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.     

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.     

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.     

Curvature-specific color aftereffects

Adaptation to convex and concave arcs in different colored light results in curvature-specific color aftereffects when arcs are later viewed in white light. In three experiments, it was shown that these color aftereffects often are partial (restricted to limited regions of the test arcs) rather than uniform, and in addition that aftereffects induced by exposure to arcs transfer to straight-line displays of particular orientation, and vice versa. These data were interpreted as evidence that arcs are processed in the visual system in relation to the orientations of local straight line approximations instead of on a global basis. In these terms, curvature-specific color aftereffects are merely complex forms of the orientation-specific color aftereffects first described by McCollough (1965).     

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.     

“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.     

The quantitative measure of pattern representation in images using orientation-specific color aftereffects

A quantitative method is developed for assessing the quality of pattern information in imagery, using the magnitude of color aftereffects as an objective index. Subjects were given instructions to project imagined bar patterns of particular width and orientation onto adapting color fields, in such a manner as to simulate standard conditions for establishing the McCollough effect. Our control procedures indicate that the resulting orientation-specific complementary color aftereffects cannot be attributed to the conditioning of particular directions of eye scanning movements to color processing during adaptation, or to other possible sources of experimental bias. Furthermore, subjects who rated themselves prior to the adaptation procedure as having relatively vivid imagery showed significantly larger aftereffects than those who reported having relatively low imagery. These results not only provide an important confirmation of our earlier finding that imagination can replace physical pattern information in the formation of basic color-feature associations in the human visual system, but also demonstrate that these aftereffects can provide a practical measure of the fidelity of pattern representation in visual images.     

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.     

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.     

Conjunction of color and form without attention: evidence from an orientation-contingent color aftereffect

According to feature-integration theory (Treisman & Gelade, 1980), separable features such as color and shape exist in separate maps in preattentive vision and can be integrated only through the use of spatial attention. Many perceptual aftereffects, however, which are also assumed to reflect the features available in preattentive vision, are sensitive to conjunctions of features. One possible resolution of these views holds that adaptation to conjunctions depends on spatial attention. We tested this proposition by presenting observers with gratings varying in color and orientation. The resulting McCollough aftereffects were independent of whether the adaptation stimuli were presented inside or outside of the focus of spatial attention. Therefore, color and shape appear to be conjoined preattentively, when perceptual aftereffects are used as the measure. These same stimuli, however, appeared to be separable in two additional experiments that required observers to search for gratings of a specified color and orientation. These results show that different experimental procedures may be tapping into different stages of preattentive vision.     

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.     

Characteristics of the indirect McCollough effect

Induction of contingent color aftereffects with a single chromatic grid sometimes results in an illusory color on a grid different from the one presented during induction. Such illusory color, contingently elicited by a noninduced grid, has been termed the indirect McCollough effect (indirect ME). We show that the indirect ME occurs only when the color complementary to the grid color is present during induction (either physically present or as a color afterimage), and that the indirect ME is seen only on gratings that are orthogonal to the induction orientation. These findings are in accord with the account of the indirect ME proposed by Humphrey, Dodwell, and Emerson (1989). We also show that characteristics of the indirect ME (seen following one-grid induction), both on induced and orthogonal orientations, are similar to those observed with the direct ME (seen following the usual two-grid induction procedure). Both procedures result in contingent aftereffects that display substantial retention and that do not display interocular transfer.     

Characteristics of the indirect McCollough effect.

Induction of contingent color aftereffects with a single chromatic grid sometimes results in an illusory color on a grid different from the one presented during induction. Such illusory color, contingently elicited by a noninduced grid, has been termed the indirect McCollough effect (indirect ME). We show that the indirect ME occurs only when the color complementary to the grid color is present during induction (either physically present or as a color afterimage), and that the indirect ME is seen only on gratings that are orthogonal to the induction orientation. These findings are in accord with the account of the indirect ME proposed by Humphrey, Dodwell, and Emerson (1989). We also show that characteristics of the indirect ME (seen following one-grid induction), both on induced and orthogonal orientations, are similar to those observed with the direct ME (seen following the usual two-grid induction procedure). Both procedures result in contingent aftereffects that display substantial retention and that do not display interocular transfer.     

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.     

Monocular-contingent and binocular-contingent aftereffects

Alternate monocular and binocular exposure to complementary stimulation can yield opposite but coexisting aftereffects that are contingent on whether the test display is viewed with one eye or two eyes. The motion aftereffect was studied by adapting each eye separately to a contracting spiral and both eyes together to an expanding spiral. The stationary test spiral subsequently appeared to be expanding when viewed monocularly, but to be contracting when it was seen with both eyes open. With respect to the McCollough effect, after monocular exposure to red-vertical and green-horizontal gratings and binocular exposure to red-horizontal and green-vertical gratings, the appearance of the color of the test gratings when viewed with one eye was different from that when viewed with both eyes. Opposite, coexisting aftereffects induced by complementary stimulation can be interpreted as evidence that there are unique binocular aspects to visual function.     

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.     

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.     

The associative basis of contingent color aftereffects.

According to a conditioning analysis of the orientation-contingent color aftereffect (McCollough effect, ME), orientation stimulus (grids) become associated with color. Contrary to this interpretation are reports that simple forms cannot be used to elicit illusory color and that the ME is not degraded by decreasing the grid-color correlation. The present results indicate: (a) Form stimuli can contingently elicit color aftereffects; (b) even a non-patterned stimulus--the lightness of a frame surrounding a colored area--can contingently elicit color aftereffects; (c) this frame lightness-contingent aftereffect, like the ME, persists for at least 24 hr; and (d) the frame lightness-contingent aftereffect can be used to demonstrate that correlational manipulations affect the ME, as they affect other types of conditional responses.     

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.