Professional Documents
Culture Documents
Abstract. We describe a neural network that enhances (1993), Heitger and von der Heydt (1993), Parent and
and completes salient closed contours in images. Our Zucker (1989), Sha’ashua and Ullman (1988), and
work is different from all previous work in three Grossberg and Mingolla (1985). In this paper we de-
important ways. First, like the input provided to scribe a neural network that enhances and completes
primary visual cortex (V1) by the lateral geniculate salient closed contours in images. Our work is different
nucleus (LGN), the input to our computation is isotro- from all previous work in three important ways. First,
pic. That is, it is composed of spots, not edges. Second, like the input provided to primary visual cortex (V1)
our network computes a well-defined function of the by the lateral geniculate nucleus (LGN), the input to
input based on a distribution of closed contours our computation is isotropic. That is, the input is
characterized by a random process. Third, even though composed of spots, not edges.1 Second, our network
our computation is implemented in a discrete network, computes a well-defined function of the input based on
its output is invariant to continuous rotations and a distribution of closed contours characterized by a
translations of the input image. random process. Third, even though our computation
is implemented in a discrete network, its output is in-
variant to continuous rotations and translations of the
input image.2
There are two important properties a computation
1 Introduction must possess if it is to be invariant to rotations and
translations, i.e., Euclidean invariant. First, the input,
Vision, whether that of humans or machines, must solve the output, and all intermediate representations must
certain fundamental information-processing problems. be Euclidean invariant. Second, all transformations of
One such problem is robustness to rotations and these representations must also be Euclidean invariant.
translations of the input image. Ideally, the output of Previous models are not Euclidean invariant, first and
a visual computation should vary continuously under foremost because their input representations are not
continuous transformation of its input. We believe that Euclidean invariant. That is, not all rotations and
an understanding of how the human visual system translations of the input can be represented equally
achieves this invariance will lead to significant improve- well. This problem is often skirted by choosing input
ments in the performance of machine vision systems. patterns that match particular choices of sampling rate
Conversely, by studying the fundamental information and phase. For example, Li (1998) used only six
processing problems common to both human and samples in orientation (including 0 ) and Heitger and
machine vision we can achieve a deeper understanding von der Heydt (1993) only twelve (including 0 , 60
of the structure and function of the human visual and 120 ). Li’s first test pattern was a dashed line of
cortex. orientation, 0 , while Heitger and von der Heydt
There is a long history of research on neural net- (1993) used a Kanizsa Triangle with sides of 0 , 60 ,
works inspired by the structure of visual cortex whose
functions have been described as contour completion,
saliency enhancement, orientation sharpening, or seg- 1
Like our model, Guy and Medioni’s (1996) can also be applied
mentation, e.g., August (2000), Li (1998), Yen and to input patterns consisting of spots.
Finkel (1997), Guy and Medioni (1996), Iverson 2
Guy and Medioni (1996) represent a distribution on R2 S 1 as
an N N array of 2 2 scatter matrices. Ignoring the issue of the
biological plausibility of such an encoding scheme, we observe that
although the representation of S 1 is clearly continuous, the rect-
Correspondence to: L.R. Williams angular grid used to sample R2 is not. It follows that their com-
(e-mail: Williams@cs.unm.edu) putation is not Euclidean invariant.
3
and 120 orientation.3 There is no reason to believe gives the probability that ðx; hÞ and ðu; /Þ are distin-
that the experimental results shown in these papers guishable edges from the boundary of a single object.
would be similar if the input images were rotated by The short-time propagator:
as little as 5 . To our knowledge, until now no one Z 1
has ever commented on this problem. P1 ðx; hju; /Þ ¼ dt½1 vðtÞ Gðx; h; tju; /; 0Þ ð3Þ
0
2 A continuum formulation of the saliency problem gives the probability that ðx; hÞ and ðu; /Þ are from the
boundary of a single object but are indistinguishable. In
The following section reviews the continuum formula- both of these propagators, vð:Þ is a cut-off function with
tion of the contour completion and saliency problem as vð0Þ ¼ 0 and limt!1 vðtÞ ¼ 1:
described in Williams and Thornber (2001). ht i
1 2
vðtÞ ¼ 1 þ atan l a : ð4Þ
2 p D
2.1 Shape distribution
The cutoff function is characterized by three parameters
Mumford (1994) observed that the probability distribu- – a, l, and D. The parameter D is the scale of the edge-
tion of object boundary shapes could be modeled by a detection process. Relative to this scale, the parameter a
Fokker-Planck equation of the following form, specifies the location of the cutoff, and l specifies its
hardness.
@p @p @p r2 @ 2 p 1
¼ cos h sin h þ p ; ð1Þ
@t @x @y 2 @h2 s
where pðx; h; tÞ is the probability that a particle is located 2.3 Eigenfunctions
at position x ¼ ðx; yÞ and is moving in direction h at time
t. This partial differential equation can be viewed as a set We use the long-time propagator, P0 , to define an
of independent advection equations in x and y (the first integral linear operator, Qð:Þ, which combines three
and second terms) coupled in the h dimension by the sources of information: (1) the probability that two
diffusion equation (the third term). The advection equa- edges belong to the same object, (2) the probability
tions translate probability mass in direction, h, with unit that the two edges are distinguishable, and (3) the
speed, while the diffusion term implements a Brownian probability that the two edges exist. It is defined as
motion in direction, with diffusion parameter r. The follows:
combined effect of these three terms is that particles tend 1 1
to travel in straight lines, but over time they drift to the Qðx; hju; /Þ ¼ bðxÞ2 P0 ðx; hju; /ÞbðuÞ2 ð5Þ
left or right by an amount proportional to r2 . Finally, where the input bias function, bðxÞ, gives the probability
the effect of the fourth term is that particles decay over that an edge exists at x. Note that, as defined, bðxÞ
time, with a half-life given by the decay constant s. depends only on the position, x, and not the direction, h,
of the edge. This reflects the fact that the input to the
2.2 The propagators computation is isotropic.4
As described in Williams and Thornber (2001), the
The Green’s function, Gðx; h; t1 ju; /; t0 Þ, gives the prob- right and left eigenfunctions, sð:Þ and sð:Þ, of Qð:Þ with
ability that a particle observed at position u and the largest positive real eigenvalue, k, play a central role
direction / at time t0 will later be observed at position in the computation of saliency:
x and direction h at time t1 . It is the solution, pðx; h; t1 Þ, Z Z Z
of the Fokker-Planck initial value problem with initial ksðx; hÞ ¼ dud/Qðx; hju; /Þsðu; /Þ ð6Þ
2 1
value pðx; h; t0 Þ ¼ dðx uÞdðh /Þ where d is the Dirac Z Z ZR S
delta function. Unlike Williams and Thornber (2001), ksðx; hÞ ¼ dud/sðu; /ÞQðu; /jx; hÞ : ð7Þ
who assumed that the location of input edges could be R2 S 1
specified with arbitrary precision, we explicitly model the
limited spatial acuity of the edge-detection process. We Because Qð:Þ is invariant under the transformation
consider two edges to be indistinguishable if they are Qðx; hju; /Þ ¼ Qðu; / þ pjx; h þ pÞ ; ð8Þ
separated by a distance smaller than some scale, D. The
Green’s function is used to define two propagators. The which reverses the order and direction of its arguments,
long-time propagator: the right and left eigenfunctions are related by
Z1 sðx; hÞ ¼ sðx; h þ pÞ : ð9Þ
P0 ðx; hju; /Þ ¼ dtvðtÞ Gðx; h; tju; /; 0Þ ð2Þ
0
4
The model could easily be extended to include nonisotropic
3
Nor are we blameless in this regard. Williams and Jacobs (1997) input by making bð:Þ a function of both x and h. This would allow
used 36 discrete orientations including 0 , 60 , and 120 , and also us to model any orientation bias present in feedforward connec-
show results on a conveniently oriented Kanizsa Triangle. tions from LGN to V1.
4
2.4 Stochastic completion field input-dependent linear operator, bð:Þ. The dynamics of
the neural network are derived from the update equation
The fact that the distribution of closed contours, or the for the standard power method for computing eigen-
stochastic completion field, can be factored into a source vectors. It is useful to draw an analogy between our
field derived from sð:Þ and a sink field derived from sð:Þ is neural network for contour completion and the models
discussed in detail in Williams and Thornber (2001). The for the emergence of orientation selectivity in V1
stochastic completion field, cðu; /Þ, gives the probability described by Somers et al. (1995) and Ben-Yishai et al.
that a closed contour containing any subset of the edges (1995). See Fig. 1. First, we can identify sð:Þ with simple
exists at ðu; /Þ. It is the sum of three terms: cells in V1 and the input-bias function, bð:Þ, which
modulates sð:Þ in the numerator of the update equation,
cðu; /Þ ¼ with the feedforward excitatory connections from the
p0 ðu; /Þ
p0 ðu; /Þ þ p0 ðu; /Þp1 ðu; /Þ þ p1 ðu; /Þp0 ðu; /Þ LGN. Second, we can identify P0 ð:Þ with the intracor-
RRR tical excitatory connections, which Somers et al. hy-
k R2 S 1 dxdh sðx; hÞsðx; hÞ
pothesize are primarily responsible for the emergence of
ð10Þ orientation selectivity in V1. As in the model of Somers
et al., these connections are highly specific and target
where pm ðu; /Þ is a source field and pm ðu; /Þ is a sink mainly cells of similar orientation preference. Third, we
field: identify the denominator of the update equation with
Z Z Z
1 the orientation-nonspecific intracortical inhibitory con-
pm ðu; /Þ ¼ dxdh Pm ðu; /jx; hÞbðxÞ2 sðx; hÞ nections, which Somers et al. hypothesize keep the level
R2 S 1
Z Z Z of activity within bounds. Finally, we identify cð:Þ, the
pm ðu; /Þ ¼
1
dxdh sðx; hÞbðxÞ2 Pm ðx; hju; /Þ : stochastic completion field, with the population of cells
2
R S 1 in V2 described by von der Heydt et al. (1984).
The purpose of writing cðu; /Þ in this way is to remove
the contribution, p1 ðu; /Þ
p1 ðu; /Þ, of closed contours at 4 A discrete implementation of the continuum formulation
scales smaller than D that would otherwise dominate the
completion field. Given the above expression for the Following Zweck and Williams (2000), the continuous
completion field, it is clear that the key problem is functions comprising the state of the computation are
computing the eigenfunction, sð:Þ, of Qð:Þ with the represented as weighted sums of a finite set of shiftable-
largest positive real eigenvalue. To accomplish this, we twistable basis functions. The weights form the coeffi-
can use the well-known power method. See Golub and cient vectors for the functions. The computation we
Van Loan (1996). In this case, the power method describe is biologically plausible in the sense that all
involves repeated application of the linear operator, transformations of state are effected by linear transfor-
Qð:Þ, to the function, sð:Þ, followed by normalization: mations (or other vector parallel operations) on the
coefficient vectors.
sðmþ1Þ ðx; hÞ ¼
RRR ðmÞ
2 1 dud/Qðx; hju; /Þs ðu; /Þ
RRR R RRRS 4.1 Shiftable-twistable bases
ðmÞ ðu; /Þ
R2 S 1 R2 S 1 dxdhdud/ Qðx; hju; /Þs
ð11Þ The input and output of the above computation are
ðmþ1Þ functions defined on the continuous space, R2 S 1 , of
In the limit, as m gets very large, s ðx; hÞ converges to
the eigenfunction of Qð:Þ, with the largest positive real
eigenvalue.5 We observe that the above computation can
be considered a continuous-state, discrete-time, recur-
rent neural network.
3 A neural implementation
C
positions in the plane, R2 , and directions in the circle, S 1 . where bð:Þ is the input, cð:Þ is the output, ! is the
Tx0 ;h0
For such computations, the important symmetry is computation, and ! is the shift-twist transformation.
determined by those transformations, Tx0 ;h0 , of R2 S 1 , Correspondingly, we define a shiftable-twistable basis6 of
that perform a shift in R2 by x0 , followed by a twist in functions on R2 S 1 to be a set of functions on R2 S 1
R2 S 1 through an angle, h0 . A twist through an angle, h0 , with the property that whenever a function, f ðx; hÞ, is in
consists of two parts: (1) a rotation, Rh0 , of R2 and (2) a their span, then f ðTx0 ;h0 ðx; hÞÞ is as well for every choice
translation in S 1 , both by h0 . The symmetry Tx0 ;h0 , which is of ðx0 ; h0 Þ in R2 S 1 . As such, the notion of a shiftable-
called a shift-twist transformation, is given by the formula twistable basis on R2 S 1 generalizes that of a shiftable-
Tðx0 ;h0 Þ ðx; hÞ ¼ ðRh0 ðx x0 Þ; h h0 Þ : ð12Þ steerable basis on R2 introduced by Freeman and
Adelson (1991) and Simoncelli et al. (1992).
The relationship between continuous shift-twist trans- Shiftable-twistable bases can be constructed as fol-
formations and computations in primary visual cortex lows. Let Wðx; hÞ be a function on R2 S 1 that is peri-
has been hypothesized by Williams and Jacobs (1997) odic (with period X ) in both spatial variables, x. In
and by Bressloff et al. (2001). The continuous shift-twist analogy with the definition of a shiftable-steerable
group characterizes the symmetry of the Green’s func- function on R2 , we say that W is shiftable-twistable on
tion of the Fokker-Planck equation described by R2 S 1 if there are integers, K and M, and interpolation
Mumford (1994): functions, ak;m ðx0 ; h0 Þ, such that for each ðx0 ; h0 Þ 2
R2 S 1 , the shift-twist of W by ðx0 ; h0 Þ is a linear com-
Gðx; h; t1 ju; /; t0 Þ ¼ bination of a finite number of basic shift-twists of W by
GðR/ ðx uÞ; h /; t1 t0 j0; 0; 0Þ : ð13Þ amounts ðkD; mDh Þ, i.e., if
X
The continuous shift-twist symmetry of the Green’s WðTx0 ;h0 ðx; hÞÞ ¼ ak;m ðx0 ; h0 ÞWðTkD;mDh ðx; hÞÞ : ð14Þ
function can be understood by considering the analo- k;m
gous discrete symmetry of highest order. An idealized
Here D ¼ X =K is the basic shift amount and Dh ¼ 2p=M
model of V1 might be based on a set of hypercolumns
is the basic twist amount. The sum in Eq. (14) is taken
arranged in a hex lattice and with six discrete orientation
over all pairs of integers, k ¼ ðkx ; ky Þ, in the range
preferences (see Fig. 2). By virtue of the shift-twist
0 kx ; ky < K, and all integers, m, in the range
symmetry of the interconnection matrix, the long-range,
0 m < M. As in Simoncelli et al. (1992) the interpo-
orientation-specific, intracortical excitatory connections
lation functions, ak;m , are sinc functions.
labeled A and B have equal weight. However, because
The Gaussian-Fourier basis is the product of a shif-
this lattice is not invariant under continuous shift-twist
table-steerable basis of Gaussians in x and a Fourier
transformations, the output of such a model (unlike
series basis in h. For the experiments in this paper, the
ours) would not be Euclidean invariant. A visual
standard deviation of the Gaussian basis function,
computation, C, on R2 S 1 is called shift-twist invariant 2 2
gðxÞ ¼ D1 ekxk =2D , equals the basic shift amount, D. We
if, for all ðx0 ; h0 Þ 2 R2 S 1 , a shift-twist of the input by
regard gðxÞ as a periodic function of period, X , which is
ðx0 ; h0 Þ produces an identical shift-twist of the output.
chosen to be much larger than D, so that gðX =2; X =2Þ
This property can be depicted in the following commu-
and its derivatives are essentially zero. For each fre-
tative diagram:
quency, x, and shift amount, D (where K ¼ X =D is an
C integer), we define the Gaussian-Fourier basis functions,
bðx; hÞ ! cðx; hÞ
Wk;x , by
# Tx0 ;h0 # Tx0 ;h0
bðRh0 ðx x0 Þ; h h0 Þ !
C
cðRh0 ðx x0 Þ; h h0 Þ; Wk;x ðx; hÞ ¼ gðx kDÞeixh : ð15Þ
Zweck and Williams (2000) showed that the Gaussian-
Fourier basis is shiftable-twistable.
(i.e., the basic step in the power method) can be 4.4 The bias operator B
implemented as a discrete linear transform in a Gauss-
ian-Fourier shiftable-twistable basis: In the continuum, the bias operator multiplies the
function, sð:Þ, by the input-bias function, bð:Þ. Our aim is
sðmþ1Þ ¼ PBsðmÞ : ð18Þ to identify an equivalent linear operator in the shiftable-
twistable basis. Suppose that both sð:Þ and bð:Þ are
represented in a Gaussian basis, gk ðxÞ ¼ gðx kDÞ.
4.3 The propagation operator P Their product is:
X X
In practice, we do not explicitly represent the matrix, P. bðxÞsðxÞ ¼ sk gk ðxÞ b‘ g‘ ðxÞ ð26Þ
This is because it is quite large, and explicitly computing XX
k ‘
the above matrix-vector product using the matrix would ¼ sk b‘ gk ðxÞg‘ ðxÞ : ð27Þ
be prohibitively expensive. Instead, we compute the k ‘
matrix-vector product using the advection-diffusion-
decay operator in the Gaussian-Fourier shiftable-twist- Now, the product of two Gaussian basis functions, gk
able basis, A D, which is derived and described in and g‘ , is a Gaussian of smaller variance (with higher-
detail in Zweck and Williams (2000). The coefficient frequency content) that cannot be represented in
vector, sðmþ1Þ , is a weighted average of solutions, qðm;nÞ , the Gaussian basis, gk . Because bðxÞsðxÞ is a linear
of a Fokker-Planck initial-value problem. The average is combination of the products of pairs of Gaussian-basis
over all positive times, t ¼ nDt. That is, functions, it cannot be represented in the Gaussian
basis, either. However, we observe that the convolu-
X
1
tion of bðxÞsðxÞ and a Gaussian, hðxÞ ½bðxÞsðxÞ,
sðmþ1Þ ¼ vðnDtÞqðm;nÞ ð19Þ
where
n¼0
1 jjxjj2 =D2
where vð:Þ is the cutoff function, and the initial value for hðxÞ ¼ e ð28Þ
q is the product of the bias operator, B, and the D2 p
coefficient vector, sðmÞ , can be represented in the Gaussian basis. It follows that
ðm;0Þ ðmÞ there exists a matrix, B, such that
q ¼ Bs : ð20Þ
X
For convenience, we introduce a variable, pðm;nÞ , to hðxÞ ½bðxÞsðxÞ ¼ ½Bsk gk ðxÞ : ð29Þ
represent the partial sum of qðm;nÞ from t ¼ 0 to t ¼ nDt. k
This leads to the following update equations: The formula for the matrix, B, is derived by first
completing the square in the exponent of the product of
qðm;nþ1Þ ¼ ðA DÞqðm;nÞ ð21Þ two Gaussians to obtain:
ðm;nþ1Þ ðm;nÞ ðm;nþ1Þ
p ¼p þ vðnDtÞq ; ð22Þ pffiffiffi D D
gk ðxÞg‘ ðxÞ ¼ gð 2ðx ðk þ ‘ÞÞÞgðpffiffiffi ðk ‘ÞÞ : ð30Þ
where A is the the advection operator and D is the 2 2
diffusion-decay operator. In the shiftable-twistable
basis, the advection operator, A, is a discrete convolu- This product is then convolved with h to obtain a
tion: function, f ðxÞ, which is a shift of the Gaussian basis
function, gðxÞ. Finally, we use the shiftability formula
ðm;nþ1Þ
X ðm;nÞ X
q‘;g 2 ¼ a^‘k;gx ðDtÞ qk;x ð23Þ gðx x0 Þ ¼ ak ðx0 Þgk ðxÞ ; ð31Þ
k;x
k
with the kernel, where ak are the interpolation functions, to express f ðxÞ
Z in the Gaussian basis. The result is
1 2p
a^k;g ðDtÞ ¼ ak ðDt½cos h; sin hT Þ expðighÞ dh ð24Þ X
2p 0 Bk;‘ ¼ bi expðjji ‘jj2 =4Þak ðDði þ ‘Þ=2Þ : ð32Þ
i
where the ak are interpolation functions. Let N be the
number of Fourier-series frequencies, x, used in the Although this formula is very general, using it to
shiftable-twistable basis, and let Dh ¼ 2p=N . The diffu- compute the matrix-vector product would be prohibi-
sion-decay operator, D, is given by the diagonal matrix- tively expensive due to the nonsparseness of the matrix
vector product: B. To overcome this problem we have developed an
7
alternative representation for the bias operator in the In the first experiment, the input-bias function, bðxÞ,
Gaussian-Fourier basis. For this representation, we consists of eight Gaussian spots spaced equally on a
assume that the input-bias function is of the form circle. The positions are real valued, i.e., they do not lie
on theRgrid of basis functions. The stochastic completion
X
N
field, S 1 cðu; /Þd/, after five iterations of the power
bðxÞ ¼ gðx xj Þ ; ð33Þ
method, is shown in Fig. 3. Although the local orien-
j¼1
tations of the circle at the locations of the spots were not
where the centers, xj , of the Gaussian spots are assumed specified in the input-bias function, they are an emergent
to be far apart from each other relative to their widths. property of the completion field.
In this case, the matrix B representing the bias operator In the second experiment, the input-bias function was
can be expressed as a sum of N outer products of vectors created by adding twenty randomly positioned Gaussian
representing values of the Gaussian-basis functions g‘ spots to twenty spots on the boundary of an avocado.
and values of the sinc interpolation functions ak at the See Fig. 4 (left). The stochastic completion field com-
locations xj of the Gaussian spots: puted using 32 iterations of the power method is shown
in Fig. 4 (right).
X
N
In the last experiment, the input-bias function from
Bk;‘ ¼ g‘ ðxj Þak ðxj Þ : ð34Þ the first experiment was rotated by 45 and translated by
j¼1
half the distance between the centers of adjacent basis
Consequently, the action of the bias operator on a functions, bðR45 ðx ½D2 ; D2 T ÞÞ. See Fig. 5 (left). The sto-
coefficient vector is given by: chastic completion field is identical (up to rotation and
" # translation) to the one computed in the first experiment.
X
N X See Fig. 5 (right). Finally, we plot the estimate of the
½Bsk ¼ s‘ g‘ ðxj Þ ak ðxj Þ : ð35Þ largest positive real eigenvalue, k, as a function of m,
j¼1 ‘ the power-method iteration for both the rotated and
nonrotated input patterns. See Fig. 6. The final value of k
In this form, the bias operator is relatively inexpensive to
compute. For this reason, it was used in the experimen-
tal results described below. However, in the continuum,
this representation of the bias operator does not agree
with the representation given in Eq. 32. To understand
how the second representation is related to the first, we
now describe how it was derived. Assume that the input-
bias function
P is simply bðxÞ ¼ gðx x0 Þ and that
sðxÞ ¼ ‘ s‘ g‘ ðxÞ. Then
X X
½Bs‘ g‘ ðxÞ ¼ bðxÞsðxÞ s‘ gðx x0 Þg‘ ðxÞ ð36Þ
‘ ‘
" #
X
’ s‘ g‘ ðx0 Þ gðx x0 Þ ; ð37Þ
‘
Fig. 3. Left: The input-bias function, bðxÞ, consists of eight Gaussian
where the final equality holds exactly in the limit that spots spaced equally on a circle. The positions are real valued, i.e.,
they do not lie on
R the grid of basis functions. Right: The stochastic
gðx x0 Þ ! dðx x0 Þ, where d denotes the Dirac delta completion field, S 1 cðu; /Þ d/, computed using 192 192 92 basis
function. The second representation of the bias operator functions
in the basis then follows by expressing gðx x0 Þ in the
Gaussian basis using the shiftability formula.
5 Experimental results
be formally characterized as a frame and its dual. Re- excitatory connections within V1 were the neural
cently, Salinas and Abbott (2000) have argued that embodiment of the P0 ð:Þ operator. The fact that these
simple-cell receptive fields do not form a tight frame. If connections are orientation-specific implies that the
they are correct, then the corresponding basis functions, basis functions that represent the source field are
which are only implicitly defined, do not have the same localized in S 1 . However, our implementation uses
shape as the receptive fields. Consequently, even under harmonic signals, which have zero locality, to represent
the strong assumption of linearity, and even when the the orientation dimension. The problem of identifying a
dimensionalities of the input and the intermediate rep- basis of shiftable-twistable functions that are localized in
resentation are the same, it is a mistake to equate re- orientation and specifying the corresponding P0 ð:Þ
ceptive fields and basis functions. operator is a topic for future research.
With regard to V1 and V2, this is a moot point, since
the assumption of linearity is known to be unrealistic.
Many researchers have demonstrated that the response 7 Conclusion
of simple cells in V1 is modulated by stimuli outside of
the classical (i.e., linear) receptive field. Furthermore, the We described a neural network that enhances and
dynamics of this modulatory activity is consistent with a completes salient closed contours. Even though the
computation implemented in a recurrent network, not a computation is implemented in a discrete network, its
simple feedforward convolution of the input from LGN. output is invariant under continuous rotations and
The assumption of linearity is even less justified in V2, translations of the input.
where von der Heydt et al. (1984) have identified neu-
rons that ‘‘respond’’ to illusory contours of specific po- Acknowledgements. L.R.W. was supported in part by Los Alamos
sition and orientation. Since these neurons do not National Laboratory. J.W.Z. was supported in part by the Albu-
respond to an isolated spot, the concept of impulse querque High Performance Computing Center. We wish to thank
Jonas August and Steve Zucker for their insightful comments.
response function is not useful.
Parent P, Zucker SW (1989) Trace inference, curvature consistency Sun J, Perona P (1998) Where is the sun? Nature Neurosci 1: 183–184
and curve detection. IEEE Trans Pattern Analy Mach Intell 11: Von der Heydt R, Peterhans E, Baumgartner G (1984) Illusory
823–889 contours and cortical neuron responses. Science 224: 1260–
Salinas E, Abbott LF (2000) Do simple cells in primary visual 1262
cortex form a tight frame? Neural Comput 12(2): 313–335 Williams LR, Jacobs DW (1997) Stochastic completion fields: a
Sha’ashua A, Ullman S (1988) Structural saliency: the detection of neural model of illusory contour shape and salience. Neural
globally salient structures using a locally connected network. Comput 9(4): 837–858
In: Proceedings of the 2nd international conference on com- Williams LR, Thornber KK (2001) Orientation, scale, and dis-
puter vision. Clearwater, FL, pp 321–327 continuity as emergent properties of illusory contour shape.
Simoncelli E, Freeman W, Adelson E, Heeger D (1992) Shiftable Neural Comput 13(8): 1683–1711
multiscale transforms. IEEE Trans Inf Theory 38(2): 587–607 Yen S, Finkel L (1997) Salient contour extraction by temporal
Somers DC, Nelson SB, Sur M (1995) An emergent model of ori- binding in a cortically-based network. Adv Neural Inf Process
entation selectivity in cat visual cortical cells. J Neurosci 15: Syst 10: 915–921
5448–5465 Zweck JW, Williams LR (2000) Euclidean group invariant com-
Sompolinsky H, Shapley R (1997) New perspectives on the mech- putation of stochastic completion fields using shiftable-twist-
anisms for orientation selectivity. Curr Opinion Neurobiol 7: able functions. In: Proceedings of European conference on
514–522 computer vision (ECCV ’00), Dublin, Ireland