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Attentionalnetworks
Michael I. P o s n e r a n d S t a n i s l a s D e h a e n e
Recent brain-imaging and neurophysiological data There has been evidence of more general mechan- Michad L Posneris at
indicate that attention is neither a properly of a singleisms related to visual selection. One way to select the DeN of
brain area, nor of the entire brain. While attentional information is to orient to it. Eye movements to Psychology,
foveate a visual stimulus is a clear case, since without
effects seem mediated by a relative amplification of blood University of Oregon,
foveation there is little ability to examine details of the Eugene, OR 97403,
flow and electrical activity in the cortical areas processing
USA, and Stanislas
visual scene. There are also brain mechanisms for
the attended computation, the details of how this is done Dehaene is at
through enhancement of attended or suppression of visual orienting that do not involve any overt changes the Laboratoire
unattended items, or both, appear to depend on the task in head or eye. In the 1970s, it was found that cells in de Sciences
the parietal lobe increased their firing rate in response
and brain-area studied. The origins of these amplifwation Co&nitivesand
to stimulation of their receptive field when monkeys
effects are to be found in specialized cortical areas of the Psycholinguistique,
attended to peripheral stimuli even when no eye
frontal and parietal lobes that have been described as the CNRS, EHESSand
anterior and posterior attention systems. These results movements were allowed ~. It was also shown that INSERM, Paris,
humans could covertly shift attention to peripheral
represent substantial progress in the effort to determine France.
how brain activity is regulated through attention. Whilestimuli and when they did so they responded more
many philosophical and practical issues remain in rapidly, at lower threshold and with enhanced elec-
developing an understanding of attentional regulation, trical activity to stimuli at the attended location.
the new tools available should provide the basis for Lesions of the parietal lobe specifically damaged this
progress. covert orienting ability on the side of space opposite
the lesion. These findings supported the idea that
The study of selective attention has been an important portions of the parietal lobe were involved in covert
area of research since the inception of psychology in orienting to visual stimuli2.
the late 1800s. However, it has remained contro- Since 1987, when we last reviewed this topic 2, it
versial whether there are any separate brain mechan- has become possible to use neuroimaging methods to
isms that subserve attention. Attention does not give observe the networks of brain areas that become
rise to a unique qualitative experience like vision or active when people perform complex tasks 3'4, Certain
touch, nor does it automatically produce motor brain areas appear active when subjects have to
responses. While we appear to be able to select orient, select or transform information in ways that
sensory stimuli, information in memory or motor would be said, by most psychological models, to
responses, this might not indicate a separate attention involve attention. These brain areas appear to fit quite
system, since all brain systems play a role in well with the idea of selection and control as properties
selection. of attention outlined in our previous article. In this
0"_!
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tasks such as serial search, the
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appropriate processing stream
might change as a function of time
(for example, the subject might
attend to each stimulus in turn or
A to all vertical items first and then to
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0 - !- ! ! all horizontal items).
Recent advances in functional
brain imagery, such as positron
emission tomography (PET) scan,
have provided new evidence con-
III 0 l 1 cerning the cortical and subcortical
l networks responsible for these
two functions of selective attention.
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A
As it turns out, the two functions
seem to be carried out by two
I_ 0_ A
V
! distinct systemsT: the posterior
attention system (superior parietal
cortex, pulvinar and superior col-
liculus) is largely responsible for
Fig. 1. Visual search for a vertical rectangle among a large number of distractors (horizontal and selecting one stimulus location
vertical ellipses and rectangles). The target is defined by a conjunction of form and orientation. In among many and for shifting from
most cases, selective attention is used to sequentially search the display until the target is found 5. one stimulus to the next, whereas
the anterior attention system (an-
article, we use the neuroimaging data to outline what terior cingulate and basal ganglia) serves a more
is known about networks of brain areas involved in executive function (attention for action) and is in-
selection during visual search and then extend the volved in the attentional recruitment and control of
discussion to mechanisms of selection and control brain areas to perform complex cognitive tasks. This
more generally. network is also involved in selecting visual objects
when the instructions emphasize properties of the
Visual search object. Ultimately, however, the effect of both
Throughout this article, we use the example of a systems is relative amplification of activity within the
simple visual-search task in order to illustrate the role cortical areas relevant to a given task.
of attention in finding objects 5. Data indicate that the
time for a subject to locate a target (for example, the Attentional amplification
vertical rectangle in Fig. 1) increases almost linearly In the visual display of Fig. 1, if you are attending to
with the number of distractors, as though the subject the correct location it is easy to see the target.
attended to each item in turn. The reaction time when Otherwise, the target is not seen. How does the brain
there is no target has twice the slope found for 'yes' implement such target detection? It appears that brain
responses, indicating that one stops as soon as the activity, in many cortical areas, can be selectively
target is found5. amplified or suppressed as a function of attentional
A simplified view of the problem faced by the ~rain set. When subjects are instructed to pay attention to
in this task is presented in Fig. 2. The visual array is one particular dimension of the stimuli, the brain areas
processed through a diverging tree of cortical areas 6. that specialize in the processing of this stimulus
Cortical cells early in the stream have narrow dimension are selectively enhanced. These findings
receptive fields and respond mechanically whenever are striking in the prestfiate areas of the visual
their preferred stimuli are present, whether they system. In one experiment, subjects viewed passively
correspond to a target or not. As one moves up in the either stationary or moving objects, presented every
hierarchy, however, receptive fields widen and the second for a minute s. When blood flow during the
cells become more sensitive to attentional and inten- stationary control condition was subtracted from that
tional influences, and react less effectively to the found in the moving condition, a prestriate area of the
passive presentation of a stimulus. midtemporal lobe was found to have a significant
The simple illustration in Fig. 2 makes it clear that increase in blood flow. This might be the human
selective attention serves at least two distinctive equivalent of area V5 or MT which, in the monkey,
functions. First, the brain must enhance the process- contains cells with a strong selectivity for moving
ing of the selected stimulus relative to other stimuli stimuli. In a different experiment 9, subjects always
present, otherwise all stimuli would be processed to a viewed moving objects of varying color and form, but
similar degree (Fig. 2, top). Second, processing of the in one set of trials they were instructed to detect
selected stimulus must be actively oriented or guided differences in velocity, whereas in a control set they
towards the cortical areas appropriate for a given had to detect changes in other stimulus parameters.
task. The diverging tree of cortical areas, each of Even though the physical stimuli were the same in the
which specializes for a distinct aspect of the stimuli, two conditions, the instruction to attend to velocity
must be selectively 'pruned' to focus only on those activated a brain area that was similar to that found in