McCollough effect: A theory based on the anatomy of the lateral geniculate body

A model network derived from anatomy and physiology of the monkey retina and lateral geniculate body (LGN) is presented. Control of the strength of concentric surrounds by receptive field center illumination leads to inhomogeneous use of different surround parts, and this occurs even in the presence of irregular eye movements (“autostroboscopic impression”). Various pattern- and movement-contingent color aftereffects, as well as threshold findings, can be explained. Anatomical studies have revealed that the output signals of LGN interneurons are presynaptically controlled at each ending by retinal afferents. This is assumed to result in the above center-surround interaction, and it implies that an interneuron producesdifferent output signals at each of its numerous output endings. The inability of the somatic metabolic machinery to cope with a multitude of different time-varying demands is proposed to be the origin of the remarkably long persistence of the McCollough effect     

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.     

McCollough effect: A neural network model based on source separation

McCollough effects (MEs) are a group of visual contingent aftereffects that involve colour and contour. These effects have been the subject of a large body of literature concerning their properties and theoretical accounts, but the mechanisms underlying the ME have never been fully clarified. We make the assumption that a general adaptive neural process tending to maintain independent dimensions in visual perception could account for the ME. The proposed neural network model generating the ME, though of minimal complexity, can reproduce various detailed experimental results (such as the tilt effect contingent to colour) and above all it accounts for the distinctive long temporal persistence of this aftereffect.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

The indirect McCollough effect: An examination of an associative account

It has been found that viewing, for example, a red-and-black vertical grating alternating with a green homogeneous field produces a color aftereffect—a McCollough effect—on a black-and-white vertical grating (i.e., green). Viewing such colored patterns also produces an aftereffect on a noninduced horizontal grating (i.e., pink)—the indirect McCollough effect. Humphrey, Dodwell, and Emerson (1989) argued that the indirect McCollough effect is caused by opponent properties of the visual system that organize the processing of contour and color along contrasting, probably orthogonal, dimensions. Recently, however, their interpretation of the indirect McCollough effect has been challenged by some findings of Eissenberg, Allan, Siegel, and Petrov (1995). These researchers have proposed that the indirect McCollough effect, like the McCollough effect, can be explained by associative principles. The results reported here question crucial aspects of the hypothesis of Eissenberg et al.     

Re-enchanting the body: overcoming the melancholy of anatomy

I argue here that Weberian disenchantment is manifest in the triumph of instrumental reason and the expansion of analytic enquiry, which now dominates not simply those sciences upon which medicine depends, but medical practice itself. I suggest ways that analytic enquiry, also referred to here as anatomical reasoning, are part of a particular ideology—a way of seeing, speaking about, and inhabiting the world—that often fails to serve the health of patients because it is incapable of “seeing” them in the moral sense described by Iris Murdoch and others. I use the work of James Elkins and Wendell Berry to call for the recovery of a way of seeing the human body as both other and more than an object of scientific enquiry and social control.     

The McCollough effect: Influence of several kinds of visual stimulation on decay rate

An experiment was conducted to determine whether the decay rate of the McCollough effect could be differentially influenced by the type of visual stimulation that followed its induction. After acquiring this effect, Os were exposed to achromatic gratings, homogeneous chromatic fields, natural visual stimulation, or complete darkness. Exposure to achromatic gratings caused a marked fading of the effect; the other types of stimulation were associated with similar and much less rapid decay.