J Physiol 589.

1 (2011) pp 4957 49

TOPICAL REVIEW

The neural basis of visual attention

James W. Bisley
Department of Neurobiology and Jules Stein Eye Institute, David Geffen School of Medicine at UCLA, and Department of Psychology and the Brain
Research Institute, UCLA, Los Angeles, CA 90095, USA

Visual attention is the mechanism the nervous system uses to highlight specific locations, objects
or features within the visual field. This can be accomplished by making an eye movement to bring
the object onto the fovea (overt attention) or by increased processing of visual information in
neurons representing more peripheral regions of the visual field (covert attention). This review
will examine two aspects of visual attention: the changes in neural responses within visual cortices
due to the allocation of covert attention; and the neural activity in higher cortical areas involved
The Journal of Physiology

in guiding the allocation of attention. The first section will highlight processes that occur during
visual spatial attention and feature-based attention in cortical visual areas and several related
models that have recently been proposed to explain this activity. The second section will focus
on the parietofrontal network thought to be involved in targeting eye movements and allocating
covert attention. It will describe evidence that the lateral intraparietal area, frontal eye field and
superior colliculus are involved in the guidance of visual attention, and describe the priority map
model, which is thought to operate in at least several of these areas.

(Received 5 May 2010; accepted after revision 27 August 2010; first published online 31 August 2010)
Corresponding author J. Bisley: Department of Neurobiology, PO Box 951763, Los Angeles, CA 90095-1763, USA.
Email: jbisley@mednet.ucla.edu

Visual attention described as looking out of the corner of your eye.

Although we are not always aware of it, visual attention
is incredibly important for visual perception and for
all the uses we put perception to, such as learning,
memory and our interactions with the visual world. This
is highlighted by behavioural protocols such as change
blindness, which require subjects to identify changes in a
scene, but remove the usual cues of these changes (such as
movement or sudden onset) by having the entire display
flash off and then on again at the time of the change
(Rensink, 2000). Subjects are remarkably poor at detecting
these changes when broadly viewing the entire scene, but
improve dramatically when their attention is allocated to
the location of the change. In addition to these types of
protocols, the importance of attention is highlighted in
patients with parietal lesions who often show signs of
attentional deficits as striking as the inability to notice
anything in a particular region of space (Husain & Nachev,
2007; Adair & Barrett, 2008).
Generally we think of focusing our attention onto
objects or features in terms of making a rapid eye
movement (a saccade) to bring the object of our attention
to the centre of gaze. However, our visual system can
also process information from selected peripheral regions
of the retina. When this is done consciously, it is often
As the eyes do not move, this is referred to as covert
attention (Posner, 1980). Both covert and overt attention
can be allocated voluntarily (commonly referred to as
top-down) or involuntarily (bottom-up). Classic examples
of bottom-up attractors of attention are flashing or moving
objects, such as the lights on top of emergency vehicles.
In contrast, top-down attention can refer to any voluntary
attention or attention that is driven by factors other than
external stimuli.
The effects of attention on neurons in visual
cortical areas
In the lab, covert attention provides a number of
behavioural benefits. Subjects attending a particular
James Bisley is an Assistant Professor of Neurobiology at UCLA.
In 1998 he received his PhD from the University of Melbourne,
Australia. His research uses physiological, psychophysical and
computational techniques to study the cognitive processing of visual
information, such as attention, short-term memory and spatial
processing. His work on the neural mechanisms underlying the
allocation of visuospatial attention has been recognized by awards
from the McKnight Foundation, the Sloan Foundation and the
Klingenstein Foundation.


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50
location have perceptual benefits when examining a
stimulus at that location, such as improved contrast
sensitivity (Cameron et al. 2002) or spatial resolution
(Yeshurun & Carrasco, 1999). Furthermore, reaction times
are improved, such that subjects respond more rapidly to
an event at an attended location than at another location
(Posner, 1980). In this section, we briefly describe the
effects of attention on neuronal responses in a number of
visual areas (light grey regions, Fig. 1) that may underlie
these behavioural benefits.
Spatial attention. Just as eye movements are aimed at
particular regions of space, covert attention can also
be focused on discrete locations in the visual scene. To
examine this, experiments have been performed in which
a single stimulus was placed in the receptive field of
the neuron being studied and one or more stimuli were
placed outside of the receptive field (Fig. 2A, left panel).
Responses were then recorded when the animal focused its
attention towards the stimulus within the receptive field or
outside of the receptive field. Generally, neurons respond
significantly more to the stimulus in the receptive field
when it is the locus of the animals attention than when
it is ignored. These results have been found in a host of
visual areas from the lateral geniculate nucleus through
to the middle temporal area (MT) and V4 (Motter, 1993;
Vidyasagar, 1998; McAdams & Maunsell, 1999b; Treue &
Maunsell, 1999; Reynolds et al. 2000; Casagrande et al.
2005; McAdams & Reid, 2005; McAlonan et al. 2008).
However, the magnitude of this effect is not particularly
strong and in some studies the effect is almost absent
in certain cortical areas (Luck et al. 1997; Seidemann &
Newsome, 1999).
Figure 1. Illustration of the primate brain
The locations of visual areas containing neurons whose responses
are modulated by visual attention are shown in light grey. Cortical
areas involved in the allocation of attention and the guidance of eye
movements are shown in dark grey. The superior colliculus, also
involved in this process, is not visible. MT, the middle temporal area;
LIP, the lateral intraparietal area; FEF, the frontal eye field.
J. W. Bisley J Physiol 589.1
Striking effects of attention are seen when two stimuli
are placed within the neurons receptive field (Fig. 2A,
right panel). The logic underlying these experiments
is that as receptive fields get bigger in progressively
higher visual areas, mechanisms must still be available
to focus attention to a discrete region of space. Thus,
these experiments generally place both a preferred and
non-preferred stimulus within the receptive field and have
the animals attend one of the two stimuli. In addition,
extra stimuli are often placed outside of the receptive
field, so that the responses to the stimuli within the
receptive field can be measured under conditions in which
attention is not focused on either. The result of such an
experiment is illustrated in Fig. 2B, which shows the mean
responses from 70 MT neurons with a single or two stimuli
within the receptive field (Lee & Maunsell, 2010). The
effects of attention under these conditions are clear; the
response of the population is biased towards its normal
response to the attended stimulus when it is presented
alone. Thus, if the attended stimulus is the non-preferred
stimulus (continuous grey trace, Fig. 2B), then the activity
is suppressed compared to the non-attended response
to both stimuli (dashed black trace, Fig. 2B), and if the
attended stimulus is the preferred stimulus, then the
response is enhanced (dotted black trace, Fig. 2B). Similar
effects have been seen in V2, V4 and in the temporal lobe
(Moran & Desimone, 1985; Chelazzi et al. 1993, 2001;
Luck et al. 1997; Reynolds et al. 1999; Treue & Maunsell,
1999; Martinez-Trujillo & Treue, 2002).
Feature-based attention. When searching for a particular
item, we are able to effectively ignore objects that do not
share any features with our target. This ability to ignore
completely irrelevant distractors while focusing attention
on categories of objects based on their composition
is termed feature-based attention. Studies examining
this form of attention have shown that neurons that
preferentially respond to a feature have elevated responses
to that feature when it is part of an object within the
neurons receptive field and when that feature is being
attended even if the specific object in the receptive field is
not the focus of attention (Motter, 1994; Treue & Martinez
Trujillo, 1999; McAdams & Maunsell, 2000; Bichot et al.
2005; Buracas & Albright, 2009). For example, if a subject
is looking for a red ball and a red star is in the receptive
field of a neuron that has a preference for red objects, then
the response will be enhanced compared to if the subject
is looking for a blue ball. In this way, the visual system is
highlighting objects that are more likely to be the goal of
the search and, thus, increasing the efficiency of search.
The response modulations under feature-based
attention and spatial attention have been examined
together in a number of studies (McAdams & Maunsell,
2000; Busse et al. 2006; Katzner et al. 2009). These studies

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J Physiol 589.1 Neural basis of visual attention 51

Figure 2. Studying the effects of attention on MT neurons

A, the two main ways that the effects of attention have been studied in visual cortices. Usually at least 1 preferred
stimulus (a direction of motion or oriented bar) and 1 non-preferred stimulus (the opposite direction of motion or
a bar rotated 90 deg from the preferred orientation) are presented on the screen. Either a single stimulus is placed
in the receptive field (left panel), with the other stimulus in an opposite location, or both stimuli are placed in
the receptive field (right panel). B, the response of a population of 70 MT neurons under 5 attentional conditions
(illustrated on right). The dashed grey oval represents the receptive field (RF), the cross represents the fixation point
(FP), and the black cone illustrates the focus of attention. Adapted from Lee & Maunsell (2010) with permission
from the Society for Neuroscience.

have found that single neurons in areas MT and V4 exhibit
similar modulations due to both feature-based attention
and spatial attention, suggesting that the mechanism used
to produce the modulation for the two processes may be
the same (Katzner et al. 2009).
Activity modulation. It is intuitive to imagine that
attention effectively shrinks the receptive field to
encompass the attended stimulus when multiple objects
are in the receptive field (continuous grey or dotted black
traces, Fig. 2B). However, understanding what is occurring
when only a single stimulus is within the RF is less obvious
because the attentional modulation varies with both the
neuronal tuning and stimulus contrast (Martinez-Trujillo
& Treue, 2002; Reynolds & Desimone, 2003). In coming
up with an explanation for the results found when a
single stimulus is presented in the receptive field, two
theories have dominated the literature. The first suggests
that attention shifts the stimulusresponse curve (Fig. 3A;
Reynolds et al. 2000). The second suggests that attention
modulates the gain of the neural response above base-
line (Fig. 3B; McAdams & Maunsell, 1999a,b; Treue &
Martinez Trujillo, 1999). One study set out to differentiate
between these two possibilities, but found that the data
were very well fitted by both models; what variance there
was in the neural data did not allow the responses to fit
unambiguously into one category or the other (Williford
& Maunsell, 2006). However, recent modelling work
has suggested that the differences in the changes to the
stimulusresponse curves may be due to the stimulus
size relative to the area over which the animal is paying
attention (Reynolds & Heeger, 2009). They found that
when the stimulus was smaller than the attended area, a
response shift was seen and when the stimulus was larger
than the attended area, a response gain was seen.
A number of recently published models have been
attempting to explain all attentional modulation using
a response normalization mechanism. These models
explain the response modulations seen with both one
and two stimuli within neurons receptive fields by
either implementing an input gain (Ghose, 2009) or a
normalization of the output signal (Lee & Maunsell, 2009;
Figure 3. Theoretical effects of attention on the contrast
response function
A, the response shift model predicts that the response to an
attended stimulus of a given contrast is the same as the response to
an unattended stimulus of higher contrast. Thus, attention is similar
to turning up the brightness of the stimulus. B, the response gain
model predicts that the response to an attended stimulus of a given
contrast, is a multiplicative increase in response to that contrast.
Thus, attention is similar to just turning up the gain of the response.
Note that the response gain model (B) suggests that the peak
response to a bright stimulus can be increased, whereas the
response shift model (A) does not. Continuous lines, unattended
contrast response functions; dashed lines, attended contrast
response functions.


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52 J. W. Bisley
Reynolds & Heeger, 2009). Although differing in subtle
ways, these models show that it is likely that a unifying
mechanism is used to modulate activity under most, if not
all, attentional conditions (Lee & Maunsell, 2010). These
models are a welcome consensus in the field as they meld
two sets of data that were treated by many as separate
problems to be solved.
Other effects of attention. In addition to modulating
firing rate, attention has been found to influence neural
activity in other ways. Focusing attention increases the
synchronization of local field potentials (LFPs; thought
to represent synaptic input) in V4 (Fries et al. 2001,
2008) and increases the synchronization between the LFP
and the spiking output (Bichot et al. 2005; Fries et al.
2008). Attention also produces more reliable responses in
putative interneurons in V4, as indicated by a reduction
in the variance (Mitchell et al. 2007). There is also
recent evidence from V4 that attention reduces inter-
neuronal correlations (Cohen & Maunsell, 2009), which
would improve the population encoding of information
by reducing correlated noise that would otherwise remain
after pooling. Finally, when attention is allocated to a
stimulus close to, but not within, the receptive field, then
the centre of the receptive field appears to shift towards the
attended location (Connor et al. 1996, 1997; Womelsdorf
et al. 2006). It is thought that this may be related to
the changes in centresurround interactions seen due
to attention (Sundberg et al. 2009) and, consistent with
the normalization models, may be due to multiplicative
attentional modulation at earlier stages within the visual
system (Womelsdorf et al. 2008).
Mechanisms underlying the allocation of attention
Thus far, we have talked about the ways in which attention
modulates the activity of neurons in visual cortices. Now
we will describe the mechanism thought to guide the
allocation of attention. Although we will focus on results
from monkeys, there is a growing literature in humans
based on lesion, TMS and functional imaging (Corbetta
et al. 2008), which shows that the animal model is
providing a good foundation for our understanding of
these processes in the human.
The parietofrontal network. In both humans and
monkeys, the guidance of attention appears to be focused
on a network involving parietal and frontal areas, as well as
the superior colliculus (SC). In the monkey, it is thought
that the lateral intraparietal area (LIP) and the frontal eye
field (FEF) are the specific regions within the parietal and
frontal cortices, respectively, which are involved in this
process (Fig. 1). Anatomically, these areas are all inter-
connected with each other and with visual cortical areas
(Stanton et al. 1988, 1995; Andersen et al. 1990; Blatt et al.
J Physiol 589.1
1990; Tanaka et al. 1990), so they are ideally suited for the
collection of visual information and the feedback to visual
areas in order to guide attention. It is notable that most
of these areas are also involved in the processing of eye
movements, which are really just overt shifts in attention.
Thus, it is likely that the same network guides both overt
and covert attention (Bisley & Goldberg, 2010).
Neural evidence of attentional allocation by the
parietofrontal network. As it is difficult to know exactly
where an animals focus of covert attention is, there are
only a limited number of studies that have explicitly related
the activity of neurons to a qualitatively defined locus of
attention. This has been done explicitly in LIP (Bisley
& Goldberg, 2003; Herrington & Assad, 2010) and SC
(Ignashchenkova et al. 2004). However, the use of visual
search tasks, in which subjects must covertly shift their
attention around an array of objects to find a target without
moving their eyes, gives an indirect measure of attention
that can be used to identify attention-related activity. Such
studies have found covert attention-related activity in LIP
(Thomas & Pare, 2007; Ipata et al. 2009), SC (McPeek
& Keller, 2002b) and FEF (Juan et al. 2004; Thompson
et al. 2005; Buschman & Miller, 2007). Indeed, one recent
study has found evidence showing what appears to be a
neural correlate of shifting covert attention in the activity
of FEF neurons during such search (Buschman & Miller,
2009). In this study, the monkeys appeared to have a
tendency to covertly scan the array in a clockwise direction
(Fig. 4A). Figure 4B shows the responses of FEF neurons
sorted based on where their receptive fields were relative
to the target location. When the target was in the neurons
preferred location, the response was elevated just prior to
the animals signal that he had found the target (indicated
by a saccade). When the target was clockwise to the
neurons preferred location, the response was elevated
earlier as if the animal checked the stimulus in the
neurons preferred location and then moved on to the
next stimulus where it found the target. A similar, but
more exaggerated effect was seen when the target was
opposite the RF as if the animal checked the stimulus in
the neurons preferred location and then checked the next
one before finding the target at the third location. This
result suggests that the activity in FEF and, by extension,
the parietofrontal network, guides covert attention on a
moment-by-moment basis.
To test whether this is the case, several studies have
recorded the activity in the parietofrontal network and in
a mid-level visual area. They have found that attention
leads to enhanced synchrony in responses across the areas
(Saalmann et al. 2007; Gregoriou et al. 2009). These data
suggest that neurons in the parietofrontal network can
modulate the activity in visual areas and are consistent
with the hypothesis that these areas guide the allocation
of covert attention.

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J Physiol 589.1 Neural basis of visual attention 53

Visual search paradigms can also be used to relate neural
activity to saccade generation and, thus, overt attention.
The principle of these studies is that if the neural activity is
used to guide eye movements, then once a target has been
selected by these neurons a set time should elapse before
an eye movement is triggered to the location represented
by the neurons. We can think of this in the following way:
once the neuron selects the target, it tells the oculomotor
system what to aim for and the remaining time is some
set time it takes for the oculomotor system to generate a
saccade. Such a relationship between saccade goal selection
and saccadic onset has been seen in LIP (Ipata et al. 2006a;
Thomas & Pare, 2007), FEF (Bichot et al. 2001; Sato et al.
2001) and SC (McPeek & Keller, 2002a; Shen & Pare, 2007).
Thus, the same areas that appear to be involved in guiding
covert attention are also involved in providing targeting
data to the oculomotor system for overt attention.
Causal evidence of attentional allocation by the
parietofrontal network. In addition to correlations of
neural activity with measures of behaviour, a growing
number of studies have found that inactivation or micro-
stimulation of single areas within the parietofrontal
network biases the allocation of attention. As a rough
tool, these methods can ask whether artificially increasing
or decreasing the activity within an area has an effect on
the allocation of attention. But, they also allow the testing
of specific hypotheses; one can make a prediction about
what would happen to the allocation of attention after
stimulation or inactivation and see if the effects match the
predictions.
Microstimulation involves injecting low current levels
into clusters of physiologically defined neurons. Micro-
stimulation of LIP has been shown to bias orienting
to a flashed stimulus (Cutrell & Marrocco, 2002), while
stimulation of the SC has biased performance in several
attention-demanding tasks (Cavanaugh & Wurtz, 2004;
Muller et al. 2005). The most impressive data from micro-
stimulation come from a series of studies in which Moore
and colleagues have stimulated FEF, using currents that
are too low to induce eye movements. These studies have
shown that microstimulation of FEF improves stimulus
detection behaviourally (Moore & Fallah, 2001) and that
microstimulation induces response changes in V4 neurons
reminiscent of the modulations seen when attention
is allocated to those neurons (Moore & Armstrong,
2003; Armstrong et al. 2006). One of the downsides of
microstimulation experiments is that it is possible that
the behavioural or neural effects are primarily due to
remote stimulation of other more relevant areas via anti-
dromic or orthodromic stimulation of neurons connecting
to the stimulated cluster, or even via stimulation of
axons that pass close to the stimulated area, but are
not connected to it. Given the robust changes elicited
by Moore and colleagues and the fact that the regions
within the parietofrontal network appear to work together,
it is generally accepted that their data are very strong
evidence for this network playing a role in the allocation
of attention.
The role of an area can also be tested by artificially
reducing the activity of neurons within that area. In this
field, it is most commonly done with muscimol, which
inhibits neurons with GABAA receptors for hours at a
time. This removes the main concern of more classical
lesion studies, which is that any main roles the area may
play could be masked by retraining or because redundant

Figure 4. Evidence of shifting covert attention in frontal eye field (FEF)

A, in this experiment, monkeys had to fixate on a small point (the fixation point, FP) and covertly find a target
amongst 3 distractors. Based on the behavioural data, the authors found that the monkeys were serially attending
the stimuli in a clockwise direction (illustrated by the dashed line). B, the responses (illustrated as z-scores by the
colour coding) of FEF neurons are aligned by the onset of the choice saccade and are sorted based on where
the target was relative to the preferred location (RF). Asterisks show significance with Bonferroni correction, dots
show uncorrected significant. Adapted from Buschman & Miller (2009) with permission from Elselvier.


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54 J. W. Bisley
circuits kick in during the recovery phase. Performance in
visual search requiring shifts in covert attention has been
affected by inactivation of LIP (Wardak et al. 2004; Balan
& Gottlieb, 2009; Liu et al. 2010), FEF (Wardak et al. 2006;
Monosov & Thompson, 2009) and SC (McPeek, 2008).
In addition, Lovejoy & Krauzlis (2010) have shown that
cued covert attention is disrupted by inactivation of SC
(Fig. 5). In this study, monkeys were cued by a red ring to
attend a particular location in space and, when a pulse of
motion occurred in that location, they had to indicate the
direction of motion. Further, the monkeys had to ignore a
distractor pulse of motion that was given in the opposite
location. When the cue was in the intact quadrant (right
panel), the animals performance was good and consistent
both before and during inactivation. On the opposite side,
performance was good before inactivation (left panel),
but when the cue was in the inactivated quadrant during
inactivation, the animal often indicated the direction of
the motion pulse in the distractor location (yellow points)
rather than the cued location (red points). This means
that during inactivation the animals often attended the
incorrect location. As the monkeys had to indicate the
direction of motion by a saccade or by a button press
(in different experiments), the authors concluded that
the attentional deficit was not related to a deficit in eye
movement generation. Thus, these data strongly suggest
that the SC is involved in allocating or maintaining guided
Figure 5. Inactivation of the superior colliculus (SC) creates a
deficit in the allocation of attention
In this experiment, monkeys were cued by a red circle to pay
attention to a stimulus in a region of visual space represented by
inactivated SC neurons (left panel) or in the opposite location (right
panel). The animals had to indicate the direction of a pulse of
motion in the cued location (red) and ignore a pulse of motion in the
opposite location (yellow). These graphs show the proportion of
choices made when the monkey correctly indicated the direction of
the cued motion pulse (red points), indicated the direction of the
motion pulse in the distractor location (yellow points) and did not
indicate either of these directions (grey) before and after an injection
of muscimol. Adapted from Lovejoy & Krauzlis (2010) with
permission from Macmillan Publishers Ltd: Nature Neuroscience,
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J Physiol 589.1
covert attention and together with the neural correlations
and microstimulation studies described above, strongly
imply that the frontoparietal network, in which we have
included SC, guides visual attention.
The priority map hypothesis. Many results from
recording, inactivation and microstimulation studies
are consistent with the idea that the frontoparietal
network may operate as a priority map. A priority
map is a representation of the visual world in which
items, objects or locations are represented by activity
that is proportional to their attentional priority. The
hypothesis is based on the saliency map model (Itti
& Koch, 2001); however, we use the term priority to
remove any implication that the responses are due to
bottom-up inputs alone (Serences & Yantis, 2006). The
attentional priority is some combination of bottom-up
input and top-down influences, which include task goals,
evaluation of importance and personal biases (Fig. 6).
The hypothesis is that eye movements are made to the
peak of the map at the time they were programmed and,
on a moment-to-moment basis, covert attention is also
allocated to the peak of the map. This latter statement,
supported by some neurophysiological data (Bisley &
Goldberg, 2003, 2006), is somewhat contentious because
subjects can generally split their attention over long
periods of time. However, it is not clear whether this
splitting is due to a single spotlight shuttling between
locations or due to an actual splitting of attentional
resources.
It is likely that a priority map operates in FEF and SC,
or within the parietofrontal network as a whole; however,
the majority of research studying this hypothesis has been
performed on LIP. These studies have shown that LIP
Figure 6. Theoretical priority map response during visual
search
A, stimulus arrangement and eye movements (white dashed lines) in
a hypothetical visual search task. B, theoretical responses on a
priority map to the search performed in A. Red stimuli are
represented by low activity, blue stimuli are represented by higher
activity. Bars oriented the same way as the target are represented by
greater activity than bars that are not. The bright yellow pop-out
stimulus is represented by elevated activity due to its inherent
salience. The middle blue bar that had just been fixated is
represented by reduced activity.

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J Physiol 589.1 Neural basis of visual attention
activity has a bottom-up component (Roitman & Shadlen,
2002; Balan & Gottlieb, 2006) and top-down influences.
The top-down inputs can enhance activity due to task
demands (Gottlieb et al. 1998; Ipata et al. 2009), reward
expectation (Platt & Glimcher, 1999; Dorris & Glimcher,
2004; Sugrue et al. 2004) and motor plans (Gnadt &
Andersen, 1988), and can suppress responses to salient,
but behaviourally distracting, stimuli (Ipata et al. 2006b)
and to stimuli that have been judged as unimportant
after being foveally examined (Mirpour et al. 2009). All
these inputs and influences appear to be summed almost
linearly (Ipata et al. 2009) and the output predicts saccadic
latency (Ipata et al. 2006a; Thomas & Pare, 2007), the
shifting time of covert attention (Bisley & Goldberg,
2003, 2006; Herrington & Assad, 2010) and monkeys
decisions on two-alternative forced choice tasks (Roitman
& Shadlen, 2002). Thus, the concept of a priority map
for visual attention is particularly appealing because it
explains the varied roles that had been assigned to LIP
(Bisley & Goldberg, 2010) while incorporating a strong
and plausible model of attentional allocation (Itti & Koch,
2001). Indeed, the model is part of the foundation for
an exciting series of experiments which appear to have
successfully aided patients with spatial impairments due
to parietal lesions (Bays et al. 2010).
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Acknowledgements
J.W.B. is supported by a Klingenstein Fellowship Award in
the Neurosciences, an Alfred P. Sloan Foundation Research
Fellowship, a McKnight Scholar Award, and the National Eye
Institute (R01 EY019273).