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Neuropsychologia 49 (2011) 1446–1457

Contents lists available at ScienceDirect

Neuropsychologia
journal homepage: www.elsevier.com/locate/neuropsychologia

A neural theory of visual attention and short-term memory (NTVA)

Claus Bundesen ∗ , Thomas Habekost, Søren Kyllingsbæk
Department of Psychology, University of Copenhagen, Øster Farimagsgade 2A, DK-1353 Copenhagen K, Denmark

a r t i c l e i n f o a b s t r a c t

Article history: The neural theory of visual attention and short-term memory (NTVA) proposed by Bundesen, Habekost,
Received 27 July 2010 and Kyllingsbæk (2005) is reviewed. In NTVA, filtering (selection of objects) changes the number of cortical
Received in revised form neurons in which an object is represented so that this number increases with the behavioural importance
10 November 2010
of the object. Another mechanism of selection, pigeonholing (selection of features), scales the level of
Accepted 2 December 2010
activation in neurons coding for a particular feature. By these mechanisms, behaviourally important
Available online 10 December 2010
objects and features are likely to win the competition to become encoded into visual short-term memory
(VSTM). The VSTM system is conceived as a feedback mechanism that sustains activity in the neurons
Keywords:
Vision
that have won the attentional competition. NTVA accounts both for a wide range of attentional effects in
Attention human performance (reaction times and error rates) and a wide range of effects observed in firing rates
VSTM of single cells in the primate visual system.
TVA © 2010 Elsevier Ltd. All rights reserved.

1. Introduction the number of neurons representing the categorization, which is

This article is a review of the neural theory of visual attention
and short-term memory (NTVA) proposed by Bundesen, Habekost,
and Kyllingsbæk (2005) (see also Bundesen & Habekost, 2008;
Kyllingsbæk, 2006). By use of the same basic equations, NTVA
accounts for a wide range of attentional effects in human per-
formance (reaction times and error rates) and a wide range of
effects observed in firing rates of single cells in the visual system
of primates. NTVA is a further development of the theory of visual
attention (TVA) proposed by Bundesen (1990), which is a formal,
computational theory that applies to attentional effects in mind
and behaviour reported in the psychological literature.
NTVA provides a neural interpretation of the central equations
of TVA: the rate and weight equations. The two equations jointly
describe two mechanisms of attentional selection: one for selection
of objects and one for selection of features or categories. Paying trib-
ute to Broadbent (1971), we use the term filtering for selection of
objects and the term pigeonholing for selection of features or cate-
gories. Selection of a feature is the same as selection of a category
formed by all those objects that share the given feature.
In NTVA, filtering changes the number of cortical neurons in
which an object is represented, whereas pigeonholing changes the
rate of firing of cortical neurons coding for particular features (see
Fig. 1). The total activation representing a visual categorization of
the form “object x has feature i” is directly proportional to both
∗ Corresponding author. Tel.: +45 3532 4858/3253 1752.
E-mail address: bundesen@psy.ku.dk (C. Bundesen).
controlled by filtering, and the level of activation of the individ-
ual neurons representing the categorization, which is controlled
by pigeonholing. The rate equation of TVA essentially expresses this
fact.
Filtering makes the number of cells in which an object is rep-
resented increase with the behavioural importance of the object.
Thus, processing is parallel with differential allocation of resources
so that important objects are represented in many cells. More
specifically, the probability that a cortical neuron represents a par-
ticular object in its classical receptive field equals the attention
weight of the object divided by the sum of the attention weights
across all objects in the receptive field.
The weight equation of TVA describes the computation of atten-
tion weights. Logically the weights must be computed before
processing resources (cells) can be distributed in accordance with
the weights. Therefore, in NTVA, a normal perceptual cycle consists
of two waves: a wave of unselective processing, in which attention
weights are computed, and a wave of selective processing, when
processing resources have been allocated in accordance with the
weights. During the first wave, cortical processing resources are
distributed at random (unselectively) across the visual field. At the
end of the first wave, an attention weight has been computed for
each object in the visual field and the weight has been stored in
a priority map. The weights are used for reallocation of attention
(visual processing capacity) by dynamic remapping of receptive
fields of cortical neurons so that the number of neurons allocated
to an object increases with the attention weight of the object. Thus,
during the second wave, cortical processing is selective in the sense
that the amount of processing resources allocated to an object (the

0028-3932/$ – see front matter © 2010 Elsevier Ltd. All rights reserved.
doi:10.1016/j.neuropsychologia.2010.12.006
 Author's personal copy
C. Bundesen et al. / Neuropsychologia 49 (2011) 1446–1457
Pigeonholing (rate of firing)
Filtering (number of cells)
Fig. 1. Attentional selection in NTVA: combined effects of filtering (selection of
objects) and pigeonholing (selection of features) on the set of cortical spike trains
representing a particular visual categorization of the form “object x has feature i.”
The 4 conditions (quadrants) correspond to the factorial combinations of 2 levels
of filtering (weak vs. strong support to object x) by 2 levels of pigeonholing (weak
vs. strong support to feature i). Filtering changes the number of cortical neurons in
which an object is represented. Pigeonholing changes the rate of firing of cortical
neurons coding for a particular feature.
From “A Neural Theory of Visual Attention: Bridging Cognition and Neurophysiol-
ogy,” by Bundesen et al. (2005), Psychological Review, 112, p. 292. Copyright 2005 by
the American Psychological Association. Adapted with permission.
number of neurons in which the object is represented) varies with
the attention weight of the object. As more processing resources
are devoted to behaviourally important objects than less important
ones, the important objects are more likely to become encoded into
visual short-term memory (VSTM). The VSTM system is conceived
as a feedback mechanism that sustains activity in the neurons that
have won the attentional competition.
In the rest of this article, we first describe TVA as a formal the-
ory of visual attention and short-term memory and summarize the
applications of TVA to human performance. We then present the
neural interpretation of TVA, NTVA, and review some examples of
applications of NTVA to single-cell studies in primates.
2. A formal theory of visual attention and short-term
memory (TVA)
2.1. Basic assumptions
In TVA, both visual recognition and selection of objects in the
visual field consist in making visual categorizations. A visual catego-
rization has the form “object x has feature i” or, equivalently, “object
x belongs to category i.” The categorization is made (or, equivalently,
selected) when the categorization is encoded into visual short-term
memory (VSTM). When the visual categorization that x belongs to i
is made, object x is said both to be selected and also to be recognized
as possessing feature i or, equivalently, as a member of category i.
When a visual categorization of an object completes processing,
the categorization enters VSTM if memory space for the categoriza-
tion is available in VSTM. Usually the capacity of VSTM is assumed
to be limited to K different objects, where K is about 4 (cf. Luck &
Vogel, 1997).
Clearing VSTM starts a race among objects in the visual field to
become encoded into VSTM. An object is encoded in VSTM if, and
only if, some categorization of the object is encoded in VSTM. Each
object x may be represented in the encoding race by all possible
categorizations of the object.
1447
2.1.1. Rate equation
By the rate equation of TVA, the rate, v(x, i), at which a particular
visual categorization, “x belongs to i,” is encoded into VSTM is given
by a product of three terms:
wx
v(x, i) = (x, i)ˇi  (1)
z∈S
wz
The first term, (x, i), is the strength of the sensory evidence
that x belongs to category i. The second term, ˇi , is a perceptual
decision bias associated with category i (0 ≤ ˇi ≤ 1). The third term
is the relative attention weight of object x—that is, the weight of
object x, wx , divided by the sum of weights across all objects in the
visual field, S.
2.1.2. Weight equation
The attention weights in the rate equation of TVA are derived
from pertinence values. Every visual category j is supposed to have
a pertinence, j , which is a nonnegative real number. The perti-
nence of category j is a measure of the momentary importance of
attending to objects that belong to category j. The attention weight
of an object x in the visual field is given by the weight equation of
TVA,

wx = (x, j)j , (2)
j∈R
where R is the set of all visual categories, (x, j) is the strength of
the sensory evidence that object x belongs to category j, and j is
the pertinence of category j. By Eq. (2), the attention weight of an
object is a weighted sum of pertinence values. The pertinence of a
given category enters the sum with a weight equal to the strength
of the sensory evidence that the object belongs to the category.
2.2. Mechanisms of selection
Taken together, the rate and weight equations of TVA describe
two mechanisms of selection: a mechanism for selection of objects
(filtering) and a mechanism for selection of categories (pigeonhol-
ing). The filtering mechanism is represented by pertinence values
and attention weights. If selection of objects with feature j is
desired, the pertinence of feature j should be high. The weight
equation implies that when feature j has a high pertinence, objects
possessing feature j get high attention weights. Accordingly, by
the rate equation, processing of objects with feature j is fast, so
objects with this feature are likely to win the processing race and
be encoded into VSTM.
The pigeonholing mechanism is represented by perceptual deci-
sion bias parameters. Pertinence values determine which objects
are selected (filtering), but perceptual biases determine how the
objects are categorized (pigeonholing). If particular types of cate-
gorizations must be reported or otherwise responded to, the bias
values of the corresponding categories should be high so that the
desired types of categorizations are likely to be made (cf. Eq. (1)).
When the selection system is coupled to a sensory system
that supplies appropriate  values, and when pertinence and bias
parameters have been set, both filtering and pigeonholing are
accomplished by an encoding race between visual categorizations
whose rate parameters are determined through the simple alge-
braic operations of the rate and weight equations. Thus the theory
yields a truly computational account of selective attention in vision.
2.2.1. Note on search templates
Many investigators have emphasized the role of search templates
in visual search (see, e.g., Duncan & Humphreys, 1989). In the biased
competition theory of Desimone and Duncan (1995), Desimone
(1999), and Duncan (1996), visual search is initiated by uploading a
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1448 C. Bundesen et al. / Neuropsychologia 49 (2011) 1446–1457
search template, which is a representation of the target, into VSTM,
which in turn yields a competition bias that facilitates subsequent
processing for sensory input that matches the template. In TVA,
visual selection is determined by , ˇ and  values. Other factors,
such as search templates, may influence the selection process, but
they do so via changes in , ˇ and  values.  values are computed by
comparing stimuli against memory representations. For example,
the strength of the sensory evidence that object x belongs to cate-
gory i, (x, i), is computed by comparing object x against a memory
representation of members of category i (the template associated
with category i). A representation in VSTM can be used as a template
for category i, that is, used for computing (x, i) by being compared
against x, but an observer does not necessarily search the visual
input for any item represented in VSTM. A short-term visual search
template can be defined as a representation in VSTM that is used for
computing  values for a category with a high  value. Long-term
templates can be given a similar definition. A long-term visual search
template can be defined as a representation in long-term memory
that is used for computing  values for a category with a high 
value. In TVA, a search template for category i need not be retained
in VSTM in order to search the visual input for members of category
i.
3. Applications of TVA to human performance
TVA has been applied to findings from a broad range of
paradigms concerned with single-stimulus identification and selec-
tion from multielement displays in normal subjects. In addition,
TVA has been applied in studies of attention deficits, and the scope
of the theory has been extended to other cognitive domains such
as memory and executive control.
3.1. Single-stimulus identification
For single-stimulus identification, TVA provides a mathemati-
cal derivation of the classical biased-choice model of Luce (1963)
(see Bundesen, 1990, 1993). The biased-choice model has been
successful in explaining many experimental findings on effects of
visual discriminability and bias (see, e.g., Townsend & Ashby, 1982;
Townsend & Landon, 1982).
3.2. Selection from multielement displays
For selection from multielement displays, TVA provides a math-
ematical derivation of the fixed-capacity independent race model
(FIRM) of Shibuya and Bundesen (1988). Fig. 2 shows a fit to
the data of Shibuya and Bundesen (1988): probability distribu-
tions of partial and whole report scores with varying numbers
of targets and distractors and varying exposure durations. Other
well-established results closely fitted by TVA include effects of
object integrality (Duncan, 1984) and findings of stochastic inde-
pendence (Bundesen, Kyllingsbæk, & Larsen, 2003; Kyllingsbæk
& Bundesen, 2007) in processing of multiple features, effects of
the spatial position of targets in studies of divided attention (e.g.,
Posner, Nissen, & Ogden, 1978; Sperling, 1967), effects of selec-
tion criterion and effects of the number of distractors in studies of
focused attention (e.g., Bundesen & Pedersen, 1983; Estes & Taylor,
1964; Treisman & Gelade, 1980; Treisman & Gormican, 1988), and
effects of consistent practice in search (Schneider & Fisk, 1982; also
see Kyllingsbæk, Schneider, & Bundesen, 2001).
3.3. Attention deficits
The principles of TVA have been extensively applied in stud-
ies of attention deficits. In the pioneering study, Duncan et al.
(1999) showed how analysis in terms of parameters defined by
TVA enables a very specific measurement of attention deficits in
visual neglect patients (see also Bublak et al., 2005; Finke et al.,
2005). TVA-based assessment has now been used in studies of
simultanagnosia (Duncan et al., 2003), integrative agnosia (Gerlach,
Marstrand, Habekost, & Gade, 2005), alexia (Habekost & Starrfelt,
2006; Starrfelt, Habekost, & Gerlach, 2010; Starrfelt, Habekost, &
Leff, 2009), developmental dyslexia (Dubois et al., 2010), Hunting-
ton’s disease (Finke, Bublak, Dose, Müller, & Schneider, 2006; Finke
et al., 2007), Alzheimer’s disease (Bublak, Redel, & Finke, 2006;
Bublak et al., 2009; Redel et al., 2010), and effects of stroke in par-
ticular parts of the brain (Habekost & Bundesen, 2003; Habekost
& Rostrup, 2006, 2007; Peers et al., 2005; see Habekost & Starrfelt,
2009, for a review). Recently the effect of pharmacological sub-
stances on the TVA parameters has also been investigated (Finke
et al., 2010).
3.4. Other cognitive domains
Logan (1996) proposed a combination of TVA with the con-
tour detector (CODE) theory of perceptual grouping by proximity
(van Oeffelen & Vos, 1982, 1983): the CODE theory of visual atten-
tion (CTVA). CTVA explains a wide range of spatial effects in visual
attention (see Logan, 1996; Logan & Bundesen, 1996; see Bundesen,
1998, for a formalization of CVTA).
Logan and Gordon (2001) extended CTVA into a theory of exec-
utive control in dual-task situations. The theory, ECTVA, accounts
for crosstalk, set-switching cost, and concurrence costs. ECTVA
assumes that executive processes control subordinate processes by
manipulating their parameters. TVA is used as the theory of subor-
dinate processes, and a task set is defined as a set of TVA parameters
that is sufficient to configure TVA to perform a task. Set switching
is viewed as a change in one or more of these parameters, and the
time taken to change a task set is assumed to depend on the number
of parameters to be changed.
Logan (2002) proposed a wide-ranging instance theory of
attention and memory (ITAM) combining ECTVA with the
exemplar-based random walk model of categorization proposed
by Nosofsky and Palmeri (1997). The exemplar-based random walk
model is itself a combination of Nosofsky’s (1986) generalized con-
text model of categorization and Logan’s (1988) instance theory of
automaticity. By integrating theories of attention, categorization,
and memory, the development of ITAM seems a major step toward
a unified account of visual cognition.
4. A neural interpretation of TVA (NTVA)
NTVA gives the equations of TVA an interpretation at the level
of individual neurons in the brain. The theory assumes that a typi-
cal neuron in the visual system is specialized to represent a single
feature. The feature for which the neuron is specialized can be
a more or less simple “physical feature” or a “microfeature” in
a distributed representation. Further, the neuron is assumed to
respond to the properties of only one object at any given time. The
object selection of the neuron occurs by dynamic remapping of the
cell’s classical receptive field so that the functional receptive field
contracts around the selected object. The probability that the neu-
ron comes to represent a particular object in this way equals the
attention weight of the object divided by the total sum of the atten-
tion weights of all objects in the classical receptive field. Dynamic
remapping of receptive fields corresponds to the filtering mecha-
nism in TVA.
NTVA defines the activation of a neuron by the appearance of
an object in its receptive field as the increase in firing rate above
a baseline rate representing the undriven activity of the neuron.
If the baseline rate is zero, the activation is just the firing rate.
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C. Bundesen et al. / Neuropsychologia 49 (2011) 1446–1457 1449

Fig. 2. Relative frequency of scores of j or more (correctly reported targets) as a function of exposure duration with j, number of targets T, and number of distractors D as
parameters in partial report experiment of Shibuya and Bundesen (1988). Data are shown for subject MP. Parameter j varies within panels; j is 1 (open circles), 2 (open
squares), 3 (solid squares), 4 (solid circles), or 5 (triangle). T and D vary among panels. Smooth curves represent a theoretical fit to the data by the FIRM model. For clarity,
observed frequencies <.02 were omitted from the figure.
Adapted from “Visual Selection From Multielement Displays: Measuring and Modeling Effects of Exposure Duration,” by Shibuya and Bundesen (1988), Journal of Experimental
Psychology: Human Perception and Performance, 14, p. 595. Copyright 1988 by the American Psychological Association.

Independently of the distribution of neurons among objects (i.e.,
independently of filtering), the activation in neurons represent-
ing particular features (say, the activation in the set of neurons
representing feature i) can be scaled up or down. This scaling of
firing rates corresponds to varying ˇi in TVA (i.e., the pigeonholing
mechanism). In this neural interpretation, the rate equation of TVA:
wx
v(x, i) = (x, i)ˇi 
w
z∈S z
describes the combined effects of filtering and pigeonholing on the
total activation of the population of neurons representing the cat-
egorization “object x has feature i.” The v value on the left-hand
side of the rate equation is the total activation of the neurons that
represent this particular categorization at the level of processing
in the brain where objects compete for entrance into VSTM. At this
level in the visual system, the classical receptive fields of neurons
are assumed to be so large that each one covers the entire visual
field. On the right-hand side of the equation, both the bias for cat-
egorizing objects as having feature i, ˇi , and the relative attention
weight of object x:
wx
 of activation of the individual neurons representing the categoriza-
z∈S
wz
range between 0 and 1. This implies that the strength of the sensory
evidence that x is an i, (x, i), equals the highest possible value of
v(x, i). Thus, (x, i) equals the total activation of the set of all neurons
coding feature i when all of these cells represent object x (say, when
x is the only object in the visual field) and when the featural bias in
favor of i is maximal (i.e., ˇi = 1). When the proportion of feature-i
coding neurons representing object x:
wx

wz
z∈S
is <1, then the total activation representing the categorization
“object x has feature i” is scaled down by multiplication with this
factor on the right-hand side of the equation. The total activation
representing the categorization also depends on the level of activa-
tion of the individual neurons representing the categorization. The
bias parameter ˇi is a scale factor that multiplies activations of all
feature-i coding neurons, so the total activation representing the
categorization “object x has feature i” is also directly proportional
to ˇi .
Thus, in the neural interpretation we propose for the rate equa-
tion of TVA, the total activation representing the categorization
“object x has feature i” is directly proportional to both the num-
ber of neurons representing the categorization as well as the level
tion. The number of neurons is controlled by the relative attention
weight of x (filtering) and the activation of the individual neurons
is controlled by ˇi (pigeonholing).
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1450 C. Bundesen et al. / Neuropsychologia 49 (2011) 1446–1457
Fig. 3. Possible distribution of visual processing across the human brain. Visual
information from the eye enters the lateral geniculate nucleus (LGN) of the tha-
lamus (1) and is transmitted to striate and extrastriate cortical areas where  values
(strengths of evidence that objects at particular scales and positions have particular
features) are computed (2). The  values are multiplied by  (pertinence) values, and
the products are transmitted from the cortex to a priority map in the pulvinar (Pul)
nucleus of the thalamus, where the products are summed up as attention weights
of the stimulus objects (3). After the first (unselective) wave of processing, cortical
processing capacity is redistributed by means of attention weight signals (w) from
the pulvinar to the cortex, so that during the second (selective) wave of processing,
objects with high attention weights are processed by many neurons (4). The result-
ing  values are multiplied by ˇ (bias) values, and the products are transmitted from
the cortex to a multiscale VSTM map of locations, which is tentatively localized in
the thalamic reticular nucleus (TRN) (5). When the VSTM map is initialized, objects
in the visual field effectively start a race to become encoded into VSTM. In this race,
each object is represented by all possible categorizations of the object, and each
possible categorization participates with an activation (v value) proportional to the
corresponding  value multiplied by the corresponding ˇ value. For the winners
of the race, the TRN gates activation representing a categorization back to some of
those cells in LGN whose activation supported the categorization (6). Thus activity
in neurons representing winners of the race is sustained by positive feedback.
From “A Neural Theory of Visual Attention: Bridging Cognition and Neurophysiol-
ogy,” by Bundesen et al. (2005), Psychological Review, 112, p. 295. Copyright 2005 by
the American Psychological Association. Adapted with permission.
NTVA is a general neurocomputational model that does not
depend critically on a particular anatomical localization of the pro-
posed computations. However we have suggested some plausible
ways in which visual processing may be distributed across the
human brain (see also Section 4.2). One possibility is illustrated in
Fig. 3. For further discussions on the functional anatomy of the pro-
cesses described in TVA, see Bundesen et al. (2005) and Habekost
and Starrfelt (2009).
4.1. The VSTM system
A main purpose of the VSTM system is to keep visual informa-
tion available for other cognitive processes – further perceptual
analysis, thinking, or permanent storage – after the immediate
sensory stimulation has disappeared. In modelling visual short-
term memory, NTVA follows a general idea dating back to Hebb
(1949). Hebb suggested that short-term memory implies that the
activity of neurons representing the selected information is sus-
tained in a positive feedback loop. This way the representation is
kept active in a reverberating circuit beyond the duration of the
original sensory impression. Hebb’s notion of short-term mem-
ory has now gained wide acceptance in cognitive neuroscience.
The feedback mechanism is often assumed to depend on interac-
tions between prefrontal and posterior cortical areas (Fuster, 1997;
Goldman-Rakic, 1995) but thalamo-cortical interaction is also a
main candidate (see Fig. 3).
In NTVA the short-term memory mechanism is based on a topo-
graphically organized map of the objects in the visual field. The map
does not in itself represent the features of the selected objects,
but rather functions as a pointer to their locations. The neurons
representing features of objects at the pointed-to locations are
kept active beyond the immediate sensory stimulation by recip-
rocal connections to the corresponding parts of the VSTM map (see
Section 4.2). Thus, NTVA assumes that visual short-term memory
is constituted by a feedback interaction between sensory neurons
and the VSTM map of locations, which makes it possible for visual
representations to outlast the original sensory stimulation.
The process of encoding information into VSTM starts each
time the map is initialized (i.e., cleared of previous activity). This
is assumed to happen immediately after attention weights have
been computed for a new sensory impression, when the visual sys-
tem is optimally tuned to select the most relevant information.
The selection process functions in a winner-take-all manner, in
which the first activated object representations in the VSTM map
block later ones from entering. The suggested type of winner-take-
all networks is described in detail in Section 4.2, but briefly, all
nodes in such a network excite themselves (when externally trig-
gered) while inhibiting the other nodes. Thus, once a node has been
triggered by external stimulation, its activity will tend to sustain
itself while inhibiting the other nodes to the point where they can
no longer be activated by external signals. In the VSTM map the
inhibitory connections are configured such that when fewer than
K objects are active, an external signal can still overcome the inhi-
bition from the objects that are already encoded. However, when K
units are active, external stimulation representing a new object (an
object at a new location) can no longer activate a new object node
(“K-winners-take-all network”). This mechanism explains the lim-
ited storage capacity of VSTM, where only K objects can be active at
the same time. Note that the network is only limited in terms of the
number of objects (or corresponding locations) in the VSTM map,
but not in the number of features related to each of the selected
objects. This means that once an object has established itself in
VSTM, many different categorizations can be attached to it.
Which type of neural signal is needed for capturing a slot in
the VSTM map? It is not currently clear what the effective signal
is. The simplest possibility is the very first spike from one of the
many sensory neurons projecting to the map. If neural processing is
viewed as very noisy, this may seem like a highly unreliable mech-
anism, but there is evidence to suggest that single spikes can in fact
have profound effects on cognitive processing (Parker & Newsome,
1998). Alternatively, a stronger signal from multiple neurons may
be needed, perhaps a volley of at least n synchronized spikes, n
being a number substantially greater than one. However the effec-
tive signal is defined, NTVA assumes that the VSTM system is
configured so that the first such impulse to the VSTM map activates
the winner-take-all network. Assuming that signals for different
categorizations as well as repeated signals for a given categoriza-
tion occur independently of each other, this leads to an exponential
race model. That is, it implies a model where selection is deter-
mined by a parallel processing race among categorizations, a race
in which processing times for individual categorizations are mutu-
ally independent, exponentially distributed random variables. Such
a model is completely in line with the original TVA model.
The description so far applies to the simple case in which the first
visual categorization arriving in VSTM is used directly by the cog-
nitive system. This mode of processing can be termed immediate
perception. However, sometimes mutually contradicting catego-
rizations of the same object are encoded into VSTM. This may for
example be the case under poor lighting conditions, where the
colour of an object is not obvious. In TVA terms, if the  value for “x is
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C. Bundesen et al. / Neuropsychologia 49 (2011) 1446–1457 1451

of times (see Logan, 1996). In a random-walk procedure, the deci-

Fig. 4. Winner-take-all network with three units. Excitatory connections are shown
by arrows. Inhibitory connections are shown by lines ending in solid circles.
From “A neural theory of visual attention: Bridging cognition and neurophysiology”
by Bundesen et al. (2005), Psychological Review, 112, 291–328. Copyright 2005 by
the American Psychological Association. Adapted with permission.
red” is not substantially different from the  value for “x is yellow”,
both categorizations may end up in VSTM. In situations like this
decisional procedures are required: a processing mode of mediate
perception. Two possible mechanisms for mediate perception have
been suggested within the TVA framework, both based on gradual
accumulation of evidence: counter procedures and random-walk
models. In a counter procedure, a decision is made when one of the
categorizations has been repeated (confirmed) a certain number
sion depends on the difference between the numbers of alternative
categorizations: the “winning” categorization must occur a certain
number of times more than its closest competitor (see Logan, 2002).
Both of these models can be given a mathematically exact form
(see Appendix B of Bundesen et al., 2005), but still await conclusive
empirical testing.
Representations in VSTM can be relatively concrete and “picto-
rial” or relatively abstract, “schematic,” and “conceptual.” The more
concrete sensation-like representations are often named mental
images. In NTVA simple mental images are regarded as faint copies of
sense impressions. A suggested implication of this sensory nature
is that a mental image includes a specification of the position of
the represented object in the visual field, although localization
may be less precise than with real sense impressions. Neurophys-
iologically, mental images should activate only a subset of the
neurons that are implicated in the corresponding sensory impres-
sions, and the activity in these neurons should be weaker (hence
the “faintness” of the representation). An observable effect in sin-
gle cell activity should be a moderate increase in the firing rate of
some neurons when a mental image is positioned in their recep-
tive fields and no external stimuli are present. Such a baseline shift
has been shown in several studies (e.g., Chelazzi, Duncan, Miller, &
Desimone, 1998; Miller, Erickson, & Desimone, 1996; Miller, Li, &
Desimone, 1993).
4.2. Computational networks
In this section we present some simple networks that can per-
form the attentional operations described in NTVA. We show how
the computations of these networks correspond to the rate and
weight equations of TVA, and we hint at where such networks may

w1

w1

2 w2

w3 w2
3
w3

w4

4 w4
w5
w5

5
w6

6 w6

Stimuli V2/V3 V4 IT
Fig. 5. Hierarchic network with processing units at three levels: V2/V3, V4, and IT. Each V2/V3 unit has one stimulus in its classical receptive field, each V4 unit has two
stimuli in its classical receptive field, and the IT unit has all of the six stimuli in its classical receptive field. By attentional gating, the effective receptive fields of V4 and IT
units are contracted so that each of the effective receptive fields contains only one object: The symbol stands for a gate which is open on only the upper or the lower line,
and stands for a gate which is open on only one of the three lines. The gates are set by competing attentional weight signals. For k = 1, . . ., 6, wk is the attention weight of
Stimulus k.
From “A neural theory of visual attention: Bridging cognition and neurophysiology” by Bundesen et al. (2005), Psychological Review, 112, 291–328. Copyright 2005 by the
American Psychological Association. Adapted with permission.
 Author's personal copy
1452 C. Bundesen et al. / Neuropsychologia 49 (2011) 1446–1457
be located in the primate brain. However, NTVA is a fairly general
neurophysiological interpretation of TVA, and it does not depend
critically on a specific anatomical localization of the proposed oper-
ations. The particular networks we present in the following may be
regarded as merely proofs of the existence of simple and biologi-
cally plausible neural networks that can implement the attentional
operations of NTVA.
4.2.1. Winner-take-all networks
A basic functional element for two central mechanisms in NTVA,
dynamic remapping of receptive fields and encoding into VSTM,
is the winner-take-all network. NTVA uses a simple network pro-
posed by Grossberg (1976, 1980), which is illustrated in Fig. 4. As
shown in the figure, it consists of a set of units (populations of
cells) where each unit excites itself and inhibits all other units in
the cluster. When the cluster is initialized (i.e., its level of activ-
ity is reset to the baseline level), an external signal of a certain
above-threshold strength is sufficient to trigger one of the units.
Once the unit has been triggered it keeps firing because of its self-
exciting connections. At the same time it inhibits the other units.
If the other units are inhibited so strongly that they can no longer
be activated by external signals, one can simply read from the state
of the cluster which of the units received the first above-threshold
trigger signal after the cluster was initialized. This way, the clus-
ter can serve as a device for recording the winner of a processing
race. If the strengths of the inhibitory signals are adjusted so that
at least K units must be active in order to block external activation
of the remaining units, the cluster works as a K-winners-take-all
network.
4.2.2. Dynamic remapping of receptive fields
Using the Grossberg winner-take-all network as a building
block, dynamic remapping of receptive fields can be carried out in
a simple hierarchical network. In this network, processing capac-
ity (i.e., neurons) can be variously allocated to different stimuli by
opening and closing gates that control communication between
lower and higher levels in the visual system. The gates can be con-
trolled by attentional weight signals in such a way that the expected
number of cells that represent a particular object at the highest level
of the visual system (which projects to VSTM) becomes propor-
tional to the relative attention weight of the object. The hierarchical
network is shown in Fig. 5. For simplicity, only three processing lev-
els are shown (named V2/V3, V4, and IT), with gates between each
level. Fig. 6 shows one of these gates in close-up. It consists of a
winner-take-all cluster of units (one unit for each line through the
gate), logical AND units (one on each line through the gate) and a
logical OR unit (which collects information from the AND units).
The winner-take-all cluster records the first occurring signal to one
of the two units in the cluster. Thus, if the upper unit receives a sig-
nal before the lower unit, the upper unit will be excited, the lower
unit will be inhibited, and only the upper line through the gate will
be open. External signals to units in the winner-take-all clusters
come from the so-called priority map, which represents attention
weights at particular locations. As indicated in Fig. 5, a winner-take-
all cluster that gates responses from cells at a lower level to a cell
at a higher level receives one attentional weight signal for every
object within the classical receptive field of the higher level cell.
Attentional weight signals for objects within the receptive field of
a given lower level cell go to that unit in the winner-take-all clus-
ter that supports communication from the lower level cell to the
higher level cell. Attentional weight signals for objects outside the
receptive field of the lower level cell (but within the receptive field
of the higher level cell) go to other, competing units in the winner-
take-all cluster. As shown below, this simple network can do object
selection according to the TVA equations.
&
w1
OR
w2
&
V2/V3 V4
Fig. 6. Close-up of the uppermost gate in Fig. 5. The gate consists of a winner-take-
all network with two units (each controlling one of the two lines through the gate),
a logical AND (&) unit on each line, and a logical OR unit transmitting information
from either AND unit. For k = 1, 2, wk is the attention weight of Stimulus k. The signal
representing the weight of a given stimulus comes from the representation of the
stimulus in a topographic priority map.
From “A neural theory of visual attention: Bridging cognition and neurophysiology”
by Bundesen, Habekost, and Kyllingsbæk, 2005, Psychological Review, 112, 291–328.
Copyright 2005 by the American Psychological Association. Adapted with permis-
sion.
Consider the probability that the receptive field of the IT unit in
the network described contracts around Object 1 rather around one
of the other objects in the visual field. This equals the probability
that both the upper one of the gates from V2/V3 to V4 as well as the
gate from V4 to IT open on their upper lines. To estimate the proba-
bility, we make some simple assumptions on the nature of parallel
processing in the visual system. Let the arrival times of attentional
weight signals for an object x be exponentially distributed with a
rate parameter equal to the attention weight of x, wx . Let attentional
weight signals for different objects be stochastically independent.
Finally, let attentional weight signals that represent the same object
x but go to different gates be stochastically independent. On these
assumptions, the probability that the upper of the gates from V2/V3
to V4 opens on its upper line is w1 /(w1 + w2 ). The probability that
the gate from V4 to IT opens on its upper line is (w1 + w2 )/wk ,
where the summation extends over Objects 1–6. The probability of
both events equals the product of the two probabilities (since they
are stochastically independent), which is w1 /wk . Next, consider a
population of IT units, each of which is similar to the one we have
considered so far. Each unit has Objects 1–6 and no other objects
within its classical receptive field. If there are N units in the popula-
tion, the expected number of IT units representing Object 1 equals
Nw1 /wk , the expected number of IT units representing Object 2
equals Nw2 /wk , and so on. In general, the expected number of IT
units representing a particular object should be proportional to the
relative attention weight of the object. This way, the network con-
verts activity representing attention weights in the priority map
directly into the number of IT cells working on the corresponding
objects.
Dynamic remapping of the receptive fields of high-level neurons
also has the interesting property of dramatically increasing their
spatial resolution. In the network model shown in Figs. 5 and 6, the
effective receptive field of an IT neuron is coded by the way the gates
in the network are set. The code formed by the gate settings can be
used for directing signals from an IT cell, whose effective recep-
tive field is contracted around an object at a particular location, to
another unit (a neuron or a population of neurons) representing
the object at the selected location. Especially, the code formed by
the gate settings can be used for directing signals to the location in
a topographic map (such as the priority or VSTM maps) that corre-
sponds to the location of the stimulus object. A simple network for
this operation is shown in Fig. 7.
 Author's personal copy
C. Bundesen et al. / Neuropsychologia 49 (2011) 1446–1457
&
&
& &
&
& w1
&
& OR
& w2
&
Inter- Topographic
V2/3 V4 IT
neurons map
Fig. 7. Network for directing signals from an IT cell, whose effective receptive field
is contracted around an object at a particular location, to a unit representing the
object at the given location in a topographic map. The network on the left side of the
figure is similar to the network in Fig. 5, but each cell in a winner-take-all cluster
in one of the gates on the left side of the figure controls not only an input line on
the route to the IT unit but also a corresponding output line from the IT unit to the
topographic map. In the depicted state of the network, the effective receptive field
of the IT unit equals the receptive field of the upper V2/V3 unit, and output from the
IT unit is directed to a place in the topographic map of objects that corresponds to
the receptive field of the upper V2/V3 unit.
From “A neural theory of visual attention: Bridging cognition and neurophysiology”
by Bundesen, Habekost, and Kyllingsbæk, 2005, Psychological Review, 112, 291–328.
Copyright 2005 by the American Psychological Association. Adapted with permis-
sion.
Consider a cell in a winner-take-all cluster in one of the gates on
the left side of Fig. 7. The cell has an excitatory connection to an AND
unit on a particular input line on the route to the IT unit. The cell
also has an excitatory connection to an AND unit on a corresponding
output line from the IT unit to a topographic map of objects. By this
arrangement, the pattern of open and closed lines on the left side
of the figure is duplicated on the right side of the figure. As a result,
signals from the IT unit are directed to the appropriate location in
the topographic map of objects. In the depicted state of the network,
the effective receptive field of the IT unit equals the receptive field
of the upper V2/V3 unit, and output from the IT unit is directed to
a place in the topographic map of objects that corresponds to this
location.
The topographic map of objects depicted in Fig. 7 may be located
at a very high level in the visual system such as the prefrontal cortex.
In this particular case, the dashed lines in Fig. 7 should symbolize
long-ranging connections from cells in gates controlling the flow of
information from the retina to the IT (gates on the left side of the
figure) to cells controlling the flow of information from IT to the pre-
frontal cortex (AND units on the right side of the figure). However,
as illustrated in Fig. 3, another possibility is that the topographic
map is found at a low level of the visual system such as the tha-
lamus. By this hypothesis, the right side of Fig. 7 depicts a flow of
information from the IT unit back towards the retina: a gate and an
AND unit connected by a dashed line are located at the same level of
the visual system, and the connections symbolized by the dashed
lines are quite local. This alternative hypothesis is illustrated in
Fig. 8, which shows a gate in close-up. The figure is an extension of
Fig. 6, and the dashed lines correspond to dashed lines in Fig. 7. The
figure illustrates how the same gates routing information from a
particular location on the retina to a certain cell at a higher level of
the visual system can be used for routing information back from the
high-level cell to a corresponding place in a map of objects found
at a low level of the visual system.
Visual pathways are generally reciprocal so that a pathway from
area A to area B is accompanied by a pathway from B to A (e.g., Zeki,
1993). From an engineering point of view, it seems simple to design
the visual system so that the two pathways in a pair open and close
1453
&
&
&
Fig. 8. Close-up of the uppermost gate in Fig. 7. The figure is an extension of Fig. 6,
and the dashed lines correspond to dashed lines in Fig. 7. The figure illustrates
how the same gates routing signals along a particular pathway from lower-level
to higher-level cells in the visual system can be used for routing signals along a par-
allel pathway in the opposite direction. The gate is bistable. In one state, both upper
lines through the gate are open and both lower lines are closed; in the other state,
both lower lines are open and both upper lines are closed. When the upper lines
are open, signals are directed from the retinal location corresponding to the pair
of cells in the upper left corner through the ascending pathway originating in the
lower member of the rightmost pair of cells. In this state, signals arriving through the
descending pathway to the upper member of the rightmost pair of cells are directed
back towards the retinal location corresponding to the pair of cells in the upper left
corner rather than the retinal location corresponding to the pair of cells in the lower
left corner.
From “A neural theory of visual attention: Bridging cognition and neurophysiology”
by Bundesen, Habekost, and Kyllingsbæk, 2005, Psychological Review, 112, 291–328.
Copyright 2005 by the American Psychological Association. Adapted with permis-
sion.
at the same time (bidirectional opening and closing). Thus, it seems
plausible that the system is designed so that information about an
object at a particular location in the visual field is processed at high
levels of the visual system, but routed back to a topographic map of
objects at a low level of the system (cf. Schneider, 1995). Evolution
may have chosen this simple solution.
4.2.3. Priority map
Fig. 9 shows a possible implementation of the priority map,
which represents the attention weights of objects in the visual
field. Computation of attention weights occurs at many different
levels of the visual system, perhaps all the way from primary visual
cortex (V1) to IT cortex. The network in Fig. 9 computes attention
weights by summing up input (activations measured in spikes per
second) to a unit (a neuron or a population of neurons) in a topo-
graphically organized priority map. The priority map is tentatively
located in the pulvinar nucleus of the thalamus. The input is gen-
erated by cortical units specialized to represent different features,
based on visual signals from the lateral geniculate nucleus (LGN).
Each input to the unit for object x in the priority map is a product
of two factors: a level of activation of a cortical neuron represent-
ing the strength of the sensory evidence that x has some feature,
j, and a pertinence factor,  j , which is proportional to the perti-
nence of j, j , and represents the current importance of attending
to feature j. The products are summed up for each object location in
the priority map to produce a topographic distribution of attention
weights, following the principle laid out in TVA’s weight equation
(for mathematical details, see Appendix A of Bundesen et al., 2005).
4.2.4. VSTM loops
NTVA assumes that a visual categorization is encoded in VSTM
when the categorization is embedded in a positive feedback loop
gated by a unit in the VSTM map of objects. Two feedback loops of
 Author's personal copy
1454 C. Bundesen et al. / Neuropsychologia 49 (2011) 1446–1457
Fig. 9. Network for computing attention weights. Signals from the lateral genicu-
late nucleus (LGN) are transmitted to striate and extrastriate cortical areas where
 values (strengths of evidence that objects at particular scales and positions have
particular features) are computed. The  values are multiplied by  (pertinence) val-
ues, and the products are transmitted from the cortex to the pulvinar nucleus of the
thalamus, where the products are summed up as attention weights of the stimulus
objects.
From “A neural theory of visual attention: Bridging cognition and neurophysiology”
by Bundesen, Habekost, and Kyllingsbæk, 2005, Psychological Review, 112, 291–328.
Copyright 2005 by the American Psychological Association. Adapted with permis-
sion.
this type are illustrated in Fig. 10. As shown in the figure, impulses
routed to a unit that represents an object at a certain location in the
VSTM map of objects are fed back to the feature units from which
they originated, provided that the VSTM unit is activated. If the
VSTM unit is inactive, impulses to the unit are not fed back. Thus,
for each feature-i neuron representing object x, activation of the
neuron is sustained by feedback when the unit representing object
x in the topographic VSTM map of objects is activated.
Fig. 11 shows how the circuits illustrated in Fig. 10 might be
implemented as thalamo-cortical feedback loops. In Fig. 11, feature
& the feature-i and the feature-j neurons allocated to object x are pro-
& categorization that “object x has (motion) feature j” are routed to
Object unit
Feature units
in VSTM map
Fig. 10. Two feedback loops gated by the same unit in the VSTM map of objects.
Activation of either feature unit is sustained by positive feedback if, and only if, the
object unit is activated.
From “A neural theory of visual attention: Bridging cognition and neurophysiology”
by Bundesen, Habekost, and Kyllingsbæk, 2005, Psychological Review, 112, 291–328.
Copyright 2005 by the American Psychological Association. Adapted with permis-
sion.
Fig. 11. Possible implementation of the circuits illustrated in Fig. 10 as thalamo-
cortical feedback loops. The feature units are located in different cortical areas,
where  values (strengths of evidence that objects at particular scales and posi-
tions have particular features) are computed. The  values are multiplied by ˇ (bias)
values and the products are projected back to the thalamus as rates of activation.
The VSTM map of objects is located in the thalamic reticular nucleus (TRN), which
lies like a thin shield between the thalamus and the cortex. When the VSTM map
is initialized, objects in the visual field effectively start a race to become encoded
into VSTM. In this race, each object is represented by all possible categorizations of
the object, and each possible categorization participates with a firing rate (v value)
proportional to the product of the corresponding  and ˇ (bias) values. For the win-
ners of the race, the TRN gates activation representing a particular categorization
back to some of those cells in LGN whose activation supported the categorization.
Thus activity in neurons representing winners of the race is sustained by positive
feedback.
From “A neural theory of visual attention: Bridging cognition and neurophysiology”
by Bundesen, Habekost, and Kyllingsbæk, 2005, Psychological Review, 112, 291–328.
Copyright 2005 by the American Psychological Association. Adapted with permis-
sion.
i is a shape feature of an object x, and feature j is a motion feature
of the same object x. Shape-selective feature-i neurons are found in
a high-level cortical area in the ventral stream of visual processing
(see Milner & Goodale, 1995; Ungerleider & Mishkin, 1982), where
their rates of activation are modulated by multiplication with the
perceptual decision bias ˇi . Motion-selective feature-j neurons are
found in a high-level cortical area in the dorsal stream of visual pro-
cessing (cf. Milner & Goodale, 1995; Ungerleider & Mishkin, 1982),
where their rates of activation are modulated by multiplication
with the perceptual decision bias ˇj . The rates of activation of both
jected back to the thalamus. The top-down impulses representing
the categorization that “object x has (shape) feature i” are routed to
some of those cells in LGN whose bottom-up activation supported
this categorization, and the top-down impulses representing the
some of those cells in LGN whose bottom-up activation supported
that categorization. In either case, the feedback occurs through an
AND unit, which is tentatively located in the LGN close to the cells
that are targeted by the feedback. To make the feedback effective,
the AND gates must be open. This means that the AND units must
receive input from a unit representing object x in the VSTM map of
objects, tentatively located in the thalamic reticular nucleus (TRN).
Effectively, the feedback loops with information about features of
object x are complete if the TRN unit representing object x in the
VSTM map of objects is active. An obvious way of activating the TRN
unit is illustrated in Fig. 11: both impulses representing “object x
has feature i” and impulses representing “object x has feature j” are
projected back, not only whence they came in the LGN, but also
 Author's personal copy
C. Bundesen et al. / Neuropsychologia 49 (2011) 1446–1457
to the unit representing object x in the TRN. If, for any reason, the
TRN unit representing object x is and remains inactive, impulses
from the cortex representing object x are not transmitted beyond
the AND units in the LGN.
5. Applications of NTVA to single-cell studies
NTVA has been applied to explain a wide range of attentional
effects observed in single-cell studies: effects on the rate of fir-
ing with multiple stimuli in the receptive field of a cell, effects
on the rate of firing with a single stimulus in the receptive field,
effects on baseline firing, and effects on neural synchronization (see
Bundesen et al., 2005; Bundesen & Habekost, 2008). A few examples
of simple, qualitative applications to classical studies of filtering
and pigeonholing are sketched below.
5.1. Neural filtering
NTVA’s notion of attentional filtering by dynamic remapping
of receptive fields was originally inspired by a study of Moran
and Desimone (1985). Moran and Desimone presented macaque
monkeys with stimuli at two locations in the visual field. The mon-
keys were trained to attend only to stimuli presented at one of the
locations and ignore any stimuli shown at the other location. The
monkeys performed a match-to-sample task, in which they were to
encode a sample stimulus that was shown at the attended location,
retain it in memory for a brief delay period, and then compare it
to a test stimulus shown at the same location. During the presen-
tation of both the sample and test displays, Moran and Desimone
recorded the responses of single neurons to the stimuli in cortical
areas V1, V4, and IT. They found that, when the target and distractor
stimuli were both present within the receptive field of the recorded
cell in visual areas V4 or IT, the cell’s rate of firing depended on the
properties of the target, but not the distractor. For example, they
recorded the responses of individual cells to a pair of stimuli con-
sisting of (a) a stimulus that elicited a high rate of firing when the
stimulus was presented alone (an effective sensory stimulus) and
(b) a stimulus that had little or no effect on the rate of firing when
the stimulus was presented alone (an ineffective sensory stimulus).
On trials in which the effective stimulus was the target and the inef-
fective stimulus was the distractor, the cell showed a high rate of
firing. However, on trials in which the ineffective stimulus was the
target and the effective stimulus was the distractor, the cell’s fir-
ing rate was low. Moran and Desimone remarked that the typical
cell responded “as if the receptive field had contracted around the
attended stimulus.”
The effect depended on both the target and the distractor being
located within the recorded neuron’s receptive field. However in
cortical area IT the receptive fields were very large, covering at least
the central 12 degrees of both the contralateral and ipsilateral visual
field. For these neurons, the distractors were always located inside
the receptive field. On the contrary, in V1 only one stimulus could be
fitted into each neuron’s receptive field. In this case, no significant
effects of attention were found. Also in V4, no effect of attention
was found when only a single stimulus was placed in the receptive
field. These negative findings fit well with the predictions of NTVA.
Overall, the results of Moran and Desimone clearly support NTVA’s
notion of filtering, and in fact suggested the interpretation in the
first place.
Reynolds, Chelazzi, and Desimone (1999) corroborated and
extended the findings of Moran and Desimone (1985). Recording
from neurons in cortical areas V2 and V4, they found that the mean
firing rate of a cell to a pair of stimuli in its receptive field approx-
imated a weighted average of the firing rates of the cell to each
of the stimuli in the pair when they were presented alone. When
1455
the monkey’s attention was directed to one of the stimuli in the
pair, this increased the weight on the target stimulus so that the
mean response of the neuron was driven (up or down) toward the
response elicited when the target stimulus was presented alone.
The results fit in with the dual conjectures that (a) at any time, a
cell was driven by only one of the stimuli in its receptive field and
(b) the probability that the cell was driven by a given stimulus was
proportional to the attention weight of the stimulus. This is exactly
what is predicted by NTVA.
5.2. Neural pigeonholing
Whereas NTVA’s filtering mechanism affects the number of
neurons in which an object x is represented, the pigeonholing mech-
anism affects the way in which the element is represented in each
of those neurons that are allocated to the object. The activation in
a neuron representing the categorization that object x has feature
i should be proportional to the multiplicative perceptual bias, ˇi ,
which is applied to neurons that are specialized for signaling fea-
ture i. Recordings from single cells in the visual system of monkeys
have provided evidence of such a mechanism of attentional selec-
tion: a feature-based mechanism of attention that selects groups
of neurons with similar stimulus preferences for a multiplicative
enhancement in response strength.
Treue and Martinez-Trujillo (1999) (also see Martinez-Trujillo
& Treue, 2004) recorded from cortical area MT, which contains
cells that are selective to the direction of motion. In the basic task,
Treue and Martinez-Trujillo presented monkeys with two coher-
ently moving random dot patterns, one placed inside the receptive
field of the neuron being recorded and the other one in the opposite
visual hemifield (see Fig. 12). At the start of each trial, the monkey
was shown a cue at one of the two locations. Following this, the
two random dot patterns appeared and the monkey was required
to detect small changes in the speed or direction of movement of
the pattern at the cued location. These motion changes occurred
after a random delay ranging between a few hundred milliseconds
and several seconds.
In one of their experiments, Treue and Martinez-Trujillo demon-
strated an effect of pigeonholing with respect to a given direction
of movement (“feature-based attention”); probably the first clear
demonstration of pigeonholing in the single-cell literature. In this
experiment, the recorded neuron’s receptive field was stimulated
by a pattern moving in the direction preferred by this particu-
lar neuron. Outside the receptive field, a pattern was presented
Fig. 12. Stimuli used by Treue and Martinez-Trujillo (1999). The random dot pat-
tern inside the receptive field (dashed circle) always moved in the cell’s preferred
direction (upward pointing arrow (1)); the stimulus outside moved in either the
same (2) or the opposite direction (3).
From “Feature-Based Attention Influences Motion Processing Gain in Macaque
Visual Cortex,” by S. Treue and J. C. M. Martinez-Trujillo, 1999, Nature, 399, p. 577.
Copyright 1999 by Nature Publishing Group. Adapted by permission from Macmillan
Publishers Ltd.
 Author's personal copy
1456 C. Bundesen et al. / Neuropsychologia 49 (2011) 1446–1457
that moved either in the same direction or in the opposite (non-
preferred) direction. When the monkey attended to the pattern
outside the receptive field, the response of the recorded neuron
varied with the direction of movement being attended. The firing
rate went up when the attended pattern (outside the receptive field
of the recorded neuron) moved in the preferred direction of the
recorded neuron. The firing rate went down when the attended
pattern moved in the direction opposite to the preferred direc-
tion of the recorded neuron. The spatial location of the attended
pattern was exactly the same in the two conditions. Thus a nonspa-
tial, feature-based mechanism of attention seemed to be at work:
pigeonholing.
The result can be explained by assuming that when the mon-
key attended to movement in a particular direction, the ˇ value
for movement in that direction was high while the ˇ value for
movement in the opposite direction was low. As the monkey was
monitoring the pattern for hundreds to thousands of milliseconds,
there should be plenty of time to adjust the ˇ values in accordance
with the display. Consider the situation in which the preferred
movement of the recorded neuron was upwards but the monkey
attended to movement in the opposite direction (downwards), try-
ing to detect a small change in the movement of the pattern to
be attended. Perceptual bias should generally reflect expectations,
and since the monkey had learned that only small changes in the
direction of movement of the target would occur, and the target pat-
tern was moving downwards, ˇupwards should be low. Because the
activation of the recorded cell should be proportional to ˇupwards ,
the recorded activation should also be low. By contrast, when the
monkey attended to movement in the preferred direction of the
recorded cell, ˇ values should be high for upwards and nearby
directions. In this case, ˇupwards being high, the activation of the
recorded neuron also should be high.
6. Concluding remarks
As exemplified in the previous section, NTVA has been applied
to explain a wide range of attentional effects observed in single-
cell studies on filtering and pigeonholing (for applications of NTVA
to brain imaging studies of visual attention and short-term mem-
ory, see Bundesen & Habekost, 2008, chapter 9). The applications
to single-cell studies have yielded substantial support to the rate
equation of TVA and the neurophysiological interpretation of this
equation made in NTVA. As emphasized in the sections above
entitled The VSTM System and Computational Networks, NTVA also
contains specific proposals for neural networks implementing not
only the attentional functions but also the visual short-term mem-
ory functions assumed in NTVA (see, in particular, Figs. 10 and 11).
The particular networks we have presented may be regarded as
merely proofs of the existence of simple and biologically plausi-
ble neural networks that can implement the operations of NTVA.
However, given the simplicity and the biological plausibility of the
networks, it would seem worthwhile to search for the suggested
types of networks in the human brain.
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