Apparent Motion Path Research Articles

Tactile interactions in the path of tactile apparent motion

Perceptual completion is a fundamental perceptual function serving to maintain robust perception against noise. For example, we can perceive a vivid experience of motion even for the discrete inputs across time and space (apparent motion: AM). In vision, stimuli irrelevant to AM perception are suppressed to maintain smooth AM perception along the AM trajectory where no physical inputs are applied. We investigated whether such perceptual masking induced by perceptual completion of dynamic inputs is general across sensory modalities by focusing on touch. Participants tried to detect a vibro-tactile target stimulus presented along the trajectory of AM induced by two other tactile stimuli on the forearm. In a control condition, the inducing stimuli were applied simultaneously, resulting in no motion percept. Tactile target detection was impaired with tactile AM. Our findings support the notion that the perceptual masking induced by perceptual completion mechanism of AM is a general function rather than a sensory specific effect.

Apparent Motion Induces Activity Suppression in Early Visual Cortex and Impairs Visual Detection.

Apparent motion (AM) is induced when two stationary visual stimuli are presented in alternating sequence. Intriguingly, AM leads to an impaired detectability of stimuli along the AM path (i.e., AM-induced masking). It has been hypothesized that AM triggers an internal representation of a moving object in early visual cortex, which competes with stimulus-evoked representations of visual stimuli on the motion path in early visual cortex of 25 human adults (16 female). We tested this hypothesis by measuring BOLD responses in early visual cortex during the process of AM-induced masking, using fMRI and population receptive field methods. Surprisingly, and counter to our hypothesis, we showed that AM suppressed, rather than increased, BOLD responses along early visual (V1 and V2) representations of the AM path, including regions that were not directly activated by the AM inducer stimuli. This activity suppression of the visual response predicted the subsequent reduction in detectability of the target that appeared in the middle of the AM path. Our data thereby provide direct empirical evidence for suppressive neural mechanisms underlying AM and suggest that illusory motion can render us blind to objects on the motion path by suppressing neural activity at the earliest cortical stages of visual perception.SIGNIFICANCE STATEMENT When two spatially distinct visual objects are presented in alternating sequence, apparent motion (AM) occurs and impairs detectability of stimuli along its path. The underlying mechanism is thought to be that increased activation in human early visual cortex evoked by AM interferes with the representation of the stimulus. Strikingly, however, we show that AM suppresses neural activity along the motion path, and the strength of activity suppression predicts the subsequent behavioral performance decrement in terms of detecting a stimulus along the AM path. Our findings provide empirical evidence for a suppressive, rather than faciliatory, mechanism underlying AM.

Suppressive Traveling Waves Shape Representations of Illusory Motion in Primary Visual Cortex of Awake Primate.

How does the brain link visual stimuli across space and time? Visual illusions provide an experimental paradigm to study these processes. When two stationary dots are flashed in close spatial and temporal succession, human observers experience a percept of apparent motion. Large spatiotemporal separation challenges the visual system to keep track of object identity along the apparent motion path, the so-called "correspondence problem." Here, we use voltage-sensitive dye imaging in primary visual cortex (V1) of awake monkeys to show that intracortical connections within V1 can solve this issue by shaping cortical dynamics to represent the illusory motion. We find that the appearance of the second stimulus in V1 creates a systematic suppressive wave traveling toward the retinotopic representation of the first. Using a computational model, we show that the suppressive wave is the emergent property of a recurrent gain control fed by the intracortical network. This suppressive wave acts to explain away ambiguous correspondence problems and contributes to precisely encode the expected motion velocity at the surface of V1. Together, these results demonstrate that the nonlinear dynamics within retinotopic maps can shape cortical representations of illusory motion. Understanding these dynamics will shed light on how the brain links sensory stimuli across space and time, by preformatting population responses for a straightforward read-out by downstream areas.SIGNIFICANCE STATEMENT Traveling waves have recently been observed in different animal species, brain areas, and behavioral states. However, it is still unclear what are their functional roles. In the case of cortical visual processing, waves propagate across retinotopic maps and can hereby generate interactions between spatially and temporally separated instances of feedforward driven activity. Such interactions could participate in processing long-range apparent motion stimuli, an illusion for which no clear neuronal mechanisms have yet been proposed. Using this paradigm in awake monkeys, we show that suppressive traveling waves produce a spatiotemporal normalization of apparent motion stimuli. Our study suggests that cortical waves shape the representation of illusory moving stimulus within retinotopic maps for a straightforward read-out by downstream areas.

Temporal ventriloquism along the path of apparent motion: speed perception under different spatial grouping principles.

The coordination of intramodal perceptual grouping and crossmodal interactions plays a critical role in constructing coherent multisensory percepts. However, the basic principles underlying such coordinating mechanisms still remain unclear. By taking advantage of an illusion called temporal ventriloquism and its influences on perceived speed, we investigated how audiovisual interactions in time are modulated by the spatial grouping principles of vision. In our experiments, we manipulated the spatial grouping principles of proximity, uniform connectedness, and similarity/common fate in apparent motion displays. Observers compared the speed of apparent motions across different sound timing conditions. Our results revealed that the effects of sound timing (i.e., temporal ventriloquism effects) on perceived speed also existed in visual displays containing more than one object and were modulated by different spatial grouping principles. In particular, uniform connectedness was found to modulate these audiovisual interactions in time. The effect of sound timing on perceived speed was smaller when horizontal connecting bars were introduced along the path of apparent motion. When the objects in each apparent motion frame were not connected or connected with vertical bars, the sound timing was more influential compared to the horizontal bar conditions. Overall, our findings here suggest that the effects of sound timing on perceived speed exist in different spatial configurations and can be modulated by certain intramodal spatial grouping principles such as uniform connectedness.

Apparent Motion Suppresses Responses in Early Visual Cortex: A Population Code Model.

Two stimuli alternately presented at different locations can evoke a percept of a stimulus continuously moving between the two locations. The neural mechanism underlying this apparent motion (AM) is thought to be increased activation of primary visual cortex (V1) neurons tuned to locations along the AM path, although evidence remains inconclusive. AM masking, which refers to the reduced detectability of stimuli along the AM path, has been taken as evidence for AM-related V1 activation. AM-induced neural responses are thought to interfere with responses to physical stimuli along the path and as such impair the perception of these stimuli. However, AM masking can also be explained by predictive coding models, predicting that responses to stimuli presented on the AM path are suppressed when they match the spatio-temporal prediction of a stimulus moving along the path. In the present study, we find that AM has a distinct effect on the detection of target gratings, limiting the maximum performance at high contrast levels. This masking is strongest when the target orientation is identical to the orientation of the inducers. We developed a V1-like population code model of early visual processing, based on a standard contrast normalization model. We find that AM-related activation in early visual cortex is too small to either cause masking or to be perceived as motion. Our model instead predicts strong suppression of early sensory responses during AM, consistent with the theoretical framework of predictive coding.

Decoding information about dynamically occluded objects in visual cortex

During dynamic occlusion, an object passes behind an occluding surface and then later reappears. Even when completely occluded from view, such objects are experienced as continuing to exist or persist behind the occluder even though they are no longer visible. The contents and neural basis of this persistent representation remain poorly understood. Questions remain as to whether there is information maintained about the object itself (i.e. its shape or identity) or non-object-specific information such as its position or velocity as it is tracked behind an occluder, as well as which areas of visual cortex represent such information. Recent studies have found that early visual cortex is activated by “invisible” objects during visual imagery and by unstimulated regions along the path of apparent motion, suggesting that some properties of dynamically occluded objects may also be neurally represented in early visual cortex. We applied functional magnetic resonance imaging in human subjects to examine representations within visual cortex during dynamic occlusion. For gradually occluded, but not for instantly disappearing objects, there was an increase in activity in early visual cortex (V1, V2, and V3). This activity was spatially-specific, corresponding to the occluded location in the visual field. However, the activity did not encode enough information about object identity to discriminate between different kinds of occluded objects (circles vs. stars) using MVPA. In contrast, object identity could be decoded in spatially-specific subregions of higher-order, topographically organized areas such as ventral, lateral, and temporal occipital areas (VO, LO, and TO) as well as the functionally defined LOC and hMT+. These results suggest that early visual cortex may only represent the dynamically occluded object's position or motion path, while later visual areas represent object-specific information.

Representations along the path of apparent motion in visual cortex

Representations along the path of apparent motion in visual cortex

Reconstructing representations of dynamic visual objects in early visual cortex

As raw sensory data are partial, our visual system extensively fills in missing details, creating enriched percepts based on incomplete bottom-up information. Despite evidence for internally generated representations at early stages of cortical processing, it is not known whether these representations include missing information of dynamically transforming objects. Long-range apparent motion (AM) provides a unique test case because objects in AM can undergo changes both in position and in features. Using fMRI and encoding methods, we found that the "intermediate" orientation of an apparently rotating grating, never presented in the retinal input but interpolated during AM, is reconstructed in population-level, feature-selective tuning responses in the region of early visual cortex (V1) that corresponds to the retinotopic location of the AM path. This neural representation is absent when AM inducers are presented simultaneously and when AM is visually imagined. Our results demonstrate dynamic filling-in in V1 for object features that are interpolated during kinetic transformations.

Color updating on the apparent motion path.

When a static stimulus appears successively at two distant locations, we perceive illusory motion of the stimulus across them-long-range apparent motion (AM). Previous studies have shown that when the apparent motion stimuli differ in shape, interpolation between the two shapes is perceived across the AM path. In contrast, the perceived color during AM has been shown to abruptly change from the color of the first stimulus into that of the second, suggesting interpolation does not occur for color during AM. Here, we report the first evidence to our knowledge, that an interpolated color, distinct from the colors of either apparent motion stimulus, is represented as the intermediate percept on the path of apparent motion. Using carefully chosen target colors-cyan, pink, and lime-that are perceptually and neurally intermediate between blue and green, orange and magenta, and green and orange respectively, we show that detection of a target presented on the apparent motion path was impaired when the color of the target was "in-between" the initial and terminal stimulus colors. Furthermore, we show that this feature-specific masking effect for the intermediate color cannot be accounted for by color similarity between the intermediate color and the color of the terminal inducer. Our findings demonstrate that intermediate colors can be interpolated over the apparent motion trajectory as in the case of shape, possibly involving similar interpolation processes for shape and color during apparent motion.

Backward position shift in apparent motion

We investigated the perceived position of visual targets in apparent motion. A disc moved horizontally through three positions from -10° to +10° in the far periphery (20° above fixation), generating a compelling impression of apparent motion. In the first experiment, observers compared the position of the middle of the three discs to a subsequently presented reference. Unexpectedly, observers judged its position to be shifted backward, in the direction opposite that of the motion. We then tested the middle disc in sequences of 3, 5, and 7 discs, each covering the same spatial and temporal extents (similar speeds). The backwards shift was only found for the three-disc sequence. With the extra discs approaching more continuous motion, the perceived shift was in the same direction as the apparent motion. Finally, using a localization task with constant static references, we measured the position shifts of all the disc locations for two-disc, three-disc and four-disc apparent motion sequences. The backward shift was found for the second location of all sequences. We suggest that the backward shift of the second element along an apparent motion path is due to an attraction effect induced by the initial point of the motion.

Perceived Causality Can Alter the Perceived Trajectory of Apparent Motion

When objects collide, human observers perceive not only motion but also causal relations, such as which object caused the other to move. In the present experiments, we investigated whether such causal interpretations can actually influence the perceived path of apparent motion. Displays contained two alternately flashing motion targets positioned at either end of a semicircular occluder. Two additional “context objects” moved in such a way that the motion targets appeared to collide with and launch them. The collision was manipulated so that it was consistent with apparent motion either along the straight path between the targets or along a curved path passing behind the occluder. Subjects almost exclusively perceived motion consistent with the implied launch, which suggests that causally coherent interpretations can influence basic perceptual processes.

Reported visual imagery and apparent motion

A previous behavioral study showed that a group of individuals with high vividness of visual imagery (High group), as determined from the score for the Vividness of Visual Imagery Questionnaire (VVIQ), could perceive the apparent motion path more strongly than a group of individuals with low vividness of visual imagery (Low group). To examine the physiological differences underlying these differences in perception, we compared the brain activity during an apparent motion task for the High and the Low groups using electroencephalography. We initially screened 60 potential participants using the VVIQ. On the basis of their scores, we invited 20 people from the lower and the higher ends of the VVIQ distribution to participate in our event-related potential study. Our results showed that individuals in both the High and the Low groups were sensitive to the apparent motion content of the task. Perception of apparent motion evoked a negative potential starting around 90 ms, followed by a positive potential beginning at 150-170 ms after the second stimulus. The scalp distributions of both negative and positive potentials for the High group were broader than those for the Low group. Moreover, the onset of positivity in the High group (150 ms) was earlier than that in the Low group (170 ms). We believe that these results may be mechanistically associated with the differences in the perception of apparent motion between individuals with high and low vividness of visual imagery.

S6-2: Neural Representation in the Apparent Motion Path

When a static stimulus appears successively at two different locations, we perceive a transition of the stimulus across them—apparent motion. Previous studies have shown that some spatio-temporal representation is reconstructed in the apparent motion path and it leads to increased activation in the region of the primary visual cortex (V1) corresponding to the apparent motion path (Muckli et al., 2005 PLoS Biology 3 1501–1510). However, little is known about whether visual properties of an object engaged in apparent motion are maintained in this representation. In order to address this question, we used fMRI and pattern classification methods to examine the neural representation created on the apparent motion path when the object changes its orientation across the apparent motion path. Two intermediate orientations (0° and 90°) on the apparent motion path were induced by presenting gratings with different orientations at separate locations. The gratings were presented in a bistable quartet sequence, and su...

Curved Apparent Motion Initiated by a Causal Launch

When objects collide, observers perceive not only the motion but also causal relations, such as which objects caused which others to move. The present study investigated whether such causal interpretations can influence the perceived path of apparent motion. We presented a display of two alternately flashing motion tokens on the ends of a semicircular occluder, and two additional “context objects” placed immediately above the tokens moved upward at each token onset. In such a display the motion token was seen as moving along the curved path behind the occluder, deviating from the default shortest straight path. This finding suggested that observers spontaneously attributed the vertical displacement of the context object to collision by the motion token, which affected the percept of the path of the colliding token. To rule out a potential alternative explanation based on motion priming, a subsequent experiment tested new displays in which the spatial or temporal pattern of context events was altered in wa...

Representation of stimulus features in V1 along the apparent motion path

Representation of stimulus features in V1 along the apparent motion path

Representation of stimulus features in V1 along the apparent motion path

Representation of stimulus features in V1 along the apparent motion path

Masking and color inheritance along the apparent motion path

Long-range apparent motion is the illusory motion that can be perceived when two static and distant stimuli are presented in succession. Within some spatiotemporal range not only is motion sensed, but it appears as if one stimulus is displaced from one place to another (termed beta or optimal motion). Several groups have found that this illusory percept can interact with perception of a physically present stimulus, but some disagree on the origin of these interactions. We know little about how suppressive effects depend on feature-similarity between a target and the stimuli in apparent motion (inducers)-which would indicate an early perceptual locus-or even about the minimal conditions under which to obtain this effect. Unlike early studies that used a two-stroke apparent motion paradigm, we were able to demonstrate that motion can mask stimuli presented at interpolated locations along the apparent motion path, as shown by the elevation of contrast thresholds compared to a control condition. Apparent motion masking depended on color similarity between target and inducers. Further, we found evidence that the color of inducers alters the apparent color of intervening gray probes, indicating some inheritance or chromatic averaging across distant locations, but no clear evidence of predictive updating. Finally, the analysis of the presentation times delivering maximal masking effects suggests a predictive interpolation process is responsible for interference by apparent motion filling-in. We discuss alternative mechanisms, in particular the possible role of apparent-motion-induced metacontrast masking in generating this pattern of results.

"Non-retinotopic processing" in Ternus motion displays modeled by spatiotemporal filters

Recently, M. Boi, H. Ogmen, J. Krummenacher, T. U. Otto, & M. H. Herzog (2009) reported a fascinating visual effect, where the direction of apparent motion was disambiguated by cues along the path of apparent motion, the Ternus-Pikler group motion, even though no actual movement occurs in this stimulus. They referred to their study as a "litmus test" to distinguish "non-retinotopic" (motion-based) from "retinotopic" (retina-based) image processing. We adapted the test to one with simple grating stimuli that could be more readily modeled and replicated their psychophysical results quantitatively with this stimulus. We then modeled our experiments in 3D (x, y, t) Fourier space and demonstrated that the observed perceptual effects are readily accounted for by integration of information within a detector that is oriented in space and time, in a similar way to previous explanations of other motion illusions. This demonstration brings the study of Boi et al. into the more general context of perception of moving objects.

Auditory Motion Capturing Ambiguous Visual Motion

In this study, it is demonstrated that moving sounds have an effect on the direction in which one sees visual stimuli move. During the main experiment sounds were presented consecutively at four speaker locations inducing left or rightward auditory apparent motion. On the path of auditory apparent motion, visual apparent motion stimuli were presented with a high degree of directional ambiguity. The main outcome of this experiment is that our participants perceived visual apparent motion stimuli that were ambiguous (equally likely to be perceived as moving left or rightward) more often as moving in the same direction than in the opposite direction of auditory apparent motion. During the control experiment we replicated this finding and found no effect of sound motion direction on eye movements. This indicates that auditory motion can capture our visual motion percept when visual motion direction is insufficiently determinate without affecting eye movements.

Contextual and Predictive Coding in V1

Primary visual cortex (V1) is often characterized by the receptive field properties of its feed-forward input (only up to 5%). We focussed our fMRI studies on nonstimulated retinotopic regions in V1 to gain a better understanding of contextual processing. We investigated activation along the nonstimulated long-range apparent motion path (1), occluded a visual quarterfield of a natural visual scene (2), or blindfolded our subjects and presented environmental sounds (3). We were able to demonstrate predictive activity along the illusory apparent motion path (1), use decoding to classify natural scenes from nonstimulated regions in V1 (2), and to decode environmental sounds from V1, V2 and V3 (3). Is this contextual processing useful to predict upcoming visual events? To investigate predictability we used apparent motion as the prime stimuli and tested with a probe stimulus along the apparent motion path to find that predicted stimuli are processed more efficiently—leading to less fMRI signal and better detectability (1). In summary, we have found brain imaging evidence consistent with the hypothesis of predictive coding in early visual areas.