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McCollough effect

The McCollough effect is a long-lasting form of orientation-contingent color aftereffect in human visual perception, in which prolonged exposure to gratings composed of black-and-white lines superimposed with specific colors—such as red vertical stripes and green horizontal stripes—causes subsequent achromatic gratings of matching orientations to appear tinted with the complementary colors, like cyan for verticals and magenta for horizontals.^[1] This illusion arises from adaptation to the combined features of color and line orientation, distinguishing it from simple negative afterimages that fade quickly and are not feature-specific.^[2] First formally described by psychologist Celeste Howard McCollough in 1965, the effect was induced through deliberate adaptation procedures using colored edge patterns, building on earlier incidental observations of similar contingent aftereffects during prism-wearing experiments in the 1930s and 1950s.^[1] Key properties include its high specificity to the inducing orientations and colors, lack of significant transfer between eyes, and resistance to disruption by sleep or darkness, with effects strengthening over repeated inductions but weakening upon frequent testing.^[2] Notably, the duration can extend from hours to several months; for instance, after 10-15 minutes of adaptation, the aftereffect persists for at least 24 hours without decay during sleep, and studies have documented minimal decline over 120 hours in untested observers, with reports of up to 3.5 months under minimal re-exposure conditions.^[3] The McCollough effect has been extensively studied to probe interactions between color and form processing in the visual system, revealing that it likely originates from adaptive changes in early cortical areas, particularly the primary visual cortex (V1), where orientation-selective neurons become contingently tuned to color via mechanisms such as selective fatigue or synaptic weakening.^[4] Evidence from neuroimaging and neuropsychological cases supports this localization, showing retinotopic organization and interactions between orientation- and color-coding pathways as early as V1, though contributions from higher areas like V4 may modulate its persistence and strength.^[5] These findings underscore the effect's utility as a tool for investigating neural plasticity and unconscious visual processing, with applications in understanding conditions like blindsight and synesthesia.^[4]

Introduction

Definition

The McCollough effect is a phenomenon of human visual perception characterized as an orientation-contingent color aftereffect, in which neutral, achromatic gratings appear tinted with colors complementary to those presented during an induction phase, with the perceived color determined by the grating's orientation.^[1] For instance, after adaptation to a horizontal grating composed of green and black stripes, a subsequent horizontal black-and-white grating will appear pinkish or reddish, whereas a vertical black-and-white grating remains colorless.^[2] This effect arises from viewers fixating on inducing stimuli consisting of colored gratings with specific orientations, resulting in a persistent association between orientation and color perception when viewing aligned neutral patterns.^[1] Unlike transient retinal afterimages, which fade quickly and are not linked to specific stimulus features such as orientation, the McCollough effect is selectively contingent on the alignment of the test grating with the induction orientation and can endure for extended periods, often minutes to hours.^[2] This contingency distinguishes it as a higher-level perceptual adaptation rather than a simple photochemical response in the retina.^[1]

Discovery and History

The McCollough effect was discovered by American psychologist Celeste McCollough in 1965 during her research on visual adaptation. She had observed it accidentally while demonstrating to her students the aftereffects produced by wearing prism spectacles that invert the visual field, using alternating exposures to orange vertical gratings and blue horizontal gratings to induce adaptation more rapidly; neutral achromatic gratings subsequently appeared tinted in complementary colors—yellowish for horizontal and bluish for vertical—persisting for several hours after adaptation. McCollough attributed it to selective fatigue in orientation- and color-tuned edge detectors within the visual system. Her findings were detailed in a seminal paper published in the journal Science.^[2] The effect garnered immediate attention and was replicated extensively in the late 1960s and 1970s, with early studies confirming its cortical locus through evidence of complete binocular transfer and independence from retinal factors, such as no decay during prolonged eye closure. For instance, Harris (1970) and Stromeyer and Klein (1974) demonstrated the phenomenon's robustness across varied induction conditions, establishing it as distinct from peripheral adaptation. These replications, including those exploring multi-orientation variants, underscored the effect's reliability as a probe for central visual processing. Research in the 1970s advanced understanding of the effect's longevity, with Jones and Holding (1975) reporting persistence up to 24 hours following 10 minutes of induction, far exceeding typical afterimages. In the 1980s, studies focused on decay dynamics revealed that the effect diminishes exponentially under patterned visual stimulation but shows minimal decline in darkness or during sleep, as extended in follow-up work building on MacKay and MacKay (1976). Notably, Skowbo et al. (1984) examined whether decay mimicked extinction in conditioned responses, reinforcing the effect's stability as a marker of enduring neural adaptation. The McCollough effect profoundly shaped visual perception research by challenging purely retinal adaptation theories and pioneering the study of contingent aftereffects, where sensory features like orientation gate color responses. This shifted emphasis toward cortical feature integration, inspiring decades of experiments on perceptual learning and plasticity.^[2]

Induction

Procedure

The standard procedure for inducing the McCollough effect involves a two-phase process: adaptation (induction) and testing. During the adaptation phase, participants alternately fixate on two orthogonally oriented gratings presented against a white background—one consisting of horizontal black lines paired with a red hue and the other vertical black lines paired with a green hue (or complementary colors such as blue and yellow). Subjects are instructed to maintain central fixation on a marked point in each grating, with eye movements often restricted using a chin rest or bite bar to minimize saccades and ensure consistent retinal stimulation; the alternation between gratings typically occurs every 10-20 seconds to sustain attention without inducing fatigue.^[6] The total induction duration is usually 5-15 minutes, with longer periods producing stronger effects.^[7] To enhance reliability and control stimulus purity, variations in presentation methods have been employed. In the original setup, colored filters (e.g., red and green gelatin filters) were placed over achromatic gratings projected onto a screen at a viewing distance of about 57 cm, ensuring equiluminant conditions between colors. Modern replications often use digital displays or computer monitors to generate the gratings, allowing precise control over color calibration and luminance matching, which reduces variability from projector inconsistencies.^[8] Precautions such as brief dark adaptation breaks (e.g., 30 seconds every few minutes) are recommended to prevent observer fatigue or bleaching of photopigments, particularly during extended sessions.^[9] Following induction, the testing phase assesses the contingent aftereffect using achromatic (black-and-white) gratings matching the orientations of the adaptation stimuli, presented immediately afterward at the same viewing distance. Participants report the perceived coloration subjectively—such as a greenish tint on horizontal gratings and reddish on vertical ones—or undergo psychophysical scaling tasks, where they adjust the amount of canceling color (e.g., via heterochromatic matching) to null the induced hue and quantify its strength. This measurement confirms the orientation-specific color perception without the inducing stimuli present.^[8]

Stimulus Parameters

The McCollough effect is typically induced using achromatic gratings oriented at 90 degrees to each other, most commonly horizontal and vertical, as these orthogonal orientations facilitate the association between color and contour direction. The effect achieves maximum strength when the test gratings precisely match the orientations of the inducing stimuli, with selectivity diminishing for rotations as small as 20 degrees and disappearing around 45 degrees.^[1]^[10] Complementary color pairs, such as red paired with green or blue with yellow, are used for the inducing gratings to maximize chromatic contrast and produce robust aftereffects, as these opponencies align with opponent-process color vision mechanisms. For instance, vertical gratings are often presented with one color (e.g., green) against black bars, while horizontal gratings use the complementary hue (e.g., magenta or red). Luminance levels between the colors must be carefully matched—typically equiluminant—to prevent confounding luminance-based aftereffects or artifacts that could mimic the orientation contingency.^[1]^[11]^[10] Spatial frequencies of the gratings are optimally tuned within 2-6 cycles per degree, aligning with peak human contrast sensitivity, to ensure reliable induction without reducing the effect's specificity to orientation. High contrast levels, such as 90% or near-maximal (e.g., black bars at low luminance against brightly colored stripes), enhance the strength of adaptation, though lower spatial frequencies can broaden selectivity at the cost of precision.^[12]^[10]^[13] Additional parameters include a neutral white or black background luminance to minimize extraneous cues during induction, with exposure durations of 5-15 minutes total—often alternating every 10-20 seconds, though some protocols use faster alternations of a few seconds to build the contingency without fatigue. In some variants, incorporating motion, such as drifting gratings at low velocities, or flicker can amplify the effect's intensity by increasing neural activation.^[1]^[13]^[10]

Properties

Duration and Stability

The McCollough effect exhibits a wide range of durations, typically lasting from minutes to several months depending on induction strength. Brief adaptation periods of 2-4 minutes produce weak effects that persist for about 1 hour or more, while prolonged inductions of 20 minutes or longer can yield effects enduring for weeks, with some cases maintaining over 50% strength after 3 months. Peak effect strength occurs within the first hour post-induction, followed by a gradual decline that can extend to several months, with studies documenting over 50% strength after 2.8 months in untested observers.^[3] With repeated inductions, persistence may be further extended, though verified reports beyond several months remain limited. Several factors influence the stability and decay of the effect. Stronger inductions through extended exposure times result in longer-lasting effects, as adaptation duration correlates positively with both initial magnitude and retention period. Testing or repeated measurement accelerates decay, often initiating a linear reduction that can eliminate the effect within days, whereas disuse without testing allows minimal decline over similar intervals. Sleep plays a role in maintenance, with normal sleep cycles preserving effect strength, while deprivation can reduce it by up to 50%; visual disuse in darkness mimics this preservation to some extent, but lacks sleep's full protective effect. Re-exposure to induction stimuli can reinstate or strengthen faded effects. Decay, particularly following testing, is often modeled as an exponential decline in strength, S(t) = S_0 e^{-t/\tau}, where S(t) is strength at time t, S_0 is initial strength, and \tau is the time constant; however, without repeated testing, the long-term component shows minimal decline over extended periods. Individual variability affects these parameters, with children exhibiting stronger and potentially longer-lasting effects than adults, and differences in attention during induction influencing initial strength. Long-term retention suggests a form of neural storage akin to imprinting, enabling effects to persist or partially recover after months without full decay. ^[1] ^[14] ^[15]

Specificity

The McCollough effect demonstrates pronounced orientational specificity, with the aftereffect strength peaking when the test grating's orientation precisely matches that used during induction and declining rapidly with angular deviations. For orthogonal orientations (90° difference), the effect is typically absent or reversed in polarity, while intermediate angles around 45° yield substantially reduced effects. Empirical measurements indicate a tolerance of approximately ±10-15° for robust induction, beyond which the aftereffect becomes rare or negligible, as shown in studies varying angular separation during adaptation. Oblique gratings further illustrate this tuning, often producing diminished or null effects compared to cardinal orientations like horizontal or vertical.^[10] Color contingency is a core feature, wherein the perceived tint on achromatic test gratings corresponds to the complementary color of the inducing stimulus and remains strictly locked to the associated orientation. For example, adaptation to red vertical gratings paired with green horizontal gratings results in vertical gratings appearing greenish and horizontal ones reddish, with no color shift observed for mismatched orientations. This selectivity extends chromatically, as the effect is most potent with opponent colors like red-green or blue-yellow, showing weaker responses to non-opponent hues such as orange or ambiguous spectral filters.^[10] Spatial specificity limits the transfer of the aftereffect across different grating frequencies, with maximal strength occurring when induction and test stimuli share the same spatial frequency. Adaptation to high-frequency gratings produces little to no effect on low-frequency tests, and vice versa, though partial transfer may occur over spans up to 2 octaves with reduced saturation. Chromatic aspects reinforce this, as the effect's color opponency does not generalize broadly across wavelengths outside the primary adaptation range.^[16]^[10] Regarding binocular aspects, the McCollough effect exhibits partial interocular transfer, allowing some adaptation induced monocularly to influence the contralateral eye, particularly under homochromatic binocular stimulation that engages shared pathways. However, the aftereffect is stronger and more consistent when both eyes receive similar adaptation, as dichoptic viewing with dissimilar stimuli can impede buildup or reduce overall potency.^[17]^[10]

Explanations

Theoretical Models

The contingent aftereffect model posits the McCollough effect as a form of learned association between stimulus orientation and color, analogous to classical conditioning, where orientation serves as a conditioned stimulus paired with color as the unconditioned stimulus.^[18] This framework, advanced in the 1970s, suggests that repeated exposure strengthens the linkage, leading to the persistent aftereffect, with decay resembling extinction of the conditioned response. Key proponents, including J.E.W. Mayhew and S.M. Anstis, drew parallels to color-contingent motion aftereffects, arguing that behavioral patterns mimic conditioning rather than simple fatigue. Similarly, P.K. Leppmann modeled it as an association between opponent-color responses and contour detectors, emphasizing psychological learning over physiological adaptation.^[18] Adaptation-level theory extends opponent-process color theory to orientation-selective mechanisms, proposing that prolonged exposure shifts response thresholds in edge detectors, causing complementary color perceptions in achromatic gratings. Originally suggested by C. McCollough, this view attributes the effect to selective fatigue in color-sensitive neurons tuned to specific orientations, with optimal effects occurring at 90° separations between gratings. Supporting evidence from S. Fidell indicated that the aftereffect magnitude depends on spatial frequency matches, aligning with adaptation in orientation-specific channels rather than global color fatigue. This theory prioritizes sensory recalibration, where the visual system adjusts adaptation levels to prolonged chromatic stimulation along contours.^[18] The dual-process model integrates transient and sustained components, attributing short-term aspects to retinal or early adaptation and long-term persistence to cortical associative processes, thereby rejecting purely peripheral explanations due to the effect's stability beyond typical retinal recovery times. L.A. Riggs and colleagues proposed a sequential analysis where color adaptation feeds into orientation detectors, explaining non-monotonic build-up and decay curves observed in experiments.^[18] G.M. Murch similarly described parallel processing of color and form, with fatigued opponent-process channels influencing higher-level orientation selectivity, forming a hybrid of adaptation and integration. Criticisms of early conditioning models emerged in the late 1970s, highlighting their failure to account for the effect's confinement to low-level visual features like orientation and spatial frequency, rather than arbitrary stimulus pairings, which undermined claims of general associative learning.^[2] Adaptation theories faced challenges in explaining durations exceeding weeks, as simple fatigue could not sustain such persistence without additional mechanisms.^[18] By the 1980s, these gaps prompted a shift toward hybrid models, as compiled in C.S. Harris's edited volume, which emphasized combined adaptation and plasticity to reconcile behavioral specificity with longevity. This evolution favored frameworks blending psychological association with channel-based recalibration, better aligning with observed properties like orientation selectivity.^[2]

Neural Mechanisms

Functional magnetic resonance imaging (fMRI) studies have demonstrated that the McCollough effect arises from adaptation in early visual cortical areas, particularly V1 and V2, where orientation-selective neurons bind color information. In one such study, participants adapted to orientation-contingent color gratings, and fMRI revealed that perception of the aftereffect correlates with activity in extrastriate areas such as the lingual and fusiform gyri.^[19] Additionally, a 2020 fMRI adaptation experiment confirmed the physiological locus in primary visual cortex, showing selective adaptation effects—reduced BOLD signals—in V1 for the McCollough aftereffect, supporting its role in maintaining the contingency.^[20] The effect's central processing is further evidenced by its lack of significant interocular transfer; adaptation induced monocularly does not substantially transfer to the other eye, even after prolonged occlusion, which rules out peripheral retinal mechanisms and indicates processing in monocular channels of the early visual cortex. Adaptation occurs at hypercolumnar levels in the primary visual cortex, where orientation columns integrate chromatic signals, leading to long-lasting shifts in neuronal responsiveness. A 2008 study using behavioral measures separated transient and permanent adaptation components, attributing the enduring McCollough effect to slow-decaying changes in V1 orientation-selective cells.^[8] At the cellular level, neural models propose that the McCollough effect involves long-term potentiation (LTP)-like modifications in orientation-selective neurons, enabling stable storage of the color-orientation association over extended periods. Neural models propose that inhibitory interneurons in V1 contribute to this by modulating lateral connections, enforcing specificity to the inducing orientation and preventing generalization.^[21] Recent neuroimaging post-2020 has linked the effect to cortical plasticity, with fMRI showing adaptation-driven changes in early visual areas that mimic synaptic strengthening. A 2023 study in Vision Research found enhanced McCollough effects in anomalous trichromats, attributed to nonlinear compressive encoding of color saturation in early visual cortex and compensatory amplification in V4, where reduced cone contrasts are normalized postreceptorally.^[22] No major breakthroughs emerged in 2024-2025, but virtual reality induction methods, such as the 2021 NIH "McCollough World" protocol, have been extended to probe neural modeling of plasticity, allowing controlled adaptation in immersive environments to study V1 dynamics.^[13]

Anti-McCollough Effect

The anti-McCollough effect is a variant of the orientation-contingent color aftereffect in which an achromatic test grating of a specific orientation appears tinted with the same hue as the inducing colored grating, rather than its complementary color as in the standard McCollough effect.^[23] For example, alternating adaptation to a red horizontal grating and an achromatic horizontal grating results in the achromatic horizontal test grating appearing reddish.^[23] Induction of the anti-McCollough effect requires interleaving presentations of a colored grating and an achromatic grating sharing the same orientation, typically alternating every 10 seconds for a total adaptation period of 20 minutes, using stimuli such as gratings with a spatial frequency of 3.4 cycles per degree.^[23] This procedure differs from the standard McCollough effect, which uses exclusive pairings of colored gratings with orientations without achromatic interleaving; the anti-effect is notably weaker in magnitude and, while stable for up to 24 hours, generally shorter-lived compared to the standard effect's potential duration of days to weeks.^[23] Additionally, the anti-McCollough effect exhibits complete interocular transfer, indicating a higher-level neural representation than the partial transfer seen in the classical version.^[23] Key findings from studies on the anti-McCollough effect highlight the brain's flexibility in forming orientation-color contingencies even when one stimulus lacks chromaticity, as the achromatic grating's orientation and spatial frequency are critical for directing the same-hue aftereffect.^[24] This variant has been used to probe the boundaries of adaptation processes, revealing that single orientation-color pairings suffice for induction and that the effect depends on specific stimulus configurations, such as alternating with orthogonal achromatic gratings for maximal strength.^[24] In terms of applications, the anti-McCollough effect aids in distinguishing genuine neural adaptation from mere sensory fatigue, as its full interocular transfer and persistence suggest involvement of central, higher-order visual processing rather than peripheral mechanisms.^[23] It also contributes to disentangling transient perceptual components from more stable learned associations in contingent aftereffects.^[24]

Other Contingent Aftereffects

Contingent aftereffects encompass a family of perceptual phenomena where adaptation to one visual feature induces an afterimage selectively tied to another feature, extending the principles observed in orientation-specific color adaptations. These higher-order effects demonstrate the visual system's ability to associatively bind features such as motion, spatial frequency, and form, often persisting for minutes to hours after exposure. Unlike simple aftereffects, contingent variants require the co-occurrence of inducing features during adaptation, revealing mechanisms for feature integration in early visual processing. A key example is the motion-contingent color aftereffect, in which prolonged exposure to colored gratings moving in opposite directions produces a directional color illusion in subsequently viewed achromatic motion. Observers adapting to green upward-moving vertical gratings alternating with red downward-moving horizontal ones report a pink afterimage for upward motion and a green one for downward motion in neutral test patterns. This effect, first reported by Stromeyer (1971), suggests adaptation at sites where color and motion signals converge, potentially in motion-sensitive areas like MT, and complements reciprocal color-contingent motion aftereffects where color adaptation biases perceived motion direction.^[25] Further studies, such as those by Ramachandran and Gregory (1973), reinforced this by showing that such bindings can last several minutes and are selective to the direction and speed of adaptation.^[26] Spatial frequency contingent aftereffects provide another illustration, where color illusions become specific to the density or bar width of grating patterns. Adaptation to red low-frequency gratings and green high-frequency ones induces complementary color tinges only on test gratings matching the respective frequencies, with minimal transfer to mismatched ones. Seminal work by May and Matteson (1976) using chromatic checkerboards demonstrated that these aftereffects align with the Fourier components of the stimuli rather than edge orientations, supporting the existence of narrow-band spatial frequency channels that process color contingent on pattern scale.^[27] This specificity underscores how the visual system parses complex scenes by linking chromatic responses to spatial details, as later confirmed in experiments showing tuning across a wide range of frequencies from 1 to 16 cycles per degree.^[28] Beyond these, contingent aftereffects include bindings like color to tilt or texture density, exemplifying multi-feature associations akin to those in the classic orientation-contingent paradigm. For instance, the tilt aftereffect can be rendered color-specific, where adaptation to a red-tilted grating and a green-orthogonal one yields a tilt illusion confined to each hue in achromatic tests. These variants, explored in studies such as Favreau and Cavanagh (1982), highlight the generality of contingency as a tool for probing feature binding.^[29] The McCollough effect has notably inspired investigations into perceptual binding, including parallels with synesthesia, where involuntary cross-feature experiences resemble induced contingent colors. Research by Rothen and Meier (2017) utilized McCollough-like inductions to test synesthetes' color associations, finding that synesthetic hues exhibit similar orientation specificity and resistance to verbal interference as contingent aftereffects, suggesting shared neural substrates for feature linkage.^[30] The specificity of these effects to the adapted feature combination, as prototyped in orientation contingencies, thus provides a framework for understanding broader associative adaptations in vision.

References

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Color Adaptation of Edge-Detectors in the Human Visual System
Celeste McColloughAuthors Info & Affiliations. Science. 3 Sep 1965. Vol 149, Issue 3688. pp. 1115-1116. DOI: 10.1126/science.149.3688.1115 · PREVIOUS ARTICLE.
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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.Missing: studies | Show results with:studies
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Much of the evidence suggests that the McCollough effect depends on neural mechanisms that are located early in the cortical visual pathways, probably in V1.
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The McCollough effect indicates that colour- and orientation-coding mechanisms interact at some point during visual processing.
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In the McCollough effect, adaptation is produced not by color alone, but by a combination of color and orientation.
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The McCollough effect (McCollough, 1965, 2000) is an orientation contingent color aftereffect—adaptation to, e.g., red vertical and green horizontal gratings ...
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Feb 15, 2012 · The McCollough effect is a contingent color after effect induced by adapting to colored gratings for several minutes.
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In a well-known study, McCollough (1965) described a color aftereffect that is contingent on pattern orientation. For example, after the viewing of a ...
[12]
The role of some spatial parameters of gratings on the McCollough ...
The number of repetitions of black and colored stripes per unit visual angle, hereafter referred to as spatial frequency, affects the strength of the McCollough.
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A Functional Magnetic Resonance Imaging Study - Sage Journals
The McCollough effect is a striking color aftereffect that is linked to the orientation of the patterns used to induce it. To produce the McCollough effect ...
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Uncovering the physiological locus of the McCollough Effect using ...
The McCollough Effect (ME) is a color afterimage produced by exposure to colored, oriented patterns. For example, following viewing of alternating vertical ...Missing: EEG | Show results with:EEG
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Nonlinear cortical encoding of color predicts enhanced McCollough ...
In this account of the enhanced anomalous McCollough effect, normal and anomalous trichromats have identical visual systems except for the visual pigment swap.
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The effect, termed the anti-McCollough effect, although weaker than the classical aftereffect, remained stable for a moderate duration of time (24 hours).
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Jul 26, 2011 · To the contrary, we argue that the anti-ME is contingent upon a number of features of the stimulus, including orientation and spatial frequency.
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After human observers alternately view green stripes moving up and red stripes moving down for periods of 1/2 to 4 hours, they see a pink aftereffect when ...
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These color-contingent motion after effects complement reports of motion-contingent color aftereffects and suggest that both may reflect adaptation of detectors ...
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Two-dimensional Fourier analysis of checkerboards reveals that major components are at a 45° angle to the check edges. After adapting to chromatic ...
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Contingent color aftereffects are associated with the Fourier components rather than the edges of such patterned stimuli at high spatial frequencies.
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Since the McCollough effect also results from oriented contours (i.e., form) evoking specific colors, we conjectured that synesthetes may experience an enhanced ...Abstract And Figures · References (25) · Recommended Publications