Spatial-frequency-contingent color aftereffects: adaptation with one-dimensional stimuli.

The McCollough effect was shown to be spatial-frequency selective by Lovegrove and Over (1972) after adaptation with vertical colored square-wave gratings separated by 1 octave. Adaptation with slide-presented red and green vertical square-wave gratings separated by 1 octave failed to produce contingent color aftereffects (CAEs). However, when each of these gratings was adapted alone, strong CAEs were produced. Adaptation with vertical colored sine-wave gratings separated by 1 octave also failed to produce CAEs, but strong effects were produced by adaptation with each grating alone. By varying the spatial frequency of the test sine wave, CAEs were found to be tuned for spatial frequency at 2.85 octaves after adaptation of 4 cycles per degree (cpd) and at 2.30 octaves after adaptation of 8 cpd. Adaptation of both vertical and horizontal sine-wave gratings produced strong CAEs, with bandwidths ranging from 1.96 to 2.90 octaves and with lower adapting contrast producing weaker CAEs. These results indicate that the McCollough effect is more broadly tuned for spatial frequency than are simple adaptation effects.     

Spatial-frequency-contingent color aftereffects: Adaptation with one-dimensional stimuli

The McCollough effect was shown to be spatial-frequency selective by Lovegrove and Over (1972) after adaptation with vertical colored square-wave gratings separated by 1 octave. Adaptation with slide-presented red and green vertical square-wave gratings separated by 1 octave failed to produce contingent color aftereffects (CAEs).However, when each of these gratings was adapted alone, strong CAEs were produced. Adaptation with vertical colored sine-wave gratings separated by 1 octave also failed to produce CAEs, but strong effects were produced by adaptation with each grating alone. By varying the spatial frequency of the test sine wave, CAEs were found to be tuned for spatial frequency at 2.85 octaves after adaptation of 4 cycles per degree (cpd) and at 2.30 octaves after adaptation of 8 cpd. Adaptation of both vertical and horizontal sine-wave gratings produced strong CAEs, with bandwidths ranging from 1.96 to 2.90 octaves and with lower adapting contrast producing weaker CAEs. These results indicate that the McCollough effect is more broadly tuned for spatial frequency than are simple adaptation effects.     

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.     

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.     

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.     

Further studies of the McCollough effect

Following prolonged viewing of black and white striped pattems in colored light, red and green aftereffects that lasted as long as 3 days were seen on the patterns, illuminated with white light. Altemate exposures of a vertical pattern of stripes in green light and a horizontal in white light (or a vertical in white light and a horizontal in red light) produced a red aftereffect on the vertical pattern and a green on the horizontal. The red and green aftereffects were also produced with a single vertical pattern. Adaptation colors that were at all greenish produced a red aftereffect on a vertical pattern and a green on a horizontal, whereas colors that were at all reddish produced a green aftereffect on a vertical pattern and a red on a horizontal. Colors near pure blue and pure yellow, which had little red or green content, produced weak aftereffects. The saturation of the aftereffects on the vertical grating varied in proportion to the red or green content of the adaptation color. Vivid red and green aftereffects were frequently obtained with the vertical and horizontal adaptation patterns paired with colors that closely bracketed pure yellow or pure blue. In all cases, the aftereffects gradually desaturated as the head was gradually tilted down to the side; the colors on each test pattern, vertical and horizontal, vanished at 45-deghead tilt and reversed beyond 45 deg.     

Frequency specific chromatic adaptation in the human visual system

Subjects were exposed to a vertical chromatic grating alternating with horizontal chromatic grating of the identical frequency. They were then tested with a series of test gratings of varied spatial frequencies to examine whether the responses were effected by the spatial frequency of the adaptation pattern in relation to the test pattern. It was found that maximum response occurred when adaptation and test gratings had the same spatial frequency, the effect was asymmetric and finally that enhancements were found at octaves. Thus the experiment further demonstrated that neural elements specific to spatial frequency exist in the human visual system.     

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.     

Overshadowing and blocking of the orientation-contingent color aftereffect: Evidence for a conditioning mechanism

Orientation-contingent color aftereffects have been interpreted by nonassociative mechanisms (adaptation of neural units that are both color and orientation specific) and by associative mechanisms (conditioning resulting from the pairing of pattern and hue). To evaluate associative accounts, contingent aftereffects were induced by exposing subjects to compound chromatic grid patterns consisting of two component gratings: one was horizontal or vertical, and the other a left- or right-learning diagonal. The ability of a component grating to elicit a color aftereffect depended on the relative salience and the aftereffect training history of the grating components. That is, orientation-contingent color aftereffects, like other conditional responses, display overshadowing and blocking. The results suggest that conditioning contributes to these aftereffects.     

Orientation-contingent color aftereffects mediated by subjective transparent structures.

We examined whether the orientation-contingent color aftereffect (the McCollough effect) could be mediated by subjective horizontal and vertical structure induced by the perception of transparency. In our experiments, red vertical bars and green horizontal bars were alternated as an adapting stimulus. After adaptation, subjects (n = 6) were asked to adjust the green and red saturation of a test pattern until they obtained a neutral gray. Horizontal and vertical stripes were combined in the test pattern in three different ways: (1) overlapping with a luminance combination that gave rise to a perception of transparent overlays of horizontal and vertical stripes (valid transparency condition), (2) overlapping with luminance combinations that did not induce a perception of transparency (invalid transparency condition) and that appeared more as a patchwork of checks, and (3) presented in adjacent, nonoverlapping areas. Our results showed that the McCollough effect was significantly greater in the valid transparency condition than in the invalid transparency conditions. The effect in the valid transparency condition was nevertheless less strong than was the effect seen with the standard test stimulus made up of nonoverlapping vertical and horizontal stripes. Our results suggest that the McCollough effect can be mediated by the subjective spatial organization (inner representation of vertical and horizontal stripes) that accompanies the perception of transparency in our stimulus.     

Orientation-contingent color aftereffects mediated by subjective transparent structures

We examined whether the orientation-contingent color aftereffect (the McCollough effect) could be mediated by subjective horizontal and vertical structure induced by the perception of transparency. In our experiments, red vertical bars and green horizontal bars were alternated as an adapting stimulus. After adaptation, subjects (n=6) were asked to adjust the green and red saturation of a test pattern until they obtained a neutral gray. Horizontal and vertical stripes were combined in the test pattern in three different ways: (1) overlapping with a luminance combination that gave rise to a perception of transparent overlays of horizontal and vertical stripes (valid transparency condition), (2) overlapping with luminance combinations that did not induce a perception of transparency (invalid transparency condition) and that appeased more as a patchwork of checks, and (3) presented in adjacent, nonoverlapping areas. Our results showed that the McCollough effect was significantly greater in the valid transparency condition than in the invalid transparency conditions. The effect in the valid transparency condition was nevertheless less strong than was the effect seen with the standard test stimulus made up of nonoverlapping vertical and horizontal stripes, Our results suggest that the McCollough effect can be mediated by the subjective spatial organization (inner representation of vertical and horizontal stripes) that accompanies the perception of transparency in our stimulus.     

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-specific aftereffects and illusions in the perception of brightness

Orientation-specific brightness aftereffects were found when vertical and horizontal gratings of the same space-average luminance were viewed following alternate exposure to vertical and horizontal gratings that differed in space-average luminance. The vertical test grating appeared bright following exposure to a dim vertical grating, and dim after a bright vertical grating had been viewed. This aftereffect did not occur when the adaptation gratings had been seen by one eye and the test gratings by the other eye. An orientation-specific illusion in the perception of brightness was also found, with the white sectors of a vertical grating appearing brighter against a background of horizontal lines than they did against a background of vertical lines. Both distortions imply that there are detectors in the human visual system that are conjointly tuned to luminance and contour orientation.     

Outflow and inflow in movement duplication

Following prolonged exposure to two vertical grating patterns differing in spatial frequency—one pattern illuminated in green light alternated with the other pattern illuminated in red light—human observers will sometimes report seeing desaturated complementary colors when presented with a neutrally illuminated test field consisting of adjacent halves of the two adapting gratings. The number of such color reports increases as the difference between the spatial frequencies of the adapting gratings increases. This frequency-specific chromatic aftereffect is similar to that obtained with orientation-specific color adaptation and may be mediated by neural “channels,” sensitive to both color and frequency input, which are similar to units known to exist in the visual systems of lower organisms.     

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.     

Pattern alternation: Effects of spatial frequency and orientation

Campbell and Howell (1972) reported an effect called “monocular pattern alternation.” They found that a pattern composed of two orthogonal sinusoidal gratings, one horizontal and the other vertical, underwent rivalry when viewed monocularly for a period of time. In the present study, it has been shown that monocular pattern alternation depends upon the orientation of the pattern and the spatial frequency of its components. Fewer reversals were found for an obliquely oriented pattern than for a pattern with components in the horizontal and vertical meridians. Alternation rate was higher when the gratings were similar in frequency but differed in orientation than when the components of the pattern differed in both dimensions. It was concluded that pattern alternation reflects an antagonistic interaction between interdependent channels in the human visual system that respond to orientation and spatial frequency.     

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.     

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.     

The effects of adaptation to square-wave gratings as a function of grating orientation

An adaptation technique was used to measure the selectivity or tuning for grating orientation in the visual system for different orientations of the inspection stimulus. Duration thresholds for grating patterns of constant luminance were determined for 13 test gratings oriented from ±5 to 90 deg away from each of five adaptation gratings: 0, 22, 45, 67, and 90 deg. Threshold data obtained for test gratings without prior adaptation indicated higher sensitivity for gratings oriented along the horizontal and vertical axis than along the oblique axis. After adaptation, thresholds increased (sensitivity was reduced) for gratings having similar orientations as the test gratings. However, the functions relating sensitivity reduction to degree of angular disparity between test and adaptation grating did not vary across the five inpsection orientations, i.e., selectivity or tuning for grating orientation appeared to be independent of the orientation of the adapting stimulus.     

A test of the spatial-frequency explanation of the Müller-Lyer illusion

Previous investigations have shown that the response of spatial-frequency-specific channels in the human visual system is differentially affected by adaptation to gratings of distinct spatial frequencies and/or orientations. A study is reported of the effects of adaptation to vertical or horizontal gratings of a high or a low spatial frequency on the extent of the Brentano form of the Müller-Lyer illusion in human observers. It is shown that the illusion decreases after adaptation to vertical gratings of low spatial frequency, but seems unaffected otherwise. These results are consistent with the notion of visual channels that are spatial-frequency and orientation specific, and support the argument that the Müller-Lyer illusion may be due primarily to lower-spatial-frequency components in the Fourier spectra of the image.