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.     

The McCollough effect: Dissociating retinal from spatial coordinates

Three experiments were conducted to dissociate the perceived orientation of a stimulus from its orientation on the retina while inducing the McCollough effect. In the first experiment, the typical contingency between color and retinal orientation was eliminated by having subjects tilt their head 90° for half of the induction trials while the stimuli remained the same. The only relation remaining was that between color and the perceived or spatial orientation, which led to only a small contingent aftereffect. In contrast, when the spatial contingency was eliminated in the second experiment, the aftereffect was as large as when both contingencies were present. Finally, a third experiment determined that part of the small spatial effect obtained in the first experiment could be traced to hidden higher order retinal contingencies. The study suggested that even under optimal conditions the McCollough effect is not concerned with real-world properties of objects or events. Implications for several classes of theories are discussed.     

Loss of wavelength selectivity in contour masking and aftereffect following dichoptic adaptation

Two experiments measured the apparent orientation (aftereffect) and the threshold for detection (masking) of a colored grating viewed by one eye after exposure to a colored grating to the same or the opposite eye (monoptic inspection) or after stimulation of one eye by color and the other eye by contours (dichoptic inspection). Under the monoptic condition, the color relationship between the inspection and test stimuli exerted control over the extent of aftereffect and masking when the two stimuli were viewed with the same eye, but not when they were seen with different eyes. Aftereffect and masking were nonselective to wavelength following dichoptic inspection, irrespective of whether the test stimulus was presented to the color-adapted or to the contour-adapted eye. The results support other claims that visual detectors with chromatic and spatial tuning have monocular specificity.     

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.     

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.     

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.     

An Associative Interpretation of the Indirect McCollough Effect

Following an induction procedure in which a coloured grid is alternated with a square of a complementary colour, subjects report colour after-effects on both the grid orientation present during induction and the orthogonal non-induced grid orientation. The after-effect reported on the induced grid orientation is called the McCollough effect (ME). The aftereffect reported on the non-induced grid orientation is called the indirect ME. There is evidence that the ME represents an instance of Pavlovian conditioning. The present results support a conditioning interpretation of the indirect ME and are inconsistent with interpretations of the indirect ME that attribute the phenomenon to special orthogonal coding mechanisms within the visual system.     

Orientation constancy of the McCollough effect

Two different techniques were used to determine the orientation constancy of orientation-contingent color aftereffects, McCollough effects. The results from both investigations agree and indicate That the aftereffect fails to exhibit orientation constancy other than that which can be explained by ocular countertorsion. Thus, the retinal rather than phenomenal orientation of the adaptation stimuli appears to be the determinant of aftereffect orientation. In fact, it is concluded that the aftereffect can be used to accurately monitor torsional eye position over long periods of time.     

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.     

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.     

Isoluminance and contingent color aftereffects

In the typical induction of the orientation-contingent color aftereffect (CCAE), the stimuli are composed of elements that differ in both color and luminance. Three experiments are reported that show that chromatic contrast between stimulus elements is insufficient for the induction of the orientation-CCAE and that luminance contrast is necessary. These experiments expand on previous research concerned with the role of luminance contrast in the induction of orientation-CCAEs by eliminating alternative explanations.     

An oblique effect of chromatic gratings measured by color-mixture thresholds

Contrast sensitivity is lower for obliquely oriented achromatic gratings than for vertical or horizontal gratings at high spatial and low temporal frequencies. Although this response is suggestive of mediation by P-pathway cortical correlates, no clear sensory (i.e. class 1) oblique effect has been demonstrated with isoluminant chromatic stimuli. In the present experiment, a two-alternative forced-choice detection task was used to measure observers' sensitivity to spatiotemporal sinusoids varying in orientation and color contrast. A maximum-likelihood method fit ellipses to the thresholds, with the length of each ellipse taken as a measure of chromatic contrast sensitivity at isoluminance, and the width as luminance contrast threshold. A chromatic oblique effect was observed at about 3 cycles deg-1 suggesting an orientation bias within the cortical stream conveying P-cell activity.     

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.     

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.     

Presence and absence of color selectivity in the motion aftereffect

It is controversial whether the magnitude of the motion aftereffect is greater when both inspection and test stimuli are the same color rather than different colors (color selectivity). The present experiments show that the extent of color selectivity in the classical motion aftereffect depends upon (1) the duration of the interval between inspection and test, and (2) the nature of the stimulation during this interval. These findings are consistent with previous reports of two phases in the motion aftereffect and are interpreted in terms of the known properties of sustained and transient cells in the human visual system.     

Cholinergic agents and the McCollough effect

The strength of the McCollough effect (ME), a pattern-contingent colour aftereffect, has been shown to be inversely related to acetylcholine, being significantly strengthened by (anticholinergic) scopolamine and weakened by (cholinergic) physostigmine delivered before adapting to the ME stimuli. The purpose of the present study was (i) to establish whether the effect of pre-adaptation scopolamine is linearly dose-dependent and (ii) to investigate the effects of scopolamine and physostigmine delivered between adaptation and testing. In experiment 1, ten healthy male volunteers who received placebo, or 0.6 mg, 1.2 mg, or 1.8 mg scopolamine before adapting to ME stimuli showed a significant linear dose-dependence over tests repeated from 10 to 70 min after adaptation. In experiment 2 twelve male volunteers adapted to ME stimuli and then received placebo, 1.2 mg oral scopolamine, or 0.75 mg subcutaneous physostigmine. On subsequent repeated testing, strength of the ME was increased by scopolamine and decreased by physostigmine relative to placebo. Both experiments were double-blind double-dummy repeated measures. These data support the view that the ME is a product of inhibitory mechanisms in the visual system rather than processes involved in associative learning.     

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.     

Interocular suppression differentially affects achromatic and chromatic mechanisms

Results from a series of psychophysical experiments show that interocular suppression produced by continuous flash suppression (CFS) differentially affects visual features of a target viewed by the other eye. When CFS stimuli are defined by luminance contrast, target color can be reliably identified but percent-correct discrimination of target orientation is near chance. When the colored target is moving, color identification deteriorates with motion speed but direction of motion discrimination improves with target speed. Color’s immunity to suppression is also weakened when interocular suppression is induced by equiluminant CFS stimuli that presumably stimulate the chromatic pathway. These results are discussed in terms of functional segregation of achromatic and chromatic processing in the visual system.