Discussion 1: Developments in Cognitive Psychology

profileReeb79
NeuroimagingasaNewToolintheToolboxodPsychologicalScience.pdf

Commentary

Neuroimaging as a New Tool in the Toolbox of Psychological Science John T. Cacioppo,1 Gary G. Berntson,2 and Howard C. Nusbaum1

1 The University of Chicago and

2 Ohio State University

ABSTRACT—During the past quarter century, advances in

imaging technology have helped transform scientific fields.

As important as the data made available by these new

technologies have been, equally important have been the

guides provided by existing theories and the converging

evidence provided by other methodologies. The field of

psychological science is no exception. Neuroimaging is an

important new tool in the toolbox of psychological science,

but it is most productive when its use is guided by psycho-

logical theories and complemented by converging

methodologies including (but not limited to) lesion, electro-

physiological, computational, and behavioral studies.

Based on this approach, the articles in this special issue

specify neural mechanisms involved in perception, atten-

tion, categorization, memory, recognition, attitudes, social

cognition, language, motor coordination, emotional regu-

lation, executive function, decision making, and depression.

Understanding the contributions of individual and func-

tionally connected brain regions to these processes benefits

psychological theory by suggesting functional representa-

tions and processes, constraining these processes, produc-

ing means of falsifying hypotheses, and generating new

hypotheses. From this work, a view is emerging in which

psychological processes represent emergent properties of a

widely distributed set of component processes.

KEYWORDS—functional magnetic resonance imaging; cog-

nitive processes; social processes; clinical processes; develop-

mental processes

New imaging technologies are having a demonstrable impact on

the landscape of scientific research. The most expensive imaging

instrument, and the most vivid example, is the Hubble Space

Telescope. The Hubble telescope was deployed in April 1990

and has undergone three major repairs and upgrades since that

time. It has also provided data and images at a resolution Galileo

Galilei could not have imagined when, early in the 17th century,

he discovered the craters on the moon, sunspots, the rings of

Saturn, and the moons of Jupiter by gazing through his first crude

telescope. The discoveries made possible by the high-resolution

data and images from the Hubble nearly four centuries later

include massive black holes at the center of galaxies, the

existence of precursors to planetary systems like our own, and

a greater quantity and distribution of dark matter than expected.

As important as were the data provided by the Hubble Telescope,

however, these discoveries were dependent on extant theories

and methodologies. The discovery of stellar black holes at the

centers of galaxies, for instance, was guided by general relativity

theory and supported by research using several converging

methodologies (e.g., Dolan, 2001).

Developments in neuroimaging during the past quarter

century have increasingly made it possible to investigate the

differential involvement of particular brain regions in normal

and disordered thought in humans. Previously, studies of the

neurophysiological structures and functions associated with

psychological states and processes were limited primarily to

animal models, postmortem examinations, electrophysiological

measures, and observations of the occasional unfortunate indi-

vidual who suffered trauma to or disorders of the brain (e.g.,

Raichle, 2003). The detailed three-dimensional color images

provided by neuroimaging, modeling statistical properties of the

working brain, have captured the imagination of the public and the

scientific community, shaped funding priorities at federal funding

agencies and foundations, and produced a dramatic growth in

scientific papers and journals in the area (Cacioppo et al., 2007).

Address correspondence to John T. Cacioppo, Center for Cognitive and Social Neuroscience, The University of Chicago, 5848 S. Uni- versity Avenue, Chicago, Illinois 60637; e-mail: cacioppo@uchicago. edu.

C U R R E N T D I R E C T I O N S I N P S Y C H O L O G I C A L S C I E N C E

62 Volume 17—Number 2Copyright r 2008 Association for Psychological Science

This special issue of Current Directions in Psychological Science

summarizes recent theoretical advances in various fields of

psychological science that are attributable in part to the use of

neuromaging technology—most prominently, functional mag-

netic resonance imaging (fMRI). Although reading these reviews

leaves the impression that neuroimaging is an important new tool

in the toolbox of psychological science, one cannot help but also

be impressed that neuroimaging—like the Hubble Space Tele-

scope—is most productive scientifically when its use is guided by

extant theories and complemented by converging methodologies.

In the typical neuroimaging study, psychological states and

processes are manipulated and activation in different brain

regions is measured. The logic of this design is best suited

for drawing inferences about the differential involvement of par-

ticular brain regions in specific psychological operations.

For instance, when Poeppel and Monahan (2008, this issue) ask

how speech signals are represented and processed in the brain,

they are using neuroimaging along with converging methods

from psychological science, and guided by the blueprint of

competing theoretical accounts for speech perception, to in-

vestigate the differential involvement of particular brain regions

in psychological states and processes. It is also possible under

certain conditions to draw reasonable inferences about psycho-

logical operations based on regions of brain activation (Cacioppo

& Berntson, in press; Cacioppo & Tassinary, 1990; Henson,

2006; Poldrack, 2006; Sarter, Berntson, & Cacioppo, 1996).

Across the articles in this special issue, evidence from lesion

studies, animal studies, neuroimaging, single-cell recording,

event-related brain potentials, transcranial magnetic stimulation,

computational modeling, and behavior is reviewed to investigate

the brain regions involved in perception, attention, categorization,

memory, recognition, attitudes, social cognition, language, motor

coordination, emotional regulation, executive function, decision

making, and depression. Together, the evidence converges on the

view that psychological states and processes are mediated by a

network of distributed, often recursively connected, interacting

brain regions, with the different areas making specific, often task-

modulated contributions (see Poeppel & Monahan, 2008).

DISTRIBUTED NETWORKS INVOLVED IN COGNITIVE

REPRESENTATIONS AND FUNCTION

Humans are visual creatures. The visual properties of scenes

drive neurons in the lateral geniculate nucleus of the thalamus,

and visual perception has been found to involve a dorsal stream,

or ‘‘where pathway,’’ and a ventral stream, or ‘‘what pathway.’’

The dorsal (where) stream includes the areas designated V1, V2,

V5/MT, and the inferior parietal lobule and is associated with

motion, representation of object locations, and control of the

eyes and arms when visual information is used to guide saccades

or reaching. The ventral (what) stream includes the areas V1,

V2, V4, and the inferior temporal lobe (areas that include

the lateral occipital complex and the fusiform gyrus) and is

associated with form recognition, object representation, face

recognition, and long-term memory (Engel, 2008, this issue).

Grill-Spector and Sayres (2008, this issue) provide evidence

that changes in the size, position, orientation, and other aspects

of physical appearance of faces activate the lateral occipital

complex; differences in the identity of individuals are related

to adaptation responses in the fusiform gyrus; and changes

in facial expression and gaze direction involve the superior

temporal sulcus.

Different theoretical representations and decompositions of

speech perception into processing components are described,

and the neural outcomes associated with each of these theoret-

ical representations are reviewed, by Poeppel and Monahan

(2008). Here, too, the evidence suggests the involvement of a

specialized, interconnected set of neural regions that are widely

distributed across the temporal, parietal, and frontal lobes.

Specifically, the early spectrotemporal analyses involve bilateral

auditory cortices and the superior temporal cortex, and phono-

logical analyses involve the middle and posterior portions of the

superior temporal sulcus. The processing streams then appear to

divide into a ventral stream—which maps auditory and phono-

logical representations onto lexical conceptual representations

and involves the middle temporal gyrus and inferior temporal

sulcus—and a dorsal stream—which maps auditory and pho-

nological representations onto articulatory and motor represen-

tations and involves the Sylvian parietotemporal area, posterior

frontal gyrus, premotor cortex, and anterior insula (Poeppel &

Monahan, 2008).

The attentional modulation of perceptual processes is influ-

enced by motivational states and goals as well as by stimulus

properties. Visual attentional control can modulate neural

activity in the lateral geniculate nucleus and superior colliculus,

as well as in the posterior parietal cortex (specifically, the re-

gions of the superior parietal lobule and the lateral intraparietal

area within the intraparietal sulcus) and the frontal eye field and

supplementary eye field within the prefrontal cortex (Yantis,

2008, this issue). These perceptual and attentional processes

contribute to the acquisition of knowledge about the world that

is organized categorically. Barsalou (2008, this issue) notes

that the dominant theory in cognitive science posits that this

knowledge is represented in an abstract, amodal fashion and

constitutes semantic memory. He shows in his review, however,

that categorical knowledge includes modal representations

using the same neural mechanisms involved in perception,

affect, and action. In Barsalou’s view, the representation of a

category involves a neural circuit distributed across the relevant

modalities, all of which can become activated during conceptual

processing. Thus, conceptual processing can be viewed as an

embodied rather than purely abstract process.

How does categorical knowledge being represented in this

distributed, modal fashion square with the neuroscientific

evidence for differences in the localization of short- and long-

term memory processes? Nee, Berman, Moore, and Jonides

Volume 17—Number 2 63

John T. Cacioppo, Gary G. Berntson, and Howard C. Nusbaum

(2008, this issue) suggest that the evidence supporting the

qualitative distinction between short- and long-term storage

processes has been misinterpreted, and they suggest that the

data instead support a unitary model of memory in which the

same regions of the brain that represent perception, action, and

affect are involved in both short- and long-term storage pro-

cesses. That is, short- and long-term memories do not differ in

representation but in the activation by attention, which in turn

involves a frontal biasing (i.e., maintenance) of representational

cortices (e.g., frontal eye fields and intraparietal sulcus for

spatial representations; superior temporal sulcus and Sylvian

parietotemporal area for phonological and articulatory repre-

sentations). For instance, damage in the presylvian region pro-

duces deficits in short- and long-term memory that depends on

phonological material, and the greater prevalence of such ma-

terial in studies of short- than of long-term storage processes may

inadvertently have led to evidence that this region was involved

uniquely in short-term memory (Nee et al., 2008). Short- and

long-term memory retrieval also activates overlapping regions of

the left lateral frontal cortex, whereas the monitoring of retrieved

information, whether from short- or long-term memory, is

associated with the anterior prefrontal activation (cf. Cabeza,

Dulcos, Graham, & Nyberg, 2002).

Neuropsychological research dating back several decades sug-

gested structures in the medial temporal lobe (e.g., the hippo-

campus) were involved in declarative rather than nondeclarative

learning. Unlike the distinction between short- and long-term

memory, the distinction between declarative and nondeclarative

learning is supported by neuroimaging research. For instance,

research by Knowlton and Foerde (2008, this issue) has shown that

when performance on a probabilistic classification task is based

on declarative memory performance the medial temporal lobe

is activated, whereas when performance on the task is based on

nondeclarative memory performance the striatum is activated.

Knowlton and Foerde (2008) also review evidence showing

that nondeclarative skill learning, at least for simple tasks, is asso-

ciated with repetition suppression—reductions in the regions of

neural activation associated with the initial performances of a task

(e.g., the premotor region)—a finding that has been interpreted as

indicating a greater efficiency of processing in the neural struc-

tures involved in novice performance. Priming-related reductions,

on the other hand, are found in perceptual and prefrontal regions,

with only the latter associated with behavioral facilitation.

Knowlton and Foerde (2008) duly note, however, that activation in

the perceptual cortices may appear to be less important in the

extant literature in part because of the type of priming paradigms

that have been used in fMRI research.

The complexities of social living, such as recognizing indi-

viduals and groups, negotiating nontransitive social hierarchies

and shifting alliances, using language to communicate and

manipulate, and engaging in social exchanges over extended

periods and locales, place special demands on the capacities of

the human brain. Mitchell (2008, this issue) reviews evidence

that thinking about thinking people (e.g., impression formation,

social causality)—in contrast, for instance, to thinking about

physical causality—is associated with activation of the medial

prefrontal cortex, the right temporo-parietal junction, and the

medial parietal region (e.g., the precuneus/posterior cingulate

cortex). Mitchell (2008) suggests that the activation of the

medial prefrontal cortex appears to be involved whenever people

are obliged to consider the psychological characteristics of

another person, whereas the temporo-parietal junction appears

to be activated when the attentional and perceptual require-

ments of taking the perspective of another are invoked. The

medial parietal region, on the other hand, is activated during

the retrieval of episodic memories and self-knowledge, as well

as during the viewing of two or more interacting people (e.g.,

Iacoboni et al., 2004).

The brain has evolved to guide behavior in contextually

flexible, coordinated, and adaptive ways, and, as with attention,

there are top-down as well as bottom-up influences on the

orchestration of motor processes. Oliveira and Ivry (2008, this

issue) focus on the top-down influences in their discussion of

goals as higher-level action representations that connect sensory

and motor processes to guide response selection and motor

coordination. They review fMRI studies showing that motor

planning and externally guided movements are associated with

activity in the posterior superior parietal region and, at least for

externally guided movements, in premotor regions; internally

generated movements are associated with activity in the basal

ganglia, the anterior cingulate cortex, and inferior frontal and

parietal cortices; and conflicting action goals and effort are

associated with activity in medial frontal areas, including the

anterior cingulate cortex and presupplementary motor areas.

One suggestion that has emerged from this area of research

is that goal representation and action planning are not imple-

mented simply as an abstract code but rather involve embodied

processes. Not unlike how Barsalou (2008) invokes modal

mechanisms, Rizzolati and Arbib (1998) and Skipper, Nusbaum,

and Small (2006) review evidence for the role of embodied

representations in categorization and language.

The fundamental idea that the motor system is important for

cognition and perception, through prior experience and mirror

neurons, has become an important contribution of neuroscience

to bolstering theoretical constructs in the psychology of em-

bodied understanding. However, much of the work on the mirror

system in cognition and understanding has been carried out with

trained nonhuman primates or with adult humans. For any theory

of adult function to be viable, it is critical to understand the

development of these mechanisms. Diamond and Amso (2008,

this issue) review work on the neural substrates underlying

cognitive development, including the mirror-neuron system

and neonatal imitative behaviors and maternal touch and

gene expression. As the authors note, a major contribution of

neuroscience to theories of cognitive development is ‘‘demon-

strating the remarkable role of experience in shaping the mind,

64 Volume 17—Number 2

Neuroimaging and Psychological Science

brain, and body’’ (p. 136). Such cross-cutting work is necessary

to begin to link biological development with learning and

experience. Moreover, as cognition can no longer be studied

in isolation from the social context of its use, this work suggests

the importance of understanding development within its social

context of parental interaction. Given the importance of social

context, then, it is important to go beyond the treatment

of specific processes to understand how such processes depend

on the goals they are directed at achieving.

To achieve one’s goals, one has to be able to represent

the likely rewards (and punishments) associated with different

decisions, encode the risk or certitude that the reward will

be obtained, update these representations, and act on the basis

of these representations. O’Doherty and Bossaerts (2008, this

issue) review evidence regarding the brain regions associated

with each of these components of decision making. Specifically,

they report that the encoding of reward expectation is associated

with activation of the orbitofrontal cortex, medial prefrontal

cortex, amygdala, and ventral striatum; recognizing greater risk

or uncertainty associated with obtaining a reward correlates with

increased activity in the anterior insula and lateral orbitofrontal

regions; updating of reward expectancies is associated with

the ventral striatum and orbitofrontal cortex; and selecting

one of several responses to obtain the greatest reward involves

the striatum, with the ventral striatum more involved in the

prediction of reward across the various options and the dorsal

striatum more involved in the selection among the alternatives

(O’Doherty & Bossaerts, 2008).

The frontal regions have long been thought to be involved

in executive functions such as formulating goals and plans;

selecting among options to achieve these goals; monitoring the

consequences of actions in light of one’s goals; and inhibiting,

switching and regulating one’s behaviors accordingly. Aron

(2008, this issue) reviews evidence that the initiation of a motor

response proceeds from the planning areas of the frontal cortex

to the putamen, globus pallidus, thalamus, primary motor cortex,

motor nucleus in the spinal cord, and finally to the muscles.

Being able to inhibit a motor response once it has been initiated

has obvious adaptive value, and Aron (2008) shows that this

inhibition involves the right inferior frontal cortex, which pro-

jects to the subthalamic nucleus (a region of the basal ganglia

that may act on the globus pallidus to block the motor response).

Monitoring for response conflicts, in turn, appears to involve the

dorsal anterior cingulate and the adjacent presupplementary

motor area, which, in turn, is connected to the right inferior

frontal cortex and subthalamic nucleus. Switching also involves

the presupplementary motor area and the right inferior frontal

cortex (Aron, 2008). This work has led to a model in which ‘‘the

[presupplementary motor area] may monitor for conflict between

an intended response and a countervailing signal . . . Then, when

such conflict is detected, the ‘brakes’ could be put on via the

connection between the right [inferior frontal cortex] and the

[subthalamic nucleus] region’’ (Aron, 2008, p. 127).

Emotional regulation is another form of executive function

in which activity of the amygdala and insula cortex, which are

involved in emotional responding, is modulated by activity in the

prefrontal cortex (e.g., BA10, ventromedial prefrontal cortex,

dorsolateral prefrontal cortex) and anterior cingulate. Ochsner

and Gross (2008, this issue) review evidence that different

components of reappraisal processing correspond to different

areas of prefrontal activation: Selective attention and working

memory components are related to dorsal portions of the pre-

frontal cortex, language or response inhibition are related to

ventral portions of the prefrontal cortex, monitoring or control

processes are related to the dorsal anterior cingulate cortex, and

reflections on one’s emotional state are related to dorsal portions

of the medial prefrontal cortex. Although the correspondences

proposed by Ochsner and Gross (2008) do not match perfectly

those articulated by Aron (2008), the overlapping role for the

anterior cingulate is noteworthy in light of the notion that the

presupplementary motor area may be especially involved in the

monitoring and control of motor conflicts.

The complexities of daily living are simplified in part by the

formation of preferences and attitudes, which can serve as

behavioral guides and simplify decision making. These attitudes

can be explicit or implicit. Stanley, Phelps, and Banaji (2008,

this issue) review evidence suggesting that the activation

of implicit attitudes toward social groups (e.g., minorities) is

associated with increased activity in the amygdala, dorsolateral

prefrontal cortex, and anterior cingulate cortex. The cumulative

evidence to date suggests that the automatic evaluation of

a stimulus (e.g., social category) is associated with amygdala

activation, the monitoring for response conflicts (e.g., the extent

to which the stimulus elicits competing impulses) is associated

with anterior cingulate activation, and the regulation of those

impulses is associated with dorsolateral prefrontal activation.

Failures of effective emotional regulation can become costly

in personal, social, and economic terms when these failures become

systemic. Depression, for instance, has been estimated to cost more

than $43 billion per year in the United States (Greenberg, Stiglin,

Finkelstein, & Berndt, 1993). Understanding the variation in

biological systems that leads to individual differences in neural

mechanisms of emotional regulation is critical to understanding

how some systemic failures become chronic and debilitating.

Gotlib and Hamilton (2008, this issue) review evidence that

depressed individuals show less activity in the dorsolateral

prefrontal cortex and greater activation of the amygdala and

subgenual anterior cingulate cortex to emotional stimuli than do

healthy controls. Parallel findings for basal activity levels in

these brain regions are also noted. These findings are consistent

with Gotlib and Hamilton’s notion that depression is in large part

a disorder of emotion regulation in which the normal inhibitory

influence of limbic structures by the anterior cingulate and

dorsolateral prefrontal cortex is disrupted, although the

subgenual anterior cingulate cortex may play an especially

critical role in this dysregulation (Gotlib & Hamilton, 2008).

Volume 17—Number 2 65

John T. Cacioppo, Gary G. Berntson, and Howard C. Nusbaum

Given the importance of the anterior cingulate and dorsolateral

prefrontal cortex in motor control, attention, and emotion,

the individual variation in function in these areas that can lead

to depression may also explain that disorder’s other associated

cognitive symptoms.

Indeed, understanding the relationship between biological

variation in neural mechanisms and psychological processes is

important beyond clinical problems. Kosslyn et al. (2002) and

Vogel and Awh (2008, this issue) have argued that, to bridge

the gap between psychological phenomena and their underlying

biological substrata, such variation should be regarded as

important data in its own right. Kosslyn et al. (2002) describe how

an idiographic approach can be used to address three types of

issues: the nature of the mechanisms that give rise to a specific

ability, the role of psychological or biological mediators of envi-

ronmental challenges, and the existence of variables that have

nonadditive effects with other variables. Vogel and Awh (2008)

extend this argument in their discussion of three additional ways

in which an idiographic approach can contribute to psychological

theory: validating neurophysiological measures, demonstrating

associations among constructs, and demonstrating dissociations

among similar constructs. Thus, an idiographic approach, which

complements the more typical nomethetic approach, can be

applied in any domain to help elucidate psychological theory.

Together, the theory and data summarized in this special issue

of Current Directions in Psychological Science highlight the

notion that encephalization and the remarkable connectivity

in the human brain provide the substrate for the integration

of inputs from widely distributed neural regions (only some of

which are amenable to current brain-imaging technology) whose

activation and organization can be contextually determined. The

distributed nature of and substantial overlap among the extant

networks calls for a revision in our thinking about basic

psychological constructs. The early reliance on introspection as

a method of identifying elemental psychological processes led to

a recognition of the category error—the intuitively appealing but

often erroneous notion that the organization of psychological

phenomena maps in a one-to-one fashion onto the organization

of underlying neural substrates. Perception, memories, emotions,

and beliefs were each once thought to be localized in distinct

sites in the brain. The contributions to this special issue clearly

indicate that psychological and behavioral concepts do not each

map onto clear and identifiable ‘‘centers,’’ but rather that each

concept is associated with a distributed, interconnected set of

neural regions. What appears at one point in time to be a singular

theoretical construct (e.g., memory), when examined in con-

junction with evidence from the brain (e.g., lesions, neuroimag-

ing), may reveal a more complex and interesting organization at

both levels (e.g., declarative vs. procedural memory processes).

Conversely, what appeared to be distinct constructs (e.g.,

short- vs. long-term memory) may need to be reconsidered in light

of new neuroscientific evidence. We suspect we are far from seeing

the last of such revisions to psychological theories. It is only

through these revisions, and corresponding refinements in our

understanding and conceptions of the underlying neural functions,

that we can reduce the category error and move toward an

isomorphism between the psychological and biological domains.

Neuroimaging and work in neuroscience more generally are

reshaping the constructs that are being used to build psycho-

logical theories. Psychological research during the 20th century

resulted in many of the basic psychological elements derived

from introspection to be recast as the product of multiple,

more specific component processes. As illustrated by the articles

in this special issue, many of these component processes in-

volve a network of distributed, often recursively connected,

interacting brain regions, with the different areas making

specific, often task-modulated contributions. Moreover, a single

neural region can often be involved in what have been treated

as very different psychological processes. One implication is that

what have been considered basic psychological or behavioral

processes are being conceptualized as manifestations of com-

putations performed by networks of widely distributed sets of

neural regions.

How might these neural components be combined to produce

distinct psychological processes? One metaphor is the Lego set,

in which the computations performed in localized neural regions

are fixed (like distinct Lego pieces), but different pieces

and configurations of these building blocks produce different

psychological processes. An alternative metaphor is the periodic

table in chemistry, in which different neural component pro-

cesses may have properties and affinities whose function (com-

putation) depends on the network of areas with which they

are combined. There is no evidence at present to favor either

perspective, but the important point here is that they suggest

very different ways of thinking about neural activity and

psychological function.

In sum, neuroimaging work is leading to a rethinking of how

psychological and neural functions are parcelled. For instance,

the close proximity of motor control, emotional appraisal,

attention, working memory, and behavioral regulation suggests

that these functions may not be as separable as they are currently

treated and studied. We may well need a new lexicon of

constructs that are neither simply anatomical (e.g., Brodmann

area 6 vs. Brodmann area 44) nor psychological (e.g., attention,

memory), as we usher in a new era of psychological theory

in which what constitutes elemental component processes

(functional elements) are tied to specific neural mechanisms

(structural elements) and in which the properties of interrelated

networks of areas may indeed be more than the sum of the parts.

CONCLUSION

Critics who say neuroimaging is costly and has contributed little

if anything to psychological theory sometime appear to expect

the images of the working brain to come with labels regarding

their cognitive functions. Although an adequate specification of

66 Volume 17—Number 2

Neuroimaging and Psychological Science

neurobiology should contribute to our understanding of cogni-

tive architecture and function, our understanding of the relevant

neurobiology is influenced strongly by our extant theoretical

models regarding cognitive architecture and function (see

Hagoort, 2008, this issue). The contributions to this special issue

demonstrate that neuroimaging is an important new tool in the

toolbox of psychological science, but one that is most productive

scientifically when its use is guided by psychological theories

and complemented by converging methodologies. This approach,

in which theory and converging methods are used hand in hand to

expand our understanding of the neural mechanisms involved in

cognition and the contributions of individual and functionally

connected brain regions to these processes, promises to advance

psychological theory by suggesting functional representations

and processes, by imposing significant constraints on these pro-

cesses, and by producing not only new behavioral hypotheses but

also new means of falsifying theoretical hypotheses.

Acknowledgments—Preparation of this paper was supported

by grants from the National Institute of Mental Health (Grant No.

P50 MH72850) and the John Templeton Foundation.

REFERENCES

Aron, A.R. (2008). Progress in executive-function research: From

tasks to functions to regions to networks. Current Directions in Psychological Science, 17, 124–129.

Barsalou, L.W. (2008). Cognitive and neural contributions to

understanding the conceptual system. Current Directions in Psychological Science, 17, 91–95.

Cabeza, R., Dulcos, F., Graham, R., & Nyberg, L. (2002). Similarities

and differences in the neural correlates of episodic memory

retrieval and working memory. Neuroimage, 16, 317–330.

Cacioppo, J.T., Amaral, D.G., Blanchard, J.J., Cameron, J.L., Carter,

C.S., Crews, D., et al. (2007). Social neuroscience: Progress and

implications for mental health. Perspectives on Psychological Science, 2, 99–123.

Cacioppo, J.T., & Berntson, G.G. (in press). Integrative neuroscience

for the behavioral sciences: Implications for inductive inference.

In G.G. Berntson & J.T. Cacioppo (Eds.), Handbook of neuroscience for the behavioral sciences. New York: Wiley.

Cacioppo, J.T., & Tassinary, L.G. (1990). Inferring psychological signifi-

cance from physiological signals. American Psychologist, 45, 16–28.

Diamond, A., & Amso, D. (2008). Contributions of neuroscience to

our understanding of cognitive development. Current Directions in Psychological Science, 17, 136–141.

Dolan, J.F. (2001). How to find a stellar black hole. Science, 292, 1079–1080.

Engel, S.A. (2008). Computational cognitive neuroscience of the visual

system. Current Directions in Psychological Science, 17, 68–72.

Gotlib, I.H., & Hamilton, J.P. (2008). Neuroimaging and depression:

Current status and unresolved issues. Current Directions in Psychological Science, 17, 159–163.

Greenberg, P.E., Stiglin, L.E., Finkelstein, S.N., & Berndt, E.R. (1993).

The economic burden of depression in 1990. Journal of Clinical Psychiatry, 54, 405–418.

Grill-Spector, K., & Sayres, R. (2008). Object recognition: Insights from

advances in fMRI methods. Current Directions in Psychological

Science, 17, 73–79.

Hagoort, P. (2008). Should psychology ignore the language of the brain?

Current Directions in Psychological Science, 17, 96–101.

Henson, R. (2006). Forward inference using functional neuroimaging:

Dissociations versus associations. Trends in Cognitive Sciences, 10,

64–69.

Iacoboni, M., Lieberman, M.D., Knowlton, B.J., Molnar-Szakacs, I.,

Mortiz, M., Throop, C.J., & Fiske, A.P. (2004). Watching social

interactions produces dorsomedial prefrontal and medial parietal

BOLD fMRI signal increases compared to a resting baseline.

Neuroimage, 21, 1167–1173.

Knowlton, B.J., & Foerde, K. (2008). Neural representations of

nondeclarative memories. Current Directions in Psychological

Science, 17, 107–111.

Kosslyn, S.M., Cacioppo, J.T., Davidson, R.J., Hugdahl, K., Lovallo,

W.R., Spiegel, D., & Rose, R. (2002). Bridging psychology

and biology: The analysis of individuals in groups. American

Psychologist, 57, 341–351.

Mitchell, J.P. (2008). Contributions of functional neuroimaging to social cog-

nition. Current Directions in Psychological Science, 17, 142–146.

Nee, D.E., Berman, M.G., Moore, K.S., & Jonides, J. (2008). Neuroscien-

tific evidence about the distinction between short- and long-term

memory. Current Directions in Psychological Science, 17, 102–106.

Ochsner, K.N., & Gross, J.J. (2008). Cognitive emotion regulation: In-

sights from social cognitive and affective neuroscience. Current

Directions in Psychological Science, 17, 153–158.

O’Doherty, J.P., & Bossaerts, P. (2008). Towards a mechanistic

understanding of human decision making: Contributions of

functional neuroimaging. Current Directions in Psychological

Science, 17, 119–123.

Oliveira, F.T.P., & Ivry, R.B. (2008). The representation of action:

Insights from bimanual coordination. Current Directions in

Psychological Science, 17, 130–135.

Poeppel, D., & Monahan, P.J. (2008). Speech perception: Cognitive

foundations and cortical implementation. Current Directions in

Psychological Science, 17, 80–85.

Poldrack, R.A. (2006). Can cognitive processes be inferred from

neuroimaging data? Trends in Cognitive Sciences, 10, 59–63.

Raichle, M.E. (2003). Functional brain imaging and human brain

function. The Journal of Neuroscience, 23, 3959–3962.

Rizzolatti, G., & Arbib, M.A. (1998). Language within our grasp. Trends in Neuroscience, 21, 188–194.

Sarter, M., Berntson, G.G., & Cacioppo, J.T. (1996). Brain imaging and

cognitive neuroscience: Toward strong inference in attributing

function to structure. American Psychologist, 51, 13–21.

Skipper, J.I., Nusbaum, H.C., & Small, S.L. (2006). Lending a helping

hand to hearing: Another motor theory of speech perception. In

M.A. Arbib (Ed.), Action to language via the mirror neuron system

(pp. 250–285). New York: Cambridge University Press.

Stanley, D., Phelps, E., & Banaji, M. (2008). The neural basis of implicit

attitudes. Current Directions in Psychological Science, 17, 164–

170.

Vogel, E.K., & Awh, E. (2008). How to exploit diversity for scientific

gain: Using individual differences to constrain cognitive theory.

Current Directions in Psychological Science, 17, 171–176.

Yantis, S. (2008). The neural basis of selective attention: Cortical

sources and targets of attentional modulation. Current Directions in

Psychological Science, 17, 86–90.

Volume 17—Number 2 67

John T. Cacioppo, Gary G. Berntson, and Howard C. Nusbaum