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.     

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.     

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.     

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.     

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.     

Mystery of the anti-McCollough effect

The McCollough Effect (ME) is a complex perceptual aftereffect that remains of interest half a century after its discovery. It is argued that a recently reported variant, dubbed the anti-McCollough effect, is not the reverse of the ME, with aftereffect colors in the same direction as the inducing stimuli. A red-horizontal stimulus leads to a reddish aftereffect not because of red-horizontal parings, but despite them. The anti-ME is a weak standard-direction ME produced by complementary afterimage colors (afterimage green with horizontal), rather than by environmental colors, first shown decades ago. It is not a new type of contingent aftereffect. The red-horizontal pair does not interfere with the afterimage green-horizontal pair it produces because a single color-orientation pairing provides more ambiguous input than does the standard two orientation-color pairings (red-horizontal, green-vertical) of the ME. It is also argued that not even one orientation-contingent color aftereffect is convincingly shown in the "anti"-ME, let alone, as has previously been suggested, two simultaneous orientation-contingent color aftereffects in opposite directions at different levels of the visual system, in which the higher-level effect suppresses the downstream effect from reaching consciousness. The "anti"-ME can be explained by existing theories of contingent aftereffects, including perceptual-learning theory.     

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.     

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.     

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.     

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.     

利用反诱导消除McCollough效应

ＭｃＣｏｌｌｏｕｇｈ效应在完全黑暗和自然视觉条件下的消退过程很缓慢,而利用同频补色条栅来进行反诱导,能在十几分钟内消除显色效应。消除所用时间与反诱导条栅的亮度、颜色和栅条间距（空间频率）有关。进一步说明导致ＭｃＣｏｌｌｏｕｇｈ效应恢复平衡的主要因素跟网膜刺激方式有关。     

Color aftereffect contingent on text

During adaptation, two different letter strings (each five or six letters) were presented to subjects alternately, one in green and the other in magenta. The extent to which these letter strings subsequently elicited a color aftereffect was assessed. In different experiments, the chromatic letter strings consisted of words and nonwords. The results indicated that letter strings that form English words can contingently elicit a color aftereffect. This was the case even when the words were anagrams. There was no evidence that nonword letter strings could contingently elicit such an aftereffect, even when the nonwords conformed to English orthography. The results are relevant to understanding other contingent color aftereffects (McCollough effects), illusory color noted by computer operators who work at monochrome (green or amber) displays, and the processing of text.     

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.     

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.     

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.     

Rapid discrimination of McCollough effects.

The McCollough effect is a colour aftereffect that is contingent on pattern orientation. Three experiments were conducted to establish whether such aftereffect colours could serve as a basis for discrimination in several rapid discrimination tasks. In the first experiment it was investigated whether aftereffect colours could act like a simple 'feature' in a visual search task involving a difficult orientation discrimination. Without McCollough adaptation, the time taken to detect a 'target' among 'distractors' increased substantially as the number of distractors increased. With adaptation, detection time was essentially independent of the number of distractors, indicating that the nature of the task changed from a difficult orientation discrimination to a simple discrimination based on differences in aftereffect colours. The second and third experiments employed a difficult four-alternative forced-choice procedure in which subjects were required to discriminate a monochromatic patch of square-wave grating oriented at 45 degrees from three others oriented at 135 degrees (and vice versa). The gratings were presented very briefly (67-333 ms) followed by a 500 ms mask. Subjects performed the task with and without McCollough adaptation. Performance was strikingly better after adaptation: colour aftereffects could be used to make the discrimination even at exposure durations as short as 67 ms. The third experiment demonstrated that this enhanced performance was indeed due to perceived colour differences (rather than a possible contrast difference). The results of the three experiments are discussed in relation to proposals about the locus of the McCollough effect.     

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.     

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.