Journal of Vision (2012) 12(2):3, 1–10 http://www.journalofvision.org/content/12/2/3 1

Polarity selectivity of spatial interactions in perceived

Department of Psychology, The University of Tokyo, Japan,& Human and Information Science Laboratory, Hiromi Sato NTT Communication Science Laboratories, NTT, Japan

Human and Information Science Laboratory, Isamu Motoyoshi NTT Communication Science Laboratories, NTT, Japan

Department of Psychology, Takao Sato The University of Tokyo, Japan

The apparent contrast of a texture is reduced when surrounded by another texture with high contrast. This contrast–contrast phenomenon has been thought to be a result of spatial interactions between visual channels that encode contrast energy. In the present study, we show that contrast–contrast is selective to luminance polarity by using texture patterns composed of sparse elongated blobs. The apparent contrast of a texture of bright (dark) elements was substantially reduced only when it was surrounded by a texture of elements with the same polarity. This polarity specificity was not evident for textures with high element densities, which were similar to those used in previous studies, probably because such stimuli should inevitably activate both on- and off-type sensors. We also found that polarity-selective suppression decreased as the difference in orientation between the center and surround elements increased but remained for orthogonally oriented elements. These results suggest that the contrast–contrast illusion largely depends on spatial interactions between visual channels that are selective to on–off polarity and only weakly selective to orientation. Keywords: polarity selectivity, spatial interactions, perceived contrast Citation: Sato, H., Motoyoshi, I., & Sato, T. (2012). Polarity selectivity of spatial interactions in perceived contrast. Journal of Vision, 12(2):3, 1–10, http://www.journalofvision.org/content/12/2/3, doi:10.1167/12.2.3.

(Cannon & Fullenkamp, 1993, 1996; Chubb et al., 1989; Introduction Ejima & Takahashi, 1985; Sagi & Hochstein, 1985). Similar mechanisms have been assumed to underlie The contrast of a texture pattern appears to be lower spatial interactions observed in detection threshold (Polat when it is surrounded by another texture with a high & Sagi, 1993; Zenger & Sagi, 1996), perceived orienta- contrast (Cannon & Fullenkamp, 1993; Chubb, Sperling, tion (Blakemore, Carpenter, & Georgeson, 1970), and & Solomon, 1989; Petrov, Carandini, & McKee, 2005; perceived spatial frequency (Klein, Stromeyer, & Ganz, Sagi & Hochstein, 1985; Solomon, Sperling, & Chubb, 1974). The potential neural correlates of such interactions 1993; Xing & Heeger, 2000). This contrast–contrast (2nd- have been reported for neurons in V1 (Blakemore & order) effect is distinguished from the classical brightness– Tobin, 1972; Knierim & van Essen, 1992; Zipser, Lamme, contrast (1st-order) effect (Chubb et al., 1989; McCourt, & Schiller, 1996), although their anatomical origin is still 2005) and has been extensively investigated to elucidate controversial (Webb, Dhruv, Solomon, Tailby, & Lennie, visual mechanisms underlying the perception of image 2005). From a computational viewpoint, these second- contrast. Past psychophysical studies have shown that the order spatial interactions are considered as useful for contrast–contrast effect depends on differences in orienta- efficient image coding (Schwartz & Simoncelli, 2001), tion and spatial frequency between the central and texture segmentation (Graham, 2011), and motion detec- surround regions (Cannon & Fullenkamp, 1993; Chubb tion (Chubb & Sperling, 1988). Many studies point out et al., 1989; Solomon et al., 1993; Xing & Heeger, 2000) that they also play a role for estimation of distal object and is observed even when the surround region is not properties such as reflectance and transparency (Adelson, consciously perceived (Cai, Zhou, & Chen, 2008; 1999; Dakin & Mareschal, 2000; Lotto & Purves, 2001; Motoyoshi & Hayakawa, 2010). These findings are Schofield, Hesse, & Rock, 2006; Solomon & Mareschal, consistent with the notion that the contrast–contrast effect unpublished draft; Spehar, Arend, & Gilchrist, 1995). is primarily caused by inhibitory lateral interactions Current models of the contrast–contrast phenomenon between low-level visual channels, i.e., spatial filters generally assume that contrast perception is mediated by

doi: 10.1167/12.2.3 Received October 18, 2011; published February 2, 2012 ISSN 1534-7362 * ARVO

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mechanisms that encode absolute (unsigned) contrast, i.e., contrast energy (Cannon & Fullenkamp, 1996; Solomon et al., 1993). This is a natural assumption since observers are asked to judge unsigned contrast, but it is not necessarily true. It has been known that the mammalian has not only neural units sensitive to unsigned contrast but also to either positive (on) or negative (off) contrast (Fiorentini, Baumgartner, Magnussen, Schiller, & Thomas, 1990; Jung, 1973; Schiller, Sandell, & Maunsell, 1986). These on- and off-type sensors have classically been thought to underlie the perception of brightness and darkness (Anstis, 1979; Blakeslee & McCourt, 2004; Jung, 1973). However, they could be involved in various judgments on spatial patterns. Psychophysical evidence shows that visual aftereffects in contrast detection threshold (Hanly & MacKay, 1979), perceived spatial frequency (De Valois, 1977), and contour shape (Gheorghiu & Kingdom, 2007) substantially depend on the combination of luminance polarity between the adapting and test stimuli. Similar segregation between on- and off-contrast information has been shown for texture discrimination (Chubb, Econopouly, & Landy, 1994; Chubb, Landy, & Econopouly, 2004; . Malik & Perona, 1990; Motoyoshi & Kingdom, 2007). Figure 1 Examples of stimuli used in Experiment 1. (a) Bright However, such a separation has not been found between center, bright surround. (b) Bright center, dark surround. (c) Dark stimuli of sine and anti-sine phases (Huang, Kingdom, & center, bright surround. (d) Dark center, dark surround. Hess, 2006; Malik & Perona, 1990). These findings lead us to the notion that the appearance of a textural pattern depends at least partially on separate representations of on basis of these results, we suggest that the contrast–contrast and off contrasts. effect involves polarity-selective, and partially orientation- Is the contrast–contrast effect selective for luminance selective, visual mechanisms. polarity? Solomon et al. (1993) addressed this question by using center–surround texture patterns composed of dense bright or dark dots. They found that perceived contrast in Experiment 1 the central region is reduced by the surrounding region regardless of whether it is composed of bright dots or dark dots, indicating no polarity selectivity. These results Methods seem to be consistent with a model of contrast–contrast Observers effect as being mediated by units encoding contrast Four naive participants (AY, KA, MH, and RY) and one energy. There is, however, a possibility that, as we shall of the authors (HS), with corrected-to-normal vision, explain later in more detail, such dense textures activate participated in the experiment. both on- and off-type sensors regardless of the elements’ luminance polarity. In the present study, we show that when texture is Apparatus composed of sparse elements that selectively stimulate on or off units, contrast–contrast shows a clear selectivity to Images were generated by a graphics card (CRS Visage) luminance contrast polarity. Figure 1 shows examples of and displayed on a 21-inch CRT (Eizo Flex Scan T962, stimuli. In Experiment 1, we demonstrated that the central SONY GDM F520R only for AY) with a refresh rate of texture appears to have lower contrast when it is surrounded 160 Hz. The pixel resolution of the CRT was 1.72 min/ pixel at a viewing distance of 1.0 m. The mean luminance of by a texture of the same polarity than when surrounded by 2 2 that of the opposite polarity. In Experiment 2, we found the homogenous field was 63 cd/m for AY and 54 cd/m that this polarity selectivity of contrast suppression for the other observers. diminished for textures made of dense elements, as previously reported by Solomon et al. (1993). In Experiment 3,wealso Stimuli showed that this polarity-selective suppression is slightly weakened as the difference in elements’ orientation was The stimuli for this experiment consisted of standard increased between the center and the surround. On the and test stimuli. The standard stimulus was a circular

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texture region surrounded by an annular inducing texture 500 ms. The circles stayed on for 500 ms after standard as shown in Figure 1. The diameter of the central field was and test stimuli disappeared. Early onset of the circle for 2.9 deg and that of the surround field was 8.6 deg. The 500 ms was useful for getting observers attend to the central field of the standard stimulus alone was presented image regions to be compared. separately and used as a test stimulus. The two stimuli The observers were asked to press one of two buttons to were presented side by side on both sides of a black indicate which of the standard or test stimulus appeared to fixation dot (0.1 Â 0.1 deg) at the center of a homogenous have a higher contrast (2AFC). They were instructed (1) not stimulus field (22.9 Â 17.2 deg). The separation from the to judge the overall brightness or darkness of the stimuli but fixation dot to the center of each stimulus was 5.7 deg. the contrast, or the strength, of elements and (2) to The texture was composed of elongated blobs defined by a concentrate on the central regions designated by the black two-dimensional Gaussian function with standard deviations circles. No feedback was given. of 0.07 deg (short axis) and 0.23 deg (long axis). All The contrast of the test stimulus was varied in elements were vertically oriented. Each element was accordance with the staircase method. It was decreased randomly placed with a minimum center-to-center separa- by 0.1 log unit when the observer indicated that the test tion of 0.64 deg. The elements within each field had the same stimulus had a higher contrast than the standard and was value of either positive or negative contrast polarity. The increased by 0.1 log unit when the observer indicated that combinations of polarities between the center and surround it had a lower contrast. Within a single session, 12 staircases were varied as shown in Figures 1a–1d. The absolute each corresponding to a different condition were randomly contrast of the texture field was defined as ªLmax j L0ª/L0 interleaved. In each session, the staircase terminated when when the polarity was positive and ªLmin j L0ª/L0 when the number of trials in all of the staircases exceeded 30. negative, where L0 is the luminance of elements’ back- Session was repeated several times, until at least 120 trials of ground and Lmax and Lmin are the positive and negative the data (83 for AY) were collected for each condition. The peaks of blob elements, respectively. The contrast of the proportion responses were fitted with a logistic function by surround field was 1.0, and that of central field was varied means of the maximum likelihood method, and the contrast between 0.125, 0.25, and 0.5. that yielded 50% probability was taken as the matched The mean luminance of each texture fieldVthe test contrast, or PSE. The standard error for each PSE was stimulus and the center and surround of the standard estimated through bootstrapping of 5000 samples. stimulusVwas equated to the mean luminance of the background homogenous field. Therefore, the background luminance (L0) within each texture field varied depending on the contrast polarity of texture elements. Because of this Results luminance difference, there was a luminance (first-order) edge between the center and surround of the standard Figure 2a shows the matched contrast of test stimulus as stimulus. The magnitude of luminance difference and the a function of the physical contrast of the standard clarity of the contour vary depending on the contrast stimulus. The left panel shows the average of all polarity. To control the clarity of the border between the observers, and the right small panels show the results for center and surround fields, we added a black circular each observer. (The average was calculated on the log contour (0.06 deg in width) having the same diameter as coordinate.) Red circles represent the results when the the central field to mask this luminance contour. This circle center had positive polarity, and blue circles represent the also helped the observers to segregate the center from the results when the polarity of the center was negative. Filled surround and prevented unnecessary crowding of elements. circles show the results when the center and surround had It was quite possible that addition of this black contour the same polarities, and open circles show the results affect the result. We checked this by a pilot experiment. The when they had opposite polarities. The perceived contrast results are basically the same between the conditions with is lower than actual when the center had the same contrast or without the black contour, but it was so much easier for polarity as the surround (filled circles). However, when observers to judge when there was a black circle. the center had an opposite polarity to the surround, the matched contrast appears almost equal to the physical contrast (open circles), indicating no suppression. Procedure Figure 2b shows the matched contrast relative to the We measured the matching point between the standard physical contrast. The same center–surround polarity and test stimuli, i.e., the point of subjective equality (PSE) (filled symbols) produced strong suppression at all con- for perceived contrast, by using a staircase method. trast levels of the center. On the other hand, the opposite Observers binocularly viewed the stimuli and fixated on center–surround polarity (open symbols) produced little, if the fixation dot through each trial. At the beginning of any, suppression except for the lowest contrast of the each trial, the fixation marker and the circles alone for center. These results suggest that contrast–contrast is both stimuli were presented. Both standard and test selective to luminance contrast polarity for a wide range stimuli appeared 500 ms later and were presented for of suprathreshold contrast levels.

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Figure 2. (a) Polarity selectivity of spatial contrast suppression. The matched contrast is plotted as a function of the physical contrast of the standard stimuli. The dashed line denotes the matched contrast equivalent to physical contrast. Filled circles show the results when the center and surround have the same polarities, and open circles show the results when they have opposite polarities. Red circles show results for a positive center, and blue circles show the results for the negative center. The left panel shows the average across observers and the right small panels show the results for individual observers. Error bars represent T1 SE. (b) The matched contrast relative to the physical contrast. Values larger than 1.0 indicate facilitation, and values smaller than 1.0 indicate suppression.

Control experiments contours, it is possible that the difference in the background In the above experiment, we equalized the mean lumi- luminance still had some other effects. nance between the center and surround regions to minimize We thus examined the contrast suppression using the effect of spatial luminance contrast across regions. stimuli with a constant background luminance regardless Because of this control, however, the background luminance of the combination of polarity. In this additional experi- within each region varied depending on the contrast of ment, the absolute contrast of the center was 0.5, and that texture elements, and this variation produced sharp lumi- of the surround was 1.0. The luminance of the element’s nance edges at the boundary between two regions. Although background was kept equal with the homogenous stimulus these edges were made invisible by presenting circular field (54 cd/m2). We found essentially the same results as

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for the main experiment. The suppression was observed effect of density by systematically manipulating spatial only for the same polarity stimulus (PSE 0.38, T0.04 SEM; separation between texture elements. 4 observers) but not at all for the opposite polarity stimulus (PSE 0.51, T0.02 SEM). In the other additional experiment, we also used textures consisting of isotropic Methods Difference-of-Gaussian (DoG) patterns with standard deviations of 0.07 deg (center Gaussian) and 0.18 deg The stimulus was a center–surround texture field as (surround Gaussian). The absolute contrast of the center used in Experiment 1. In order to control the element was 0.3, and that of the surround was 1.0. We found again density, we varied the minimum separation between that suppression occurred only with the same polarity elements for 0.29, 0.36, 0.50, and 0.64 deg (Figure 3). stimulus (PSE 0.22, T0.02 SEM; 4 observers) and not with The contrast of the central field was 0.5, and that of the the opposite polarity stimulus (PSE 0.30, T0.02 SEM). surround was 1.0. Four observers (HS, KA, MH, and RY) These results indicate that the effect of luminance differ- participated in the experiment. The mean luminance of the ence produced by equalization was not the main source of homogenous field was 54 cd/m2 for all observers. The contrast suppression found in the main experiment. In mean luminance was equalized between the center and addition, the results with circular texture elements (DoG surround regions. The methods are the same except for patterns) indicate that the polarity selectivity does not these points. depend on whether textures consist of oriented or circular elements. Results

Figure 4 shows matched contrast relative to physical Experiment 2 contrast as a function of density. When texture had relatively high density (inter-element separations of 0.29 and 0.36 deg), perceived contrast was suppressed The results of Experiment 1 showed a polarity selectiv- regardless of the combination of polarities. When the center ity for contrast suppression. As previously mentioned, had relatively low density (0.50 and 0.64 deg), however, these results are not consistent with those of Solomon suppression occurred only when the center and surround et al. (1993). The disagreement could be caused by the had the same polarities. We further examined the effect difference in the stimulus. While we used sparse textures, by using a dense texture (inter-element separation of Solomon et al. used dense textures. We thus examined the 0.29 deg) with relatively low contrast (center and surround

Figure 3. Examples of stimuli used in Experiment 2. The center-to-center separation between adjacent elements was 0.29, 0.36, 0.50, and 0.64 deg from the left, respectively. The absolute contrast of the center was 0.5, and that of the surround was 1.0 under all conditions. The mean luminance was equalized between the center and surround regions.

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Figure 4. Effect of texture element density. The matched contrast relative to the physical contrast (0.5) is plotted as a function of the distance between texture elements. Values larger than 1.0 indicate enhancement and values smaller than 1.0 indicate suppression. The other specifications of the plots are the same as Figure 2b.

contrasts were 0.125 and 0.25) and obtained essentially the Methods same results (PSE 0.09, ÈT0.01 SEM for same polarity; PSE 0.09, ÈT0.01 SEM for opposite polarity; 2 observers). The stimulus was a texture composed of oriented blobs as Therefore, the lack of polarity selectivity at high density used in the previous experiments. The orientation of observed in the main experiment is not due to the high elements in the center was always vertical, while the contrast used there. orientation of surround was tilted clockwise or counter- Spatial suppression seems to be polarity selective for clockwise by 0, 30, 60, or 90 deg (Figure 5). The contrast of textures of sparse elements but not for textures of dense the central field was 0.5, that of the surround was 1.0, and elements. The results indicate that lack of polarity selectivity the inter-element separation between elements was 0.64 or in the contrast–contrast reported in the previous study 0.36 deg. Three observers (HS, KA, and MH) participated (Solomon et al., 1993) is mainly caused by the use of high- in the experiment. The mean luminance of the homogenous density textures. In addition, we found in our pilot field was 54 cd/m2 for all observers. Mean luminance was observations that the suppression is substantially weakened equalized between the center and surround regions. The when the center and surround had different element sizes or methods except for these points were the same as in densities. This implies a selectivity for spatial frequency Experiment 1. The data for bilaterally symmetric conditions and/or texture density of the polarity-dependent suppression. (e.g., 30 deg and j30 deg) were pooled in the analysis.

Results Experiment 3 Figure 6a shows the matched contrast relative to the physical contrast as a function of the center–surround The results of the above experiments show that contrast orientation difference, as obtained for low-density textures. suppression is specific to luminance polarity, and this When the center had the same polarity as the surround, the polarity selectivity is not evident for textures with high suppression slightly decreased as the orientation difference element densities. It is known that contrast suppression increased. The suppression occurred even with orthogo- depends on the difference in orientation between center nally oriented elements. As we fitted a Gaussian function in and surround elements (Petrov et al., 2005; Solomon et al., the orientation domain to the power spectrum of a texture 1993; Xing & Heeger, 2000). However, it is still not clear with sparse elements (at the peak spatial frequency), the how orientation dependency relates to polarity selectivity. estimated standard deviation of a Gaussian was È20 deg. It Solomon et al. (1993) examined both orientation and is thus unlikely that the suppression at the orthogonal polarity dependency, but their stimuli were composed of condition (94 sigma distant) was caused by the overlap in high-density elements. Hence, it remains possible that the orientation energy of elements. On the other hand, for orientation dependency is peculiar to dense stimuli. the condition that the center had an opposite polarity to the Thus, in this experiment, we examined whether contrast surround, little or no suppression was observed over all suppression with sparse stimuli is also orientation selective. orientation differences. Figure 6b shows the results for

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Figure 5. Examples of stimuli used in Experiment 3. Orientation of the surround elements was 0, 30, 60, and 90 deg from the left, respectively. The center-to-center separation between adjacent elements is 0.64 deg in these examples.

high-density texture. The suppression occurred regardless threshold level, but those of off units would not. As a of the center–surround polarity combinations and slightly result, this texture selectively activates on units only. On decreased as the center–surround orientation difference the other hand, when the elements are densely placed, the increased. filtered outputs of off units to the background in between the bright elements can also exceed the threshold. As a result, a high-density texture inevitably activates both on and off units. This would be one of the main reasons for Discussion why we and Solomon et al. (1993) did not find polarity selectivity for high-density textures. A clear polarity selectivity in the contrast–contrast The present study shows that spatial interactions in the (2nd-order) effect may be considered as a class of brightness/ contrast–contrast effect depend on the congruency of darkness (1st-order) contrast between the elements. That is, luminance polarity between the center and surround texture the apparent brightness of each element in the center is regions. The perceived contrast of the central texture was reduced by high luminance incremental elements in the markedly reduced (up to 30–40%) by a surrounding texture surround, it would cause a reduction in apparent contrast. with the same luminance polarity but not at all by a texture Likewise, if the high luminance incremental surround blobs with the opposite polarity. The results support the idea that reduce the brightness of the decremental luminance test field contrast–contrast is largely mediated by independent elements, this would cause an increase in apparent contrast spatial interactions between visual units sensitive to on- (or at least would not produce a contrast reduction). The lack and off-type contrast rather than interactions between units of polarity selectivity at high density may reflect the sensitive to unsigned contrast. difficulty of segregating elements from the background and The present results are apparently inconsistent with a represent a shift from first-order to second-order induction previous study that showed no polarity selectivity of (see McCourt, 2005). However, it should be noted that this spatial interactions. However, the results of Experiment 2 interpretation hypothesizes the existence of an additional clearly show that this discrepancy is caused by a differ- process that segregates elements from background. There- ence in the density of texture elements. Polarity selectivity fore, we believe that the present results can be explained was evident for textures with sparse elements but absent more parsimoniously by assuming interactions between on- for textures with dense elements. This is a natural and off-type spatial filters. consequence of the spatial response profile of on- and We also found that the suppression effect is selective, off-type visual units, which are commonly modeled as though weakly, for orientation as well as for contrast band-pass spatial filters followed by half-wave rectifica- polarity. Similar weak orientation tunings have also been tion and threshold. Suppose there is a texture pattern reported for gratings and for line textures (Cannon & composed of bright elements on a dark background. When Fullenkamp, 1993; Solomon et al., 1993; Xing & Heeger, the elements are sparsely placed in the texture, the filtered 2000). Notably, the significant suppression remained when outputs of on units would be large enough to exceed the elements in the center had an orientation orthogonal to

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Figure 6. Orientation selectivity. The matched contrast relative to the physical contrast (0.5) is plotted as a function of the orientation difference between center and surround. Panels in (a) show the results for textures of low density and those in (b) show the results for textures of high density. The other specifications of the plots are the same as Figure 2b.

those in the surround. This seems to indicate the involve- ment of spatial suppression of both orientation-selective Conclusion and non-selective (isotropic) units. The critical role of polarity-selective isotropic units has also been proposed for The present study showed that the contrast–contrast texture discrimination (Motoyoshi & Kingdom, 2007; illusion is selective for luminance polarity except for Sharan, Adelson, Motoyoshi, & Nishida, 2007). This idea textures with very high density. The results indicate is consistent, at least partially, with the recent physiological separate contributions of on/off responses for the percep- findings that there are two types in surround suppressions tion of image contrast. of V1 neural activities, one based on interactions between orientation-tuned cortical neurons and the other based on inhibitory inputs from LGN neurons with broader orienta- tion tunings that operate when the center and surround have Acknowledgments the same contrast polarity (Soodak, Shapley, & Kaplan, 1987; Webb et al., 2005). The other possible neural This study was done when H.S. was a trainee at NTT substrate for the non-orientation-selective component is Communication Science Laboratories. The authors thank intrinsic interactions among cells in CO blobs of V1 (e.g., Dr. J. Solomon for helpful comments on the draft and Mr. Ts’o & Gilbert, 1988). T. Kusano for technical supports. A part of the results was

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