Induction of the McCollough effect I: Figural variables

The effect of line orientation and line configuration on the induction of orientation-specific negatively colored aftereffects was investigated in three separate studies. In the first study, subjects viewed magenta-and-black vertical gratings with one eye, alternating with green-and-black vertical gratings to the other. Monocular tests revealed complementary aftereffects in each eye which disappeared when the test patterns were viewed with both eyes together. In Study 2, imposing a single colored bar against a black background induced negatively colored aftereffects in a white bar against a black background and in a black-and-white grating, while imposing a single black bar against a colored background was ineffective. In Study 3, presenting a magenta square outline elicited green aftereffects in vertical and horizontal bars and gratings as well as in outlines of squares and diamonds, while pairing the magenta square with a green cross had no effect. It was concluded that the induction mechanism responsible for the McCollough effect is sensitive to line orientation but not to shape. This specificity appears incompatible with a simple conditioning model.     

Storage of spatially specific threshold elevation

The decay of several visual aftereffects may be prolonged by interposing a period of light-free or pattern-free viewing between adaptation and testing. We demonstrate that this storage phenomenon can be observed using the threshold elevation aftereffect that follows inspection of a high-contrast grating pattern. Control experiments comparing thresholds for vertical and horizontal gratings after adaptation to a vertical grating reveal that the stored aftereffect, like its unstored counterpart, is pattern-selective. Storage is equally pronounced with stimuli that are detected by pattern-analyzing or movement-analyzing visual channels. Unlike other aftereffects, the threshold-elevation aftereffect requires that the storage period be light-free; no storage is seen if a blank field is inspected between adaptation and testing. The results are discussed with respect to the nature of visual aftereffects, and possible cognitive or physiological models of storage.     

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.     

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.     

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.     

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.     

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.     

Effects of spatial filtering and lack of effects of visual imagery on pattern-contingent color aftereffects

Checkerboards contain fundamental two-dimensional Fourier components oriented 45° from the edges of individual checks. Previous studies have shown that contingent color aftereffects following adaptation to chromatic checkerboard stimuli were associated with the fundamental components rather than the edges, In the present experiments, we measured contingent color aftereffects, using the method of constant stimuli, after subjects adapted to unfiltered checkerboards and checkerboards with the fundamental Fourier components removed. The adaptation stimuli were magenta (or green) squares and green (or magenta) diamonds; the test stimuli were vertical or oblique sine-wave gratings with different saturations, After adaptation to unfiltered checkerboards, aftereffects contingent on the fundamental components were obtained. In contrast, after adaptation to filtered stimuli, aftereffects of smaller magnitude were found to be aligned with the edges. The data support the previous findings of spatial-frequency-contingent color after-effects with checkerboard adaptation stimuli and indicate that the aftereffects can be associated with edges if the fundamental components of adaptation stimuli are removed by spatial filtering. We reexamined the possibility of color aftereffects induced by imagery of checkerboards. Contrary to the previous reports, no significant aftereffects were obtained.     

Color aftereffects contingent on perceptual organization

The present experiment demonstrated that (a) cognitive organizational factors influence an orientation-contingent color (McCollugh) aftereffect, and (b) adaptation to colored gratings affected the perceptual organization of a reversible figure. Following adaptation to colored gratings, color aftereffects were reported in only one of two organizations of a reversible figure, and the colors conformed to the subjective contours of the test pattern rather than exclusively to the regions defined by horizontal and vertical pattern elements. In addition, an organization of the reversible figure that segregated vertical and horizontal pattern elements was dominant subsequent to adaptation.     

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.     

Variations in apparent spatial frequency with stimulus orientation: L Incidence of the effect in the general population

The human visual system is anisotropic not only for stimulus detection and discrimination but also for stimulus appearance. The apparent length of a stationary stimulus and the apparent velocity of a moving stimulus vary with orientation. These variations are predictable from a model assuming a compression of the horizontal spatial meridian relative to the vertical meridian. In this paper, the apparent spatial frequency of a suprathreshold grating is shown to depend on the grating orientation. Specifically, horizontal gratings appear coarser than vertical gratings of equivalent spatial frequency. Additional data collected from the same group of observers show that horizontal lines appear shorter than vertical lines of the same extent. The presence of the spatial frequency and length illusions, although both predictable from the compression model, were not found to be correlated within observers. Implications of these results for other data in the literature are discussed.     

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.     

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.     

Pattern-contingent color aftereffects on noninduced patterns

In a series of experiments, we found that in addition to expected reports of color aftereffects on patterns viewed during induction, reliable and predictable reports of color were given by subjects to patterns they did not view during induction. These reports to noninduced patterns were generally to patterns that were orthogonal to the patterns seen during induction. Induction with, for example, a red vertical grating led to appropriate aftereffects (i.e., green) on that vertical pattern and to the complementary aftereffect (i.e., pink) on a horizontal grating. We suggest that such color aftereffects on noninduced patterns are based on a shift in the activity of orientation coding mechanisms as a result of viewing the inducing patterns. We further propose that the results are consistent with the Lie transformation group theory of neuropsychology and that they add to a growing body of research demonstrating the applicability of this theory to the understanding of pattern-contingent color aftereffects.     

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.     

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.     

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.