Discussion 2: Attention
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Research on Attention Networks as a Model for the Integration of Psychological Science Michael I. Posner and Mary K. Rothbart1
Psychology Department, University of Oregon, Eugene, Oregon 97403-1291; email: mposner@darkwing.uoregon.edu, maryroth@darkwing.uoregon.edu
Annu. Rev. Psychol. 2007. 58:1–23
First published online as a Review in Advance on October 9, 2006
The Annual Review of Psychology is online at http://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
1 We appreciate the invitation of the editors of the Annual Review of Psychology to submit a prefatory essay to this year’s volume. We have taken the opportunity to propose a unified basis for psychological science based upon an effort to combine experimental and differential approaches to the field. This article is an improved and expanded version of an earlier one along these lines (Posner & Rothbart 2004), and its developmental aspects are further expanded in a book (Posner & Rothbart 2007).
Key Words
attention, candidate genes, orienting, neural networks, temperament
Abstract As Titchener pointed out more than one hundred years ago, at- tention is at the center of the psychological enterprise. Attention research investigates how voluntary control and subjective experi- ence arise from and regulate our behavior. In recent years, attention has been one of the fastest growing of all fields within cognitive psy- chology and cognitive neuroscience. This review examines attention as characterized by linking common neural networks with individual differences in their efficient utilization. The development of atten- tional networks is partly specified by genes, but is also open to specific experiences through the actions of caregivers and the culture. We believe that the connection between neural networks, genes, and socialization provides a common approach to all aspects of human cognition and emotion. Pursuit of this approach can provide a basis for psychology that unifies social, cultural, differential, experimen- 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.
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Attention: the regulating of various brain networks by attentional networks involved in maintaining the alert state, orienting, or regulation of conflict
Neural networks: a number of brain areas that when orchestrated carry out a psychological function
Contents
INTRODUCTION . . . . . . . . . . . . . . . . . 2 HEBB’S NETWORK
APPROACH . . . . . . . . . . . . . . . . . . . . 4 New Tools . . . . . . . . . . . . . . . . . . . . . . . 5
IMAGING ATTENTION NETWORKS . . . . . . . . . . . . . . . . . . . 6 Simulating Attention Networks . . . 8
INDIVIDUAL EFFICIENCY . . . . . . . 9 NETWORK DEVELOPMENT . . . . 11
Socialization and Culture . . . . . . . . . 13 PATHOLOGY OF ATTENTION
NETWORKS . . . . . . . . . . . . . . . . . . . 15 Plasticity . . . . . . . . . . . . . . . . . . . . . . . . . 16
INTEGRATION OF PSYCHOLOGICAL SCIENCE . . 17 Generality . . . . . . . . . . . . . . . . . . . . . . . 17
INTRODUCTION
Can psychology be a unified science, or must it remain fragmented and subject to transient research interests rather than cumulative de- velopment? Is psychology teachable or must psychology textbooks remain encyclopedias of often unrelated findings? In this article, we examine research on attention to suggest ways of looking at psychological questions and findings that might lead to hopeful answers to these questions. We offer a model based on Hebbian psychology that reaches out from biological roots to tackle questions of emo- tion and thought as well as behavior. We, with many of our colleagues, see different levels of analysis in psychology as informing each other, with each level of equal scientific valid- ity. Bridging these levels can allow a higher level of understanding and prediction.
We argue that D.O. Hebb (see Figure 1), beginning with his monograph in 1949 and continuing through a series of introductory textbooks (1958, 1966), has convincingly pre- sented the basis for such integration. This in- tegration lies in understanding how genes and experience shape neural networks underlying human thoughts, feelings, and actions.
At the time Hebb wrote his monograph, relatively little was known about how the structure and organization of the central ner- vous system contribute to the functions ob- served in psychological studies. This led Hebb to talk in terms of the conceptual nervous system, that is, ideas about its structure that might be imagined or inferred from psycho- logical studies. We suggest that the meth- ods available to Hebb, mostly animal research and behavioral human experiments, were not sufficient to provide empirical methods for linking his conceptual nervous system to real events in the human brain. This methodology has now been provided by neuroimaging. Al- though Hebb also recognized the importance of studying individual differences in intelli- gence and affect, at that time there were no methods for exploring the specific genes that are an important source of these differences. The human genome project has provided new methods for exploring this issue.
In Hebb’s time, the idea of a network (cell assembly or phase sequence) was a rather vague verbal abstraction that did not allow for models that could produce specific pre- dictions. As a result of the rapid changes in cellular biology (Bullock et al. 2005) and in the mathematics of multilevel networks, this too has changed (Rumelhart & McClelland 1986). Although early versions of these net- works were inspired by simple versions of neurons as all-or-none elements, more recent versions (O’Reilly & Munakata 2000) have be- gun to use the details of neuroanatomy and cellular structure as provided by imaging and cellular studies to develop networks that take more realistic advantage of the structure of the human brain. Hebb’s basic idea, together with the new methodological tools and new disciplines (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 strategy advocated 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
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Figure 1 Photograph of Professor Donald O. Hebb.
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the theoretical questions that need to be ad- dressed by neuroimaging and genetic studies are 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 when a 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 was immediately recognized as providing the po- tential for an integrated psychology. One reviewer (Attneave 1950, p. 633) wrote:
I believe The Organization of Behavior to be the most important contribution to psycho- logical theory in recent years. Unlike those of his contemporaries who are less interested in 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 determined facts of behavior, as they are, and account for them in terms of events within the central nervous system.
The most important basic idea that Hebb presented was the cell assembly theory out- lined in chapters 4 and 5 of his book (Goddard 1980, Harris 2005). Hebb argued that ev- ery psychological event, sensation, expecta- tion, emotion, or thought is represented by the flow of activity in a set of interconnected neurons. Learning occurs by a change in synaptic strength when a synapse conducts excitation at the same time the postsynap- tic neuron discharges. This provided a basis for the modification of synapses and showed how neural networks might be organized un- der the influence of specific experiences. The Hebb synapse plays a central role in modern neuroscience (see Kolb 2003, Milner 2003, Sejnowski 2003). There are important new developments in the study of synapses and in the discovery of other influences among neu-
rons and between neurons and other brain cells (Bullock et al. 2005). These develop- ments have reduced the gap between networks revealed in imaging studies and the complex intracellular activity that underlies them. In particular, they show that learning may reflect the activity of interactions at many time scales and may be modified by aspects of the organ- ism’s overall state.
Hebb also introduced the concept of the phase sequence involved in the coordination of multiple cell assemblies. He recognized the importance of the temporal correspondence between assemblies. In recent years these ideas have been supported by studies show- ing that synchronization of brain areas may be critical to detecting stimuli (Womelsdorf et al. 2006) and for transfer of information between remote areas (Nikolaev et al. 2001). In later years, Hebb (1958) developed an introductory psychology textbook that integrated much of psychology using the framework provided by his 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 approach to emotions in their social contexts, individ- ual differences in intelligence, and abnormal psychology.
Despite his efforts and those of his fol- lowers at McGill and elsewhere, Hebb’s work was 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 the major problems with Hebb’s framework was that 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 a conceptual nervous system inferred from psy- chological studies of motivation. This idea
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remains of key importance. Despite our growing understanding of the function and structure 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 only within an integrated field of psychology can knowledge at all levels be used to develop and constrain these hypotheses.
New Tools
Several major late-twentieth century events give improved prospects for an integration of psychological science around the ideas in- troduced by Hebb. Cell assemblies and phase sequences 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 a much better position to develop detailed the- ories integrating information from physiolog- ical, cognitive, and behavioral studies.
In addition, new neuroimaging methods now allow us to examine neuronal activity in terms of localized changes in blood flow or metabolism in positron emission tomogra- phy (PET) or changes in blood oxygenation by functional magnetic resonance imaging (fMRI). By using tracers that bind to different transmitters, PET can also be used to examine transmitter density. By measuring electrical (electroencephalogram, or EEG) and mag- netic (magnetoencephalogram) signals out- side the skull, the time course of activation of different brain areas localized by fMRI can be measured (Dale et al. 2000). Pathways of activation can also be imaged by use of dif- fusion tensor imaging, a form of MRI that traces white matter tracts. In addition to the study of naturally occurring lesions, interrupt- ing information flow by transcranial magnetic stimulation (TMS) can produce temporary functional lesions of pathways (see Toga & Mazziotta 1996 for a review of these and other methods). These methods provide a toolkit that can be used either alone or together to
PET: positron emission tomography
fMRI: functional magnetic resonance imaging
Temperament: relatively enduring biological characteristics of individuals that include both reactive and self-regulatory (attentional) aspects
make human brain networks accessible for de- tailed physiological study.
Some have thought that the influence of imaging has been merely to tell us where in the brain things have happened (Utall 2001). Certainly many, perhaps even most, imaging studies have been concerned with anatomical issues. As Figure 2 illustrates, several func- tions of attention have been shown to involve specific anatomical areas that carry out impor- tant functions. However, imaging also probes neural networks that underlie all aspects of human thought, feelings, and behavior. The full significance of imaging for (a) viewing brain networks, (b) examining their compu- tation in real time, (c) exploring how they are assembled in development, and (d ) their plas- ticity following physical damage or training is a common theme in current research that is just beginning to reach its potential.
A third development, the mapping of the human genome (Venter et al. 2001), offers the potential for an increased understanding of the physical basis for individual differences, including individual differences in tempera- ment and personality. Many genes exhibit a number of relatively high-frequency variants (polymorphisms) that can code for different physical 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 different efficiency in binding to dopamine and thus differences in underlying neural networks. In a number of cases, it has been possible to relate these genetic differences to individual per- formance in tasks involving the network (see Fossella & 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 all members of the species.
We recognize that not all topics of psy- chology have been sufficiently explored to il- lustrate this framework and that limitations in 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 2 Anatomy of three attentional networks: alerting, orienting, and executive attention (from Posner & Rothbart 2007).
temperament, areas of our study, can be used to consider how a large number of psycho- logical questions can be explored using the neural network framework outlined by Hebb. Attention serves as a basic set of mechanisms that underlie our awareness of the world and the voluntary regulation of our thoughts and feelings. 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 networks related to other aspects of human behavior.
Any approach based on neural networks raises the issue of crude reductionism. Many agree 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, and self-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 within a brain network framework, as Hebb (1955) illustrated in his Textbook of Psychology.
IMAGING ATTENTION NETWORKS
Functional neuroimaging has allowed many cognitive tasks to be analyzed in terms of the brain areas they activate, and studies of atten- tion have been among those most often ex- amined (Corbetta & Shulman 2002, Driver et 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 out the 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 in Table 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 executive attention involves mechanisms for monitor- ing and resolving conflict among thoughts, feelings, and responses. The alerting system has been associated with thalamic as well as frontal and parietal regions of the cortex (Fan et al. 2005). A particularly effective way to vary alertness has been to use warning signals prior to targets. The influence of warning signals on the level of alertness is thought to be due to modulation of neural activity by the neu- rotransmitter norepinephrine (Marrocco & Davidson 1998).
Orienting involves aligning attention with a 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 orienting system for visual events has been associated with posterior brain areas, including the superior parietal lobe and temporal parietal junction, and in addition, the frontal eye fields (Corbetta & Shulman 2002). Orienting can be manipulated by presenting a cue indicating where in space a target is likely to occur, thereby directing attention to the cued loca- tion (Posner 1980). It is possible to determine the anatomy influenced by the cue separately from that influenced by the target by using MRI to trace changes in the blood that specif- ically follow the cue. This method is called event-related functional magnetic resonance imaging, and its use has suggested that the superior parietal lobe is associated with orienting following the presentation of a cue (Corbetta & Shulman 2002). The superior parietal lobe in humans is closely related
Orienting: the interaction of a brain network with sensory systems designed to improve the selected signal
Table 1 A summary of the anatomy and chemical modulators involved in the alerting, orienting, and executive attention networks.
Function Structures Modulator Orient Superior parietal Acetylcholine
Temporal parietal junction Frontal eye fields Superior colliculus
Alert Locus coeruleus Norepinephrine Right frontal Parietal cortex
Executive attention Anterior cingulate Dopamine Lateral ventral Prefrontal Basal ganglia
to the lateral intraparietal area (LIP) in monkeys, which is involved in the production of eye movements (Andersen 1989). When a target occurs at an uncued location and attention has to be disengaged and moved to a new location, there is activity in the temporal parietal junction (Corbetta & Shulman 2002). Lesions of the temporal parietal junction lobe and superior temporal lobe have been consistently related to difficulties in orienting (Karnath et al. 2001).
Executive control of attention is often studied by tasks that involve conflict, such as various versions of the Stroop task. In the Stroop task, subjects must respond to the color of ink (e.g., red) while ignoring the color word name (e.g., blue) (Bush et al. 2000). Re- solving conflict in the Stroop task activates midline frontal areas (anterior cingulate) and lateral prefrontal cortex (Botvinick et al. 2001, Fan et al. 2005). There is also evidence for the activation of this network in tasks involv- ing conflict between a central target and sur- rounding flankers that may be congruent or incongruent with the target (Botvinick et al. 2001, Fan et al. 2005).
Recently, the role of the anterior cingulate in modulating sensory input has been demon- strated experimentally by showing enhanced connectivity between the anterior cingulate cortex and the sensory modality to which the
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ANT: attention network test
RT: reaction time
ACT-R: adaptive control of thought-rational
person is asked to attend (Crottaz-Herbette & Menon 2006). This finding supports the gen- eral idea that anterior cingulate cortex activity regulates 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 executive attention network (MacDonald et al. 2000). In accord with the findings in recent neuroimag- ing studies (Beauregard et al. 2001, Ochsner et al. 2001), we have argued that the exec- utive attention network is involved in self- regulation of positive and negative affect as well as a wide variety of cognitive tasks under- lying intelligence (Duncan et al. 2000). This idea suggests an important role for attention in 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 network approach is that it lends itself to the devel- opment of precise computer models that al- low both summarization of many findings in the field and prediction of new findings. Cur- rently, symbolic models, such as rule-based systems (Newell 1990), appear to be a good way to capture data from reaction time and other psychological findings. In these models, cognitive functions are represented as chains of production rules and can be identified with the mental operations postulated by cogni- tive studies. On the other hand, subsymbolic models, such as connectionist models (e.g., O’Reilly & Munakata 2000), permit a more biologically realistic implementation of the operations and closer links to imaging studies.
We have developed the attention network test (ANT) to examine individual differences in 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 reaction time (RT) between conditions to measure the efficiency of each network. Each trial begins with a cue (or a blank interval, in the no-cue
condition) that informs the participant either that a target will occur soon or where it will occur, or both. The target always occurs either above or below fixation, and consists of a cen- tral arrow, surrounded by flanking arrows that can point either in the same direction (con- gruent) or in the opposite direction (incon- gruent). Subtracting RTs for congruent from incongruent target trials provides a measure of conflict resolution and assesses the efficiency of the executive attention network. Subtract- ing RTs obtained in the double-cue condition that provides information on when but not where the target will occur from RTs in the no-cue condition gives a measure of alerting due 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 the spatial cue, but not the central cue, provides valid information about where a target will occur.
The attention network task has now been simulated in the framework of the adaptive control 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 of the direction of the arrow in the center and in- volves an attention switch and response initia- tion. Each of these operations is implemented by a chain of production rules. The operations are similar to those discussed in most psycho- logical studies and localized in neurological studies. For example, the switching of atten- tion based on the peripheral cue or target is thought to be implemented by the temporal parietal 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 and associative, biologically realistic algorithm (LEABRA) (O’Reilly & Munakata 2000). The subroutines of ACT-R are now replaced by specific connections between hypothesized neurons. These neurons are somewhat real- istic, and are designed to represent the known
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properties of specific brain areas. Thus, the orienting network can be designed to reflect the known properties of the frontal eye fields, superior parietal lobe, and temporal parietal junction, and they can be connected within the simulation. As shown in Figure 3, sim- ulations do a reasonable job of fitting with the known ANT data, although improvement can be expected in the future. The symbolic model makes contact with the mental opera- tions related to imaging, whereas the connec- tionist framework allows a strong treatment of the underlying biology. Together they illus- trate how network views provide a computa- tional means for summarizing many findings within psychology, allowing novel predictions reflecting properties of the network.
INDIVIDUAL EFFICIENCY
Psychology is often divided into two ap- proaches that are almost completely separate in the literature (but see Gardner 1983 for an effort at integration). The discussion above focused on general features of the human mind, such as the ability to attend. Another approach 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 the emotions. Almost all studies of attention have been concerned either with the general abili- ties involved or with effects of brain injury or pathology on attention. However, it is clear that normal individuals differ in their ability to attend to sensory events, and it is even clearer that they differ in their ability to concentrate for 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, each of 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 be involved in the development of utilization of attentional networks
The ability to measure differences in at- tention among adults raises the question of the degree to which attention is heritable. To explore this issue, the ANT was used to assess attention in monozygotic and dizygotic same- sex twins (Fan et al. 2001). Strong heritabil- ity was found for the executive network, some heritability for the orienting network, and no apparent 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 neuromodulator dopamine (see Table 1) as a way of search- ing for candidate genes that might relate to the efficiency of the networks (Fossella et al. 2002). To do this, 200 persons performed the ANT and were genotyped to examine frequent polymorphisms in genes related to dopamine. We found significant association of the dopamine 4 receptor and monoamine oxidase A genes. We then conducted a neu- roimaging experiment in which persons with different alleles of these two genes were com- pared while they performed the ANT (Fan et al. 2003). Groups with different alleles of these genes showed differences in perfor- mance on the ANT and also produced sig- nificantly different activations in the anterior cingulate, a major node of the executive atten- tion network.
Recent studies have confirmed and ex- tended these observations to other dopamine genes and to the orienting network. In two dif- ferent studies employing other conflict tasks, the catecholamine-O-methyltransferase gene was linked to the mental operations re- lated to resolving conflict (Blasi et al. 2005, Diamond et al. 2004). Different alleles of cholinergic genes were also related to perfor- mance on orienting tasks such as visual search (Parasuraman et al. 2005), thus confirming the link between orienting and the neuromodula- tor acetylcholine (see Table 1).
Hebb (1955) thought that most of the networks involved in higher functions were shaped primarily through experience. We now know that there is a great deal in common
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Output Executive 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 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.
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among humans in the anatomy of these high- level networks, and thus that they must have a basis within the human genome. It seems likely that the same genes that are related to individual differences in attention are also im- portant in the development of the attentional networks that are common among people. Some of these networks are also common to nonhuman animals. By examining these net- works in animals, it should be possible to test these assumptions further and to understand the role of genes in shaping networks.
Of importance for this effort is the de- velopment of methods to manipulate relevant genes in specific anatomical locations that are important nodes of a particular network. Usu- ally genes are expressed at multiple locations so that changes (e.g., gene knockouts) are not specific 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 of network development will become a very pro- ductive link between genes and both normal and pathological psychological function.
NETWORK DEVELOPMENT
Every parent of more than one child recog- nizes that from birth, infants are individuals with their own distinct characteristics and dis- positions. These include reactive traits such as emotionality, activity level, and orienting to sensory events, and regulatory traits like at- tention focusing and shifting and inhibitory control. We believe that these early devel- oping temperamental differences reflect the maturation of particular neural networks. We have studied individual differences in atten- tion and related these differences to emotional and behavioral control (Rothbart & Rueda 2005, Ruff & Rothbart 1996).
A major advantage of viewing attention as an organ system is to trace the ability of chil- dren and adults to regulate their thoughts and feelings. 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 (Harman et 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 showed how attentional orienting calms that distress while 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 distress remains present and is held by networks in the amygdala. Parents often use manipulation of the infant’s orienting to control distress, and the infant also exhibits coping behaviors in- volving orienting to form the basis for one aspect 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 temperamental effortful control (Gerardi-Caulton 2000). Ef- fortful control is a broad variable identified in temperament research; it includes the ability to inhibit a dominant response in order to pro- duce subdominant response and to detect and correct 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 in a 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 and one to the right. Each key displayed a picture, and on every trial, a picture identical to one of the pair appeared on either the left or right side of the screen. Children were rewarded for responding to the identity of the stimu- lus, regardless of its spatial compatibility with the matching response key. Reduced accuracy and slowed reaction times for spatially incom- patible trials relative to spatially compatible trials reflected the effort required to resist the dominant response and to resolve conflict between these two competing dimensions. Performance on this task produces a clear in- terference effect in adults and activates the anterior cingulate (Fan et al. 2003). Children 24 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, but like adults, responded more slowly and with reduced 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) by using a version of the ANT adapted for chil- dren (Table 2). In some respects, results were remarkably similar to those found for adults using the adult version of the task. Reaction times for the children were much longer, but
they showed similar independence among the three networks. Children had much larger scores for conflict and alerting, suggesting that they have trouble in resolving conflict and in maintaining the alert state when not warned of the new target. Rather surprisingly, the ability to resolve conflict in the flanker task, as measured by the ANT, remains about the same from age eight to adulthood (see Table 2).
There is considerable evidence that the executive 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 that draw 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 the mind, for example, by formal disciplines like mathematics or Latin, does not generalize be- yond the specific domain trained (Thorndike 1903, Simon 1969). However, attention may be an exception to this idea. Attention involves specific brain mechanisms, as we have seen, but its function is to influence the operation of other brain networks (Posner & Rothbart 2007). Anatomically, the network involving resolution of conflict overlaps with brain ar- eas related to general intelligence (Duncan et al. 2000). Training of attention either ex- plicitly or implicitly is sometimes a part of the school curriculum (Posner & Rothbart 2007), but additional studies are needed to deter- mine exactly how and when attention training can best be accomplished and its long-lasting importance.
Socialization and Culture
Cognitive measures of conflict resolution have been linked to aspects of children’s self- control in naturalistic settings. Children who are relatively less affected by spatial conflict also received higher parental ratings of tem- peramental effortful control and higher scores on laboratory measures of inhibitory control (Gerardi-Caulton 2000). Questionnaires have shown that the effortful control factor, defined in terms of scales measuring attentional focus- ing, inhibitory control, low intensity pleasure, and perceptual sensitivity (Rothbart & Rueda 2005), 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 impulsive approach tendencies.
Empathy is strongly related to effortful control, 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 distress or pleasure. Imaging work shows that sad faces activate the amygdala. As sadness increases, this activation is accompanied by activity in the anterior cingulate as part of the attention network (Blair et al. 1999). It seems likely that the cingulate activity represents the basis for our attention to the distress of others.
Developmental studies find two routes to successful socialization. A strongly reac- tive amygdala would provide the signals of distress 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 relatively easy to socialize. In the absence of this form of control, development of the cingu- late would allow appropriate attention to the signals provided by amygdala activity. Con- sistent with its influence on empathy, effort- ful control also appears to play a role in the development 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, internalized conscience is facilitated for children high in effortful control (Kochanska et al. 1996). Two separable control systems, one reactive (fear) and one self-regulative (effortful control), appear to regulate the development of conscience.
Individual differences in effortful control are also related to some aspects of metacog- nitive knowledge, such as theory of mind, which is the knowledge that people’s behavior is guided by their beliefs, desires, and other mental states (Carlson & Moses 2001). Tasks that require the inhibition of a prepotent re- sponse are related to theory of mind tasks even when other factors, such as age, intelli- gence, and working memory, are factored out (Carlson & Moses 2001). The mechanisms of self-regulation and of theory of mind share a similar developmental time course, with advances in both areas between the ages of 2 and 5.
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Efforts to determine the neural network involved in theory of mind tasks reveal some of the reasons for the common developmen- tal time course in self-regulation. Theory of mind tasks activate a network that includes areas of the anterior cingulate that are also in- volved in self-regulation, as well as temporal lobe areas (Gallagher & Frith 2003). These anatomical links provide further support for efforts to integrate psychological topics at a network level.
Emotion, thought, and behavior form a cluster of temporally associated processes in specific situations as experienced by the child. Single and repeated life experiences can thus shape connections between elicited emotion, conceptual understanding of events, and use of coping strategies to deal with these events. Several theorists have made contributions to this approach (e.g., Epstein 1998, Mischel & Ayduk 2004), but the overall framework is in keeping with the idea of Hebbian learning through network activation. Mischel and his colleagues have recently developed a cogni- tive affective personality theory, making use of cognitive affective units (CAUs) seen to oper- ate within a connectionist network (Mischel & Ayduk 2004). In their model, CAUs are variables encoding the features of situations, which include environmental features as well as self-initiated thoughts.
When they are repeatedly exercised, tem- porally linked clusters of thoughts, emotions, and action tendencies to a particular situation can become highly likely to reoccur and dif- ficult to change. Some of the processes may be conscious and others unconscious; in most cases, the thoughts about affectively signifi- cant material will be self-referent. Research on the distinction between conscious and un- conscious processes has shown that special networks are active only when items are con- scious (Dehaene et al. 2003). Studies of self- reference have also suggested activation of specific networks of neural areas (Gusnard et al. 2001).
In applying this approach to the control of distress, one basic question is how to weaken
the mental connection between the situation and its component reactions. In the eastern tradition, this is done partly through medita- tion, when ideas can be brought up in a con- text of calmness and safety, and partly through changing the view of the self so that situations will become less threatening. Links between thoughts and emotions or action tendencies are also weakened. Western therapy similarly works through the client’s patterns of reac- tion, attempting to rework previously consoli- dated patterns and to provide new frameworks for meaning. From a developmental view, one would attempt to give the child the kinds of experiences that will form favorable and non- injurious mental habits.
Clusters of reactions are found within the young child’s temperament. Later clusters of thought, emotion, and behavior will also be based on individual differences in personal- ity, including emotions, expectancies, beliefs, values, goals, self-evaluations, appraisals, and thoughts about the situation, self, and/or oth- ers. These clusters will be influenced by tem- peramental predispositions, but they will also be influenced by socialization and experience. Coping and the application of effortful con- trol operate when “the subjective meaning of the situation, including its self-relevance and personal importance, are appraised. The appraisal itself activates a cascade of other cognitive-affective representations within the system—expectations and beliefs, affective re- actions, values and goals” (Mischel & Ayduk 2004, p. 105).
Different coping strategies follow and may be consolidated or rejected in the future, de- pending in part on their consequences. To de- velop this idea, Mischel & Ayduk (2004) give the example of individual differences in rejec- tion sensitivity. When persons are particularly prepared to perceive rejection from others, at- tention is likely to be narrowly focused on this possibility, and it has been demonstrated that rejection 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 serve to fend off feelings of rejection. These strate- gies in turn can also provide further support for the idea that the self is unworthy of pos- itive social relationships. Different levels of generality of these clusters are possible. Re- jection sensitivity, for example, might extend to a wide range of human relationships, but the sensitivity may also be more specific, so that only rejection by the child’s peers, but not by adults, is sensitized. The reaction may in fact be limited to a single person in a sin- gle kind of situation, in which case a particular person, but not other persons, elicits rejection sensitivity.
Socialization in western cultures strongly emphasizes the individual, promoting the pur- suit of individual security, satisfaction of indi- vidual desires, and achievement of a positive self-concept. In other cultures, the shaping of the child’s experienced world can be quite dif- ferent. Mascolo et al. (2003) suggest, for ex- ample, that the biological systems on which pride and shame are based are the same across cultures, yet can be shaped in quite differ- ent directions. In the United States, pride re- actions develop as parents and others praise the child’s accomplishments; shame reactions occur when there is self-referent failure. In China, however, parents downplay or criti- cize children’s performance while other adults praise it, leading to more moderate prideful affect. Shame, on the other hand, is directly encouraged by parents and others when chil- dren do not fulfill their obligations to family.
The biological equipment or temperament is thus similar across cultures, but the men- tal habits and representation of self created as a result of the child’s actions varies from culture to culture (Ahadi et al. 1993). By the time a child is a well-socialized member of the society, biologically based responses will have been shaped into a set of values, goals, and representations of the self and others. These representations specify what is good and bad for the person. Even for the child who is not well socialized, cultural socializa- tion may have an effect. In the United States,
for example, a child may pursue a positive view of the self even when achievements result from following goals and values that are not socially acceptable.
PATHOLOGY OF ATTENTION NETWORKS
Much of modern psychology is involved in the diagnosis and treatment of mental illness or disturbance, and the network framework may be an ideal one for incorporating ideas related to clinical remediation. An excellent example of this approach has been in the re- cent studies of Mayberg (2003) on clinical in- terventions for depression based upon a neural network model. Treatments involved drugs or cognitive behavioral therapy and both forms of therapy were about equally effective, based on the percentage of persons showing im- provement. Imaging data, however, indicated that the two therapies involved very differ- ent brain networks. The drugs remediated a largely subcortical network of brain areas that might be difficult to control voluntarily. The cognitive-behavioral therapy affected cortical networks, 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 in evaluating the outcome of clinical trials.
A similar story may emerge in the study of dyslexia. Many forms of dyslexia involve a difficulty in phonological processing that can be remediated by an intervention targeting the ability to convert visual letters to sound (McCandliss et al. 2003). Following remedi- ation, there is normal activation of a brain region at the boundary between the tem- poral and parietal lobes related to phonol- ogy. However, although these students can now decode words, they do not show flu- ent reading. This may require development of the visual word form system, which in- volves an extrastriate visual region (fusiform gyrus) quite distinct from the phonological areas (McCandliss et al. 2003). It is likely that time spent reading is one way to develop
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Table 3 Disorders that have been related to attentional networks (from Rothbart & Posner 2006)
Alerting Normal aging Attention deficit disorder
Orienting Autism
Executive control Alzheimer’s Borderline personality disorder Schizophrenia 22Q11 deletion syndrome
this brain area, but it may also be possible to create special training exercises that target the area, as has been done for phonological intervention.
The possibility that aspects of brain net- works involved in depression and dyslexia might be remediated by therapies based on training illustrates the close connection be- tween therapeutic interventions designed to correct 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 and education can represent similar approaches to improving network efficiency.
The ANT has been applied to a number of forms of pathology in adults, the aging, adolescents, and children. Table 3 presents classification of a number of disorders that have 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 ANT and other conflict-related tasks provides a ba- sis for the ability of children to regulate their behavior through the use of effortful control. Executive attention has a well-defined neu- roanatomy, and much is known about the role of genes in modulating its efficiency. Diffi- culties in effortful control provide the basis for problems in child socialization and in a
number of disorders of children and adults (Rothbart & Bates 2006). Executive atten- tion represents a neurodevelopmental process in children and adolescents, the alteration of which 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 train monkeys to perform tasks during space mis- sions (Rumbaugh & Washburn 1995), which resulted in training monkeys to resolve con- flict in a Stroop-like task (Washburn 1994).
Our exercises began by training the child to control the movement of a cat on a com- puter screen by using a joystick and to predict where an object would move on the screen, given its initial trajectory. Other exercises em- phasized the use of working memory to retain information for a matching-to-sample task and the resolution of conflict.
We have tested the efficacy of a very brief five days of attention training with groups of 4- and 6-year-old children (Rueda et al. 2005). The children were brought to the laboratory for seven days for sessions lasting approxi- mately 40 minutes, conducted over a two- to three-week period. The first and last days involved assessment of the effects of train- ing by use of the ANT, a general test of in- telligence (the K-BIT; Kaufman & Kaufman 1990), 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 and time 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 amount of training to influence the development of networks that develop over many years. Nonetheless, we found a general improve- ment in intelligence in the experimental group as measured by the K-BIT. This was
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due to improvement of the experimental group in performance on the nonverbal por- tion of the IQ test. Our analysis of the brain networks using EEG recording further sug- gested that the component most closely re- lated to the anterior cingulate in prior adult studies changed significantly in the trained children to more closely resemble what is found 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, appeared to be related to differences in at least one dopamine gene (Rueda et al. 2005). As the number of children who undergo our train- ing increases, we can examine aspects of their temperament and genotype to help us under- stand who might most benefit from attention training. Since those with the poorest initial attention seemed to show the most benefit, it is possible that better results will be ob- tained with children who have more severe attentional disorders.
We are beginning to examine the precur- sors of executive attention in even younger children, less than one year of age, with the goal of understanding the origin of executive attention. We are also genotyping children in an effort to examine the candidate genes found previously to be related to the efficiency of the executive attention networks. There is already some evidence in the literature with older children 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 in general intelligence (Kerns et al. 1999, Klingberg et al. 2002, Shavlev et al. 2003). As a result, we are also working with other groups to carry out attention training in children with learning-related problems such as ADHD and autism. These projects will test whether the programs are efficacious with children who have special difficulties with attention as part of their disorder. We also hope to have some preschools adopt attention training as a spe- cific part of their preschool curriculum. This would allow training over more extensive pe-
riods and testing of other forms of training such as those designed for social groups (Mills & Mills 2000).
We believe that the evidence we have ob- tained for the development of specific brain networks during early childhood provides a strong rationale for sustained efforts to see if we can improve the attentional abilities of children. In addition, it will be possible to de- termine how well such methods might gen- eralize to the learning of the wide variety of skills that must be acquired during school.
INTEGRATION OF PSYCHOLOGICAL SCIENCE
In this article, we have tried to cover top- ics from many areas of psychology, including cognitive, physiological, developmental, indi- vidual differences, social, clinical, and quanti- tative areas. Each of these areas can be shown to be involved as we examine attention net- works. Below we argue that most, if not all, of the topics of psychological science can also benefit from a common network approach.
Generality
Fifteen years of cognitive studies using neu- roimaging have laid out large-scale networks underlying many cognitive and emotional tasks (see Table 4 for a partial list). In several fields of research, each area of activity (node) can be associated with a particular function or mental operation (see Posner 2004 for a review). A number of these nodes are ac- tive during a given task. The organization of these nodes in real time constitutes a network roughly like Hebb’s cell assembly, but involv- ing brain areas rather than individual cells. Of course, not all authors agree on exactly what the function of each node is, and some feel that parts of the brain (e.g., frontal lobes or subcortical areas) are more likely to carry out a number of functions.
We have also attempted to show that the study of attention involves all the branches of psychology. Attention has a detailed anatomy
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Table 4 Some networks studied by neuroimaging∗
Function Selected references Arithmetic Dehaene (1997, figure 8.5) Autobiographical memory Fink et al. 1996 Faces 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. 2005 Spatial 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 details of cellular structure. Nevertheless, the func- tions of attention are best studied at the cogni- tive level, which reveals their role in tasks that approximate those of daily life. Networks de- velop under the influence of social and cultural factors and of genes. The networks support not only the general functions of attention common to all people, but also the individual differences that relate to aspects of temper- ament and intelligence. We argue that what is true of attention is also true of the many other networks that have now been studied by imaging.
The network approach is general in that it covers the topics of cognitive and emotional mechanisms, and as we have shown above for attention, it allows an approach to issues of socialization and cultural influence, genetics, clinical diagnosis and remediation and their relative individual differences—in short, to all of the traditional areas of psychology.
Results of neuroimaging research also pro- vide an answer to the old question of whether thought processes are localized. Although the network that carries out cognitive tasks is dis- tributed, the mental operations that consti- tute the elements of the task are localized. A good example is the orienting network. This network carries out a very simple function of providing a priority to a particular source of
sensory input, for example, to the particular location at which a visual event will occur. To carry out this function, the superior parietal lobe, temporal parietal junction, frontal eye fields, superior colliculus, and thalamus are all involved. However, they carry out quite different functions, not all of which are fully understood. For example, the temporal pari- etal junction is involved in interrupting a fo- cus of attention while the superior colliculus and frontal eye fields appear to move the in- dex of attention, with or without eye move- ments. Building upon the idea of localization of underlying operations, imaging methods are now being applied to studies of the cir- cuitry, plasticity, and individual development of neural networks. Working together with cellular and genetic methods, there is move- ment toward a more unified view of the role of the human brain in supporting the human mind.
Another general distinction separates con- scious and unconscious processes central to psychological thinking. It has been shown that special networks are active only when items are conscious (Dehaene et al. 2003). Studies of self-reference have also suggested activa- tion of specific networks of neural areas when the self is the object of thought (Gusnard et al. 2001).
The study of human and animal genetics is also illuminating the basis for individual- ity. A view of brain networks developing un- der combined genetic and experiential control provides a systematic basis for understanding how common networks can be linked through the study of genetic polymorphisms and the socialization of individual differences. By re- lating task performance to self- and other re- port questionnaires, as we have done in our work, 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 brain activity in terms of networks that could com- pute the various functions underlying human thoughts and feelings. He also attempted to treat all the issues that would normally be
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included in elementary psychology textbooks in terms of this common framework. With the addition of new methods, we believe that the task of integrating all aspects of scientific psy- chology in a common framework is even more feasible than when Hebb undertook it.
At a scientific level, Hebb’s dream of an integrated psychology is coming about. Sternberg & Grigorenko (2001) laid out the political and social impediments that any ef-
fort at an integrated psychology faces, such as identifications of scholars with psychologi- cal subdisciplines (e.g., clinical, social) rather than with the phenomena they study, cre- ating unnecessary boundaries. These factors may unfortunately lead to a failure to realize the old dream of an integrated science, but the opportunities are clearly there. New tools are helping, and Hebb’s efforts have led the way.
ACKNOWLEDGMENTS
We acknowledge the support and aid of many people at the University of Oregon, the Sackler Institutes, and Washington University in this work. Grant support was provided by NIMH and the McDonnell and Dana Foundations.
LITERATURE CITED
Ahadi SA, Rothbart MK, Ye R. 1993. Children’s temperament in the U.S. and China: similarities and differences. Eur. J. Personal. 7:359–78
Andersen RA. 1989. Visual eye movement functions of the posterior parietal cortex. Annu. Rev. Neurosci. 12:377–403
Attneave F. 1950. Book review of D.O. Hebb’s Organization of Behavior. Am. J. Psychol. 63:633– 35
Beauregard M, Levesque J, Bourgouin P. 2001. Neural correlates of conscious self-regulation of emotion. J. Neurosci. 21(RC165)1–6
Blair RJR, Morris JS, Frith CD, Perrett DI, Dolan RJ. 1999. Dissociable neural responses to facial expression of sadness and anger. Brain 1222:883–93
Blasi G, Mattay GS, Bertolino A, Elvevåg B, Callicott JH, et al. 2005. Effect of catechol-O- methyltransferase val158met genotype on attentional control. J. Neurosci. 25(20):5038–45
Botvinick MM, Braver TS, Barch DM, Carter CS, Cohen JD. 2001. Conflict monitoring and cognitive control. Psychol. Rev. 108:624–52
Bullock TH, Bennett MVL, Johnston D, Josephson R, Marder E, Fields RD. 2005. The neuron doctrine, redux. Science 310:791–93
Bush G, Luu P, Posner MI. 2000. Cognitive and emotional influences in the anterior cingulate cortex. Trends Cogn. Sci. 4(6):215–22
Carlson ST, Moses LJ. 2001. Individual differences in inhibitory control in children’s theory of mind. Child Dev. 72:1032–53
Provides an excellent review of the neural areas involved in orienting of attention.
Corbetta M, Shulman GL. 2002. Control of goal-directed and stimulus-driven attention in the brain. Nat. Neurosci. Rev. 3:201–15
Crottaz-Herbtte S, Menon V. 2006. Where and when the anterior cingulate cortex modulates attentional response: combined fMRI and ERP evidence. J. Cogn. Neurosci. 18:766–80
Dale AM, Liu AK, Fischi BR, Buckner R, Beliveau JW, et al. 2000. Dynamic statistical pa- rameter mapping: combining fMRI and MEG for high-resolution cortical activity. Neuron 26:55–67
Dehaene S. 1997. The Number Sense. Oxford, UK: Oxford Univ. Press
www.annualreviews.org • Research on Attentional Networks as a Model 19
A nn
u. R
ev . P
sy ch
ol . 2
00 7.
58 :1
-2 3.
D ow
nl oa
de d
fr om
a rj
ou rn
al s.
an nu
al re
vi ew
s. or
g by
U ni
ve rs
it y
of W
is co
ns in
- M
ad is
on o
n 11
/2 6/
08 . F
or p
er so
na l
us e
on ly
.
ANRV296-PS58-01 ARI 17 November 2006 1:10
Dehaene S, Artiges E, Naccache L, Martelli C, Viard A, et al. 2003. Conscious and sublimi- nal conflicts in normal subjects and patients with schizophrenia: the role of the anterior cingulate. Proc. Natl. Acad. Sci. USA 100(23):13722–27
Diamond A, Briand L, Fossella J, Gehlbach L. 2004. Genetic and neurochemical modulation of prefrontal cognitive functions in children. Am. J. Psychiatry 161:125–32
Driver J, Eimer M, Macaluso E. 2004. Neurobiology of human spatial attention: modulation, generation, and integration. See Kanwisher & Duncan 2004, pp. 267–300
Dumas T, Hostick U, Wu H, Spaltenstein J, Ghatak C, et al. 2005. Maximizing the anatomical specificity of native neuronal promoters by a subtractive transgenic technique. Soc. Neurosci. Abstr.
Duncan J, Seitz RJ, Kolodny J, Bor D, Herzog H, et al. 2000. A neural basis for general intelligence. Science 289:457–60
Eisenberger NI, Lieberman MD, Williams KD. 2003. Does social rejection hurt? An fMRI study of social exclusion. Science 302:290–92
Epstein S. 1998. Cognitive-experiential self-theory: a dual process personality theory with implications for diagnosis and psychotherapy. In Empirical Perspectives on the Psychoanalytic Unconscious, ed. RF Bornstein, JM Masling, 7:99–140. Washington, DC: Am. Psychol. Assoc.
Fan J, Flombaum JI, McCandliss BD, Thomas KM, Posner MI. 2002. Cognitive and brain mechanisms of conflict. Neuroimage 18:42–57
Fan J, Fossella JA, Summer T, Posner MI. 2003. Mapping the genetic variation of executive attention onto brain activity. Proc. Natl. Acad. Sci. USA 100:7406–11
Describes the brain areas activated for each of the three attention networks using fMRI data.
Fan J, McCandliss BD, Fossella J, Flombaum JI, Posner MI. 2005. The activation of attentional networks. Neuroimage 26:471–79
Fan J, McCandliss BD, Sommer T, Raz M, Posner MI. 2002. Testing the efficiency and inde- pendence of attentional networks. J. Cogn. Neurosci. 3(14):340–47
Fan J, Wu Y, Fossella J, Posner MI. 2001. Assessing the heritability of attentional networks. BMC Neurosci. 2:14
Fink GR, Markowitsch HJ, Reinkemeier H, Bruckbauer T, Kessler J, Heiss WD. 1996. Cere- bral representation of one’s own past: neural networks involved in autobiographical mem- ory. J. Neurosci. 16(13):4275–82
Fossella J, Posner MI. 2004. Genes and the development of neural networks underlying cogni- tive processes. The Cognitive Neurosciences III, ed. MS Gazzaniga, pp. 1255–66. Cambridge, MA: MIT Press. 3rd ed.
Fossella J, Sommer T, Fan J, Wu Y, Swanson JM, et al. 2002. Assessing the molecular genetics of attention networks. BMC Neurosci. 3:14
Gallagher HL, Frith CD. 2003. Functional imaging of “theory of mind.” Trends Cogn. Sci. 7:77–83
Gardner F. 1983. Frames of Mind. New York: Basic Books Gerardi-Caulton G. 2000. Sensitivity to spatial conflict and the development of self-regulation
in children 24–36 months of age. Dev. Sci. 3(4):397–404 Goddard GV. 1980. Component properties of the memory machine: Hebb revisited. In The
Nature of Thought: Essays in Honor of D.O. Hebb, ed. PW Jusczyk, RM Klein, pp. 231–47. Hillsdale, NJ: LEA
Grill-Spector K. 2004. The functional organization of the visual ventral pathway and its relation to object recognition. See Kanwisher & Duncan 2004, pp. 169–93
Gusnard DA, Akbudak E, Shulman GL, Raichle ME. 2001. Medial prefrontal cortex and self- referential mental activity: relation to a default mode of brain function. Proc. Natl. Acad. Sci. USA 98:4259–64
20 Posner · Rothbart
A nn
u. R
ev . P
sy ch
ol . 2
00 7.
58 :1
-2 3.
D ow
nl oa
de d
fr om
a rj
ou rn
al s.
an nu
al re
vi ew
s. or
g by
U ni
ve rs
it y
of W
is co
ns in
- M
ad is
on o
n 11
/2 6/
08 . F
or p
er so
na l
us e
on ly
.
ANRV296-PS58-01 ARI 17 November 2006 1:10
Harman C, Rothbart MK, Posner MI. 1997. Distress and attention interactions in early infancy. Motiv. Emot. 21:27–43
Harris KD. 2005. Neural signatures of cell assembly organization. Nat. Neurosci. Rev. 6:399–407 Haxby JV. 2004. Analysis of topographically organized patterns of response in fMRI data:
distributed representation of objects in the ventral temporal cortex. See Kanwisher & Duncan 2004, pp. 83–97
Presents the original theory of how cell assemblies and phase sequences relate to psychological functions.
Hebb DO. 1949. Organization of Behavior. New York: Wiley. Hebb DO. 1955. Drives and the C.N.S. (conceptual nervous system). Psychol. Rev. 62:243–54 Hebb DO. 1958. A Textbook of Psychology. Philadelphia, PA: Saunders Hebb DO. 1966. A Textbook of Psychology. Philadelphia, PA: Saunders. 2nd ed. Johnson SC, Schmitz TW, Kawahara-Baccus TN, Rowley HA, Alexander AL, et al. 2005.
The cerebral response during subjective choice with and without self-reference. J. Cogn. Neurosci. 17(12):1897–906
Kanwisher N, Duncan J, eds. 2004. Attention and Performance XX: Functional Brain Imaging of Visual Cognition. London: Oxford Univ. Press
Karnath HO, Ferber S, Himmelbach M. 2001. Spatial awareness is a function of the temporal not the posterior parietal lobe. Nature 411:950–53
Kaufman AS, Kaufman NL. 1990. Kaufman Brief Intelligence Test Manual. Circle Pines, MN: Am. Guidance Serv.
Kerns KA, Esso K, Thompson J. 1999. Investigation of a direct intervention for improving attention in young children with ADHD. Dev. Neuropsychol. 16:273–95
Klingberg T, Forssberg H, Westerberg H. 2002. Training of working memory in children with ADHD. J. Clin. Exp. Neuropsychol. 24:781–91
Knutson B, Fong GW, Bennett SM, Adams CM, Homme D. 2003. A region of mesial prefrontal cortex tracks monetarily rewarding outcomes: characterization with rapid event-related fMRI. Neuroimage 18(2):263–72
Kochanska G. 1995. Children’s temperament, mothers’ discipline, and security of attachment: multiple pathways to emerging internalization. Child Dev. 66:597–615
Kochanska G, Murray K, Jacques TY, Koenig AL, Vandegeest KA. 1996. Inhibitory control in young children and its role in emerging internationalization. Child Dev. 67:490–507
Kolb B. 2003. The impact of the Hebbian learning rule on research in behavioural neuroscience. Can. Psychol. 44:14–16
Levitin D. 2006. This Is Your Brain on Music. London: Penguin. In press MacDonald AW, Cohen JD, Stenger VA, Carter CS. 2000. Dissociating the role of the dorso-
lateral prefrontal and anterior cingulate cortex in cognitive control. Science 288:1835–38 Marrocco RT, Davidson MC. 1998. Neurochemistry of attention. In The Attentive Brain, ed.
R Parasuraman, pp. 35–50. Cambridge, MA: MIT Press Mascolo MR, Fischer KW, Li J. 2003. Dynamic development of component systems of emo-
tions: pride, shame and guilt in China and the United States. In Handbook of Affective Sciences, ed. RJ Davidson, KR Scherer, HH Goldsmith, pp. 375–408. New York: Oxford Univ. Press
Mayberg HS. 2003. Modulating dysfunctional limbic-cortical circuits in depression: towards development of brain-based algorithms for diagnosis and optimized treatment. Br. Med. Bull. 65:193–207
McCandliss BD, Beck IL, Sandak R, Perfetti C. 2003. Focusing attention on decoding for chil- dren with poor reading skill: design and preliminary test of the word building intervention. Sci. Stud. Read. 7(1):75–104
McCandliss BD, Cohen L, Dehaene S. 2003. The visual word form area: expertise for reading in the fusiform gyrus. Trends Cogn. Sci. 7:293–99
www.annualreviews.org • Research on Attentional Networks as a Model 21
A nn
u. R
ev . P
sy ch
ol . 2
00 7.
58 :1
-2 3.
D ow
nl oa
de d
fr om
a rj
ou rn
al s.
an nu
al re
vi ew
s. or
g by
U ni
ve rs
it y
of W
is co
ns in
- M
ad is
on o
n 11
/2 6/
08 . F
or p
er so
na l
us e
on ly
.
ANRV296-PS58-01 ARI 17 November 2006 1:10
Mills D, Mills C. 2000. Hungarian Kindergarten Curriculum Translation. London: Mills Milner P. 2003. The relevance of D.O. Hebb’s neural network learning rule to today’s psy-
chology. Can. Psychol. 44:5–9 Mischel W, Ayduk O. 2004. Willpower in a cognitive-affective processing system: the dynamics
of delay of gratification. In Handbook of Self-Regulation: Research, Theory, and Applications, ed. RF Baumeister, KD Vohs, pp. 99–129. New York: Guilford
Newell A. 1990. Unified Theories of Cognition. Cambridge, MA: Harvard Univ. Press Nikolaev AR, Ivanitsky GA, Ivanitsky AM, Abdullaev YG, Posner MI. 2001. Short-term corre-
lation between frontal and Wernicke’s areas in word association. Neurosci. Lett. 298:107–10 Ochsner KN, Ludlow DH, Knierim K, Hanelin J, Ramachandran T, et al. 2006. Neural
correlates of individual differences in pain-related fear and anxiety. Pain 129(1–2):69–77 Ochsner KN, Kossyln SM, Cosgrove GR, Cassem EH, Price BH, et al. 2001. Deficits in
visual cognition and attention following bilateral anterior cingulotomy. Neuropsychologia 39:219–30
O’Reilly RC, Munakata Y. 2000. Computational Explorations of Cognitive Neuroscience. Cambridge, MA: MIT Press
Parasuraman R, Greenwood PM, Kumar R, Fossella J. 2005. Beyond heritability— neurotransmitter genes differentially modulate visuospatial attention and working mem- ory. Psychol. Sci. 16:200–7
Posner MI. 1980. Orienting of attention. The seventh Sir F.C. Bartlett Lecture. Q.J. Exp. Psychol. 32:3–25
Posner MI. 2004. The achievements of brain imaging: past and present. See Kanwisher & Duncan 2004, pp. 505–28
Posner MI, Fan J. 2007. Attention as an organ system. In Neurobiology of Perception and Com- munication: From Synapse to Society. De Lange Conference IV, ed. J Pomerantz. London: Cambridge Univ. Press. In press
Provides an overview of neuroimaging in relation to psychological concepts.
Posner MI, Raichle ME. 1996. Images of Mind. New York: Sci. Am. Posner MI, Rothbart MK. 2004. Hebb’s neural networks support the integration of psycho-
logical science. Can. Psychol. 45:265–78
Relates the implications of neural network development for issues of educating children.
Posner MI, Rothbart MK. 2007. Educating the Human Brain. Washington, DC: Am. Psychol. Assoc.
Rothbart MK, Ahadi SA, Hershey KL. 1994. Temperament and social behavior in childhood. Merrill-Palmer Q. 40:21–39
Rothbart MK, Ahadi SA, Hershey KL, Fisher P. 2001. Investigations of temperament at three to seven years: The Children’s Behavior Questionnaire. Child Dev. 72(5):1394–408
Comprehensively reviews temperament research.
Rothbart MK, Bates JE. 2006. Temperament in children’s development. In Handbook of Child Psychology: Vol. 3, Social, Emotional, and Personality Development, book ed. W Damon, R Lerner, vol. ed. N Eisenberg, pp. 99–166. New York: Wiley. 6th ed.
Rothbart MK, Posner MI. 2006. Temperament, attention, and developmental psychopathol- ogy. In Handbook of Developmental Psychopathology, Revised, ed. D Cicchetti, DJ Cohen, pp. 167–88. New York: Wiley
Describes the effortful control temperament variable and how it relates to behavior and underlying networks.
Rothbart MK, Rueda MR. 2005. The development of effortful control. In Develop- ing Individuality in the Human Brain: A Festschrift Honoring Michael I. Posner, ed. U Mayr, E Awh, SW Keele, pp. 167–88. Washington, DC: Am. Psychol. Assoc.
Rothbart MK, Ziaie H, O’Boyle CG. 1992. Self-regulation and emotion in infancy. In Emotion and Its Regulation in Early Development: New Directions for Child Development No. 55: The Jossey-Bass Education Series, ed. N Eisenberg, RA Fabes, pp. 7–23. San Francisco: Jossey- Bass
22 Posner · Rothbart
A nn
u. R
ev . P
sy ch
ol . 2
00 7.
58 :1
-2 3.
D ow
nl oa
de d
fr om
a rj
ou rn
al s.
an nu
al re
vi ew
s. or
g by
U ni
ve rs
it y
of W
is co
ns in
- M
ad is
on o
n 11
/2 6/
08 . F
or p
er so
na l
us e
on ly
.
ANRV296-PS58-01 ARI 17 November 2006 1:10
Rueda MR, Fan J, McCandliss BD, Halparin JD, Gruber DB, et al. 2004. Development of attentional networks in childhood. Neuropsychologia 42:1029–40
Rueda MR, Rothbart MK, McCandliss BD, Saccomanno L, Posner MI. 2005. Training, matu- ration, and genetic influences on the development of executive attention. Proc. Natl. Acad. Sci. USA 102(41):14931–36
Ruff HA, Rothbart MK. 1996. Attention in Early Development: Themes and Variations. New York: Oxford Univ. Press
Rumbaugh DM, Washburn DA. 1995. Attention and memory in relation to learning: a com- parative adaptation perspective. In Attention, Memory and Executive Function, ed. GR Lyon, NA Krasengor, pp. 199–219. Baltimore, MD: Brookes
Rumelhart DE, McClelland JL. 1986. Parallel Distributed Processing. Cambridge, MA: MIT Press
Sejnowski TJ. 2003. The once and future Hebb synapse. Can. Psychol. 44:17–20 Shavlev L, Tsal Y, Mevorach C. 2003. Progressive attentional training program: effective direct
intervention for children with ADHD. Proc. Cogn. Neurosci. Soc. NY, pp. 55–56 Shelton AL, Gabrieli JDE. 2002. Neural correlates of encoding space from route and survey
perspectives. J. Neurosci. 22(7):2711–17 Simon HA. 1969. The Sciences of the Artificial. Cambridge, MA: MIT Press Smith EE, Jonides J, Marshuetz G, Koeppe RA. 1998. Components of verbal working memory.
Proc. Natl. Acad. Sci. USA 95:876–82 Sternberg RJ, ed. 2004. Unity in Psychology: Prospect or Pipedream? Washington, DC: APA Books Sternberg RJ, Grigorenko EL. 2001. Unified psychology. Am. Psychol. 56(12):1069–79 Thorndike EL. 1903. Educational Psychology. New York: Teachers College Toga AW, Mazziotta JC, eds. 1996. Brain Mapping: The Methods. San Diego: Academic Ungerleider LG, Courtney SM, Haxby HV. 1998. A neural system for human visual working
memory. Proc. Natl. Acad. Sci. USA 95:883–90 Utall WR. 2001. The New Phrenology. Cambridge, MA: MIT Press Venter JC, Adams MD, Myers EW, Li PW, Mural RJ, et al. 2001. The sequence of the human
genome. Science 291:1304–35 Wang H, Fan J, Johnson TR. 2004. A symbolic model of human attentional networks. Cogn.
Syst. Res. 5:119–34 Wang H, Fan J. 2006. Human attentional networks: a connectionist model. Manuscript in
preparation Washburn DA. 1994. Stroop-like effects for monkeys and humans: processing speed or strength
of association? Psychol. Sci. 5(6):375–79 Womelsdorf T, Fries P, Mitra PP, Desimone R. 2006. Gamma-band synchronization in visual
cortex predicts speed of change detection. Nature 439:733–36
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u. R
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nl oa
de d
fr om
a rj
ou rn
al s.
an nu
al re
vi ew
s. or
g by
U ni
ve rs
it y
of W
is co
ns in
- M
ad is
on o
n 11
/2 6/
08 . F
or p
er so
na l
us e
on ly
.
Contents ARI 8 November 2006 21:2
Annual Review of Psychology
Volume 58, 2007
Contents
Prefatory
Research on Attention Networks as a Model for the Integration of Psychological Science Michael I. Posner and Mary K. Rothbart � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 1
Cognitive Neuroscience
The Representation of Object Concepts in the Brain Alex Martin � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 25
Depth, Space, and Motion
Perception of Human Motion Randolph Blake and Maggie Shiffrar � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 47
Form Perception (Scene Perception) or Object Recognition
Visual Object Recognition: Do We Know More Now Than We Did 20 Years Ago? Jessie J. Peissig and Michael J. Tarr � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 75
Animal Cognition
Causal Cognition in Human and Nonhuman Animals: A Comparative, Critical Review Derek C. Penn and Daniel J. Povinelli � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 97
Emotional, Social, and Personality Development
The Development of Coping Ellen A. Skinner and Melanie J. Zimmer-Gembeck � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 119
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Contents ARI 8 November 2006 21:2
Biological and Genetic Processes in Development
The Neurobiology of Stress and Development Megan Gunnar and Karina Quevedo � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 145
Development in Societal Context
An Interactionist Perspective on the Socioeconomic Context of Human Development Rand D. Conger and M. Brent Donnellan � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 175
Culture and Mental Health
Race, Race-Based Discrimination, and Health Outcomes Among African Americans Vickie M. Mays, Susan D. Cochran, and Namdi W. Barnes � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 201
Personality Disorders
Assessment and Diagnosis of Personality Disorder: Perennial Issues and an Emerging Reconceptualization Lee Anna Clark � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 227
Social Psychology of Attention, Control, and Automaticity
Social Cognitive Neuroscience: A Review of Core Processes Matthew D. Lieberman � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 259
Inference, Person Perception, Attribution
Partitioning the Domain of Social Inference: Dual Mode and Systems Models and Their Alternatives Arie W. Kruglanski and Edward Orehek � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 291
Self and Identity
Motivational and Emotional Aspects of the Self Mark R. Leary � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 317
Social Development, Social Personality, Social Motivation, Social Emotion
Moral Emotions and Moral Behavior June Price Tangney, Jeff Stuewig, and Debra J. Mashek � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 345
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Contents ARI 8 November 2006 21:2
The Experience of Emotion Lisa Feldman Barrett, Batja Mesquita, Kevin N. Ochsner,
and James J. Gross � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 373
Attraction and Close Relationships
The Close Relationships of Lesbian and Gay Men Letitia Anne Peplau and Adam W. Fingerhut � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 405
Small Groups
Ostracism Kipling D. Williams � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 425
Personality Processes
The Elaboration of Personal Construct Psychology Beverly M. Walker and David A. Winter � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 453
Cross-Country or Regional Comparisons
Cross-Cultural Organizational Behavior Michele J. Gelfand, Miriam Erez, and Zeynep Aycan � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 479
Organizational Groups and Teams
Work Group Diversity Daan van Knippenberg and Michaéla C. Schippers � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 515
Career Development and Counseling
Work and Vocational Psychology: Theory, Research, and Applications Nadya A. Fouad � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 543
Adjustment to Chronic Diseases and Terminal Illness
Health Psychology: Psychological Adjustment to Chronic Disease Annette L. Stanton, Tracey A. Revenson, and Howard Tennen � � � � � � � � � � � � � � � � � � � � � � � � � � � 565
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Research Methodology
Mediation Analysis David P. MacKinnon, Amanda J. Fairchild, and Matthew S. Fritz � � � � � � � � � � � � � � � � � � � � � 593
Analysis of Nonlinear Patterns of Change with Random Coefficient Models Robert 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 the present) may be found at http://psych.annualreviews.org/errata.shtml
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