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.     

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.     

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.     

Contour specificity of the McCollough effect

Subjective estimates of McCollough aftereffect strength are significantly reduced when certain spatial features of the line grating patterns are manipulated. Results are dependent upon whether the spatial parameters of the test or inspection patterns are altered. Changing the angular slant, contour sharpness, or contour completeness of the inspection gratings does not affect aftereffect strength, but changing the spatial frequency, contour sharpness, or contour completeness of the test gratings does. The implications of these results are discussed with regard to theories offered to explain the McCollough effect.     

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.     

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.     

Negative afterimages and the McCollough effect

Four experiments were conducted to test earlier claims about the relationship between the negative afterimage and the McCollough effect. The first claim (Hansel & Mahmud, 1978) is that long-lasting afterimages occur when induced by the same alternating-stimulus procedure as that used to induce the McCollough effect. The second claim (Murch & Hirsch, 1972) is that afterimages can themselves induce McCollough effects if they are induced and paired sequentially with grating patterns. In testing these claims, a reliable computer-controlled color-cancellation technique developed earlier was used to measure the apparent color of both afterimages and McCollough effects objectively. No support was found for the first claim following alternative presentation of two homogeneously colored regions for total periods of 5 min (Experiment 1) and 20 min (Experiment 2). The second claim was fully supported: After an induction period of 7.3 min, a McCollough effect occurred for a red-vertical pairing but not for a green-horizontal pairing (Experiment 3); but after an induction period of 20 min, McCollough effects occurred strongly for both pairings (Experiment 4). The theoretical implications of these outcomes are considered in the context of recent theories of color and pattern processing in the visual system.     

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.     

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.     

A functional theory of the McCollough effect

Pattern-contingent color aftereffects, or McCollough effects (MEs), are used to probe the visual brain's operations psychophysically. Their neural substrate is unknown, however, and theories about them are weak. Our theory proposes a strong functional role for MEs and a neuropsychological basis that accounts for "top-down" (global) constraints ignored by other theories. The functional aspect of the theory is based on the concept of contingent adaptation level, following Helson (1964), and on the "error-correcting device" of Andrews (1964), which tracks and adjusts internal representation to external-world contingencies. The neuropsychological part of the theory postulates that global properties are the result of MEs being generated not at the individual detector level but in vectorfields of which the detectors are elements. It is an implementation of Lie transformation group theory (Hoffman, 1966). Evidence for this model is assessed.     

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.     

Delayed one-trial extinction of the McCollough effect

Initial tests of the McCollough effect were made on separate groups at delays of 4, 24 or 96 h after induction, with little apparent decrement. Subsequent tests were made at 4, 24 and 96 h after each initial test delay, again using separate subject groups in order to avoid test-induced extinction. All tests consisted of magentagreen nulling of the aftereffect. A single test session, whenever administered, seems to initiate a decay or extinction process lasting several days, resembling the time course shown by repeated testing. These results are difficult to reconcile with a perceptual conditioning model of the McCollough effect.     

The McCollough effect across the menstrual cycle

The McCollough effect (ME) has been shown to be sensitive to cholinergic agents, being strengthened by hyoscine (antagonist) and weakened by physostigmine (agonist), and possibly to more generalized changes in CNS arousal. We therefore expected the ME to be sensitive to hormonal changes during the menstrual cycle, being strongest in the postovulatory phases when arousal is low. In two experiments we found a highly significant effect of menstrual phase for the normally cycling women, but not for oral contraceptive users: ME strength gradually increased across the cycle, reaching a premenstrual peak. These findings may be explained in terms of hormonally mediated changes in arousal across the menstrual cycle.     

Extremely long-term persistence of the McCollough effect.

The question has been raised whether an associative mechanism is responsible for the persistence of the McCollough effect. Since current estimates of its rate of decay are derived from procedures in which subjects are repeatedly tested, it was hypothesized that the measured effects might be attenuated by the testing process. Accordingly, a comparison was made between repeated testing and time-elapse testing. A conventional group of 16 subjects had repeated testing at 0, 8, 24, 56, and 120 hr. after induction. Five other groups of 16 were run, each at one of the time delays, with no intervening tests; an additional measure was taken where appropriate at 120 hr. A magenta-green nulling procedure was used to assess the aftereffect. The repeated-test group showed a linear decrease of effect against the stated delays, reaching zero at 120 hr. In contrast, the time-elapse groups showed little decline up to 120 hr. Those groups retested at 120 hr. showed declines due to prior testing. When four more groups were subsequently tested at intervals up to 2,040 hr., the effect remained at better than half strength.     

List-strength effect: II. Theoretical mechanisms

Ratcliff, Clark, and Shiffrin (1990) examined the list-strength effect: the effect of strengthening (or weakening) some list items upon memory for other list items. The list-strength effect was missing or negative in recognition, missing or positive in cued recall, and large and positive in free recall. We show that a large number of current models fail to predict these findings. A variant of the SAM model of Gillund and Shiffrin (1984), involving a differentiation hypothesis, can handle the data. A variant of MINERVA 2 (Hintzman, 1986, 1988) comes close but has some problems. Successful variants of a variety of composite and network models were not found (e.g., Ackley, Hinton, & Sejnowski, 1985; Anderson, 1972, 1973; Metcalfe Eich, 1982; Murdock, 1982; Pike, 1984). The results suggest constraints on the future development of such models.     

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.     

Two mechanisms in the stimulus suffix effect

The stimulus suffix is a redundant item presented immediately after a stimulus list. Its effect is the selective impairment of recall of the final items in a serially recalled, auditorily presented list of unconnected items. Two experiments indicate that there was no difference between the effects of suffixes .5 and 1.0 sec after the end of a digit list presented at a rate of one digit/sec. This suggests that the effect of the suffix, in this case the vowel sound "ah," is not a simple function of its time of arrival after the final digit, as has been thought. The possibility of more complex factors was supported in a further experiment which showed a slight reduction in the size of the suffix effect by repeating the suffix three times.