Classifying Illusory Contours: Edges Defined by "Pacman" and Monocular Tokens
Gerald Westheimer and
Wu Li
Division of Neurobiology, University of California, Berkeley, California 94720-3200
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ABSTRACT |
Westheimer, Gerald and Wu Li. Classifying illusory contours: edges defined by "pacman" and monocular tokens. J. Neurophysiol. 77: 731-736, 1997. Thresholds for the discrimination of orientation were measured in the human fovea for figures and borders delineated by solid lines and by "pacman" tokens as introduced by Kanizsa, as well as by contours induced by monocular tokens giving a stereoscopic depth illusion of a knife edge. Orientation discrimination of these illusory contours is poorer by a factor of ~2 than that of equivalent contours made of solid lines and is not much better than that for their supporting structures if taken alone. It is concluded that these kinds of illusory borders do not address the "border" or "edge" mechanism in the same way as real lines. Orientation discrimination and simultaneous orientation contrast (tilt illusion) were compared for a variety of illusory borders. The more robust the borders, i.e., the more sensitive to changes in orientation, the less their susceptibility to the tilt illusion.
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INTRODUCTION |
To be perceived as contours, borders need not be drawn out explicitly but can be sketched in by a variety of means. Virtual, subjective, or illusory borders differ in their conspicuity, but this is not a property easily expressed empirically. On the other hand, orientation, a prominent attribute of a straight border, can be approached via a discrimination threshold, which gives excellent quantitative measures. For example, a simple line just covering the human fovea (~30 arcmin) can have the direction of its tilt with respect to the vertical detected when it is half a degree of orientation or less. We have recently used this orientation discrimination threshold to show that some illusory borders are inferior to solid straight lines in several respects: not only do they display higher thresholds, but they take longer to process and are not robust to target motion or masking.
In the earlier paper (Westheimer and Li 1996
) we studied illusory contours made of orthogonal stacks of lines with and without a gap. Here we extend the investigation to two further types of illusory contours, those delineated by "pacman" tokens in the manner of Kanizsa figures and the startling "knife-edge" depth borders engendered by monocular features described by Nakayama and Shimojo (1990)
. We then go on to compare the orientation discrimination of a variety of kinds of illusory contours and relate it to their susceptibility to accept an orientation shift induced by surrounding tilted lines.
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METHODS |
Patterns made of thin lines of medium photopic intensity were created under computer control on high-quality flat display monitors (HP1345A). Pacman tokens were drawn by closely spaced lines of the requisite length. All edges were given orientation changes by recalculating the coordinates of all generating lines according to a rotation around the geometric center of the line or border, or, in the case of the data in Fig. 1, the whole figure. Unfortunately it is impossible to avoid secondary cues in these experiments. If the pacman tokens had remained fixed, rotation of the illusory borders would have necessitated recognizable changes in their internal configuration. Instead we chose to leave the pacman tokens invariant in shape while giving them the combination of rotation and displacement that results from use of the middle of the contour, or the whole figure, as the center of rotation. Whether displacements act as a secondary cue is a question that can be answered independently (see the DISCUSSION section).

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| FIG. 1.
Threshold for the orientation discrimination around the vertical of (1) a square, (2) a square outlined by both solid lines and 4 "pacman" tokens, (3) a square outlined by 4 pacman tokens, and (4) a square outlined by 4 pacman tokens with occluded corners. The edges were 32 arcmin long in all cases and the exposure duration was 300 ms. All pacman disks had a diameter of 16 arcmin. To produce orientation changes, the patterns were rotated around their center. Orientation changes of explicitly delineated squares could be detected about twice as well as an illusory figure, and occultation of the latter had no further deteriorating effect.
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The two authors as well as one naive subject served as observers. They faced the monitor screens in a dim room at a distance of 3.75 m, with binocular vision, wearing the needed refractive correction. The experiments whose results are illustrated in Fig. 5 were conducted in a stereoscopic setup, in which each eye saw only its own monitor by means of a system of first-surface mirrors.

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| FIG. 5.
Orientation discrimination of monocular borders made up of 3 short lines or a long solid line under 3 conditions. In 1 and 4 the monocular borders are embedded in a field of random binocular short lines with a difference in disparity between the right and left sides; in 2 and 5 all binocularly seen short lines are in the fixation plane, and in 3 and 6 the field is empty except for the monocular border. In all cases there appears a depth edge to 1 side of the monocular features and this edge is particularly pronounced when there are many binocular features. The illusory depth border does not, however, manifest itself in an improved orientation discrimination threshold. Exposure duration: 2 s.
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Thresholds were obtained by the psychophysical method of constant stimuli and probit analysis and are expressed in angle of orientation. They indicate the deviation from the vertical for which the observer can correctly identify the direction (clockwise or counterclockwise) on 75% of occasions. Details are contained in earlier papers from this laboratory (e.g., Westheimer and Li 1996
). Data were gathered in blocks of 140 responses to the same kind of pattern. All threshold measures given in this paper are based on at least two such blocks obtained on different days. No error feedback was given. Unless otherwise indicated, exposure duration was always 300 ms. A measure of the magnitude of simultaneous orientation contrast (tilt illusion) was obtained by finding, in the presence of surrounding tilted contours or lines, the actual tilt of the test border when the observer calls it "vertical." To obviate bias, it was identified by halving the difference between the 50% point on the probit curves for clockwise and counterclockwise inducing contours.
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RESULTS |
In Fig. 1 we show the threshold for orientation discrimination of four squares, all having side lengths of 32 arcmin. If the square has its four sides explicitly drawn, thresholds are good, and the presence of pacman tokens at the corner has no influence. A square outlined only by pacman tokens, which is a well-known Kanizsa figure and paradigmatic of illusory figures, needs about twice as large an orientation difference for threshold discrimination. Occlusion of the open sectors, the traditional way of eliminating the percept of an illusory figure, does not materially affect the threshold. Thus the first and major point of this study has been established: although there may be a compelling percept of an illusory square, its orientation cannot be discriminated nearly as well as that of an explicitly drawn square.
We analyze the situation further in the experiments illustrated in Fig. 2. Here the orientation discrimination around the vertical has been determined for a variety of edges, all 32 arcmin long. Explicitly drawn edges are best, and pacman delineated edges, occluded or not, are not nearly as good. Then we used two short aligned line segments, 8 arcmin long with 16-arcmin inner separation. These are in fact the vertical contours of the pacman sectors and they demonstrated about the same orientation discrimination when shown alone as when they were part of the illusory configuration. Finally, we gave a 3-arcmin horizontal offset to the pacman tokens and also to the simple line segments. On the whole, taken over the three observers, there was a slight further deterioration of performance.

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| FIG. 2.
Threshold for the orientation discrimination of a variety of real and illusory edges, all 32 arcmin long: (1) a single line; (2) 2 pacman tokens connected by a line; (3) 2 aligned pacman tokens, in the manner of a Kanizsa figure; (4) 2 aligned pacman tokens with occluded openings; (5) 2 aligned 8-arcmin line segments, providing the same vertical contours as in 2-4 but in isolation; (6) 2 pacman tokens as in 3 but offset horizontally by 3 arcmin; (7) same as 5, but offset horizontally by 3 arcmin. Exposure duration: 300 ms. In conditions 6 and 7, the offset remained fixed throughout the experiment, and the observer's task was to judge the direction of the change in orientation that resulted from rotation of the whole configuration around its middle.
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Orientation discrimination of lines improves with line length. To test whether this is also the case for illusory edges outlined by pacman tokens, we obtained the orientation discrimination threshold for three line lengths for equivalent real and illusory edges. As can be seen in Fig. 3, only when there is a real line does the performance improve with increasing length.

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| FIG. 3.
Orientation discrimination thresholds as a function of border length for 2 edges. Solid symbols: edge defined by pacman tokens. Open symbols: explicitly drawn line edge plus pacman tokens. Pacman disks were 16 arcmin in diameter. For solid lines, orientation discrimination, expressed in angles of orientation, progressively improves with line length. The illusory edge does not follow this pattern. Exposure duration: 300 ms.
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Illusory contours made of abutting offset lines without luminance gradient have an orientation discrimination threshold that is not only high but also rises further with short exposures (Westheimer and Li 1996
). The question arises, therefore, whether this is also the case for Kanizsa edges. Figure 4 shows that this is not the case. The real and illusory edges to which this report is mainly devoted can have their orientation discriminated just as well for short exposures as for long ones.

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| FIG. 4.
Orientation discrimination thresholds as a function of exposure duration for a variety of edge configurations, all 32 arcmin in length. Thresholds are substantially invariant with exposure duration in the 50- to 1,000-ms range except for the illusory contours made up of offset abutting lines without luminance gradient.
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There is yet a different kind of illusory contour pointed to in detail by Nakayama and Shimojo (1990)
. Panum (1858)
and Hering (1862)
examined the situation in which there is a monocular contour in the vicinity of a binocular one. The most relevant example is the description by Hering of the fusion of a thick vertical line in one eye with a thin one in the other. There is a compelling percept of a sharp edge with a stereoscopic depth discontinuity. Nakayama and Shimojo used the term knife edge, and we might also describe it as a depth cliff. Nakayama and Shimojo showed that this occurs also when there is just a single monocular line in the presence of binocular features and even when the monocular line is just sketched in by three aligned short vertical line segments. (There is no stereoscopic depth unless there are at least some binocular features; the phenomenon is not experienced when 1 eye is closed.) We find this illusory depth contour very compelling and wondered whether it conferred additional benefits to orientation discrimination beyond that shown by the broken line alone. Consequently we measured the orientation discrimination threshold for the sketched-in line and for a real line under three conditions: when shown monocularly by themselves, when shown monocularly within a field of random binocular lines, and when the random binocular lines were segregated into right and left depth planes in the manner of what Nakayama and Shimojo call "da Vinci stereopsis," i.e., a situation geometrically equivalent to that in which monocularity of the test border is caused by its being shielded from one eye because of occlusion by a partial anterior depth plane. To ensure fully developed stereopsis, a 2-s exposure duration was used in these experiments. The data in Fig. 5 illustrate that in no case does the evoked stereoscopic cliff improve orientation discrimination and, if anything, simulation of da Vinci stereopsis (cases 1 and 4) further impairs performance. This supports our previous conclusion that stereopsis does not yet enter at the neural stage processing contour orientation.
Finally we address a related question. All oriented configurations are subject to simultaneous orientation contrast or, as it is commonly called, the tilt illusion; that is, their perceived orientation is affected by superimposed or neighboring tilted contours shown either simultaneously or previously as adapting stimuli (the tilt aftereffect). We examined the magnitude of this simultaneous orientation contrast in a variety of real and illusory borders by the technique described earlier (Westheimer 1990
). Observers reported whether a given contour appeared tilted clockwise or counterclockwise, but the contours were shown flanked by lines with a 20° tilt, randomly clockwise or counterclockwise in long series of presentations. Results were tallied separately for the two directions of inducing tilt. The two psychometric curves are shifted
in the clockwise direction for counterclockwise inducing lines and vice versa; half the difference between their 50% points may be taken to be the mean orientation shift induced by a 20° tilt of the flanks. This measure was obtained for the contours shown in the Fig. 6, inset. Because lengths and distances were the same in all cases, it may be assumed that the numbers represent the susceptibility of these contours to tilt induction. There is an approximately inverse relationship to the orientation discrimination threshold, followed by all contours except those delineated by pacman tokens.

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| FIG. 6.
Magnitude of the tilt illusion for a variety of real and illusory borders. The length, position, and tilt of the inducing contours were the same in all cases. There is an approximately inverse relationship between the sensitivity to orientation difference and susceptibility to the tilt illusion.
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DISCUSSION |
Figures delineated by illusory contour have a very prominent perceptual presence (see Petry and Meyer 1987
for a comprehensive review; also Spillman and Dresp 1995). They can have induced illusory brightness differences (Day 1987
), are said to be detected in parallel rather than serially (Davis and Driver 1994
), and may even seem to capture depth (Ramachandran and Cavanagh 1985
). It is therefore tempting to assign their elaboration to an early stage of visual processing. Luminance increment thresholds in positions occupied by illusory borders do not indicate any special changes that cannot be accounted for by the presence of the inducers (Dresp and Bonnet 1993
), but that does not yet deny early cortical processing, because it is generally believed that luminance increment thresholds reflect retinal sensitivity and there is no reason to believe that the retina is involved in any direct way in the generation of subjective contours.
On the other hand, the "orientation" attribute of borders is widely believed to have its neural substrate in the primary visual cortex (Gilbert and Wiesel 1990
; Vogels and Orban 1990
). Orientation discrimination is a good tool for probing this mechanism, with the general expectation that the more a stimulus addresses the concerned neural apparatus, the smaller the difference in orientation that can be detected. Insofar as different kinds of contours or edges show differences in their orientation discrimination, it may be inferred that they do not have equal access to the apparatus. On the basis of this premise, we conclude that illusory borders of the Kanizsa and monocular types, although undoubtedly prominent perceptually, do not have the benefit of having their orientation processed by the orientation-selective apparatus early in the cortical visual stream. The same applies to the illusory borders generated by offset abutting lines without luminance gradients, which have previously been assigned to a V2 apparatus (Peterhans and von der Heydt 1989
; van der Zwan and Wenderoth 1995
; von der Heydt and Peterhans 1989
). Further evidence for this view is seen in Fig. 3 above, where line length dependency is quite different for real lines and illusory contours.
A distinction needs to be drawn between the detection of contour orientation and of the inevitable displacement of the outer terminators of the contours that accompanies rotation. Thus, when a vertical line is rotated counterclockwise, is the observer's judgment made on the basis of the line's changed orientation or on the fact that the top end of the line is now displaced to the left with respect to the bottom? Fortunately it is possible to differentiate between these two judgments not merely by instruction to the observers but also by their difference in thresholds. When a line is longer than ~10 arcmin in the fovea, orientation thresholds for lines are considerably better than position difference thresholds for their two endpoints, so that in such cases the end displacement cue has been eliminated; this also applies to most of the data in this paper. The data in Fig. 3 above constitute an example of this phenomenon that has recently been subjected to a fuller analysis (Westheimer 1996
) and that is at the heart of the arguments developed in the present study: because pacman-induced illusory contours have worse orientation thresholds than explicitly drawn lines, they do not utilize fully the orientation-defining neural mechanism.
Taken as a whole, the evidence presented in this paper leads one to the conclusion that insofar as judgment of their orientation is concerned, illusory contours induced by Kanizsa and monocular tokens depend on the fragments of real borders that they contain and that perceptual completion or filling in does not manifest itself in the improved performance that would be displayed by an equivalent explicit line stimulus. Moreover, occlusion of the pacman sector (Figs. 1 and 2, case 4, and Fig. 4, case 2), which abolishes the sense of an illusory figure, has essentially no additional effect.
But it would be too facile to make the distinction just between explicit and illusory borders, for there seem to be real differences between the latter, even when light intensity, line length, and exposure duration are factored out. In an earlier study (Westheimer and Li 1996
) we argued that simple illusory contours generated by interrupted lines or stacked terminators of orthogonal lines behave more or less the same as real lines, if the right conditions of length and separation are obeyed. The same can probably be said for the detection of the direction of motion of a single dot (Westheimer and Wehrhahn 1994
). But there is no question that edges delineated by abutting lines with offsets (Fig. 4, case 5) do not fall into the same category, because not only do they have a higher threshold, but they also take longer to process, are more subject to masking, and are not robust to motion. The illusory contours induced by Kanizsa and monocular depth tokens studied in this paper fall into an intermediary category, because although their orientation threshold is higher and does not improve with contour length, it is robust to shortening of exposure time and (as we have ascertained in an experiment not further described here) to image motion. Moreover, there is a small indication that some contour completion occurs. Specifically, bringing the two pacman tokens that define a contour out of alignment by only 3 arcmin (Fig. 2, cases 6 and 7) does raise the threshold in all of our three observers, in a manner that is reminiscent of the reduction in conspicuity of a Kanizsa triangle when the tokens are misaligned (Fahle and Koch 1995
).

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| FIG. 7.
Rank order for various kinds of illusory contours according to 2 criteria, sensitivity for orientation discrimination and magnitude of simultaneous orientation contrast (tilt illusion), on the basis of data from this paper and from Westheimer (1990) , Westheimer and Wehrhahn (1994) , and Westheimer and Li (1996) . The experimental conditions of contour length, foveal vision, exposure duration, position of inducing contours, etc., are essentially identical in all experiments and allow the conclusion to be drawn that there is an approximately inverse relationship between the sensitivity for orientation discrimination and the magnitude of the tilt illusion: the firmer the border, i.e., the more sensitive it is to perturbations of its orientation, the less its orientation can be influenced.
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Another way of approaching the task of ordering the magnitude of the "contour" attribute of our borders is to examine the effect on them of flanking stimuli of contrasting orientation. That illusory borders can support the tilt illusion and tilt aftereffect has been well established (Berkley et al. 1994
; Paradiso et al. 1989
; Vogels and Orban 1987
). But there are indications that they do so to different extents (Westheimer 1990
). The data in the latter paper and in Fig. 6 above, for which at least one observer is in common, permit an ordering of the extent to which tilted inducers can influence the perceived orientation of the various kinds of illusory contours. The experimental conditions of contour length, position, and orientation of inducing lines are essentially the same in all these studies and allow one to place illusory contours in a sequence according to decreasing susceptibility to induced orientation shifts. When this order is compared with the threshold for orientation discrimination, it is found that there is a remarkable agreement: the poorer a contour is in orientation discrimination, the more it allows its perceived orientation to be affected by interaction from surrounding lines (Fig. 7). And, curiously, the Kanizsa border does not quite fit this order: it shows a smaller tilt illusion than might be expected if the sequences matched exactly. Perhaps this can be read as another indication that there is indeed a small amount of border completion (see also Fig. 2), but on the whole we have shown that only explicitly drawn lines and closely spaced interrupted contours support good orientation discrimination; illusory contours in general share this capability only to an extent that is limited and that varies with the type.
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ACKNOWLEDGEMENTS |
This research was supported by National Eye Institute Grant EY-00220.
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FOOTNOTES |
Address for reprint requests: G. Westheimer, 321 Life Sciences Addition, University of California, Berkeley, CA 94720-3200.
Received 28 May 1996; accepted in final form 3 October 1996.
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