Contrast and frequency competition for orientation-contingent color aftereffects

A “competition” paradigm was developed to examine separately the effects of pattern contrast and spatial frequency characteristics on the strength of orientation-contingent color aftereffects (McCollough effects). After adapting to alternately presented red/black and green/black square-wave gratings (one horizontal, one vertical), 11 subjects viewed seven different kinds of test patterns. Unlike Standard McCollough effect test stimuli, the present patterns had variable luminance profiles running both horizontally and vertically within each test pattern area. Forced choice responses were used to determine which aftereffect color (red or green) appeared, as characteristics of vertical and horizontal luminance profiles were varied separately among test stimulus types. We conclude that pattern contrast and human contrast sensitivity account for aftereffect colors in such stimuli. When contrast is taken into consideration, aftereffects are not predicted by similarity between adaptation and test pattern Fourier characteristics, nor are they predicted by the width, per se, of pattern elements.     

Color-luminance relationships and the McCollough effect

The McCollough effect is an orientation-specific color aftereffect induced by adapting to colored gratings. We examined how the McCollough effect depends on the relationships between color and luminance within the inducing and test gratings and compared the aftereffects to the color changes predicted from selective adaptation to different color-luminance combinations. Our results suggest that the important contingency underlying the McCollough effect is between orientation and color-luminance direction and are consistent with sensitivity changes within mechanisms tuned to specific color-luminance directions. Aftereffects are similar in magnitude for adapting color pairs that differ only in S cone excitation or L and M cone excitation, and they have a similar dependence on spatial frequency. In particular, orientation-specific aftereffects are induced for S cone colors even when the grating frequencies are above the S cone resolution limit. Thus, the McCollough effect persists even when different cone classes encode the orientation and color of the gratings.     

Color—luminance relationships and the McCollough effect

The McCollough effect is an orientation-specific color aftereffect induced by adapting to colored gratings. We examined how the McCollough effect depends on the relationships between color and luminance within the inducing and test gratings and compared the aftereffects to the color changes predicted from selective adaptation to different color—luminance combinations. Our results suggest that the important contingency underlying the McCollough effect is between orientation and color—luminance direction and are consistent with sensitivity changes within mechanisms tuned to specific color—luminance directions. Aftereffects are similar in magnitude for adapting color pairs that differ only in S cone excitation or L and M cone excitation, and they have a similar dependence on spatial frequency. In particular, orientation-specific aftereffects are induced for S cone colors even when the grating frequencies are above the S cone resolution limit. Thus, the McCollough effect persists even when different cone classes encode the orientation and color of the gratings.     

Absence of binocular interaction between spatial and color attributes of visual stimuli

The hypothesis that induction of the McCollough effect (spatially selective color aftereffects) entails adaptation of monocularly driven detectors tuned to both spatial and color attributes of the visual stimulus was examined in four experiments. The McCollough effect could not be generated by displaying contour information to one eye and color information to the other eye during inspection, even in the absence of binocular rivalry. Nor was it possible to induce depth-specific color aftereffects following an inspection period during which random-dot stereograms were viewed, with crossed and uncrossed disparity seen in different colored light. Masking and aftereffect in the perception of stereoscopic depth were also nonselective to color; in both cases, perceptual distortion was controlled by stereospatial variables but not by the color relationship between the inspection and test stimuli. The results suggest that binocularly driven spatial detectors in human vision are insensitive to wavelength.     

Establishment and decay of orientation-contingent aftereffects of color

We have used a null method to measure the orientation-contingent aftereffects of color first described by McCollough. After alternately inspecting, for example, a green horizontal line grating and a magenta vertical line grating, the Os report that in achromatic test gratings the horizontal lines appear pinkish and the vertical lines appear greenish. We have used a special color-mixing projector to add variable amounts of green and magenta light to the test gratings until they appear matched and nearly achromatic. The colorimetric purity needed to achieve this null setting is a quantitative measure of the strength of the colored aftereffect. Following inspections of the colored patterns ranging from 15 sec to 150 min, six Os showed aftereffects lasting from a few minutes to 7 or more days. The indices of colorimetric purity increase with inspection time and decline with time after inspection. The decay function is not quite linear either on semilog or on log-log coordinates. The rate of decay is mainly dependent on the magnitude of the effect built up during inspection. We conclude that the buildup and decay of these aftereffects show some of the time characteristics usually associated with central adaptability rather than sensory adaptation.     

Movement-induced modulation of orientation-specific color aftereffects

The intensity of the McCollough effect is modified when, following exposure to the inducing chromatic stimuli, the achromatic test gratings are seen oscillating orthogonally to their orientations. Green aftereffect seen on stationary test gratings is enhanced by oscillations, while pink aftereffect present on the stationary gratings fades upon oscillation of the test stimulus. These opponent changes are tentatively accounted for in terms of an interaction between Fechner-Benham type induced color and processes that mediate the orientation-specific chromatic aftereffects.     

Checkerboard-specific color aftereffects: A failure to find effects of perceptual organization

Two experiments investigated the effects of differing perceptual organizations of reversible figures on McCollough aftereffects. Experiment 1 used colored checkerboard inducing stimuli and achromatic grating test stimuli. While some subjects tended to organize the checkerboards into rows and/or columns and others to organize them into obliques, these variations did not result in differences in aftereffect direction or magnitude. Experiment 2 induced an aftereffect with colored gratings and tested with checkerboards, gratings, and a reversible concentric octagon pattern. Perceptual organization had no effect on results for checkerboards, but was related to aftereffect strength for the octagon pattern. Indirect evidence suggests that, in the latter case, differences in aftereffect strength may have influenced the perceived organization, rather than vice versa. Finally, regardless of the specific organization perceived, spontaneous viewing of all test stimuli produced stronger aftereffects than were found when subjects reorganized the pattern. This may have resulted from a viewing strategy associated with reorganization, since similarly small aftereffects were found when subjects concentrated their attention on a single pattern element.     

Perception of the Mccolloch Effect Correlates with Activity in Extrastriate Cortex: A Functional MagneticResonance Imaging Study

The McCollough effect is a striking color aftereffect that is linked to the orientation of the patterns used to induce it. To produce the McCollough effect, two differently oriented grating patterns, such as a red-and-black vertical grating and a green-and-black horizontal grating, are viewed alternately for a few minutes. After such colored gratings are viewed, the white sections of avertical black-and-white test grating appear to be tinged with green, and the white sections of a horizontal grating appear to be tinged with pink. We present evidence from a functional magnetic resonance imaging study that the perception of the McCollough effect correlates with increased activation in the lingual and fusiform gyriùextrastriate visual areas that have been implicated in color perception in humans.     

Orientation specificity in chromatic adaptation of human “edge-detectors”

The McCollough effect, a pattern-specific complementary hue aftereffect, has usually been demonstrated with horizontal and vertical adapting and test patterns. The present study measured the strength of the effect produced by adapting patterns of various angular separations. The effect decreased with decreasing angular separations until it was minimal at 11 deg of separation. The results are considered to be consistent with an edge-detector interpretation of this aftereffect. With vertical and horizontal adapting patterns, the reddish aftereffect was 17% and the greenish aftereffect 9%of colorimetric purity.     

Decline and revival of McCollough effects following inspection of achromatic gratings

We have investigated the reduction in estimated strength of McCollough effects that results from inspection of achromatic gratings. Longer inspection times were found to produce greater decreases in our assessments than shorter times. Following a rest period, increments in our strength index were observed, with greatest gains associated with shortest inspection times, and least gains with longest times. Our results can be construed as supportive of a learning model of McCollough effects, or, alternatively, as suggesting the existence of achromatic McCollough effects.     

The role of some spatial parameters of gratings on the McCollough effect

Ss were alternately adapted to vertical and horizontal gratings that consisted of black bars and colored slits. The slits of one grating were green and of the other, magenta. The widths of the black bars and the colored slits were varied independently during adaptation and testing. This design separates the relative influence of bar width, slit width, and spatial frequency on an orientation specific color aftereffect known as the McCollough effect. Black bar width had the major influence on the strength of the aftereffect, suggesting that the neurophysiological mechanism underlying the McCullough effect might consist of orientation specific units that are sensitive to both the widths of black bars and the chromatic characteristics of their surrounds.     

The indirect McCollough effect: An examination of an associative account

It has been found that viewing, for example, a red-and-black vertical grating alternating with a green homogeneous field produces a color aftereffect—a McCollough effect—on a black-and-white vertical grating (i.e., green). Viewing such colored patterns also produces an aftereffect on a noninduced horizontal grating (i.e., pink)—the indirect McCollough effect. Humphrey, Dodwell, and Emerson (1989) argued that the indirect McCollough effect is caused by opponent properties of the visual system that organize the processing of contour and color along contrasting, probably orthogonal, dimensions. Recently, however, their interpretation of the indirect McCollough effect has been challenged by some findings of Eissenberg, Allan, Siegel, and Petrov (1995). These researchers have proposed that the indirect McCollough effect, like the McCollough effect, can be explained by associative principles. The results reported here question crucial aspects of the hypothesis of Eissenberg et al.     

Orientation-specific luminance aftereffects

Prolonged viewing of bright vertical (horizontal) gratings alternating with dim horizontal (vertical) gratings generates negative brightness aftereffects that are contingent on the orientation of orthogonal test gratings. The effect is measured by a brightness cancellation technique, similar to the color cancellation technique used in measuring McCollough effects. Like the latter, brightness aftereffects appear to persist for long periods. The magnitude of these aftereffects is a positive monotonic function of the luminance difference between the inducing gratings, and it depends on the conditions of induction; monocular induction generates larger aftereffects than binocular induction does. The aftereffect transfers interocularly, although its magnitude in the contralateral eye is substantially attenuated; binocular measurement, following monocular induction, results in even smaller aftereffects. An attempt to understand these findings within the computational model of brightness perception developed by Grossberg and Mingolla (1985a, 1985b) is presented.     

Orientation selectivity of the McCollough. effect: Analysis by equivalent contrast transformation

Using a color-cancellation technique, the strength of the McCollough effect was measured in units of excitation purity. The strength was studied both as a function of the contrast of the adapting gratings and as a function of the angle × between the axes of the test and the adapting gratings. Results were well described as a linear function of the contrast of the adapting gratings and as a cos(2×) function of the angle. Both functions were combined to express an equivalent contrast transformation which converts the measurements of orientation tuning into a unit comparable to that used for other kinds of orientation-specific aftereffects. The orientation tuning was found to be very broad with a half-width at half amplitude of approximately 27°. This estimate is considered to be a substantial underestimate of the actual tuning of the aftereffect’s substrate.     

Spatial contingency and the McCollough effect

On the basis of a conditioning analysis of the orientation-contingent color aftereffect (McCollough effect, ME), orientation stimuli become associated with simultaneously presented chromatic stimuli. This account suggests that decreasing the contingency between the grid orientation and color should decrease the strength of the aftereffect. Results of previous research indicate that decreasing the temporal contingency (by presenting homogeneous chromatic stimuli between presentations of chromatic grids) does not decrease the ME. However, it has been suggested that the appropriate contingency-degradation procedure would involve decreasing spatial (rather than temporal) contingency. That is, the illusion should be attenuated by extending the color beyond the confines of the grid. Contrary to this hypothesis, the results of the present experiments provide no evidence that decreasing the spatial contingency between grid and color decreases the ME; rather, the aftereffect is increased by such a manipulation.     

“Extinction” of the McCollough effect does not transfer interocularly

A McCollough effect was induced in subjects by having them view typical adapting stimuli binocularly for 5 min. In the control condition, the strength of the McCollough effect was measured 20 min after the end of the adaptation. The strength was measured during monocular and binocular viewing of a test pattern via a color cancellation technique. Monocular strengths for the two eyes of a given subject were equal to each other and slightly weaker than the binocular strength. In the test condition, 15 min of the 20 min between adaptation and testing were spent monocularly viewing black and white gratings of the same orientation and spatial frequency as the adapting gratings. The strength of the effect as measured ipsilaterally was markedly decreased from that in the control condition. The strength of the effect as measured with the contralateral eye showed only a small decrease from that of the control condition. This finding is relevant to various models of the McCollough effect and related color aftereffects, especially those that posit a “learning” type of mechanism between achromatic spatial channels (which exhibit clear interocular transfer of various achromatic effects) and monocular color channels.     

Hue difference contours can be used in processing orientation information

In most studies of orientation processing, chromatic information and achromatic information have been combined or confounded. The present experiments investigated the relative sizes of tilt aftereffect induced by these two types of information. In these experiments, the tilt aftereffect is the error in adjusting a test contour to vertical, following the scanning of an inspection contour. For inspection and test contours identical except for orientation, the tilt aftereffects varied with inspection contour orientation but not with chromatic or achromatic condition. Smaller tilt aftereffects were obtained when the inspection contour was produced by a hue difference (chromatic information) and the test contour was produced by a luminance difference (achromatic information), or vice versa. These results indicate that achromatic and chromatic information is processed in a similar manner with respect to orientation. Furthermore, there is substantial, but incomplete, pooling of chromatic and achromatic orientation information.     

Luminance as a factor in the ability of achromatic gratings to interfere with MeCollough effects

McCollough effects were reduced in assessed strength by having observers view achromatic gratings that varied in space-average luminance. Whether the gratings were presented before or after induction of McCollough effects, high luminance gratings interfered with the effects more than did low luminance gratings. If considered within the context of a conditioning model, this finding may be interpreted as inconsistent with other characteristics of McCollough effects.     

Brightness selectivity in the motion aftereffect

Eighteen Ss were required to track the apparent motion of a stationary grating viewed after prolonged inspection of a moving grating. Measures were obtained with the inspection and test gratings identical in contrast but different in space-average luminance, or with luminance held constant and contrast varied. The aftereffect was reduced as the gratings differed in space-average luminance, but contrast exerted less uniform influence as a variable. Brightness-selectivity in the motion aftereffect is interpreted within the selective adaptation model of aftereffects as evidence that some detectors in human vision are conjointly tuned to space-average luminance and image motion.     

Induction of the McCollough effect II: Two different mechanisms

An orientation-specific chromatic aftereffect was observed when a single colored grating was used as an induction stimulus. The magnitude of the aftereffect was compared to that obtained when alternating orthogonal gratings in complementary hues were used as induction stimuli. The two-stimulus condition produced a stronger aftereffect than a single-stimulus condition. This facilitation was also obtained when a colored plain square with no grating was substituted for the second colored grating in the two-stimulus condition. The results suggest that the McCollough effect involves adaptation of two different mechanisms, one which is orientation-specific and one which is not.