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Test stimulus characteristics determine the perceived speed of the dynamic motion aftereffect.

Using a speed-matching task, we measured the speed tuning of the dynamic motion aftereffect (MAE). The results of our first experiment, in which we co-varied dot speed in the adaptation and test stimuli, revealed a speed tuning function. We sought to tease apart what contribution, if any, the test stimulus makes towards the observed speed tuning. This was examined by independently manipulating dot speed in the adaptation and test stimuli, and measuring the effect this had on the perceived speed of the dynamic MAE. The results revealed that the speed tuning of the dynamic MAE is determined, not by the speed of the adaptation stimulus, but by the local motion characteristics of the dynamic test stimulus. The role of the test stimulus in determining the perceived speed of the dynamic MAE was confirmed by showing that, if one uses a test stimulus containing two sources of local speed information, observers report seeing a transparent MAE; this is despite the fact that adaptation is induced using a single-speed stimulus. Thus while the adaptation stimulus necessarily determines perceived direction of the dynamic MAE, its perceived speed is determined by the test stimulus. This dissociation of speed and direction supports the notion that the processing of these two visual attributes may be partially independent.

Figural Aftereffect↗

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.

Adult↗

The tilt aftereffect in plaids and gratings: channel codes, local signs and "patchwise" transforms.

The perceived structure of a suprathreshold plaid made from two sinusoidal gratings tilted +/- 45 deg from vertical usually resembles a blurred checkerboard. Physically increasing the tilt of the components away from vertical elongates the pattern horizontally, yielding rectangular checks. We asked whether illusory tilt of the components induced by adaptation to a vertical grating would also give apparent elongation of the checks. Using a staircase method, we found that after adaptation the component orientations had to be set 3-4 deg closer to vertical to maintain a square, checkerboard-like appearance, implying that the +/- 45 deg plaid did appear elongated. This result suggests that the tilt aftereffect adaptation not only distorts the orientation of 1-D patterns but can also change the relative location of features, even when their orientation is not altered. This is not easy to explain within filter models that (often implicitly) assume each receptive field carries a fixed "local sign" indicating its position within the retinal array. We consider a representation in which localized patches of the image are encoded by the coefficients of a 2-D Fourier-like transform. With a simple subtractive model of adaptation, we show that the spatial information carried by each patch would be distorted in just the way observed for plaids and gratings. The fact that perceived structure is both distorted and coherent, not fragmented, after adaptation suggests an additional, more global process whereby local patches are combined to form a coherent, composite "neural image". We offer a simple principle that could establish this coherence. If the positions of local patches are adjusted so as to maximize the contrast energy of the composite image, then the spatial distortions of the patches (induced by selective adaptation) are carried through to the global structure of the composite. Thus, the conflict between channel codes and local signs is resolved, but perceptual distortion is the result.

Adaptation, Ocular↗

Cross-fixation transfer of motion aftereffects with expansion motion.

It has been shown that motion aftereffect (MAE) not only is present at the adapted location but also partially transfers to nearby non-adapted locations. However, it is not clear whether MAE transfers across the fixation point. Since cells in area MSTd have receptive fields that cover both sides of the fixation point and since many MSTd cells, but not cells in earlier visual areas, prefer complex motion patterns such as expansion, we tested cross-fixation transfer of MAE induced by expanding random-dots stimuli. We also used rightward translational motion for comparison. Subjects adapted to motion patterns on a fixed side of the fixation point. Dynamic MAE was then measured with a nulling procedure at both the adapted site and the mirror site across the fixation point. Subjects' eye fixation during stimulus presentation was monitored with an infrared eye tracker. At the adapted site, both the expansion and the translation patterns generated strong MAEs, as expected. However, only the expansion pattern, but not translation pattern, generated significant MAE at the mirror site. This remained true even after we adjusted stimulus parameters to equate the strengths of the expansion MAE and translation MAE at the adapted site. We conclude that there is cross-fixation transfer of MAE for expansion motion but not for translational motion.

Figural Aftereffect↗

Orientation-contingent color aftereffects mediated by subjective transparent structures.

We examined whether the orientation-contingent color aftereffect (the McCollough effect) could be mediated by subjective horizontal and vertical structure induced by the perception of transparency. In our experiments, red vertical bars and green horizontal bars were alternated as an adapting stimulus. After adaptation, subjects (n = 6) were asked to adjust the green and red saturation of a test pattern until they obtained a neutral gray. Horizontal and vertical stripes were combined in the test pattern in three different ways: (1) overlapping with a luminance combination that gave rise to a perception of transparent overlays of horizontal and vertical stripes (valid transparency condition), (2) overlapping with luminance combinations that did not induce a perception of transparency (invalid transparency condition) and that appeared more as a patchwork of checks, and (3) presented in adjacent, nonoverlapping areas. Our results showed that the McCollough effect was significantly greater in the valid transparency condition than in the invalid transparency conditions. The effect in the valid transparency condition was nevertheless less strong than was the effect seen with the standard test stimulus made up of nonoverlapping vertical and horizontal stripes. Our results suggest that the McCollough effect can be mediated by the subjective spatial organization (inner representation of vertical and horizontal stripes) that accompanies the perception of transparency in our stimulus.

Adult↗

A population decoding framework for motion aftereffects on smooth pursuit eye movements.

Both perceptual and motor systems must decode visual information from the distributed activity of large populations of cortical neurons. We have sought a common framework for understanding decoding strategies for visually guided movement and perception by asking whether the strong motion aftereffects seen in the perceptual domain lead to similar expressions in motor output. We found that motion adaptation indeed has strong sequelae in the direction and speed of smooth pursuit eye movements. After adaptation with a stimulus that moves in a given direction for 7 sec, the direction of pursuit is repelled from the direction of pursuit targets that move within 90 degrees of the adapting direction. The speed of pursuit decreases for targets that move at the direction and speed of the adapting stimulus and is repelled from the adapting speed in the sense that the decrease either becomes greater or smaller (eventually turning to an increase) when tracking targets move slower or faster than the adapting speed. The effects of adaptation are spatially specific and fixed to the retinal location of the adapting stimulus. The magnitude of adaptation of pursuit speed and direction is uncorrelated, suggesting that the two parameters are decoded independently. Computer simulation of motion adaptation in the middle temporal visual area (MT) shows that vector-averaging decoding of the population response in MT can account for the effects of adaptation on the direction of pursuit. Our results suggest a unified framework for thinking, in terms of population decoding, about motion adaptation for both perception and action.

Adaptation, Physiological↗

A gain-control model relating nulling results to the duration of dynamic motion aftereffects.

Strength of the motion aftereffect (MAE) is most often quantified by its duration, a high-variance and rather 'subjective' measure. With the help of an automatic gain-control model we quantitatively relate nulling-thresholds, adaptation strength, direction discrimination threshold, and duration of the dynamic MAE (dMAE). This shows how the nulling threshold, a more objective two-alternative forced-choice measure, relates to the same system property as MAE-durations. Two psychophysical experiments to test the model use moving random-pixel-arrays with an adjustable luminance signal-to-noise ratio. We measure MAE-duration as a function of adaptation strength and compare the results to the model prediction. We then do the same for nulling-thresholds. Model predictions are strongly supported by the psychophysical findings. In a third experiment we test formulae coupling nulling threshold, MAE-duration, and direction-discrimination thresholds, by measuring these quantities as a function of speed. For the medium-to-high speed range of these experiments we found that nulling thresholds increase and dMAE-durations decrease about linearly, whereas direction discrimination thresholds increase exponentially with speed. The model description then suggests that the motion-gain decreases, while the noise-gain and model's threshold increase with speed.

Adaptation, Physiological↗

Motion over the retina and the motion aftereffect.

The motion aftereffect (MAE) was measured with retinally moving vertical gratings positioned above and below (flanking) a retinally stationary central grating (experiments 1 and 2). Motion over the retina was produced by leftward motion of the flanking gratings relative to the stationary eyes, and by rightward eye or head movements tracking the moving (but retinally stationary) central grating relative to the stationary (but retinally moving) surround gratings. In experiment 1 the motion occurred within a fixed boundary on the screen, and oppositely directed MAEs were produced in the central and flanking gratings with static fixation; but with eye or head tracking MAEs were reported only in the central grating. In experiment 2 motion over the retina was equated for the static and tracking conditions by moving blocks of grating without any dynamic occlusion and disclosure at the boundaries. Both conditions yielded equivalent leftward MAEs of the central grating in the same direction as the prior flanking motion, ie an MAE was consistently produced in the region that had remained retinally stationary. No MAE was recorded in the flanking gratings, even though they moved over the retina during adaptation. When just two gratings were presented, MAEs were produced in both, but in opposite directions (experiments 3 and 4). It is concluded that the MAE is a consequence of adapting signals for the relative motion between elements of a display.

Attention↗

Topography of evoked potentials associated with illusory motion perception as a motion aftereffect.

Motion aftereffect (MAE) is a type of motion illusion. After visual focusing on an object moving in one direction, an illusory perception of motion in the opposite direction occurs while the object suddenly stops moving. In this study we explored components and distribution of evoked potentials related to this motion illusion using MAE caused by motion of concentric rings. When a single array of moving rings was placed to straddle right and left visual fields, a significant bilateral increase of a positive component at about 160 ms (P160) was observed in the occipitotemporal region at the time subjects perceived the motion illusion; this increase was most prominent in the right posterior temporal region. Thus, an early positive component P160 occurs in relation to motion illusion, in agreement with previous results concerning perception of actual motion. When stimuli were presented to produce MAE limited to either the right or left visual hemifield, we also observed a P160 distributed mainly in the right temporal and parietal region. A significant increase in this component was observed in the right posterior temporal region with left hemifield stimulation, while no significant increase was observed with right hemifield stimulation. The right hemispheric dominance of P160 seemed to result partly from functional specialization of the right hemisphere, but hemispheric differences in attentional mechanisms also might contribute to the asymmetric distribution of P160.

Adult↗

Storage for free: a surprising property of a simple gain-control model of motion aftereffects.

If a motion aftereffect (MAE) for given adaptation conditions has a duration T s, and the eyes are closed after adaptation during a waiting period tw=T s before testing, an unexpected MAE of a 'residual' duration TrT s is experienced. This effect is called 'storage' and it is often quantified by a storage factor sigma=TrT/T, which can reach values up to about 0.7-0.8. The phenomenon and its name have invited explanations in terms of inhibition of recovery during darkness. We present a model based on the opposite idea, that an effective test stimulus quickens recovery relative to darkness or other ineffective test stimuli. The model is worked out in mathematical detail and proves to explain 'storage' data from the literature, on the static MAE (sMAE: an MAE experienced for static test stimuli). We also present results of a psychophysical experiment with moving random pixel arrays, quantifying storage phenomena both for the sMAE and the dynamic MAE (dMAE: an MAE experienced for a random dynamic noise test stimulus). Storage factors for the dMAE are lower than for the sMAE. Our model also gives an excellent description of these new data on storage of the dMAE. The term 'storage' might therefore be a misnomer. If an effective test stimulus influences all direction tuned motion sensors indiscriminately and thus speeds up equalization of gains, one gets the storage phenomenon for free.

Adaptation, Ocular↗

Psychohysical hallucinations of orientation and spatial frequency.

After inspection of vertical sinusoidal gratings at least three distinct types of subjective or "hallucinated" patterns can be seen on a uniform test field. One type, here called horizontal streaming (H), is already well-known from the work of MacKay. A second type (V) looks like aroughly sinusoidal grating about 1-5 octaves above the adapting spitial frequency. Under optimal conditions a second vertical component appears at about 2 octaves below the adapting frequency. The third category of aftereffect consists of diagonal lines (D) at two orientations (about +/-40 degrees from vertical). The spatial-frequency band at these two orientations appears to be fairly broad, but roughly similar to the adapting frequency. The duration and strength of D increased, while V declined, at higher adapting spatial frequencies. D and V were increasing functions of adapting contrast, while H appeared abruptly only after the highest adapting contrast. H, D, and V are thus all functionally distinct. A schematic model of cortical organization is proposed to account for these phenomena. Pattern channels selective for a given orientation are grouped together with movement channels selective for the orthogonal direction. Antagonism between channels within such "modules" accounts for the streaming effect (H). Inhibition between modules tuned to different orientations and spatial frequencies accounts for the D and V effects: after adaptation of channels in one module, neighbouring module(s) are released from inhibition to produce a spurious response which is seen as a grating-like object in the adapted part of the visual field. During flickering adaptation a "halluncinated" lattice can be seen superimposed on the adapting grating. It apparently consists of Fourier components more remote from the adapting pattern than D and V are. This disinhibitory effect is strong confirmation of the inhibitory model. The regular and highly organized matrix of channels implied by these experiments may constitute a cortical hypercolumn conducting a coarse, piecewise Fourier transformation of the retinal image.

Figural Aftereffect↗

Behavioral determination of the spatial selectivity of contrast adaptation in cats: some evidence for a common plan in the mammalian visual system.

An aftereffects paradigm was used to behaviorally measure contrast sensitivity of cats to gratings of three different test spatial frequencies after adaptation to gratings of various spatial frequencies, contrasts, and durations. Post-adaptation reductions in sensitivity occurred even after short periods of adaptation (less than 7 s) and could be as large as 1.0 log unit under some conditions. The magnitude of the adaptation effect varied monotonically with (1) adaptation grating contrast, (2) duration, and (3) the contrast sensitivity for the test grating. Average half-width (at half-height) of the spatial-frequency tuning curves constructed from the data was 1.4 octaves, and was not dependent upon the level of adaptation or the spatial frequency of the test grating. Post-adaptation psychometric functions of the cats showed reduced slopes and maxima suggesting that, unlike humans, in cats apparent contrast grows more slowly with increases in physical contrast after contrast adaptation. All of the characteristics observed are in excellent agreement with electrophysiologically measured properties of neurons in striate cortex of cats. In addition, there was a remarkable similarity of the cat tuning functions, both in shape and bandpass, to those measured in man with a similar paradigm suggesting that (1) the two visual systems are sufficiently similar to make the cat a useful spatial vision model and (2) there is a common functional plan to all mammalian visual systems despite significant anatomical differences between species.

Adaptation, Ocular↗

Extra-retinal adaptation of cortical motion-processing areas during pursuit eye movements.

Repetitive eye movement produces a compelling motion aftereffect (MAE). One mechanism thought to contribute to the illusory movement is an extra-retinal motion signal generated after adaptation. However, extra-retinal signals are also generated during pursuit. They modulate activity within cortical motion-processing area MST, helping transform retinal motion into motion in the world during an eye movement. Given the evidence that MST plays a key role in generating MAE, it may also become indirectly adapted by prolonged pursuit. To differentiate between these two extra-retinal mechanisms we examined storage of the MAE across a period of darkness. In one condition observers were told to stare at a moving pattern, an instruction that induces a more reflexive type of eye movement. In another they were told to deliberately pursue it. We found equally long MAEs when testing immediately after adaptation but not when the test was delayed by 40 s. In the case of the reflexive eye movement the delay almost completely extinguished the MAE, whereas the illusory motion following pursuit remained intact. This suggests pursuit adapts cortical motion-processing areas whereas unintentional eye movement does not. A second experiment showed that cortical mechanisms cannot be the sole determinant of pursuit-induced MAE. Following oblique pursuit, we found MAE direction changes from oblique to vertical. Perceived MAE direction appears to be influenced by a subcortical mechanism as well, one based on the relative recovery rate of horizontal and vertical eye-movement processes recruited during oblique pursuit.

Adaptation, Ocular↗

Colored aftereffects contingent upon global transformations?

Dodwell and Humphrey (Psychol. Rev. 97, 78-79, 1990) have argued that colored aftereffects (CAEs) are contingent on the global geometries of local orientation and edge components in induction patterns. The spatiotopic dimensions underlying these geometries are derived from previous applications of the mathematics of continuous transformations (Lie groups) to the stimulus dynamics of perceptual invariance. In these terms, complementary CAEs induced by inspecting alternating colored bullseyes and spokes are attributed to the adaptation of orthogonal spatiotopic channels; the former characterized by vectorfields dealing with object invariance under rotation, and the latter by vectorfields dealing with object invariance under dilation. Importantly, the pattern-contingent CAEs induced in this pair of pattern channels are independent of CAEs induced in channels characterized by vectorfields dealing with object invariance under vertical and horizontal translations (e.g. vertical and horizontal gratings). The present investigation examined whether CAEs contingent on the stimulus dynamics originally used to define these spatiotopic channels exhibit comparable properties. It is confirmed that pairs of complementary CAEs can be induced simultaneously with radially expanding and contracting bullseyes, and clockwise- and counter-clockwise-rotating spokes (Experiment 1); and it is shown that these CAEs combine linearly in generalizing to and from the radial and rotational components of spiral motion (Experiments 2 and 3 respectively). In direct contrast to findings for CAEs induced with stationary versions of these patterns, however, it is demonstrated that complementary pairs of CAEs induced by expanding and contracting bullseyes interact with CAEs induced by left- and right-moving vertical gratings, suggesting that the components of motion underlying these contingent effects are locally and not globally determined (Experiment 4).

Color Perception↗

Bivectorial transparent stimuli simultaneously adapt mechanisms at different levels of the motion pathway.

The motion aftereffect (MAE) to drifting bivectorial stimuli, such as plaids, is usually univectorial and in a direction opposite to the pattern direction of the plaid. This is true for plaids that are perceived as coherent, but also for other plaids which are seen as transparent for most or all of the adaptation period. The underlying mechanisms of this MAE are still not well understood. In order to assess these mechanisms further, we measured static and dynamic MAEs and their interocular transfer (IOT). Adaptation stimuli were plaids with small (coherent) and large (transparent) angles between the directions of the component gratings and a horizontal grating, which were adjusted in spatial frequency and drift velocity so that the pattern speed and vertical periodicity remained constant. Test stimuli were horizontal static or counterphasing gratings with the same periodicity as the adaptation stimuli. MAE duration was measured for monocular, binocular and IOT conditions. All static MAEs were smallest for the transparent plaid and largest for the grating, while all dynamic MAEs were constant across adaptation stimuli. IOT was twice as big for dynamic MAEs as for static MAEs, and did not vary with the adaptation stimuli. Other adaptation stimuli were plaids that differed in intersection luminance, contrast or spatial frequency, resulting in different amounts of perceived coherence. MAEs and IOT did not vary with perceived coherence. The results suggest that the MAE for bivectorial stimuli consists of low-level adaptation (dependent on local component properties, small IOT), as well as high-level adaptation (dependent on global integrated pattern properties, large IOT), which can be measured independently with static and dynamic test stimuli.

Adaptation, Ocular↗

Recovery of fMRI activation in motion area MT following storage of the motion aftereffect.

We used functional magnetic resonance imaging (fMRI) during storage of the motion aftereffect (MAE) to examine the relationship between motion perception and neural activity in the human cortical motion complex MT+ (including area MT and adjacent motion-selective cortex). MT+ responds not only to physical motion but also to illusory motion, as in the MAE when subjects who have adapted to continuous motion report that a subsequent stationary test stimulus appears to move in the opposite direction. In the phenomenon of storage, the total decay time of the MAE is extended by inserting a dark period between adaptation and test phases. That is, when the static test pattern is presented after a storage period equal in duration to the normal MAE, the illusory motion reappears for almost as long as the original effect despite the delay. We examined fMRI activation in MT+ during and after storage. Seven subjects viewed continuous motion, followed either by an undelayed stationary test (immediate MAE) or by a completely dark storage interval preceding the test (stored MAE). Like the perceptual effect, activity in MT+ dropped during the storage interval then rebounded to reach a level much higher than after the same delay without storage. Although MT+ activity was slightly enhanced during the storage period following adaptation to continuous motion (compared with a control sequence in which the adaptation grating oscillated and no MAE was perceived), this enhancement was much less than that observed during the perceptual phenomenon. These results indicate that following adaptation, activity in MT+ is pronounced only with the presentation of an appropriate visual stimulus, during which the MAE is perceived.

Figural Aftereffect↗

The motion aftereffect of transparent motion: two temporal channels account for perceived direction.

Adaptation to orthogonal transparent patterns drifting at the same speed produces a unidirectional motion aftereffect (MAE) whose direction is opposite the average adaptation direction. If the patterns move at different speeds, MAE direction can be predicted by an inverse vector average, using the observer's motion sensitivity to each individual pattern as vector magnitudes. These weights are well approximated by the duration of each pattern's MAE, as measured with static test patterns. However, previous efforts to use the inverse-vector-average rule with dynamic test patterns have failed. Generally, these studies have used spatially and temporally broadband test stimuli. Here, in order to gain insight into the possible contribution of temporal channels, we filtered our test pattern in the temporal domain to produce five ideal, octave-width pass-bands. MAE durations were measured for single-component stimuli drifting at various adaptation speeds and tested at a range of temporal frequencies. Then, two components with orthogonal directions and different speeds were combined and the direction of the resulting MAE was measured. The key findings are that: (i) for a given adaptation speed, the duration of a single component's MAE is dependent on test temporal frequency; (ii) the direction of MAEs produced by transparent motion (i.e., bivectorial adaptation) also varies strongly as a function test temporal frequency (by up to 90 degrees for some speed pairings); and (iii) the inverse-vector-average rule predicts the direction of the transparent MAE provided the MAE durations used to weight the vector combination were obtained from stimuli matched in adaptation speed and test temporal frequency. These results are discussed in terms of the number and shape of temporal channels in our visual system.

Adaptation, Physiological↗