearch investigates how voluntary control and subjective experi-ence arise from and regulate our behavior. In recent years, attentionhas been one of the fastest growing of all fields within cognitive psy-chology and cognitive neuroscience. This review examines attentionas characterized by linking common neural networks with individualdifferences in their efficient utilization. The development of atten-tional networks is partly specified by genes, but is also open to specificexperiences through the actions of caregivers and the culture. Webelieve that the connection between neural networks, genes, andsocialization provides a common approach to all aspects of humancognition and emotion. Pursuit of this approach can provide a basisfor psychology that unifies social, cultural, differential, experimen-tal, and physiological areas, and allows normal development to serveas 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 dealtwith all of the topics of psychology in a common framework. Use ofa common network approach to psychological science may allow afoundation for predicting and understanding human behavior in itsvaried forms.

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Attention: theregulating of variousbrain networks byattentional networksinvolved inmaintaining the alertstate, orienting, orregulation of conflict

Neural networks: anumber of brainareas that whenorchestrated carryout a psychologicalfunction

Contents

INTRODUCTION. . . . . . . . . . . . . . . . . 2HEBB’S NETWORK

APPROACH . . . . . . . . . . . . . . . . . . . . 4New Tools . . . . . . . . . . . . . . . . . . . . . . . 5

IMAGING ATTENTIONNETWORKS . . . . . . . . . . . . . . . . . . . 6Simulating Attention Networks . . . 8

INDIVIDUAL EFFICIENCY. . . . . . . 9NETWORK DEVELOPMENT . . . . 11

Socialization and Culture . . . . . . . . . 13PATHOLOGY OF ATTENTION

NETWORKS . . . . . . . . . . . . . . . . . . . 15Plasticity . . . . . . . . . . . . . . . . . . . . . . . . . 16

INTEGRATION OFPSYCHOLOGICAL SCIENCE. . 17Generality . . . . . . . . . . . . . . . . . . . . . . . 17

INTRODUCTION

Can psychology be a unified science, or mustit remain fragmented and subject to transientresearch interests rather than cumulative de-velopment? Is psychology teachable or mustpsychology textbooks remain encyclopediasof often unrelated findings? In this article,we examine research on attention to suggestways of looking at psychological questions andfindings that might lead to hopeful answers tothese questions. We offer a model based onHebbian psychology that reaches out frombiological roots to tackle questions of emo-tion and thought as well as behavior. We, withmany of our colleagues, see different levelsof analysis in psychology as informing eachother, with each level of equal scientific valid-ity. Bridging these levels can allow a higherlevel of understanding and prediction.

We argue that D.O. Hebb (see Figure 1),beginning with his monograph in 1949 andcontinuing through a series of introductorytextbooks (1958, 1966), has convincingly pre-sented the basis for such integration. This in-tegration lies in understanding how genes andexperience shape neural networks underlyinghuman thoughts, feelings, and actions.

At the time Hebb wrote his monograph,relatively little was known about how thestructure and organization of the central ner-vous system contribute to the functions ob-served in psychological studies. This led Hebbto talk in terms of the conceptual nervoussystem, that is, ideas about its structure thatmight be imagined or inferred from psycho-logical studies. We suggest that the meth-ods available to Hebb, mostly animal researchand behavioral human experiments, were notsufficient to provide empirical methods forlinking his conceptual nervous system to realevents in the human brain. This methodologyhas now been provided by neuroimaging. Al-though Hebb also recognized the importanceof studying individual differences in intelli-gence and affect, at that time there were nomethods for exploring the specific genes thatare an important source of these differences.The human genome project has provided newmethods for exploring this issue.

In Hebb’s time, the idea of a network (cellassembly or phase sequence) was a rathervague verbal abstraction that did not allowfor models that could produce specific pre-dictions. As a result of the rapid changes incellular biology (Bullock et al. 2005) and inthe mathematics of multilevel networks, thistoo has changed (Rumelhart & McClelland1986). Although early versions of these net-works were inspired by simple versions ofneurons as all-or-none elements, more recentversions (O’Reilly & Munakata 2000) have be-gun to use the details of neuroanatomy andcellular structure as provided by imaging andcellular studies to develop networks that takemore realistic advantage of the structure ofthe human brain. Hebb’s basic idea, togetherwith the new methodological tools and newdisciplines (e.g., cognitive, affective, and so-cial neuroscience), all based on network views,give abundant evidence of the value of em-ploying the converging operations strategyadvocated by Sternberg (Sternberg 2004).

It is important to recognize the need for in-tegrating cognitive, affective, and social neu-roscience with psychology because many of

2 Posner · Rothbart

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Figure 1Photograph of Professor Donald O. Hebb.

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

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the theoretical questions that need to be ad-dressed by neuroimaging and genetic studiesare exactly those that a century of psychol-ogists have explored. The neuroscience ap-proach provides crucial constraints for psy-chological theories, but it also benefits whena closer connection is made with the psycho-logical level of analysis.

HEBB’S NETWORK APPROACH

In 1949, D.O. Hebb published his epic work,The Organization of Behavior. His book wasimmediately recognized as providing the po-tential for an integrated psychology. Onereviewer (Attneave 1950, p. 633) wrote:

I believe The Organization of Behavior to bethe most important contribution to psycho-logical theory in recent years. Unlike thoseof his contemporaries who are less interestedin psychology than in some restricted as-pect thereof to which their principles con-fine them, Hebb has made a noteworthy at-tempt to take the experimentally determinedfacts of behavior, as they are, and account forthem in terms of events within the centralnervous system.

The most important basic idea that Hebbpresented was the cell assembly theory out-lined in chapters 4 and 5 of his book (Goddard1980, Harris 2005). Hebb argued that ev-ery psychological event, sensation, expecta-tion, emotion, or thought is represented bythe flow of activity in a set of interconnectedneurons. Learning occurs by a change insynaptic strength when a synapse conductsexcitation at the same time the postsynap-tic neuron discharges. This provided a basisfor the modification of synapses and showedhow neural networks might be organized un-der the influence of specific experiences. TheHebb synapse plays a central role in modernneuroscience (see Kolb 2003, Milner 2003,Sejnowski 2003). There are important newdevelopments in the study of synapses and inthe discovery of other influences among neu-

rons and between neurons and other braincells (Bullock et al. 2005). These develop-ments have reduced the gap between networksrevealed in imaging studies and the complexintracellular activity that underlies them. Inparticular, they show that learning may reflectthe activity of interactions at many time scalesand may be modified by aspects of the organ-ism’s overall state.

Hebb also introduced the concept of thephase sequence involved in the coordinationof multiple cell assemblies. He recognized theimportance of the temporal correspondencebetween assemblies. In recent years theseideas have been supported by studies show-ing that synchronization of brain areas may becritical to detecting stimuli (Womelsdorf et al.2006) and for transfer of information betweenremote areas (Nikolaev et al. 2001). In lateryears, Hebb (1958) developed an introductorypsychology textbook that integrated much ofpsychology using the framework provided byhis 1949 theory. He applied his network the-ory to heredity, learning and memory, motiva-tion, perception, thought, and development.In later editions, he extended his approachto emotions in their social contexts, individ-ual differences in intelligence, and abnormalpsychology.

Despite his efforts and those of his fol-lowers at McGill and elsewhere, Hebb’s workwas unsuccessful in providing a fully integra-tive psychology, and many still seek to de-velop such an integration (Sternberg 2004,Sternberg & Grigorenko 2001). One of themajor problems with Hebb’s framework wasthat it left no clear empirical path for the ac-quisition of new knowledge about how hu-man brain networks develop, how they dif-fer among individuals, why they break down,and how they can be restored to functioning.Most of the research by Hebb and his asso-ciates was performed on nonhuman animals,while most of psychology concerns human be-havior, brain, and mind.

In 1955, Hebb argued for the utility of aconceptual nervous system inferred from psy-chological studies of motivation. This idea


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remains of key importance. Despite ourgrowing understanding of the function andstructure of the nervous system, psycholog-ical knowledge needs to be used to pro-pose hypotheses about how the central ner-vous system (CNS) works to support feelings,thoughts, and behaviors. We argue that onlywithin an integrated field of psychology canknowledge at all levels be used to develop andconstrain these hypotheses.

New Tools

Several major late-twentieth century eventsgive improved prospects for an integrationof psychological science around the ideas in-troduced by Hebb. Cell assemblies and phasesequences are names for aspects of neural net-works. Now, thanks to work on the compu-tational properties of neural networks (i.e.,Rumelhart & McClelland 1986), we are in amuch better position to develop detailed the-ories integrating information from physiolog-ical, cognitive, and behavioral studies.

In addition, new neuroimaging methodsnow allow us to examine neuronal activityin terms of localized changes in blood flowor metabolism in positron emission tomogra-phy (PET) or changes in blood oxygenationby functional magnetic resonance imaging(fMRI). By using tracers that bind to differenttransmitters, PET can also be used to examinetransmitter density. By measuring electrical(electroencephalogram, or EEG) and mag-netic (magnetoencephalogram) signals out-side the skull, the time course of activationof different brain areas localized by fMRI canbe measured (Dale et al. 2000). Pathways ofactivation can also be imaged by use of dif-fusion tensor imaging, a form of MRI thattraces white matter tracts. In addition to thestudy of naturally occurring lesions, interrupt-ing information flow by transcranial magneticstimulation (TMS) can produce temporaryfunctional lesions of pathways (see Toga &Mazziotta 1996 for a review of these and othermethods). These methods provide a toolkitthat can be used either alone or together to

PET: positronemissiontomography

fMRI: functionalmagnetic resonanceimaging

Temperament:relatively enduringbiologicalcharacteristics ofindividuals thatinclude both reactiveand self-regulatory(attentional) aspects

make human brain networks accessible for de-tailed physiological study.

Some have thought that the influence ofimaging has been merely to tell us where inthe brain things have happened (Utall 2001).Certainly many, perhaps even most, imagingstudies have been concerned with anatomicalissues. As Figure 2 illustrates, several func-tions of attention have been shown to involvespecific anatomical areas that carry out impor-tant functions. However, imaging also probesneural networks that underlie all aspects ofhuman thought, feelings, and behavior. Thefull significance of imaging for (a) viewingbrain networks, (b) examining their compu-tation in real time, (c) exploring how they areassembled in development, and (d ) their plas-ticity following physical damage or training isa common theme in current research that isjust beginning to reach its potential.

A third development, the mapping of thehuman genome (Venter et al. 2001), offers thepotential for an increased understanding ofthe physical basis for individual differences,including individual differences in tempera-ment and personality. Many genes exhibit anumber of relatively high-frequency variants(polymorphisms) that can code for differentphysical configurations. These in turn can al-ter the efficiency of a network. For exam-ple, different types of genes (alleles) form-ing dopamine receptors can lead to differentefficiency in binding to dopamine and thusdifferences in underlying neural networks. Ina number of cases, it has been possible to relatethese genetic differences to individual per-formance in tasks involving the network (seeFossella & Posner 2004 for a review).Genetics-based research also provides an im-portant approach to the development of char-acteristics of neural networks common to allmembers of the species.

We recognize that not all topics of psy-chology have been sufficiently explored to il-lustrate this framework and that limitationsin our knowledge prevent us from explor-ing all of the areas where these develop-ments can be applied. However, attention and


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Figure 2Anatomy of three attentional networks: alerting, orienting, and executive attention (from Posner &Rothbart 2007).

temperament, areas of our study, can be usedto consider how a large number of psycho-logical questions can be explored using theneural network framework outlined by Hebb.Attention serves as a basic set of mechanismsthat underlie our awareness of the world andthe voluntary regulation of our thoughts andfeelings. The methods used to understand at-tentional networks in terms of anatomy, indi-vidual differences, development, and plastic-ity can be applied readily to explore networksrelated to other aspects of human behavior.

Any approach based on neural networksraises the issue of crude reductionism. Manyagree that all human behavior must ulti-mately be traceable to brain activity, but cor-rectly argue for the importance of cogni-

tive experiments, behavioral observations, andself-report as important elements of psycho-logical science. We hope to illustrate in this ar-ticle how important such psychological meth-ods are and how they can be integrated withina brain network framework, as Hebb (1955)illustrated in his Textbook of Psychology.

IMAGING ATTENTIONNETWORKS

Functional neuroimaging has allowed manycognitive tasks to be analyzed in terms of thebrain areas they activate, and studies of atten-tion have been among those most often ex-amined (Corbetta & Shulman 2002, Driveret al. 2004, Posner & Fan 2007). Imaging


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data have supported the presence of three net-works related to different aspects of attention(Fan et al. 2005). These networks carry outthe functions of alerting, orienting, and exec-utive attention (Posner & Fan 2007). A sum-mary of the anatomy and chemical modula-tors involved in the three networks is shown inTable 1.

Alerting is defined as achieving and main-taining a state of high sensitivity to incom-ing stimuli; orienting is the selection of in-formation from sensory input; and executiveattention involves mechanisms for monitor-ing and resolving conflict among thoughts,feelings, and responses. The alerting systemhas been associated with thalamic as well asfrontal and parietal regions of the cortex (Fanet al. 2005). A particularly effective way to varyalertness has been to use warning signals priorto targets. The influence of warning signalson the level of alertness is thought to be dueto modulation of neural activity by the neu-rotransmitter norepinephrine (Marrocco &Davidson 1998).

Orienting involves aligning attention witha source of sensory signals. This may be overt,as when eye movements accompany move-ments of attention, or may occur covertly,without any eye movement. The orientingsystem for visual events has been associatedwith posterior brain areas, including thesuperior parietal lobe and temporal parietaljunction, and in addition, the frontal eye fields(Corbetta & Shulman 2002). Orienting canbe manipulated by presenting a cue indicatingwhere in space a target is likely to occur,thereby directing attention to the cued loca-tion (Posner 1980). It is possible to determinethe anatomy influenced by the cue separatelyfrom that influenced by the target by usingMRI to trace changes in the blood that specif-ically follow the cue. This method is calledevent-related functional magnetic resonanceimaging, and its use has suggested that thesuperior parietal lobe is associated withorienting following the presentation of a cue(Corbetta & Shulman 2002). The superiorparietal lobe in humans is closely related

Orienting: theinteraction of a brainnetwork with sensorysystems designed toimprove the selectedsignal

Table 1 A summary of the anatomy and chemical modulatorsinvolved in the alerting, orienting, and executive attentionnetworks.

Function Structures ModulatorOrient Superior parietal Acetylcholine

Temporal parietal junctionFrontal eye fieldsSuperior colliculus

Alert Locus coeruleus NorepinephrineRight frontalParietal cortex

Executive attention Anterior cingulate DopamineLateral ventralPrefrontalBasal ganglia

to the lateral intraparietal area (LIP) inmonkeys, which is involved in the productionof eye movements (Andersen 1989). Whena target occurs at an uncued location andattention has to be disengaged and moved to anew location, there is activity in the temporalparietal junction (Corbetta & Shulman 2002).Lesions of the temporal parietal junctionlobe and superior temporal lobe have beenconsistently related to difficulties in orienting(Karnath et al. 2001).

Executive control of attention is oftenstudied by tasks that involve conflict, such asvarious versions of the Stroop task. In theStroop task, subjects must respond to thecolor of ink (e.g., red) while ignoring the colorword name (e.g., blue) (Bush et al. 2000). Re-solving conflict in the Stroop task activatesmidline frontal areas (anterior cingulate) andlateral prefrontal cortex (Botvinick et al. 2001,Fan et al. 2005). There is also evidence forthe activation of this network in tasks involv-ing conflict between a central target and sur-rounding flankers that may be congruent orincongruent with the target (Botvinick et al.2001, Fan et al. 2005).

Recently, the role of the anterior cingulatein modulating sensory input has been demon-strated experimentally by showing enhancedconnectivity between the anterior cingulatecortex and the sensory modality to which the


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ANT: attentionnetwork test

RT: reaction time

ACT-R: adaptivecontrol ofthought-rational

person is asked to attend (Crottaz-Herbette &Menon 2006). This finding supports the gen-eral idea that anterior cingulate cortex activityregulates other brain areas, at least for sen-sory areas. Experimental tasks may also pro-vide a means of fractionating the contribu-tions of different areas within the executiveattention network (MacDonald et al. 2000). Inaccord with the findings in recent neuroimag-ing studies (Beauregard et al. 2001, Ochsneret al. 2001), we have argued that the exec-utive attention network is involved in self-regulation of positive and negative affect aswell as a wide variety of cognitive tasks under-lying intelligence (Duncan et al. 2000). Thisidea suggests an important role for attentionin moderating the activity of sensory, cogni-tive, and emotional systems.

Simulating Attention Networks

Quantification has had a high value in psycho-logical research. An advantage of the networkapproach is that it lends itself to the devel-opment of precise computer models that al-low both summarization of many findings inthe field and prediction of new findings. Cur-rently, symbolic models, such as rule-basedsystems (Newell 1990), appear to be a goodway to capture data from reaction time andother psychological findings. In these models,cognitive functions are represented as chainsof production rules and can be identified withthe mental operations postulated by cogni-tive studies. On the other hand, subsymbolicmodels, such as connectionist models (e.g.,O’Reilly & Munakata 2000), permit a morebiologically realistic implementation of theoperations and closer links to imaging studies.

We have developed the attention networktest (ANT) to examine individual differencesin the efficiency of the brain networks of alert-ing, orienting, and executive attention dis-cussed above (Fan et al. 2002, Rueda et al.2004). The ANT uses differences in reactiontime (RT) between conditions to measure theefficiency of each network. Each trial beginswith a cue (or a blank interval, in the no-cue

condition) that informs the participant eitherthat a target will occur soon or where it willoccur, or both. The target always occurs eitherabove or below fixation, and consists of a cen-tral arrow, surrounded by flanking arrows thatcan point either in the same direction (con-gruent) or in the opposite direction (incon-gruent). Subtracting RTs for congruent fromincongruent target trials provides a measure ofconflict resolution and assesses the efficiencyof the executive attention network. Subtract-ing RTs obtained in the double-cue conditionthat provides information on when but notwhere the target will occur from RTs in theno-cue condition gives a measure of alertingdue to the presence of a warning signal. Sub-tracting RTs to targets at the cued location(spatial cue condition) from trials using a cen-tral cue gives a measure of orienting, since thespatial cue, but not the central cue, providesvalid information about where a target willoccur.

The attention network task has now beensimulated in the framework of the adaptivecontrol of thought-rational (ACT-R) theory(Wang et al. 2004). The ANT task is di-vided into subroutines. Cue processing in-volves switching of attention to the cued loca-tion. Target processing involves detection ofthe direction of the arrow in the center and in-volves an attention switch and response initia-tion. Each of these operations is implementedby a chain of production rules. The operationsare similar to those discussed in most psycho-logical studies and localized in neurologicalstudies. For example, the switching of atten-tion based on the peripheral cue or target isthought to be implemented by the temporalparietal junction (Corbetta & Shulman 2002).

A connectionist simulation of the ANT(H. Wang & J. Fan, manuscript in prepara-tion) is based upon a local error-driven andassociative, biologically realistic algorithm(LEABRA) (O’Reilly & Munakata 2000). Thesubroutines of ACT-R are now replaced byspecific connections between hypothesizedneurons. These neurons are somewhat real-istic, and are designed to represent the known


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properties of specific brain areas. Thus, theorienting network can be designed to reflectthe known properties of the frontal eye fields,superior parietal lobe, and temporal parietaljunction, and they can be connected withinthe simulation. As shown in Figure 3, sim-ulations do a reasonable job of fitting withthe known ANT data, although improvementcan be expected in the future. The symbolicmodel makes contact with the mental opera-tions related to imaging, whereas the connec-tionist framework allows a strong treatmentof the underlying biology. Together they illus-trate how network views provide a computa-tional means for summarizing many findingswithin psychology, allowing novel predictionsreflecting properties of the network.

INDIVIDUAL EFFICIENCY

Psychology is often divided into two ap-proaches that are almost completely separatein the literature (but see Gardner 1983 for aneffort at integration). The discussion abovefocused on general features of the humanmind, such as the ability to attend. Anotherapproach deals with differences among indi-viduals. These differences may involve cogni-tion, as in the measurement of intelligence,or they may involve temperamental differ-ences, many of which relate to energetic fac-tors such as the expression and control of theemotions. Almost all studies of attention havebeen concerned either with the general abili-ties involved or with effects of brain injury orpathology on attention. However, it is clearthat normal individuals differ in their ability toattend to sensory events, and it is even clearerthat they differ in their ability to concentratefor long periods on internal trains of thought.

We used the attention network task to ex-amine the individual efficiency of the alerting,orienting, and executive networks (Fan et al.2002). In a sample of 40 normal persons, eachof these scores was reliable over repeated pre-sentations. In addition, we found no correla-tion among the orienting, alerting, and exec-utive scores.

Candidate genes:genes that may beinvolved in thedevelopment ofutilization ofattentional networks

The ability to measure differences in at-tention among adults raises the question ofthe degree to which attention is heritable. Toexplore this issue, the ANT was used to assessattention in monozygotic and dizygotic same-sex twins (Fan et al. 2001). Strong heritabil-ity was found for the executive network, someheritability for the orienting network, and noapparent heritability for the alerting network.These data support a search for genes in exec-utive attention and in orienting of attention.

We then used the association of the ex-ecutive network with the neuromodulatordopamine (see Table 1) as a way of search-ing for candidate genes that might relate tothe efficiency of the networks (Fossella et al.2002). To do this, 200 persons performedthe ANT and were genotyped to examinefrequent polymorphisms in genes related todopamine. We found significant associationof the dopamine 4 receptor and monoamineoxidase A genes. We then conducted a neu-roimaging experiment in which persons withdifferent alleles of these two genes were com-pared while they performed the ANT (Fanet al. 2003). Groups with different allelesof these genes showed differences in perfor-mance on the ANT and also produced sig-nificantly different activations in the anteriorcingulate, a major node of the executive atten-tion network.

Recent studies have confirmed and ex-tended these observations to other dopaminegenes and to the orienting network. In two dif-ferent studies employing other conflict tasks,the catecholamine-O-methyltransferase genewas linked to the mental operations re-lated to resolving conflict (Blasi et al. 2005,Diamond et al. 2004). Different alleles ofcholinergic genes were also related to perfor-mance on orienting tasks such as visual search(Parasuraman et al. 2005), thus confirming thelink between orienting and the neuromodula-tor acetylcholine (see Table 1).

Hebb (1955) thought that most of thenetworks involved in higher functions wereshaped primarily through experience. Wenow know that there is a great deal in common


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OutputExecutive Control

Network

Orienting Network Object Pathway

Alerting Network Primary Visual Cortex

Visual Input

1. Fixation &Cue

Expectation(2)

2. Cue or Stimulus?

4. Stimulus Expectation

(1)

5. Stimulus Processing

(18)

3. Cue Processing

(7)

Next Trial?

Stimulus

6. Response(6)

State Switching(2)

EXPERIMENTAL RESULTS 47 51 84

ACT-R SIMULATION 55 45 86

LEABRA SIMULATION 70 33 67

ALERT ORIENT CONFLICT

Figure 3Experimental 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 arein milliseconds.


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among humans in the anatomy of these high-level networks, and thus that they must havea basis within the human genome. It seemslikely that the same genes that are related toindividual differences in attention are also im-portant in the development of the attentionalnetworks that are common among people.Some of these networks are also common tononhuman animals. By examining these net-works in animals, it should be possible to testthese assumptions further and to understandthe role of genes in shaping networks.

Of importance for this effort is the de-velopment of methods to manipulate relevantgenes in specific anatomical locations that areimportant nodes of a particular network. Usu-ally genes are expressed at multiple locationsso that changes (e.g., gene knockouts) are notspecific to one location. However, using sub-tractive genomics, a method is currently be-ing developed to do this (Dumas et al. 2005).We believe that this kind of genetic analysis ofnetwork development will become a very pro-ductive link between genes and both normaland pathological psychological function.

NETWORK DEVELOPMENT

Every parent of more than one child recog-nizes that from birth, infants are individualswith their own distinct characteristics and dis-positions. These include reactive traits such asemotionality, activity level, and orienting tosensory events, and regulatory traits like at-tention focusing and shifting and inhibitorycontrol. We believe that these early devel-oping temperamental differences reflect thematuration of particular neural networks. Wehave studied individual differences in atten-tion and related these differences to emotionaland behavioral control (Rothbart & Rueda2005, Ruff & Rothbart 1996).

A major advantage of viewing attention asan organ system is to trace the ability of chil-dren and adults to regulate their thoughts andfeelings. Over the early years of life, the regu-lation of emotion is a major issue of develop-ment. The ability of attention to control dis-

tress can be traced to early infancy (Harmanet al. 1997). Early in life, most regulation de-pends on the caregiver providing ways to con-trol infant reactions. In our study, we first in-duced a mild level of distress, and then showedhow attentional orienting calms that distresswhile the infant remains engaged with the ob-ject. When the orienting is broken, distress re-turns to the level present prior to the introduc-tion of the object. It is likely that the distressremains present and is held by networks in theamygdala. Parents often use manipulation ofthe infant’s orienting to control distress, andthe infant also exhibits coping behaviors in-volving orienting to form the basis for oneaspect of early self-regulation (Rothbart et al.1992).

Developmental changes in executive atten-tion found during the third year of life are cor-related with parent reports of temperamentaleffortful control (Gerardi-Caulton 2000). Ef-fortful control is a broad variable identified intemperament research; it includes the abilityto inhibit a dominant response in order to pro-duce subdominant response and to detect andcorrect errors (Rothbart & Rueda 2005). Be-cause children of this age do not read, the loca-tion and identity rather than the word mean-ing and ink color served as the dimensions ina spatial conflict task. In one study (Gerardi-Caulton 2000), children sat in front of two re-sponse keys, one located to the child’s left andone to the right. Each key displayed a picture,and on every trial, a picture identical to oneof the pair appeared on either the left or rightside of the screen. Children were rewardedfor responding to the identity of the stimu-lus, regardless of its spatial compatibility withthe matching response key. Reduced accuracyand slowed reaction times for spatially incom-patible trials relative to spatially compatibletrials reflected the effort required to resistthe dominant response and to resolve conflictbetween these two competing dimensions.Performance on this task produces a clear in-terference effect in adults and activates theanterior cingulate (Fan et al. 2003). Children24 months of age tended to perseverate on a


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Table 2

single response, whereas 36-month-old chil-dren performed at high accuracy levels, butlike adults, responded more slowly and withreduced accuracy to incompatible trials.

We have traced the development of ex-ecutive attention into the preschool and pri-mary school periods (Rueda et al. 2004) byusing a version of the ANT adapted for chil-dren (Table 2). In some respects, results wereremarkably similar to those found for adultsusing the adult version of the task. Reactiontimes for the children were much longer, but

they showed similar independence among thethree networks. Children had much largerscores for conflict and alerting, suggestingthat they have trouble in resolving conflictand in maintaining the alert state when notwarned of the new target. Rather surprisingly,the ability to resolve conflict in the flankertask, as measured by the ANT, remains aboutthe same from age eight to adulthood (seeTable 2).

There is considerable evidence that theexecutive attention network is of great


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importance in the acquisition of school sub-jects such as literacy (McCandliss et al. 2003)and in a wide variety of other subjects thatdraw upon general intelligence (Duncan et al.2000). It has been widely believed by psychol-ogists that training involves only specific do-mains, and that more general training of themind, for example, by formal disciplines likemathematics or Latin, does not generalize be-yond the specific domain trained (Thorndike1903, Simon 1969). However, attention maybe an exception to this idea. Attention involvesspecific brain mechanisms, as we have seen,but its function is to influence the operationof other brain networks (Posner & Rothbart2007). Anatomically, the network involvingresolution of conflict overlaps with brain ar-eas related to general intelligence (Duncanet al. 2000). Training of attention either ex-plicitly or implicitly is sometimes a part of theschool curriculum (Posner & Rothbart 2007),but additional studies are needed to deter-mine exactly how and when attention trainingcan best be accomplished and its long-lastingimportance.

Socialization and Culture

Cognitive measures of conflict resolutionhave been linked to aspects of children’s self-control in naturalistic settings. Children whoare relatively less affected by spatial conflictalso received higher parental ratings of tem-peramental effortful control and higher scoreson laboratory measures of inhibitory control(Gerardi-Caulton 2000). Questionnaires haveshown that the effortful control factor, definedin terms of scales measuring attentional focus-ing, inhibitory control, low intensity pleasure,and perceptual sensitivity (Rothbart & Rueda2005), is inversely related to negative affect.This is in keeping with the notion that at-tentional skill may help attenuate negative af-fect while also serving to constrain impulsiveapproach tendencies.

Empathy is strongly related to effortfulcontrol, with children high in effortful con-trol showing greater empathy (Rothbart et al.

1994). To display empathy toward others re-quires that we interpret their signals of distressor pleasure. Imaging work shows that sad facesactivate the amygdala. As sadness increases,this activation is accompanied by activity inthe anterior cingulate as part of the attentionnetwork (Blair et al. 1999). It seems likely thatthe cingulate activity represents the basis forour attention to the distress of others.

Developmental studies find two routesto successful socialization. A strongly reac-tive amygdala would provide the signals ofdistress that would easily allow empathic feel-ings toward others and a hesitancy to per-form behaviors that might cause harm re-lated to fear. These children are relativelyeasy to socialize. In the absence of thisform of control, development of the cingu-late would allow appropriate attention to thesignals provided by amygdala activity. Con-sistent with its influence on empathy, effort-ful control also appears to play a role in thedevelopment of conscience. The internaliza-tion of moral principles appears to be facili-tated in fearful preschool-aged children, espe-cially when their mothers use gentle discipline(Kochanska 1995). In addition, internalizedconscience is facilitated for children high ineffortful control (Kochanska et al. 1996). Twoseparable control systems, one reactive (fear)and one self-regulative (effortful control),appear to regulate the development ofconscience.

Individual differences in effortful controlare also related to some aspects of metacog-nitive knowledge, such as theory of mind,which is the knowledge that people’s behavioris guided by their beliefs, desires, and othermental states (Carlson & Moses 2001). Tasksthat require the inhibition of a prepotent re-sponse are related to theory of mind taskseven when other factors, such as age, intelli-gence, and working memory, are factored out(Carlson & Moses 2001). The mechanisms ofself-regulation and of theory of mind sharea similar developmental time course, withadvances in both areas between the ages of2 and 5.


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Efforts to determine the neural networkinvolved in theory of mind tasks reveal someof the reasons for the common developmen-tal time course in self-regulation. Theory ofmind tasks activate a network that includesareas of the anterior cingulate that are also in-volved in self-regulation, as well as temporallobe areas (Gallagher & Frith 2003). Theseanatomical links provide further support forefforts to integrate psychological topics at anetwork level.

Emotion, thought, and behavior form acluster of temporally associated processes inspecific situations as experienced by the child.Single and repeated life experiences can thusshape connections between elicited emotion,conceptual understanding of events, and useof coping strategies to deal with these events.Several theorists have made contributions tothis approach (e.g., Epstein 1998, Mischel &Ayduk 2004), but the overall framework is inkeeping with the idea of Hebbian learningthrough network activation. Mischel and hiscolleagues have recently developed a cogni-tive affective personality theory, making use ofcognitive affective units (CAUs) seen to oper-ate within a connectionist network (Mischel& Ayduk 2004). In their model, CAUs arevariables encoding the features of situations,which include environmental features as wellas self-initiated thoughts.

When they are repeatedly exercised, tem-porally linked clusters of thoughts, emotions,and action tendencies to a particular situationcan become highly likely to reoccur and dif-ficult to change. Some of the processes maybe conscious and others unconscious; in mostcases, the thoughts about affectively signifi-cant material will be self-referent. Researchon the distinction between conscious and un-conscious processes has shown that specialnetworks are active only when items are con-scious (Dehaene et al. 2003). Studies of self-reference have also suggested activation ofspecific networks of neural areas (Gusnardet al. 2001).

In applying this approach to the control ofdistress, one basic question is how to weaken

the mental connection between the situationand its component reactions. In the easterntradition, this is done partly through medita-tion, when ideas can be brought up in a con-text of calmness and safety, and partly throughchanging the view of the self so that situationswill become less threatening. Links betweenthoughts and emotions or action tendenciesare also weakened. Western therapy similarlyworks through the client’s patterns of reac-tion, attempting to rework previously consoli-dated patterns and to provide new frameworksfor meaning. From a developmental view, onewould attempt to give the child the kinds ofexperiences that will form favorable and non-injurious mental habits.

Clusters of reactions are found within theyoung child’s temperament. Later clusters ofthought, emotion, and behavior will also bebased on individual differences in personal-ity, including emotions, expectancies, beliefs,values, goals, self-evaluations, appraisals, andthoughts about the situation, self, and/or oth-ers. These clusters will be influenced by tem-peramental predispositions, but they will alsobe influenced by socialization and experience.Coping and the application of effortful con-trol operate when “the subjective meaningof the situation, including its self-relevanceand personal importance, are appraised. Theappraisal itself activates a cascade of othercognitive-affective representations within thesystem—expectations and beliefs, affective re-actions, values and goals” (Mischel & Ayduk2004, p. 105).

Different coping strategies follow and maybe consolidated or rejected in the future, de-pending in part on their consequences. To de-velop this idea, Mischel & Ayduk (2004) givethe example of individual differences in rejec-tion sensitivity. When persons are particularlyprepared to perceive rejection from others, at-tention is likely to be narrowly focused on thispossibility, and it has been demonstrated thatrejection by a social group can influence ar-eas of the anterior cingulate related to exec-utive attention and pain (Eisenberger et al.2003). Defensive behaviors such as anger or


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preventative rejection of the other may serveto fend off feelings of rejection. These strate-gies in turn can also provide further supportfor the idea that the self is unworthy of pos-itive social relationships. Different levels ofgenerality of these clusters are possible. Re-jection sensitivity, for example, might extendto a wide range of human relationships, butthe sensitivity may also be more specific, sothat only rejection by the child’s peers, butnot by adults, is sensitized. The reaction mayin fact be limited to a single person in a sin-gle kind of situation, in which case a particularperson, but not other persons, elicits rejectionsensitivity.

Socialization in western cultures stronglyemphasizes the individual, promoting the pur-suit of individual security, satisfaction of indi-vidual desires, and achievement of a positiveself-concept. In other cultures, the shaping ofthe child’s experienced world can be quite dif-ferent. Mascolo et al. (2003) suggest, for ex-ample, that the biological systems on whichpride and shame are based are the same acrosscultures, yet can be shaped in quite differ-ent directions. In the United States, pride re-actions develop as parents and others praisethe child’s accomplishments; shame reactionsoccur when there is self-referent failure. InChina, however, parents downplay or criti-cize children’s performance while other adultspraise it, leading to more moderate pridefulaffect. Shame, on the other hand, is directlyencouraged by parents and others when chil-dren do not fulfill their obligations to family.

The biological equipment or temperamentis thus similar across cultures, but the men-tal habits and representation of self createdas a result of the child’s actions varies fromculture to culture (Ahadi et al. 1993). By thetime a child is a well-socialized member ofthe society, biologically based responses willhave been shaped into a set of values, goals,and representations of the self and others.These representations specify what is goodand bad for the person. Even for the childwho is not well socialized, cultural socializa-tion may have an effect. In the United States,

for example, a child may pursue a positive viewof the self even when achievements result fromfollowing goals and values that are not sociallyacceptable.

PATHOLOGY OF ATTENTIONNETWORKS

Much of modern psychology is involved inthe diagnosis and treatment of mental illnessor disturbance, and the network frameworkmay be an ideal one for incorporating ideasrelated to clinical remediation. An excellentexample of this approach has been in the re-cent studies of Mayberg (2003) on clinical in-terventions for depression based upon a neuralnetwork model. Treatments involved drugs orcognitive behavioral therapy and both formsof therapy were about equally effective, basedon the percentage of persons showing im-provement. Imaging data, however, indicatedthat the two therapies involved very differ-ent brain networks. The drugs remediated alargely subcortical network of brain areas thatmight be difficult to control voluntarily. Thecognitive-behavioral therapy affected corticalnetworks, including areas involved in atten-tion that would be more easily subject to vol-untary control. These findings show how im-portant network approaches are likely to be inevaluating the outcome of clinical trials.

A similar story may emerge in the studyof dyslexia. Many forms of dyslexia involve adifficulty in phonological processing that canbe remediated by an intervention targetingthe ability to convert visual letters to sound(McCandliss et al. 2003). Following remedi-ation, there is normal activation of a brainregion at the boundary between the tem-poral and parietal lobes related to phonol-ogy. However, although these students cannow decode words, they do not show flu-ent reading. This may require developmentof the visual word form system, which in-volves an extrastriate visual region (fusiformgyrus) quite distinct from the phonologicalareas (McCandliss et al. 2003). It is likelythat time spent reading is one way to develop


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Table 3 Disorders that have beenrelated to attentional networks (fromRothbart & Posner 2006)

AlertingNormal agingAttention deficit disorder

OrientingAutism

Executive controlAlzheimer’sBorderline personality disorderSchizophrenia22Q11 deletion syndrome

this brain area, but it may also be possibleto create special training exercises that targetthe area, as has been done for phonologicalintervention.

The possibility that aspects of brain net-works involved in depression and dyslexiamight be remediated by therapies based ontraining illustrates the close connection be-tween therapeutic interventions designed tocorrect deficits and education designed to im-prove the performance of people in general.From the perspective of improving neural net-works through specific training, therapy andeducation can represent similar approaches toimproving network efficiency.

The ANT has been applied to a numberof forms of pathology in adults, the aging,adolescents, and children. Table 3 presentsclassification of a number of disorders thathave been related to specific nodes of at-tentional network; a review by Rothbart &Posner (2006) provides a fuller account.

Plasticity

Executive attention as measured by the ANTand other conflict-related tasks provides a ba-sis for the ability of children to regulate theirbehavior through the use of effortful control.Executive attention has a well-defined neu-roanatomy, and much is known about the roleof genes in modulating its efficiency. Diffi-culties in effortful control provide the basisfor problems in child socialization and in a

number of disorders of children and adults(Rothbart & Bates 2006). Executive atten-tion represents a neurodevelopmental processin children and adolescents, the alteration ofwhich could affect the propensity for the de-velopment of a number of disorders.

It is thus important to link efforts at reme-diation to the training of underlying brain net-works. A central aspect of the executive atten-tion network is the ability to deal with conflict.We used this feature to design a set of exer-cises for children adapted from efforts to trainmonkeys to perform tasks during space mis-sions (Rumbaugh & Washburn 1995), whichresulted in training monkeys to resolve con-flict in a Stroop-like task (Washburn 1994).

Our exercises began by training the childto control the movement of a cat on a com-puter screen by using a joystick and to predictwhere an object would move on the screen,given its initial trajectory. Other exercises em-phasized the use of working memory to retaininformation for a matching-to-sample taskand the resolution of conflict.

We have tested the efficacy of a very brieffive days of attention training with groups of4- and 6-year-old children (Rueda et al. 2005).The children were brought to the laboratoryfor seven days for sessions lasting approxi-mately 40 minutes, conducted over a two-to three-week period. The first and last daysinvolved assessment of the effects of train-ing by use of the ANT, a general test of in-telligence (the K-BIT; Kaufman & Kaufman1990), and a temperament scale (the Chil-dren’s Behavior Questionnaire or CBQ;Rothbart et al. 2001). During administra-tion of the ANT, we recorded 128 chan-nels of EEG to observe the amplitude andtime course of activation of brain areas associ-ated with executive attention in adult studies(Rueda et al. 2005).

Five days is of course a minimal amountof training to influence the developmentof networks that develop over many years.Nonetheless, we found a general improve-ment in intelligence in the experimentalgroup as measured by the K-BIT. This was


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due to improvement of the experimentalgroup in performance on the nonverbal por-tion of the IQ test. Our analysis of the brainnetworks using EEG recording further sug-gested that the component most closely re-lated to the anterior cingulate in prior adultstudies changed significantly in the trainedchildren to more closely resemble what isfound in adults (Rueda et al. 2005).

We also found evidence that prior to train-ing, performance on the ANT, as well as ac-tivity of the underlying network, appearedto be related to differences in at least onedopamine gene (Rueda et al. 2005). As thenumber of children who undergo our train-ing increases, we can examine aspects of theirtemperament and genotype to help us under-stand who might most benefit from attentiontraining. Since those with the poorest initialattention seemed to show the most benefit,it is possible that better results will be ob-tained with children who have more severeattentional disorders.

We are beginning to examine the precur-sors of executive attention in even youngerchildren, less than one year of age, with thegoal of understanding the origin of executiveattention. We are also genotyping children inan effort to examine the candidate genes foundpreviously to be related to the efficiency of theexecutive attention networks. There is alreadysome evidence in the literature with olderchildren who suffer from attention deficit hy-peractivity disorder (ADHD) that using at-tention training methods can produce im-provement in the ability to concentrate and ingeneral intelligence (Kerns et al. 1999,Klingberg et al. 2002, Shavlev et al. 2003). Asa result, we are also working with other groupsto carry out attention training in children withlearning-related problems such as ADHD andautism. These projects will test whether theprograms are efficacious with children whohave special difficulties with attention as partof their disorder. We also hope to have somepreschools adopt attention training as a spe-cific part of their preschool curriculum. Thiswould allow training over more extensive pe-

riods and testing of other forms of trainingsuch as those designed for social groups (Mills& Mills 2000).

We believe that the evidence we have ob-tained for the development of specific brainnetworks during early childhood provides astrong rationale for sustained efforts to seeif we can improve the attentional abilities ofchildren. In addition, it will be possible to de-termine how well such methods might gen-eralize to the learning of the wide variety ofskills that must be acquired during school.

INTEGRATION OFPSYCHOLOGICAL SCIENCE

In this article, we have tried to cover top-ics from many areas of psychology, includingcognitive, physiological, developmental, indi-vidual differences, social, clinical, and quanti-tative areas. Each of these areas can be shownto be involved as we examine attention net-works. Below we argue that most, if not all,of the topics of psychological science can alsobenefit from a common network approach.

Generality

Fifteen years of cognitive studies using neu-roimaging have laid out large-scale networksunderlying many cognitive and emotionaltasks (see Table 4 for a partial list). In severalfields of research, each area of activity (node)can be associated with a particular functionor mental operation (see Posner 2004 for areview). A number of these nodes are ac-tive during a given task. The organization ofthese nodes in real time constitutes a networkroughly like Hebb’s cell assembly, but involv-ing brain areas rather than individual cells. Ofcourse, not all authors agree on exactly whatthe function of each node is, and some feelthat parts of the brain (e.g., frontal lobes orsubcortical areas) are more likely to carry outa number of functions.

We have also attempted to show that thestudy of attention involves all the branches ofpsychology. Attention has a detailed anatomy


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Table 4 Some networks studied by neuroimaging∗

Function Selected referencesArithmetic Dehaene (1997, figure 8.5)Autobiographical memory Fink et al. 1996Faces Ochsner et al. (2006)Fear Haxby (2004)Music Levitin (2006)Object perception Grill-Spector (2004)Reading and listening Posner & Raichle (1996)Reward Knutson (2003)Self reference Johnson et al. 2005Spatial navigation Shelton & Gabreli (2002)Working memory Smith et al. 1998, Ungerleider et al. 1998

∗References cited are illustrative and are not meant to be comprehensive.

that involves large-scale brain areas and detailsof cellular structure. Nevertheless, the func-tions of attention are best studied at the cogni-tive level, which reveals their role in tasks thatapproximate those of daily life. Networks de-velop under the influence of social and culturalfactors and of genes. The networks supportnot only the general functions of attentioncommon to all people, but also the individualdifferences that relate to aspects of temper-ament and intelligence. We argue that whatis true of attention is also true of the manyother networks that have now been studiedby imaging.

The network approach is general in that itcovers the topics of cognitive and emotionalmechanisms, and as we have shown above forattention, it allows an approach to issues ofsocialization and cultural influence, genetics,clinical diagnosis and remediation and theirrelative individual differences—in short, to allof the traditional areas of psychology.

Results of neuroimaging research also pro-vide an answer to the old question of whetherthought processes are localized. Although thenetwork that carries out cognitive tasks is dis-tributed, the mental operations that consti-tute the elements of the task are localized. Agood example is the orienting network. Thisnetwork carries out a very simple function ofproviding a priority to a particular source of

sensory input, for example, to the particularlocation at which a visual event will occur. Tocarry out this function, the superior parietallobe, temporal parietal junction, frontal eyefields, superior colliculus, and thalamus areall involved. However, they carry out quitedifferent functions, not all of which are fullyunderstood. For example, the temporal pari-etal junction is involved in interrupting a fo-cus of attention while the superior colliculusand frontal eye fields appear to move the in-dex of attention, with or without eye move-ments. Building upon the idea of localizationof underlying operations, imaging methodsare now being applied to studies of the cir-cuitry, plasticity, and individual developmentof neural networks. Working together withcellular and genetic methods, there is move-ment toward a more unified view of the roleof the human brain in supporting the humanmind.

Another general distinction separates con-scious and unconscious processes central topsychological thinking. It has been shown thatspecial networks are active only when itemsare conscious (Dehaene et al. 2003). Studiesof self-reference have also suggested activa-tion of specific networks of neural areas whenthe self is the object of thought (Gusnard et al.2001).

The study of human and animal geneticsis also illuminating the basis for individual-ity. A view of brain networks developing un-der combined genetic and experiential controlprovides a systematic basis for understandinghow common networks can be linked throughthe study of genetic polymorphisms and thesocialization of individual differences. By re-lating task performance to self- and other re-port questionnaires, as we have done in ourwork, it is possible to examine the role of cul-tural and other social processes in the devel-opment of these networks.

Hebb provided a basis for viewing brainactivity in terms of networks that could com-pute the various functions underlying humanthoughts and feelings. He also attempted totreat all the issues that would normally be


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included in elementary psychology textbooksin terms of this common framework. With theaddition of new methods, we believe that thetask of integrating all aspects of scientific psy-chology in a common framework is even morefeasible than when Hebb undertook it.

At a scientific level, Hebb’s dream ofan integrated psychology is coming about.Sternberg & Grigorenko (2001) laid out thepolitical and social impediments that any ef-

fort at an integrated psychology faces, suchas identifications of scholars with psychologi-cal subdisciplines (e.g., clinical, social) ratherthan with the phenomena they study, cre-ating unnecessary boundaries. These factorsmay unfortunately lead to a failure to realizethe old dream of an integrated science, butthe opportunities are clearly there. New toolsare helping, and Hebb’s efforts have led theway.

ACKNOWLEDGMENTS

We acknowledge the support and aid of many people at the University of Oregon, the SacklerInstitutes, and Washington University in this work. Grant support was provided by NIMHand the McDonnell and Dana Foundations.

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Contents ARI 8 November 2006 21:2

Annual Review ofPsychology

Volume 58, 2007

Contents

Prefatory

Research on Attention Networks as a Model for the Integration ofPsychological ScienceMichael I. Posner and Mary K. Rothbart � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 1

Cognitive Neuroscience

The Representation of Object Concepts in the BrainAlex Martin � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 25

Depth, Space, and Motion

Perception of Human MotionRandolph Blake and Maggie Shiffrar � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 47

Form Perception (Scene Perception) or Object Recognition

Visual Object Recognition: Do We Know More Now Than We Did 20Years Ago?Jessie J. Peissig and Michael J. Tarr � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 75

Animal Cognition

Causal Cognition in Human and Nonhuman Animals: A Comparative,Critical ReviewDerek C. Penn and Daniel J. Povinelli � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 97

Emotional, Social, and Personality Development

The Development of CopingEllen A. Skinner and Melanie J. Zimmer-Gembeck � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 119

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Biological and Genetic Processes in Development

The Neurobiology of Stress and DevelopmentMegan Gunnar and Karina Quevedo � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 145

Development in Societal Context

An Interactionist Perspective on the Socioeconomic Context ofHuman DevelopmentRand D. Conger and M. Brent Donnellan � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 175

Culture and Mental Health

Race, Race-Based Discrimination, and Health Outcomes AmongAfrican AmericansVickie M. Mays, Susan D. Cochran, and Namdi W. Barnes � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 201

Personality Disorders

Assessment and Diagnosis of Personality Disorder: Perennial Issuesand an Emerging ReconceptualizationLee Anna Clark � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 227

Social Psychology of Attention, Control, and Automaticity

Social Cognitive Neuroscience: A Review of Core ProcessesMatthew D. Lieberman � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 259

Inference, Person Perception, Attribution

Partitioning the Domain of Social Inference: Dual Mode and SystemsModels and Their AlternativesArie W. Kruglanski and Edward Orehek � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 291

Self and Identity

Motivational and Emotional Aspects of the SelfMark R. Leary � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 317

Social Development, Social Personality, Social Motivation,Social Emotion

Moral Emotions and Moral BehaviorJune Price Tangney, Jeff Stuewig, and Debra J. Mashek � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 345

viii Contents

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The Experience of EmotionLisa Feldman Barrett, Batja Mesquita, Kevin N. Ochsner,

and James J. Gross � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 373

Attraction and Close Relationships

The Close Relationships of Lesbian and Gay MenLetitia Anne Peplau and Adam W. Fingerhut � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 405

Small Groups

OstracismKipling D. Williams � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 425

Personality Processes

The Elaboration of Personal Construct PsychologyBeverly M. Walker and David A. Winter � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 453

Cross-Country or Regional Comparisons

Cross-Cultural Organizational BehaviorMichele J. Gelfand, Miriam Erez, and Zeynep Aycan � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 479

Organizational Groups and Teams

Work Group DiversityDaan van Knippenberg and Michaéla C. Schippers � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 515

Career Development and Counseling

Work and Vocational Psychology: Theory, Research,and ApplicationsNadya A. Fouad � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 543

Adjustment to Chronic Diseases and Terminal Illness

Health Psychology: Psychological Adjustmentto Chronic DiseaseAnnette L. Stanton, Tracey A. Revenson, and Howard Tennen � � � � � � � � � � � � � � � � � � � � � � � � � � � 565

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Research Methodology

Mediation AnalysisDavid P. MacKinnon, Amanda J. Fairchild, and Matthew S. Fritz � � � � � � � � � � � � � � � � � � � � � 593

Analysis of Nonlinear Patterns of Change with Random CoefficientModelsRobert Cudeck and Jeffrey R. Harring � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 615

Indexes

Cumulative Index of Contributing Authors, Volumes 48–58 � � � � � � � � � � � � � � � � � � � � � � � � � � � 639

Cumulative Index of Chapter Titles, Volumes 48–58 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 644

Errata

An online log of corrections to Annual Review of Psychology chapters (if any, 1997 to thepresent) may be found at http://psych.annualreviews.org/errata.shtml

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