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Chromatic Chutes and Ladders:

How the McCollough effect may involve feedback loops in the visual system.

“Vision is the art of seeing things invisible.”

-- Jonathan Swift

Bryan Kennedy

Psychology 126
November 2002
TA: Christopher Cantor

Chromatic Chutes and Ladders:

How the McCollough effect may involve feedback loops in the visual system

It seems as though our eyes are always playing tricks on us: at times we see a larger

moon on the horizon than high in the sky or an artistic staircase that appears to rise infinitely. It

is unlikely that our visual systems, having evolved over millions of years, would engage in such

petty games for the delight of misleading us. Instead, these “mistaken” perceptions are likely the

result of systems that, at other times, are employed to good effect. The McCollough effect (ME)

is such an illusion, where color-tinged bars are seen on a black and white screen. The results of

studies linking the McCollough effect to processes involving occlusion and chromatic aberration,

along with recent data from fMRI brain scans, underline the possibility that this compelling

illusion involves both the lower and higher visual areas of the brain.

The McCollough effect is a convincing illusion first demonstrated by Celeste

McCollough in 1965. In order to experience it, one is first shown, and becomes adapted to, two

alternating colored bar gratings for five minutes. The gratings might be composed of vertical

black-and-green bars, and horizontal black-and-red bars, for example. When an individual

subsequently views a screen with identical black and white bars, he or she will see color hazing

around the edges of the black bars. Even more incredible to observe, these illusionary colors are

complementary to those of the adaptation, in that the horizontal bars will now appear fringed

with green, whereas they were originally red. The effects of the adaptation can be seen for days

or weeks1.

When the effect was first introduced by McCollough, it was proposed that the early visual

area V1 played a central role. The visual cortex is generally believed to be striated into at least

The inexperienced and curious reader is encouraged to participate in an online demonstration of the effect
produced by the author, located at

five main functional levels, from V1 through V5, layered like a stack of pancakes. Visual

information is first delivered from the retinas to the area V1, where it is then fed through the

remaining levels for various stages of processing. For the purposes of this discussion, we will

only concern ourselves with this first area and area V4, which is believed to be central in the

perception of color.

Information has since accumulated that supports the original theory that the McCollough

effect involves activity in V1; for example, people who have suffered severe damage to V1 do

not experience the effect. Even more convincing was the discovery that the illusionary tinge was

complementary to the wavelength of the adapting stimulus, rather than the perceived color. And

since V4 is thought to play a central role in the color constancy system that converts color

wavelengths to perceived colors, it was supposed that the effect relied on regions before area V4

(Humphrey et al., 1999). It would seem then that the argument for V1 was all but decided, but a

series of findings began to suggest that higher visual areas were involved as well.

A study conducted by Watanabe (1994) supports this new theory that the McCollough

effect may involve higher-level visual areas, and more specifically, those responsible for figuring

out occlusion. When an object is in front of another, it is said to cover or occlude it. Our visual

system is able to extract information about the covered object and conclude that it is indeed one

object rather than two disconnected halves. In Watanabe’s experiment, subjects were adapted to

normal McCollough adaptation gratings, but were shown test gratings that were partially

occluded by rectangles. It was hypothesized that subjects would see the McCollough effect

despite the occlusion, and that this would suggest that the subjects were mentally connecting the

disparate bars in the test grid. The results supported this hypothesis, and it was proposed that the

McCollough effect not only relied on the higher visual areas responsible for occlusion, but likely

involved some sort of feedback loop to V1. A subsequent study takes this assumption a step

further by actually recording McCollough-effect-related brain activity in V4, the area responsible

for color perception.

In 1999, Humphrey et al. conducted an fMRI brain scanning experiment to locate the

region responsible for McCollough adaptation. Based on earlier research evidence linking ME

with the early visual pathway, the experimenters expected to find brain activity to be mostly

centered in V1. The study involved six subjects and three test periods. The experimenters used

three main types of slides during the tests: an adaptation grating slide, similar to the example

gratings described previously in this paper, and two test slides, one with bar gratings that were

congruent with the adaptation bars (which should incite the illusion), and one with non-

congruent bars (which should not). The first of the three tests was a pretest, where subjects were

shown only the alternating test slides without first being adapted. This pretest was intended to

determine subject bias, and indeed, one subject’s results were disregarded. In the second test

period, subjects were shown the adaptation gratings minus coloration, followed by the test

gratings. This test was used as a control to ensure that subjects did not show activity based only

on contrast adaptation. The final test period involved the actual adaptation to the McCollough

gratings, followed again by the two test grids.

As expected, all subjects reported experiencing the McCollough effect when viewing the

congruent test grating. Scans taken during the viewing of the congruent and non-congruent test

slides were compared to determine if the illusion was causing measurable brain activity. Contrary

to what was originally expected, only one subject actually showed activity in V1, whereas all of

the subjects showed significant activity in area V4. If nothing else, this shows that V4 plays an

important role in the McCollough effect.


In their discussion of these unexpected results, the researchers suggested a variety of

confounders that could explain the unanticipated activity in V4. One such proposal was that the

activity in V4 might be associated with the perception of the color in the illusion itself, rather

than any actual adaptation. However, this is a moot point – for even if V4’s activity was related

to the perception of the colors of the illusion, it would still be related to the illusion, whether or

not it is considered central to it. A non-fMRI study has furthered these surprising findings, by

suggesting that V4 does indeed play a role in this intriguing visual illusion.

In their study, Broerse et al. (1998) suggest that the McCollough effect may be related to

the system in V4 that compensates for chromatic aberrations in the human eye. Chromatic

aberration, a phenomenon well known to photographers, causes color irregularities to appear on

an image, and is due to slight imperfections in the lens. In studies conducted using optical prisms

to induce chromatic aberrations, subjects reported that the color irregularities (blue tinges on

straight edges) mysteriously disappeared after extended observation. When the prisms were

subsequently removed, subjects reported seeing colors that were complementary to the original

chromatic irregularities. This finding suggests that the visual cortex is actively compensating for

chromatic aberrations, presumably as a routine method of perfecting the visual image from an

imperfect lens (Broerse et al., 1998).

Broerse et al. conducted a series of five experiments to determine if the McCollough

effect was related to that system. For the purposes of this discussion, only the first two will be

examined. In the first experiment, the researchers found that the McCollough effect could still be

induced by adapting subjects to black bars that were merely framed with fine edges of green and

red. The subjects were adapted to the novel color-fringed gratings and shown the test gratings. As

hypothesized, all subjects experienced the McCollough effect. The researchers theorized the

slight color fringes shown during adaptation were similar to that of the fringes experienced in the

original prism experiment, and thus must involve similar mechanisms of correction. They then

conducted a virtually identical follow-up study using the more traditional filled-color bars, to

ensure that the illusion in the first experiment was indeed the McCollough effect. They found

that the illusionary colors of both experiments were the same, and so concluded that the fringes

were properly inducing the McCollough effect.

This association, between chromatic distortion compensation and the McCollough effect,

could explain evolutionarily why the effect persists for a greater amount of time than other visual

aftereffects. Chromatic aberrations in the lens would be more long-term problems associated

with eye growth – thus, a system designed to account for them would optimally operate slowly. A

possible follow-up study might compare the differences in illusion strength and persistence

between children and adults, assuming that such a system would be more active and plastic

during the period of rapid eye growth in childhood.

While it is obvious that further research in this area is necessary to arrive at any definitive

conclusions, the evidence linking the McCollough effect to the higher visual areas is mounting.

Could it be that the illusion involves many layers of the visual cortex working in parallel, like a

game piece moving among the various levels of the board game “Chutes and Ladders”?

Regardless of whether or not the answer is ever found, it is clear that examining illusions such as

the McCollough effect will lead us to better a understanding of our visual system as a whole.


Broerse J., Vladusich, T., O’Shea, R. P. (1999). Colour at edges and colour spreading in
McCollough effects. Vision Research, 39(7), 1305-1320.

Humphrey, G. K., James, T. W., Gati, J. S., Menon, R. S., Goodale, M. A. (1999). Perception of
the McCollough Effect Correlates with Activity in Extrastriate Cortex: A Functional Magnetic
Resonance Imaging Study. Psychological Science, 10(5), 444-448.

Watanabe, T. (1995). Orientation and Color Processing for Partially Occluded Objects. Vision
Research, 35(5), 647-655.