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How the McCollough effect may involve feedback loops in the visual system.
Bryan Kennedy
Psychology 126
November 2002
TA: Christopher Cantor
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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.
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
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The inexperienced and curious reader is encouraged to participate in an online demonstration of the effect
produced by the author, located at http://www.electricfox.com/mccollough
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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
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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
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
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
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
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
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
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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
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
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.
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References
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.