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Directional harmonic theory: A computational Gestalt model to account for

illusory contour and vertex formation

Article  in  Perception · February 2003

DOI: 10.1068/p5011 · Source: PubMed

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Perception, 2003, volume 32, pages 423 ^ 448

DOI:10.1068/p5011

Directional harmonic theory: A computational Gestalt

model to account for illusory contour and vertex formation

Steven Lehar
Schepens Eye Research Institute, 20 Staniford Street, Boston, MA 02114-2500, USA;
e-mail: slehar@cns.bu.edu; http://cns-alumni.bu.edu/slehar/
Received 28 August 2001, in revised form 8 January 2002; published online 29 January 2003

Abstract. Visual illusions and perceptual grouping phenomena offer an invaluable tool for probing
the computational mechanism of low-level visual processing. Some illusions, like the Kanizsa
figure, reveal illusory contours that form edges collinear with the inducing stimulus. This kind of
illusory contour has been modeled by neural network models by way of cells equipped with
elongated spatial receptive fields designed to detect and complete the collinear alignment. There
are, however, other illusory groupings which are not so easy to account for in neural network
terms. The Ehrenstein illusion exhibits an illusory contour that forms a contour orthogonal to
the stimulus instead of collinear with it. Other perceptual grouping effects reveal illusory contours
that exhibit a sharp corner or vertex, and still others take the form of vertices defined by the
intersection of three, four, or more illusory contours that meet at a point. A direct extension of
the collinear completion models to account for these phenomena tends towards a combinatorial
explosion, because it would suggest cells with specialized receptive fields configured to perform
each of those completion types, each of which would have to be replicated at every location
and every orientation across the visual field. These phenomena therefore challenge the adequacy
of the neural network approach to account for these diverse perceptual phenomena.
I have proposed elsewhere an alternative paradigm of neurocomputation in the harmonic
resonance theory (Lehar 1999, see website), whereby pattern recognition and completion are
performed by spatial standing waves across the neural substrate. The standing waves perform a
computational function analogous to that of the spatial receptive fields of the neural network
approach, except that, unlike that paradigm, a single resonance mechanism performs a function
equivalent to a whole array of spatial receptive fields of different spatial configurations and of
different orientations, and thereby avoids the combinatorial explosion inherent in the older
paradigm. The present paper presents the directional harmonic model, a more specific develop-
ment of the harmonic resonance theory, designed to account for specific perceptual grouping
phenomena. Computer simulations of the directional harmonic model show that it can account
for collinear contours as observed in the Kanizsa figure, orthogonal contours as seen in the
Ehrenstein illusion, and a number of illusory vertex percepts composed of two, three, or more
illusory contours that meet in a variety of configurations.

1 Introduction
Visual illusions and perceptual grouping phenomena offer an invaluable tool for probing
the computational mechanism of low-level visual processing. Some illusions, like the
Kanizsa figure shown in figure 1a, reveal a process of collinear illusory-contour forma-
tion, whereby the illusory contour spans the gap between stimulus edges that are both
parallel, and spatially aligned. This kind of illusory-contour formation has been modeled
by neural network models that make use of `cooperative cells' equipped with elongated
spatial receptive fields designed to detect and, by top ^ down feedback, to complete
the collinear alignment with a line of neural activation along the perceived illusory
contour (Grossberg and Mingolla 1985; Zucker et al 1989). There are, however, other
illusory groupings which are not so easy to account for in neutral network terms. For
example, the Ehrenstein illusion, shown in figure 1b, exhibits an illusory contour orthog-
onal to the stimulus line segments instead of collinear with them. Grossberg and
Mingolla (1985) account for this kind of illusory contour with a competitive interaction
between cooperative cells at any particular spatial location whose orientations differ
424 S Lehar

(a) (b) (c)

(d)
Figure 1. (a) The Kanizsa illusory triangle demonstrates illusory contour formation collinear
with the inducing stimulus edges. (b) The Ehrenstein illusion demonstrates illusory contour
formation orthogonal to the inducing stimulus edges. (c) The dot ^ triangle demonstrates illusory
contours that meet at a sharp corner or vertex. (d) A circle of dots becomes, with increased
curvature, a polygon of dots with an illusory vertex at each dot location.

by about 908. In other words, the system is hard-wired to perform both collinear and
orthogonal completion. But even if this model successfully accounts for these phenom-
ena (an assertion which is not entirely unproblematic), it is difficult to imagine how
this concept could possibly generalize to perceptual completion through sharp angles
or vertices, as seen, for example, in the dot ^ triangle of figure 1c, for this case is exactly
intermediate between collinear and orthogonal completion. But the problem is still more
serious, because there are a number of illusory grouping phenomena which involve
three, four, or more illusory contours that meet at a perceived vertex, as discussed
below. Although these different forms of illusory-contour formation exhibit distinct
characteristics, there is compelling evidence that they are, nevertheless, different mani-
festations of the same underlying mechanism. For example, Kanizsa (1987) observes
that the collinear grouping percept due to a circle of dots gives way to a polygonal
percept when the number of dots in the circle is reduced, as shown in figure 1d. In
other words, the collinear grouping contour passing through each dot gives way to a
vertex grouping percept, as if the collinear contour kinks like a drinking straw that is
bent beyond its elastic limit. The key factor here concerns the angular deviation of
the illusory contour through each dot rather than the size of the circle of dots, the size
of the circle being varied merely to maintain the same spacing between dots. This
phenomenon appears to be related to a similar abrupt transition observed in the per-
ception of curvature (Wilson and Richards 1989). This calls into question the adequacy
of neural network models to account for these various perceptual grouping phenom-
ena, because a neural network model to account for all of these diverse phenomena
would suggest receptive field configurations specialized to perform collinear, orthogonal,
and sharp vertex completion, as well as for illusory vertices composed of three, four,
or more illusory contours that meet at a point, each of which would require a set of
hard-wired receptive fields of the appropriate configuration. In fact, the spatial pattern
of the neural receptive field is not different in principle from a template model, a
concept whose limitations are well known.
Elsewhere I have proposed a harmonic resonance theory (Lehar 1994a, 1994b, 1999,
2002) as an alternative to the neural network paradigm. I propose that the spatial
patterns of illusory grouping phenomena are mediated not by hard-wired spatial recep-
tive fields, but by standing waves of electrochemical oscillations in the neural substrate.
A standing wave offers a much more flexible and adaptive computational mechanism
than the spatial receptive fields of neural network theory. Unlike a neural receptive field,
Directional harmonic theory 425

a standing-wave pattern is not hard-wired to a particular spatial configuration, but can

adapt flexibly to the input stimulus like an elastic template that can kink and fold into a
variety of different configurations, all by way of a single computational mechanism.
The merits of the harmonic resonance paradigm as a principle of perceptual computation
are particularly clear in the case of the more complex illusory grouping phenomena
presented below.
1.1 Dot grouping phenomena
Figure 2a shows a pattern of dots that are grouped in pairs, ie an illusory grouping
line is observed to connect the two dots of the pair as suggested schematically by the
gray shading in the magnified depiction on the right of the figure. In other words, each
dot projects a single illusory contour extending out in one direction only. A different
pattern is observed in figure 2b which shows a collinear grouping of dots in columns,
each dot being connected by an illusory grouping contour that extends out from that
dot in opposite directions, as shown schematically to the right of the figure. Figure 2c
shows a hexagonal grouping pattern in which each dot defines the center of a three-
way vertex, as suggested schematically to the right in the figure. Finally figure 2d
depicts a grid-like percept in which each dot defines the center of a four-way vertex,
as suggested schematically to the right in the figure.

(a)

(b)

(c)

(d)
Figure 2. Amodal illusory contours, or grouping percepts, define vertices composed of (a) one,
(b) two, (c) three, and (d) four intersecting illusory contours, as suggested schematically to the right,
where the gray lines represent the perceived contours in the figures to the left.

What is interesting in these perceptual phenomena is not so much the perceived

grouping that occurs between neighboring dots, ie a grouping by the Gestalt law of
proximity, but there is a more subtle and complex inhibitory effect whereby a nearer
grouping is seen to suppress a more distant grouping. For example, the horizontal
separation between columns of dots in figure 2b is the same as that in the grid pattern
of figure 2d. But the closer vertical spacing within each column in figure 2b appears
to suppress the horizontal grouping between those dots. Similarly, the vertical and
horizontal grid grouping percept of figure 2d appears to suppress an equally valid
diagonal dot grouping, because each dot is located at the intersection of two diagonal
426 S Lehar

rows of dots as well as on vertical and horizontal columns and rows of dots. But since
the vertical and horizontal grouping has a closer spacing than the diagonal group-
ing, the diagonal grouping percept is entirely suppressed in this dot pattern. Similarly
the hexagonal grouping percept shown in figure 2c suppresses an equally present verti-
cal and horizontal grouping percept, because each dot is located on the intersection
of a vertical column and a horizontal row of dots in the stimulus, although this pattern
is not apparent in the grouping percept. These complex spatial interactions between
different grouping patterns offer a detailed manifestation of the specific computational
interactions in perception, that goes well beyond the simplistic collinear and orthog-
onal grouping phenomena which are the usual focus of psychophysical studies. It is
this secondary subtle pattern of inhibitory effects which provides the principal evidence
for the directional harmonic model.
1.2 Line-segment grouping phenomena
A similar parametric variation between different perceptual grouping patterns can be
seen in patterns composed of line segments shown in figure 3. For example, the lines in
figure 3a group into columns by collinearity, ie the illusory contour forms parallel to
the inducing line segments, as suggested schematically to the right in figure 3a. With
a closer horizontal spacing, however, the percept becomes one of an orthogonal group-
ing, as shown in figure 3b, and as suggested schematically to the right in the figure.
This orthogonal grouping is similar in principle to the Ehrenstein illusion of figure 1b.
Again, it is interesting that the closer horizontal spacing seems to suppress the alter-
native vertical grouping percept, and vice-versa. A third diagonal grouping percept can
also be obtained with the proper arrangement of line segments, as shown in figure 3c,

(a)

(b)

(c)
Figure 3. Amodal illusory contours that form in (a) collinear, (b) orthogonal, and (c) diagonal
configurations with respect to their inducing line segment stimuli, as suggested schematically to
the right, where the gray lines represent the perceived contours in the figures to the left.
Directional harmonic theory 427

and as suggested schematically to the right in the figure. This percept is considerably
less salient than the collinear and orthogonal grouping percepts, and is complicated
by the fact that it is not entirely clear whether the grouping lines connect adjacent
line endings directly, as suggested schematically to the right in the figure, or whether
diagonal rows of line segments form intersecting diagonal `streets', ie with longer
grouping lines that extend from the top of one line segment to the top of the next and
on to the top of the next, rather than from the top of one line ending to the bottom
of the next. This illusion is further complicated by the fact that the percept is some-
what bistable or rivalrous between a percept of parallel diagonal streets from lower
left to upper right, in competition with diagonal streets from upper left to lower right.
However, there is clearly a diagonal component to the percept that is clearly distinct
from the collinear and orthogonal percepts of figures 3a and 3b, and this percept
appears to involve a completion by illusory vertex formation with a Y vertex at the tip
of each line segment.
The grouping percepts in figures 2 and 3 are primarily of an amodal nature,
although there is perhaps also a faint modal or surface brightness component to them.
But the principal focus of the present analysis is on the pattern of amodal grouping
observed in these stimuli, regardless of whether or not those amodal contours also
promote a corresponding surface brightness percept. The shaded grouping lines shown
schematically to the right in figures 2 and 3 therefore represent the amodal component
of the grouping percept, and therefore these patterns of gray lines represent the amodal
output image that should be produced by an adequate computational model of these
perceptual grouping phenomena.

2 Harmonic resonance theory: An alternative paradigm of neurocomputation

The harmonic resonance theory (Lehar 1994a, 1994b, 1999, 2002) offers a computational
paradigm with the holistic global properties identified by Gestalt theory. The most
remarkable property of harmonic resonance is the sheer number of different unique
patterns that can be obtained in even the simplest resonating system. A pioneering
study of more complex standing-wave patterns was presented by Chladni (1787) who
demonstrated the resonant patterns produced by a vibrating steel plate. The technique
introduced by Chladni was to sprinkle sand on top of the plate, and then to set the
plate into vibration by bowing with a violin bow. The vibration of the plate causes the sand
to dance about randomly except at the nodes of vibration where the sand accumulates,
thereby revealing the spatial pattern of nodes. This technique was refined by Waller (1961)
who used a piece of dry ice pressed against the plate, where the escaping gas due to the
sublimation of the ice sets the plate into resonance, resulting in a high-pitched squeal
as the plate vibrates. Figure 4 (adapted from Waller 1961, page 69) shows some of the
patterns that can be obtained by vibrating a square steel plate clamped at its midpoint.
The lines in the figure represent the patterns of nodes obtained by vibration at various
harmonic modes of the plate, each node forming the boundary between portions
of the plate moving in opposite directions, ie during the first half-cycle alternate seg-
ments deflect upward while neighboring segments deflect downward, and these motions
reverse during the second half-cycle of the oscillation. The different patterns seen in
figure 4 can be obtained by touching the plate at a selected point while bowing at the
periphery of the plate, which forms a node of oscillation at the damped location, as well
as at the clamped center point of the plate. The plate emits an acoustical tone when bowed
in this manner, and each of the patterns shown in figure 4 corresponds to a unique
temporal frequency, or musical pitch, the lowest tones being produced by the patterns
with fewer large segments shown at the upper-left of figure 4, while higher tones are
produced by the higher harmonics depicted towards the lower-right in the figure.
428 S Lehar

0 1 2 3 4 5 6 7
0

Figure 4. Chladni figures for a square steel plate (adapted from Waller 1961) demonstrate the
fantastic variety of standing-wave patterns that can arise from a simple resonating system. A square
steel plate is clamped at its midpoint and sprinkled with sand. It is then set into vibration
either by bowing with a violin bow, or by pressing dry ice against it. The resultant standing-
wave patterns are revealed by the sand that collects at the nodes of the oscillation where the
vibration is minimal.

The higher harmonics represent higher energies of vibration, and are achieved by damp-
ing closer to the central clamp point, as well as by more vigorous bowing.
The utility of standing-wave patterns as a representation of spatial form is demon-
strated by the fact that nature makes use of a resonance representation in another
unrelated aspect of biological function, that of embryological morphogenesis, or the
development of spatial structure in the embryo. After the initial cell divisions following
fertilization, the embryo develops into an ellipsoid of essentially undifferentiated tissue.
Then, at some critical point, a periodic banded pattern is seen to emerge, as revealed by
appropriate staining techniques, shown in figure 5a. This pattern indicates an alternat-
ing pattern of concentration of morphogensöchemicals that permanently mark the
underlying tissue for future development. This pattern is sustained despite the fact that
the morphogens are free to diffuse through the embryo. The mechanism behind the
emergence of this periodic pattern is a chemical harmonic resonance known as reaction
Directional harmonic theory 429

(a)

(b)

(c)
Figure 5. (a) A periodic banded pattern revealed by chemical staining emerges in a developing
embryo, due to a chemical harmonic resonance whose standing waves mark the embryonic tissue
for future growth. (b) This chemical harmonic resonance has been identified as the mechanism
behind the formation of patterns in animal skins, as well as for the periodicity of the vertebrae
of vertebrates, the bilateral symmetry of the body plan, as well as the periodicity of the bones
in the limbs and fingers. (c) Murray shows the connection between chemical and vibrational
standing waves by replicating the patterns of leopard spots and zebra stripes in the standing-
wave resonances in a vibrating steel sheet cut in the form of an animal skin.

diffusion (Turing 1952; Prigogine and Nicolis 1967; Winfree 1974; Welsh et al 1983), in
which a continuous circular chemical reaction produces periodic patterns of chemical
concentration in a manner that is analogous to the periodic patterns of a resonating
steel plate. The chemical harmonic resonance in the embryo can thereby define a
spatial addressing scheme that identifies local cells in the embryonic tissue as belonging
to one or another part of the global pattern in the embryo by way of the relative
concentration of certain morphogens. The fact that nature employs a standing-wave
representation in this other unrelated biological function offers a proof that harmonic
resonance both can and does serve as a spatial representation in biological systems,
and that representation happens to exhibit the same holistic Gestalt properties that
have been identified as prominent properties of perception and behavior.
2.1 Resonance in the brain
Oscillations and temporal resonances are familiar enough in neural systems and are
observed at every scale, from long-period circadian rhythms, to the medium-period
rhythmic movements of limbs, all the way to the very rapid rhythmic spiking of the
single cell, or the synchronized spiking of groups of cells. Harmonic resonance is also
observed in single-celled organisms like the paramecium in the rhythmic beating of
flagella in synchronized travelling waves. Similar waves are observed in multicellular
 430 S Lehar

invertebrates, such as the synchronized wave-like swimming movements of the hydra

and the jellyfish, whose decentralized nervous systems consist of a distributed network
of largely undifferentiated cells. The muscle of the heart provides perhaps the clearest
example of synchronized oscillation, for the individual cells of the cardiac muscle are
each independent oscillators that pulse at their own rhythm when separated from the rest
of the tissue in vitro. However, when connected to other cells, they synchronize with each
other to define a single coupled oscillator. The fact that such unstructured neural
architectures can give rise to such structured behavior suggests a level of computa-
tional organization below that of the switching and gating functions of the chemical
synapse. The idea of oscillations in neural systems is not new. However, the proposal
advanced here is that nature makes use of such natural resonances not only to define
rhythmic patterns in space and time, but also to define static spatial patterns in the
form of electrical standing waves, for the purpose that is commonly ascribed to spatial
receptive fields. There is plenty of neurophysiological evidence which has accumulated
over the last few decades suggestive of harmonic resonances in the brain (Gerard and
Libet 1940; Bremer 1953; Eckhorn et al 1988; Sompolinsky et al 1990; Murthy and Fetz
1992; Hashemiyoon and Chapin 1993; Nicolelis et al 1995). However, it has been hard to
interpret the significance of that evidence in the absence of a paradigmatic framework
to suggest what function that resonance might serve in perception. I will show that, as
a paradigm for defining spatial pattern, the standing wave offers a great deal more
flexibility and adaptiveness to local conditions than the alternative receptive-field model,
and that a single resonating system can replace a whole array of hard-wired receptive
fields in a conventional neural model.
2.2 Invariance in harmonic resonance
One of the most interesting aspects of harmonic resonance as a representational
principle in the brain is that it exhibits certain invariances which are also characteristic
of perception (Lehar 1994a, 1994b, 1999, 2002). Figure 6 shows the Chladni figures for
a circular steel plate. This system exhibits two kinds of periodicity: a radial periodicity
in the form of concentric rings, and a circumferential or directional periodicity in the
Number of diameters
0 1 2 3 4

1
Number of circles

Figure 6. Chladni figures for circular plate, sorted by number of diameters and circles in
each pattern. These patterns can appear at any orientation on the plate. Each distinct pattern
has a unique vibration frequency. The vibration frequency therefore offers a rotation-invariant
representation of the pattern present on the plate.
Directional harmonic theory 431

form of radial lines, and these two types of periodicity appear in a variety of combina-
tions. However, owing to the circular symmetry of the plate, each of these patterns
can actually appear on the plate at any orientation. This is a very powerful feature,
because, if a standing-wave pattern does indeed function as a spatial template in the
brain, then any one of these patterns of standing waves corresponds not only to a
single template in an equivalent neural network model, but to a whole array of them,
ie with each pattern replicated at every possible orientation. Given that all of the
different patterns in figure 6 are produced by a single mechanism, this one circular
plate, and its various standing-wave patterns represent the computational equivalent of
a whole array of different spatial templates in a neural network model, each one
replicated at every possible orientation. It is this invariance feature of a harmonic
resonance representation that offers an escape from the combinatorial problem inherent
in the neural network paradigm. Furthermore, not only does the circular Chladni plate
represent a whole array of equivalent neural receptive fields, but also the cooperative
or competitive interactions between them, because the various harmonics of the
plate interact with one another in lawful ways, and these interactions make specific
predictions about the behavior of a harmonic resonance model in response to certain
patterns of input.

3 Directional harmonic model

The phenomenon of harmonic resonance is immensely complex, involving parallel
interactions in all directions simultaneously through a homogeneous continuum in a
manner that defies complete mathematical characterization or accurate numerical
simulation in all but its simplest aspects. That very complexity, however, is exactly why
harmonic resonance holds such great potential as a principle of computation and
representation in the brain. The focus of this paper will be restricted to a single aspect
of harmonic resonance: the tendency for standing waves to form patterns of circumfer-
ential, or directional periodicity, like the patterns of radial node lines seen in figure 6,
as suggested originally by Lehar (1994a, 1994b). For the dot grouping patterns presented
in figure 2 indicate a periodic basis set of different vertex types, expressed in terms of
directional periodicity, which are suggestive of these patterns of standing waves. As in the
case of a Fourier representation, any pattern of vertices can be represented in a directional
harmonic code to arbitrary precision, by the appropriate combinations of harmonic
coefficients. However, in a physical system, the higher-order terms require higher vibra-
tional energies, as is the case for the Chladni figures. A physical harmonic-resonance
representation would therefore necessarily be band-limited to the lower harmonics,
with a cut-off at some highest harmonic of directional periodicity. This low-pass cut-off
introduces a certain granularity or quantization in the representation, limiting the
complexity of the kind of vertex completion patterns to some finite set of low-order
primitives. In fact, it is this granularity in the directional harmonic code which
accounts for the geometric regularity of the illusory grouping percepts observed in the
different dot grouping patterns, as will be shown below.
The dot grouping patterns observed in figure 2 can be explained by a system that
promotes local standing-wave patterns at every dot location in the feedback layer, in
response to the pattern of influence felt from neighboring dots in adjacent regions.
Initially, each dot stimulates a point of activation at the corresponding location in the
feedback layer, and that activation propagates radially outward by passive diffusion in all
directions from each point. The diffusing activation from neighboring points of activation
in turn impinges back on the original point from different directions, and the reciprocal
exchange of energy back and forth between these active points across the feedback layer
promotes the emergence of a pattern of standing waves of directional harmonic resonance
at each active point as described below. Although harmonic resonance is a dynamic
432 S Lehar

process that proceeds to equilibrium, a simplified static model of the process is sufficient
to account for many of the observed grouping effects. This is analogous to the heat
equation which describes the dynamic propagation of heat along a conductor from a
localized source. Given a regular rate of heat loss along the conductor, the heat equation
can be solved at equilibrium to produce a temperature gradient along the conductor
declining with distance from the localized source, as a static model of the equilibrium
state of a dynamic process. Similarly, the pattern of activation in the feedback layer
of the directional harmonic model due to the presence of stimulus dots is assumed to
produce at equilibrium a static gradient of activation, declining outward from each
stimulus point with a Gaussian profile, as a static approximation to the equilibrium
state of a dynamic diffusion process. The patterns of standing waves of directional
periodicity are then computed at each stimulus dot location in response to this static
input field from adjacent dots as described below.
For clarity, this calculation is divided into two stages: an input stage and a resonance
stage. The input signal at each dot location is a circular signal, somewhat like a trace
on the scope of a radar that scans the horizon in a circular sweep, producing peaks
in every direction in which other dots are detected from that location. For example,
figure 7a shows a pattern of dots around a central dot (circled), and figure 7b shows
the circular input signal Iy at that central dot for every direction y from that dot,
expressed in degrees clockwise from the vertical. The neighboring dot in the 12 o'clock
direction in figure 7a produces a peak in the input response at 12 o'clock, or 3608,
as shown in figure 7b, and the other dots produce similar peaks in the corresponding
directions. The magnitude of the input signal fades as a Gaussian function of radial
distance r between points, owing to the passive diffusion, and this fading is modeled
by the Gaussian function
 
1 r2
gr …r† ˆ exp ÿ 2 , (1)
2psr 2sr
where sr is the standard deviation of the radial Gaussian function. This spatial decay
explains why the dot at the 9 o'clock direction in figure 7a produces a smaller peak
in the input signal in the 9 o'clock direction in figure 7b, because of the larger distance
to that dot. The larger dashed circle shown in figure 7a depicts the radius corre-
sponding to two standard deviations (2sr ) of the radial Gaussian input function used

‡1
0
ÿ1

Iy
(a) (b)
Figure 7. Computer simulation of the circular input signal at a central dot location due to the
presence of adjacent dots. (a) A pattern of dots around a central dot (circled). The larger dotted
circle indicates a radial distance which is twice the standard deviation of the radial Gaussian term.
This pattern of input dots produces (b) a circular input function at the central dot location, showing
how each adjacent dot produces a positive peak in the corresponding direction, the magnitude
of the peaks being modulated by the distance from the central dot.
Directional harmonic theory 433

in these simulations. If the activation due to a dot were confined to a singular point,
the input peak due to that dot would actually appear as an impulse function, ie a
singularity in orientation. But the passive diffusion through the feedback layer would
be expected to spread that singular peak somewhat across neighboring directions. In
the simulations, therefore, the input signal Iy was modulated by an angular Gaussian
function of the form
 
1 r2
gy …y† ˆ exp ÿ 2 , (2)
2psy 2sy
ie this is the same Gaussian function as used in the radial Gaussian term, except that
it operates in the angular dimension, with a standard deviation sy of 2p60:05 or 188,
in order to spread that impulse response into a more manageable peak of finite size,
as seen in this plot. Therefore the equation for the input signal due to one neighboring
dot at a bearing of a degrees from the central dot is given by
Iy ˆ gr …r†gy …y ÿ a† . (3)
The input signal is additive in each direction, so the three dots shown in the
3 o'clock direction in figure 7a together produce a stronger peak in figure 7b than any
one of them would produce by itself. The three dots near the 6 o'clock direction, on
the other hand, are each in slightly different directions, and therefore they produce
three individual input peaks through the 6 o'clock direction as seen in figure 7b. There-
fore the full equation for the input signal Iy due to a set of n neighboring dots di ,
i ˆ 1 ... n, at distances ri , and at angular bearings of ai , is given by
X n
Iy ˆ gr …ri †gy …y ÿ ai † . (4)
i ˆ1

Although the input signal is plotted only for the central dot in figure 7a, a similar
input signal is computed at every dot location in the simulations presented below.
This circular input signal is then used to compute the circular harmonic resonance
response at each dot location, as described below.
Figure 8a depicts a circular harmonic series of directional periodicity, whose nodes,
or stationary points (depicted as radial lines), represent the various edges that meet at
the vertex. The first harmonic of directional periodicity exhibits a single node extending
outward from the center in one direction. This harmonic corresponds to an end-stop
feature, or unilateral vertex, as seen in the dot grouping pattern of figure 2a. The second
harmonic exhibits two nodes separated 1808, which corresponds to a collinear vertex,
or collinear grouping percept, as seen in figure 2b. The third harmonic represents a
three-way or Y vertex composed of three edges that meet at 1208 as seen in figure 2c,
and the fourth harmonic represents a ‡ or X vertex with edges separated by 908
as seen in figure 2d. There is also a zeroth harmonic, like the DC term in a Fourier
code, which represents the energy across all directional frequencies simultaneously.
The zeroth harmonic corresponds to a vertex composed of edges extending in all
directions simultaneously, which, in the limit, is essentially equivalent to no edges at
all. Figure 8b shows the amplitude function Ay of each of these harmonics plotted as
a pattern of nodes and antinodes around the circle from y ˆ 0 to 2p, ie the height of
the plot represents the amplitude of the vibration as a function of angle through the
circle, which is given by
Ayh ˆ j sin…12 yh†j (5)
for harmonics h ˆ 1 to 4. Actually this figure shows a double plot, with upper and
lower traces showing the positive and negative of the amplitude, representing a vibra-
tion alternately upward and downward from zero, like the pattern of vibration of a
 434 S Lehar

standing wave amplitude function nodal pattern circular plot

‡1
‡1 ‡1 0
0 ÿ1
0 0
ÿ1 ÿ1

‡1 ‡1
1 0 0
ÿ1 ÿ1
Harmonic, h

‡1 ‡1
2 0 0
ÿ1 ÿ1

‡1 ‡1
3 0 0
ÿ1 ÿ1
‡1 ‡1
0 0
4
ÿ1 ÿ1

0 y 2p 0 y 2p
Ay ˆ j sin…12 yh†j Ny ˆ c ÿ j sin…12 yh†j
(a) (b) (c) (d)
Figure 8. Directional harmonic representation. (a) Various patterns of nodes on a circular plate
corresponding to the different harmonics of directional periodicity of the plate. The black lines
represent the nodes, or stationary points of the standing wave, which in turn correspond to various
configurations of edges that meet at the center, to define a sequence of vertex types. (b) The
amplitude function, or variation of the amplitude of vibration as a function of angle around
the circular plate. (c) The nodal pattern, or complement of the amplitude function, to produce
positive peaks in place of the nodes seen in the amplitude function. (d) A circular plot of the
nodal pattern, where the inner circle represents the value ÿ1, the outer circle represents ‡1, and
the middle circle represents zero.

guitar string, to emphasize that the phase of the vibration is irrelevant, and what is
significant is the pattern of nodes and antinodes. Since it is the nodes of the vibration
which represent perceived edges or grouping percepts in the model, a more convenient
form to express these waveforms mathematically is as nodal functions Ny which are
given by
Nyh ˆ 1 ÿ j sin…12 yh†j , (6)
as shown in figure 8c. In other words, these functions are computed by subtracting the
waveforms in figure 8b from unity, because this encodes the features, ie the edges, as
positive values rather than as the absence of positive values. The positive peaks in these
waveforms now represent the patterns of perceived edges or grouping percepts as shown
in figure 8a. An offset value c was added to these nodal functions in order to shift them
half-way into the negative region as shown in the plot, with the offset value chosen so
as to make the nodal functions sum to zero, ie to produce equal areas under positive
and negative regions of the curve. This was done so that, when used as convolution
filters, they do not impose a bias on the output. The normalized nodal functions are
given by
Nyh ˆ c ÿ j sin…12 yh†j , (7)
with c ˆ 1:63662. Figure 8d shows these nodal waveforms on a circular plot, whose
outer ring represents the value ‡1, the inner ring represents the value ÿ1, and the
middle ring represents the value zero.
Directional harmonic theory 435

The circular harmonic response Ryh to the input signal at each dot location is then
computed by a circular convolution of the circular input signal Iy with each of these
circular harmonic nodal filters Nyh in turn, for each harmonic h ˆ 1 to 4, as given by
… 2p
Ryh ˆ I…y‡r† Nrh , (8)
r

where (y ‡ r) is computed modulo 2p to wrap around the full circle. This produces a
set of harmonic responses Ryh to the input, one for each harmonic h, each response
being a circular function through y ˆ 0 to 2p, which represents the response to the
input for that particular harmonic. As is the case with any convolution, the magnitude
of each directional harmonic response is a function of the similarity, or degree of
match between the functional form of that particular harmonic and the pattern of the
input. For example, an input function I with a single peak extending in one direction
will produce a strong response R 1 to the first-harmonic filter N 1, and the peak of
that response function will be aligned with the peak in the input, while an input
function with two peaks separated by 1808 will produce a strong response R 2 to the
second-harmonic filter N 2, etc. The four harmonic responses interact with each other
by constructive and destructive interference as calculated by summation, producing a
total harmonic response Ry across all four harmonics which is computed as
X h
Ry ˆ Ry (9)
h

for h ˆ 1 to 4. In other words, wherever positive peaks from different harmonics coincide,
they summate by constructive interference to produce a larger positive peak, whereas
positive and negative peaks from different harmonic responses cancel each other by
destructive interference to produce the total or resultant harmonic response. It is this total
response to all harmonics of directional periodicity which corresponds to the predicted
perceptual grouping at each dot location in response to the presence of adjacent dots.
3.1 Dot pattern grouping simulations
I will now present computer simulations of very simple dot stimuli composed of only two
or three dots, to demonstrate how the harmonic response is calculated for these simple
cases, before proceeding to more interesting cases involving more complex patterns
of stimulus dots. Figure 9 demonstrates the computation of the circular harmonic
individual harmonic
responses
hˆ1
hˆ2 total harmonic
hˆ3
dots stimulus input signal hˆ4 response
08
3308 308

3008 608

2708 908

2408 1208

2108 1508
Iy 1808 Ry
harmonic Ryh
(a) (b) (c) (d)
Figure 9. Harmonic response to a single vertically adjacent dot (a) is computed by a circular
convolution of the circular input signal (b) with a set of circular nodal functions. This produces
a set of circular harmonic-response functions (c). The final perceptual grouping is computed as a
sum of these response functions, as shown in (d), where the positive portion (shaded) represents
the actual grouping percept.
436 S Lehar

response at a central dot location in response to a single neighboring dot in the

12 o'clock direction, as shown in figure 9a. The input signal Iy due to that dot exhibits
a single peak in the 12 o'clock direction, as shown in figure 9b. Each of the filters Nyh
of figure 8d is convolved with the input signal Iy to produce the set of responses Ryh
shown in figure 9c as defined in equation (8). The first harmonic Ry1 (solid-line plot)
produces a positive response in the 12 o'clock direction, with a sharp positive peak at
08, and a broad negative trough through 1808. The second harmonic Ry2 (dashed-line
plot) produces a double-peaked response, with sharp positive peaks at 08 and 1808, and
broad negative troughs through 908 and 2708. The third harmonic Ry3 (dash-dot-line
plot) produces a three-peaked response, with positive peaks at 08, 1208, and 2408, and
negative troughs in between, and the forth harmonic Ry4 (dotted-line plot) produces
four positive peaks at 08, 908, 1808, and 2708, with negative troughs at the diagonals.
The harmonic response to this single-peaked input signal is similar to the impulse
response, or point-spread function of the system, which is why the response of each
harmonic to this particular input is virtually identical to the shape of the waveform of
the harmonic itself. The total harmonic response to this input, Ry , is then calculated
by summing all of the four harmonic responses at every direction as defined in equa-
tion (9), which results in the combined harmonic response shown in figure 9d. The
positive portion of this response (shaded) points upward in the direction of the input
dot, and this response in turn represents a grouping percept extending upward from the
lower dot towards the upper dot. By symmetry, the harmonic response at the upper dot
(not plotted) would be identical, except rotated by 1808, ie with a grouping percept
extending downward towards the first dot. If the calculation were extended to include
higher harmonics, the irregular response profile of figure 9d would become progres-
sively smoother in the negative portion and sharper in the positive peak, eventually
matching the shape of the input function itself. In other words, the irregular pattern of
peaks in the negative range in figure 9d reflects the quantization due to the fact that
only the lowest four harmonics were computed in the simulation.
Figure 10 introduces a spatial plotting convention to give a more intuitive depiction
of the perceptual grouping predicted by the model, with input and harmonic response
functions displayed for every dot in the stimulus rather than only for one central dot.
Figure 10a shows two vertically adjacent dots, as before. Figure 10b shows the input
signal at each dot, plotted as before, but this time overlaid on the spatial plot of that
same input signal. In the spatial plot, the magnitude of the circular input signal at
each dot is depicted as a gray shading extending radially outward from the dot, the
darkness of the gray shading representing the magnitude of the input signal in each
direction. The shading fades with distance from the dot by the same Gaussian function
as that used in computing the input signal for that dot. For example, the lower dot in
figure 10b has a strong peak in its input function at the 12 o'clock direction, and this
is depicted in the spatial shading convention as a region of dark shading projecting
from the dot in the 12 o'clock direction, and fading with distance from the dot. The
upper dot exhibits a similar input signal projecting downward in the 6 o'clock direc-
tion. Figures 10d, 10e, and 10f present the same data as in figures 10a, 10b, and 10c,
except this time showing only the spatial shaded plot without the circular plot overlay.
The gray shading in the input plot of figures 10b and 10e suggests the pattern of
perceptual grouping which would be predicted by a grouping-by-proximity model, in
which the strength of grouping between any pair of dots is a simple Gaussian function
of the distance between them. Figures 10c and 10f show the harmonic response function
computed for each dot as above, and displayed with the spatial shading convention.
Since it is only the positive values of the harmonic response function which correspond
to predicted perceptual grouping, only positive values are plotted in the spatial plot.
It is in this plot that the subtle and interesting predictions of the directional harmonic
Directional harmonic theory 437

dots stimulus input signal harmonic response

(a) (b) (c)

(d) (e) (f )
Figure 10. A spatial plotting convention to give a more intuitive depiction of the depicted percep-
tual grouping, showing the harmonic responses for all dots in the stimulus simultaneously.
(a) A pattern of two adjacent dots. (b) The input signal at each dot location due to the pres-
ence of the other dot, produces a peak in the input plot in the direction of the other dot, and
that peak is displayed both as a circular plot, and as a gray radial shading. (c) The harmonic
response for each dot in the stimulus, again plotted both as a circular plot, and as a gray shad-
ing in the direction of the positive peaks of the plots. In this simple case the harmonic response
is very similar to the input signal. (d), (e), and (f) The same as plots (a), (b), and (c) except this
time showing only the gray shading, without the circular plot overlay. The harmonic response
shown in (f ) represents the predicted grouping percept for this configuration of dots, in this
case predicting a first-harmonic, or end-stop feature grouping at each dot location.

model manifest themselves, although in this simple case there is no significant difference
between the prediction of the harmonic resonance model and a simple grouping-by-
proximity model.
Figure 11 depicts a slightly more complex stimulus, with three dots in a vertical
line. Figure 11a depicts the stimulus dot pattern relative to the central dot (circles), and
figure 11b depicts the input response at the central dot. Figure 11c depicts the response
of the first four harmonics of directional periodicity to this input pattern. In this case
the response is dominated by the second harmonic, with positive peaks in the 6 and
12 o'clock directions. There is a weaker response of the fourth harmonic, with positive
peaks at 6 and 12 o'clock, but the absence of dots in the 3 and 9 o'clock directions keeps
this response weaker than the second-harmonic response. The third harmonic produces
only a very weak response, because its positive and negative peaks are separated by 1808,
so when the positive lobe of the filter is aligned with one input peak, the negative lobe is
aligned with the other input peak. The same is true also for the first harmonic, which
also produces a very weak response. Figure 11d depicts the total harmonic response,
which produces positive peaks in the 6 and 12 o'clock directions due to both the second
and fourth harmonics, and small peaks in the 3 and 9 o'clock directions due to
the fourth harmonic, but since these peaks are opposed by negative peaks in the
second harmonic, the total harmonic response remains negative in those directions.
The grouping percept predicted by the directional harmonic model in response to this
stimulus therefore is a second harmonic or collinear grouping, with grouping lines
projecting upward and downward towards the adjacent dots. Figures 11e, 11f, and 11g
depict the spatial plot for this same stimulus. Note that the harmonic responses at the
438 S Lehar

individual harmonic total harmonic

dots stimulus input signal responses response

(a) (b) (c) (d)

dots stimulus input signal harmonic response

(e) (f ) (g)
Figure 11. (a) through (d) Computer simulation of the harmonic response of a dot flanked by two
neighboring dots in a straight line. (a) The pattern of dots in the stimulus. (b) The input response
at the central dot, showing peaks at 6 and 12 o'clock. (c) The individual harmonic responses to
this input at the central dot location, showing a strong second-harmonic response, and a
weaker fourth-harmonic response, and still weaker first- and third-harmonic responses. (d) The
total harmonic response for this stimulus at the central dot location, showing positive peaks
at 6 and 12 o'clock, dominated by the second-harmonic or collinear grouping percept at the
central dot. (e) through (g) A computer simulation of the grouping between three dots in a vertical
column, showing (e) the input dot stimulus, (f ) the input function, and (g) the total harmonic
response at each dot location with the use of the spatial plotting convention. This simulation
therefore predicts a collinear grouping through the middle of this columns of dots, with end-stop
groupings at the top and bottom dots.

upper and lower dots are dominated by the first harmonic, ie with an illusory grouping
line projecting downward and upward, respectively, towards the central dot, similar to
the grouping seen in figure 10. Again, in this simple case the prediction of the direc-
tional harmonic model is not very different from the input signal itself, which is the
kind of grouping which would be predicted by a simple grouping-by-proximity model.
Figure 12 shows the computer simulations for all of the dot grouping patterns of
figure 2. The three columns in figure 12 represent the dot patterns used in the simula-
tion, the input signal computed for each dot, and the directional harmonic response
due to that input signal computed at every dot location. Figure 12a shows the grouping
between pairs of dots, which produces primarily a first-harmonic response, as seen in
figure 2a. Figure 12b shows the collinear grouping along vertical lines of dots, as seen
in figure 2b. This grouping is due to a second-harmonic response at each dot loca-
tion, although a lateral or fourth-harmonic response is also in evidence. The terminal
dots at the top and bottom of each column of dots exhibit a first-harmonic response.
Figure 12c simulates the hexagonal grouping percept observed in figure 2c, due to a
third-harmonic response at each dot location. It is in this more complex stimulus that
the directional harmonic model demonstrates its predictive power. Prominently absent
from the harmonic response are the vertical, horizontal, and diagonal grouping percepts
that are in evidence in the input response at each dot location. Figure 12d shows the
Directional harmonic theory 439

dots stimulus input signal harmonic response

(a)

(b)

(c)

(d)
Figure 12. Computer simulations of the four dot grouping phenomena in figure 4, showing
for each dot pattern the stimulus configuration, the input signal at each dot location, and
the harmonic response at each dot location with the use of the spatial plotting convention.
(a) The pairs of dots form end-stop grouping percepts to the adjacent dot. (b) The columns of
dots promote a vertical collinear grouping along the columns. (c) The hexagonal dot grouping
dominated by a third-harmonic response at each dot location. (d) A grid-like percept due
to dominance of the fourth-harmonic grouping.

four-way or orthogonal grouping percept corresponding to figure 2d, due to a fourth-

harmonic response at each dot location. Again, prominently absent from the harmonic
response is the diagonal grouping which is in evidence in the input signal for this stimulus.
When a vertical column dot stimulus like that in figure 2b is varied parametrically
by shifting alternate rows of dots to the right, the perceptual experience due to that
grid of dots exhibits characteristic transitions, sometimes abrupt, between different
perceptual grouping patterns. Figure 13 shows these transitions as a spatial rather than
a temporal sequence. The alternate rows of dots in figure 13 have been shifted by a
440 S Lehar

0 0.1 0.2 0.3 0.4 0.5

Shift value

(a) linear (b) zig-zag (c) cross-hatch

Figure 13. Rectangular dot grid pattern in which alternative rows are shifted to the right, where
the amount of shift varies continuously from the left to the right side of the figure. Perceptually,
this shifting segments the percept into three distinct regions, that exhibit a (a) vertical linear,
(b) zig-zag, and (c) cross-hatch grouping percept as a function of shift.

shift value of zero on the left side of the figure (ie no shift at all) to a value of 0.5 on
the right side of the figure, expressed in units of the horizontal dot spacing, ie a shift
of 0.5 means that the alternate rows of dots have been shifted half-way to the next
adjacent column. The resulting perceptual experience can be categorized as a vertical
column or linear grouping percept, as seen in figure 13a, where the shift value is very
small. This then gives way to a zig-zag grouping as seen in figure 13b, in which each
column is perceived to be composed of a series of sharp angles. As the shift value is
further increased the percept becomes somewhat ambiguous, before finally settling
into a more stable diagonal grouping, or cross-hatch pattern, as seen in figure 13c.
Figure 14 shows computer simulations of these various grouping patterns. The abrupt
transitions between distinct grouping percepts seen in this figure reveal the influence
of the directional harmonic resonance, because these transitions appear only in the
harmonic response image, whereas the input signal exhibits only a gradual or continu-
ous transition between these arrangements of the stimulus. The abruptness of these
perceptual transitions therefore mark a significant difference between the predictions
of the directional harmonic model and a simple grouping-by-proximity model.
The different grouping patterns seen in figure 13 can be explained by the directional
harmonic model as the successive dominance of different harmonics of the grouping
mechanism at different shift values. Figure 15 shows a computer simulation of the
directional harmonics at a central dot location between two adjacent dots, as the flank-
ing dots are progressively deflected from a collinear configuration. In figure 15a, the
flanking dots are deflected 158 downward from the horizontal, ie the three dots form
an internal angle of 1508. At this small angle of deflection the harmonic response is
still dominated by the second harmonic, ie the dots are perceived to be in a collinear
alignment. In figure 15b the deflection has been increased to 308, so the internal angle
between the dots is now 1208. In this configuration the response is dominated by the
third harmonic, ie the dots are perceived to form an obtuse angle. The third-harmonic
response has a third branch, besides the two aligned with the flanking dots, which
suggests a tendency for a perceptual grouping line to emerge in that direction.
However, this tendency is balanced by the first harmonic, which exhibits a positive
peak downward, and a negative trough upward in figure 15b, as well as by a negative
trough in the fourth harmonic, which is why the combined harmonic response shows
no positive peak in the 12 o'clock direction. Figure 15c shows the angle of deflection
now increased to 458, so that the dots now define a right-angled corner. This in turn
promotes the fourth harmonic as the dominant response of the system. The fourth-
harmonic response exhibits two additional peaks besides the two aligned with the
Directional harmonic theory 441

dots stimulus input signal harmonic response

(a)

(b)

(c)
Figure 14. Computer simulation of the three perceptual groupings observed in figure 13.
(a) Collinear grouping of columns, dominated by the second-harmonic grouping. (b) Wavy-live
grouping where each dot marks the center of a two-armed vertex. (c) Cross-hatch grouping in
which every dot marks the center of a diagonal fourth-harmonic or X grouping percept.

neighboring dots, which are almost positive in the combined harmonic response shown
in figure 15c. The presence of adjacent dots in exactly those directions in figure 14c
is enough to boost those peaks to positive values, resulting in a fourth-harmonic or
four-way grouping percept at each dot.
The competition between different harmonics in response to various dot configura-
tions also offers an explanation for the abrupt kinking of a line or circle of dots, as
seen in figure 1d, which is observed to occur just as the angle between three adjacent
dots approaches 1208, the angle which favors the third-harmonic response. Again, these
quantized, or abrupt, perceptual transitions in response to a continuous parametric
variation of the stimulus are due to the loss of the higher harmonics due to impedance,
which in turn results in the perceptual characterization of these stimuli in terms of
various combinations of the lower-order terms, which serve as a basis set of geomet-
rical primitives for encoding the perceived forms.
Figure 16 demonstrates a different parametric variation of a dot grid pattern, this
time obtained by shifting alternate pairs of rows by a variable amount. This leads to
distinct perceptual grouping patterns which can be categorized as linear, wavy, and
hexagonal grouping percepts with progressively increasing shift value. Figure 17 shows
how these patterns too can be explained by the directional harmonic model. The
wavy pattern gives way to the hexagonal pattern at the point where the third leg of the
third-harmonic grouping percept at each apex (see in figure 17b) comes into alignment
442 S Lehar

dots stimulus individual harmonics harmonic response

(a)

(b)

(c)
Figure 15. Simulation of kinking of a perceived line of dots with increased curvature, as seen in
figure 1d. (a) Three dots in a line with a slight downward deflection (dashed lines) promotes a
predominantly second-harmonic or collinear grouping percept. (b) Three dots with a greater
deflection produce harmonic responses in which the third-harmonic response dominates.
(c) When the angle between the dots approaches 908, the fourth harmonic dominates, with a
tendency to form illusory grouping lines in all four directions of the fourth-harmonic vertex.

0 0.1 0.2 0.3 0.4 0.5

Shift value

(a) linear (b) wavy (c) hexagonal

Figure 16. Rectangular dot grid in which alternate pairs of rows are shifted horizontally to produce
(a) linear, (b) wavy-line, and (c) hexagonal grouping percepts.
Directional harmonic theory 443

dots stimulus input signal harmonic response

(a)

(b)

(c)
Figure 17. Computer simulation of (a) the collinear grouping, (b) the wavy-line percept, and
(c) the hexagonal grouping percept seen in figure 16.

with those on the adjacent column of dots, forming a bridge that spans the gap
between the columns, resulting in the hexagonal grouping percept of figure 17c.
3.2 Line pattern grouping simulations
In the dot grouping simulations presented above, the input to the system at each dot
location was assumed to energize the directional harmonic resonance at that location
with equal energy at all orientations. The final pattern of resonance therefore results
exclusively from the configuration of neighboring dots, as communicated through the
angular and radial Gaussian input functions. A line segment stimulus, on the other hand,
provides an oriented input signal along the line segment, corresponding to a second
harmonic or collinear grouping at every point along that line. This raw input is expected
to overwhelm the resonance at every point along that line, resulting in a strong second-
harmonic response along the stimulus lines, corresponding to a collinear `grouping'
percept, or the veridical percept of the stimulus line, as a line of the appropriate orien-
tation. The interesting grouping effects observed in the line-segment stimuli are observed
at the line endings, where each line ending behaves somewhat like a dot stimulus,
except with an oriented bias, or strong oriented input in the direction of the line
segment. The line pattern grouping simulations of the directional harmonic model
are therefore performed similar to the dot pattern groupings, in that the harmonic
resonance is computed only at the points where the line segments terminate, ie at the
line endings, as shown in figure 18a. The resonance at the line ending is therefore biased by
a fixed oriented input signal from the line of which it is the terminus. For example,
444 S Lehar

input signal input signal

(a) (b)
Figure 18. (a) The directional harmonic simulations of line segment stimuli are performed similar
to those of the dot patterns, with the harmonic response being calculated only at the location
of the line endings. The input signal at each line ending is presumed to have a permanent input
signal from the direction of the line of which that point is the terminus. For example, the top
end of a vertical line segment is presumed to have a permanent input signal from the 6 o'clock
direction, while the bottom end would have a permanent input signal from the 12 o'clock direc-
tion as shown here. (b) In the case of multiple line segments, the harmonics are computed at
each vertex, where a permanent input signal is presumed in the direction of each of the compo-
nent line segments as shown here.
line stimulus input signal harmonic response

(a)

(b)

(c)
Figure 19. Computer simulations of perceptual grouping between line segment stimuli showing:
(a) collinear grouping, (b) orthogonal grouping, and (c) diagonal grouping percepts at each line ending.

the point at the top end of a vertical line segment is assumed to have a permanent input
signal from the 6 o'clock direction, in addition to any other influences from adjacent line
endings, whereas the bottom end of a vertical line segment has a permanent input from
the 12 o'clock direction, as shown in figure 18a. The model can also be used to simulate
Directional harmonic theory 445

vertices, as shown in figure 18b, in which case the input signal at the vertex is assumed
to have permanent inputs from the directions of the component line segments of that
vertex, as shown in figure 18b. With this simple addition, the directional harmonic
model now also accounts for the perceptual line segment grouping phenomena shown
in figure 3.
Figure 19 shows the directional harmonic simulation for collinear, orthogonal, and
diagonal grouping of the line-segment stimuli, as observed perceptually in figure 3.
The collinear grouping percept shown in figure 3a is explained by a second-harmonic
response at each line ending, as shown in figure 19a. The significant difference between
the resonance response and the input signal (equivalent to the prediction of a grouping-
by-proximity model) is not so much the predominantly vertical grouping, which is
observed in both responses, but in the suppression of the alternative horizontal and
diagonal groupings observed in the input signal. The orthogonal grouping percept of
figure 3b is explained by a fourth-harmonic grouping at each line ending in figure 19b,
with attenuated grouping percepts in the collinear and diagonal directions, whereas the
diagonal grouping percept of figure 3c is explained by a third-harmonic or Y-vertex
response at each line ending, as seen in figure 19c, with a suppression of the horizontal,
and extraneous diagonal groupings observed in the input signal for that stimulus.

4 Discussion
Ever since Santiago Ramon y Cajal discovered the cellular basis of the nervous system,
the idea of the nervous system as an assembly of quasi-independent processors, some-
times called the neuron doctrine (Barlow 1972, 1995), has become firmly established as the
dominant paradigm of neurocomputation. Many different arrangements of these simple
integrate-and-threshold elements have been tried in computer simulations in an attempt
to coax some kind of interesting or suggestive behavior from them as an explanation
for perceptual processing. But certain aspects of perceptual function have remained so
elusive as to cast doubt on the entire enterprise. Particularly problematic have been the
enigmatic properties of perception identified by Gestalt theory. The principle of invari-
ance is ubiquitous in many aspects of perception, including invariance in the recognition
of simple shapes to rotation, translation, and scale. And yet this kind of invariance is
very difficult to achieve in conventional neural network models, for it leads to a
combinatorial explosion in the required number of receptive fields. The principle of
emergence, identified by Gestalt theory, is also problematic for neural network models,
because it suggests a relaxation to equilibrium of a massively parallel dynamic system
through many iterations, a concept which is difficult for the neuron doctrine given the
time delays inherent in the chemical synapse. There is also a dimensional mismatch
between the continuous field-like nature of perceptual experience identified by Gestalt
theory, and the constellation of discrete activations of individual neurons in the neuron
doctrine (Lehar 2002). I propose therefore to return all the way to the debate between
Golgi and Cajal, and to reconsider the question whether the essential principles of
neurocomputation are best described as an atomistic system of simple quasi-independent
local processors as suggested by neural network theory, or as a holistic continuous-
field-like principle as suggested by Gestalt theory. I don't propose to question the
neurophysiological and histological evidence for the cellular basis of the nervous system,
or for the properties of the chemical synapse. However, I do contest the significance
of that evidence for the essential principles of neurocomputation.
In fact, the problems with the neuron doctrine date as far back as the development of
the first brain-recording technique, the electroencephalogram (EEG). From the outset the
EEG detected a global oscillation across the whole brain, and that global synchrony
was found to correlate with particular functional states. However, the significance of this
global resonance remains to this day as mysterious as it was when it was first discovered.
446 S Lehar

Now, with the introduction of multiple-unit recordings, this global resonance is even
beginning to manifest itself in electrophysiological data. Although it is not much
discussed in the contemporary literature, this kind of global resonance is problematic
for the neuron doctrine. For the phasic spiking of a neuron is unlikely to survive across
the chemical synapse, owing to the multiple parallel paths in each axonal and dentritic
arborization, each with variable path lengths and dendritic diameters which would intro-
duce random time delays across all those parallel paths, not to mention the temporal
averaging of the signal in each of the 10 000 or so parallel chemical synapses that connect
a typical pair of neurons. After the third or fourth such transmission along a line of
cells, the phasic pulses of the neural signal would surely diffuse into a featureless blur,
which would preclude the kind of global synchrony observed in the EEG signal across the
cortex as a whole.
Another Achilles heel in the neuron doctrine that receives little mention today is the
fact that the cell membrane, although capable of insulating electrical charge inside the cell,
does not insulate alternating currents. For, like the insulating dielectric in a capacitor,
the cell wall transmits alternating-current signals perfectly well, and blocks only direct
current. That is why single-cell electrode recording can be performed extracellularly,
just as well as intra-cellularly. This in turn suggests that the alternating-current spiking
signal of the neuron is likely to propagate directly from cell to cell across the extracellular
matrix, unhindered by the insulation of the cell membrane. Pribram (1971) proposes that
the overt spiking of the neuron is not the causal origin, but merely a secondary mani-
festation or response of the cell to a graded potential oscillation that pervades the body
of neural tissue, because that oscillation has been observed to continue even after the
spiking cell falls below threshold and ceases to fire. Like whitecaps on ocean waves,
the spiking neuron merely reveals the presence of a more subtle underlying resonance that
pervades the bulk neural tissue. The reason that this graded potential oscillation has
received so little attention in electrophysiological circles has been for lack of a theoret-
ical framework to give meaning to those resonances, or to explain how harmonic
resonance can perform a computational function in the brain.
I propose that the global synchrony observed in the EEG recordings, and now in
the synchronous activity of cortical neurons, is exactly what it appears to be, ie it is a
manifestation of a global harmonic resonance that pervades the entire cortex, and that
this resonance subserves a computational function which is central to the principle of
operation of the brain (Lehar 1994a, 1994b, 1999, 2002). The purpose of this resonance
is to set up a pattern of standing waves, and these standing waves serve a function
which is normally ascribed to spatial receptive fields in neural network models, ie as
a mechanism for encoding spatial patterns in the brain both for recognition and for
perceptual completion.
The computational function performed by these standing waves is highlighted in
the directional harmonic model by comparison with the functionally equivalent neural
network models that it replaces. The second-harmonic standing wave of directional
periodicity mirrors the functionality of a collinear completion model as proposed by
Grossberg and Mingolla (1985, 1987), Walters (1987), and Zucker et al (1989), except that
functionality no longer requires separate spatial receptive fields replicated at every
location and every orientation across the visual field, but, rather, the second-harmonic
resonance emerges spontaneously at the location and orientation determined by the stim-
ulus. Furthermore, this same dynamic resonance also performs perceptual completion
through a range of different vertex types corresponding to the various harmonics of
directional periodicity, such as Y and ‡ or X vertices, with no additional hardware
required. But even that does not exhaust the computational repertoire of the direc-
tional harmonic model, for, as in a Fourier representation, these harmonics are also
used in combination to produce compound standing-wave patterns corresponding to
Directional harmonic theory 447

L and T and V vertices. This is functionally equivalent to a neural network model

capable of merging different receptive field profiles into new compound receptive fields
on the fly, and using those compound fields to perform completion of spatial patterns.
Finally, there is one further aspect of a harmonic resonance theory that deserves
mention, and that is its relation to emergence and reification in Gestalt illusions. The
neural network models of Grossberg and Mingolla (1985, 1987), and Zucker et al
(1989) incorporate an explicit top ^ down feedback from what I have called the `feed-
back layer' back down to the layer of local oriented edge detectors, so that the higher-level
groupings computed in the feedback layer are expressed back at the oriented edge layer
as explicit edge percepts. This is done in order to account for the perceptual experience
of a sharp high-resolution contour in figures like the Kanizsa triangle, at a location
where there is no edge present in the stimulus. Although this issue was not addressed
explicitly in the directional harmonic model, it is in the very nature of harmonic
resonances in adjacent subsystems to tend to couple with each other by mutual
reciprocal feedback to produce a single combined resonance. A resonance model there-
fore does not have to add top ^ down feedback of this sort as an explicit mechanism of
the model, because it is already a natural property of the resonance itself (Lehar
1999). In fact, this natural tendency of individual local resonances to couple into a
single global resonance is, I propose, the central operational principle behind the holistic
global nature of Gestalt phenomena. In other words, harmonic resonance offers a
computational principle that exhibits the holistic global aspects of perception identified
by Gestalt theory, not as specialized mechanisms or architectures contrived to achieve
those properties, but as natural properties of the resonance itself.
The principal value of the directional harmonic theory therefore is not so much
as evidence for some kind of resonance in the brain, for that evidence already
exists from a wide variety of diverse sources. But, rather, the directional harmonic
theory offers a specific and detailed model of exactly how standing waves might per-
form a computational function in perception normally ascribed to spatial receptive
fields, and demonstrates how that function circumvents some fundamental limitations
inherent in the template-like concept of the neural receptive field, by an emergent
holistic process consistent with Gestalt theory.
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