Abstract: When visual input is inconclusive, does previous experience aid the visual system in attaining an accurate perceptual interpretation? Prolonged viewing of a visually ambiguous stimulus causes perception to alternate between conflicting interpretations. When viewed intermittently, however, ambiguous stimuli tend to evoke the same percept on many consecutive presentations. This perceptual stabilization has been suggested to reflect persistence of the most recent percept throughout the blank that separates two presentations. Here we show that the memory trace that causes stabilization reflects not just the latest percept, but perception during a much longer period. That is, the choice between competing percepts at stimulus reappearance is determined by an elaborate history of prior perception. Specifically, we demonstrate a seconds-long influence of the latest percept, as well as a more persistent influence based on the relative proportion of dominance during a preceding period of at least one minute. In case short-term perceptual history and long-term perceptual history are opposed (because perception has recently switched after prolonged stabilization), the long-term influence recovers after the effect of the latest percept has worn off, indicating independence between time scales. We accommodate these results by adding two positive adaptation terms, one with a short time constant and one with a long time constant, to a standard model of perceptual switching.
Abstract: During prolonged fixation, visual objects presented in the periphery of the visual field often fade from awareness. This phenomenon, known as the Troxler effect, has been largely attributed to adaptation of neurons responding to peripheral targets. Here, we hypothesized that perceptual disappearance might result from degeneration of feedback from attention-related cortical areas to early visual areas, and that visual transients disrupt the feedback loop sustaining low-level signals and thereby trigger perceptual fading. We examined this hypothesis by briefly disrupting the functions of attention-related regions in the intraparietal sulcus (IPS) using transcranial magnetic stimulation (TMS). The hypothesis predicted that temporary disruption of IPS would trigger perceptual disappearances. We measured perceptual disappearance using a task in which participants were asked to discriminate the presence or absence (fading) of a peripheral green target immediately after a TMS pulse. On half of trials the target remained on the screen until the end of a trial, and on the other half of trials it gradually faded. The results of this experiment show that brief disruption of the IPS with a single pulse TMS is sufficient to trigger perceptual disappearance. TMS over the IPS resulted in a fewer reports of continuous percepts (i.e. more fading) without changes in the perception of physically fading stimuli. Our control experiment shows the loss of sensitivity is not due to suppression of microsaccades in response to the sound produced by TMS. Our present study supports the idea that conscious visual perception requires a coupling between the early visual areas representing sensory data and the parietal areas subserving spatial localisation functions.
Abstract: BACKGROUND: The timing at which sensory input reaches the level of conscious perception is an intriguing question still awaiting an answer. It is often assumed that both visual and auditory percepts have a modality specific processing delay and their difference determines perceptual temporal offset. METHODOLOGY/PRINCIPAL FINDINGS: Here, we show that the perception of audiovisual simultaneity can change flexibly and fluctuates over a short period of time while subjects observe a constant stimulus. We investigated the mechanisms underlying the spontaneous alternations in this audiovisual illusion and found that attention plays a crucial role. When attention was distracted from the stimulus, the perceptual transitions disappeared. When attention was directed to a visual event, the perceived timing of an auditory event was attracted towards that event. CONCLUSIONS/SIGNIFICANCE: This multistable display illustrates how flexible perceived timing can be, and at the same time offers a paradigm to dissociate perceptual from stimulus-driven factors in crossmodal feature binding. Our findings suggest that the perception of crossmodal synchrony depends on perceptual binding of audiovisual stimuli as a common event.
Abstract: People often fail to select and encode the second of two targets presented within less than 500ms in rapid serial visual presentation (RSVP), an effect known as the attentional blink. We investigated how report of the two targets is affected when one of them is maintained in working memory for a secondary, memory-search task. The results showed that report of either target was impaired when it was a member of the memory set relative to when it was not. This effect was independent of both the temporal interval separating the RSVP target from the presentation of the memory set and the interval separating the targets. We propose that the deficit in recall occurs because the association between a target and the memory-search task interferes with the formation of a new association between that target and the following RSVP task, with the result that observers may be biased to ascribe the target only to the memory set.
Abstract: When the two eyes are presented with sufficiently different stimuli, the stimuli will engage in binocular rivalry. During binocular rivalry, a subject's perceptual state alternates between awareness of the stimulus presented to the right eye and that presented to the left eye. There are instances in which competition is not eye-based, but instead takes place between stimulus features, as is the case in flicker and switch rivalry (F&S). Here we investigate another such instance, interocular grouping, using a Diaz-Caneja type stimulus in conjunction with synchronous stimulus flicker. Our results indicate that stimulus flicker increases the total duration of interocularly bound percepts, and that this effect occurs for a range of temporal flicker frequencies. Furthermore, the use of contrast-inversion flicker causes a decrease of total dominance duration of the interocularly bound percepts. We argue that different flickering regimes can be used to differentially stimulate lower and higher levels of visual processing involved in binocular rivalry. We propose that the amount of interocularly combined pattern-completed percept can be regarded as a measure of the level at which binocular rivalry is resolved.
Abstract: We show that previewing one half image of a binocular rivalry pair can cause it to gain initial dominance when the other half is added, a novel phenomenon we term flash facilitation. This is the converse of a known effect called flash suppression, where the previewed image becomes suppressed upon rivalrous presentation. The exact effect of previewing an image depends on both the duration and the contrast of the prior stimulus. Brief, low-contrast prior stimuli facilitate, whereas long, high-contrast ones suppress. These effects have both an eye-based component and a pattern-based component. Our results suggest that, instead of reflecting two unrelated mechanisms, both facilitation and suppression are manifestations of a single process that occurs progressively during presentation of the prior stimulus. The distinction between the two phenomena would then lie in the extent to which the process has developed during prior stimulation. This view is consistent with a neural model previously proposed to account for perceptual stabilization of ambiguous stimuli, suggesting a relation between perceptual stabilization and the present phenomena.
Abstract: Motion is defined as a sequence of positional changes over time. However, in perception, spatial position and motion dynamically interact with each other. This reciprocal interaction suggests that the perception of a moving object itself may dynamically evolve following the onset of motion. Here, we show evidence that the percept of a moving object systematically changes over time. In experiments, we introduced a transient gap in the motion sequence or a brief change in some feature (e.g., color or shape) of an otherwise smoothly moving target stimulus. Observers were highly sensitive to the gap or transient change if it occurred soon after motion onset (< or =200 ms), but significantly less so if it occurred later (> or = 300 ms). Our findings suggest that the moving stimulus is initially perceived as a time series of discrete potentially isolatable frames; later failures to perceive change suggests that over time, the stimulus begins to be perceived as a single, indivisible gestalt integrated over space as well as time, which could well be the signature of an emergent stable motion percept.
Abstract: Visual rivalry is thought to be a distributed process that simultaneously takes place at multiple levels in the visual processing hierarchy. Also, the different types of rivalry, such as binocular and monocular rivalry, are thought to engage shared underlying mechanisms. We hypothesized that the amount of perceptual suppression during rivalry as measured by the total duration of fully exclusive perceptual dominance is determined by a distance in a neurally represented feature space. This hypothesis can be contrasted with the possibility that the brain constructs an internal model of the outside world using full-fledged object representations, and that perceptual suppression is due to an appraisal of the likelihood of the particular stimulus configuration at a high, object-based level. We applied color and stereo-depth differences between monocular rivalry stimulus gratings, and manipulated color and eye-of-origin information in binocular rivalry using the flicker & switch presentation paradigm. Our data show that exclusivity in visual rivalry increases with increased difference in feature space without regard for real-world constraints, and that eye-of-origin information may be regarded as a segregating feature that functions in a manner similar to color and stereo-depth information. Moreover, distances defined in multiple feature dimensions additively and independently increase the amount of perceptual exclusivity and coherence in both monocular and binocular rivalry. We conclude that exclusivity in visual rivalry is determined by a distance in feature space that is subtended by multiple stimulus features.
Abstract: Colour has been shown to facilitate the recognition of scene images, but only when these images contain natural scenes, for which colour is 'diagnostic'. Here we investigate whether colour can also facilitate memory for scene images, and whether this would hold for natural scenes in particular. In the first experiment participants first studied a set of colour and greyscale natural and man-made scene images. Next, the same images were presented, randomly mixed with a different set. Participants were asked to indicate whether they had seen the images during the study phase. Surprisingly, performance was better for greyscale than for coloured images, and this difference is due to the higher false alarm rate for both natural and man-made coloured scenes. We hypothesized that this increase in false alarm rate was due to a shift from scrutinizing details of the image to recognition of the gist of the (coloured) image. A second experiment, utilizing images without a nameable gist, confirmed this hypothesis as participants now performed equally on greyscale and coloured images. In the final experiment we specifically targeted the more detail-based perception and recognition for greyscale images versus the more gist-based perception and recognition for coloured images with a change detection paradigm. The results show that changes to images are detected faster when image-pairs were presented in greyscale than in colour. This counterintuitive result held for both natural and man-made scenes (but not for scenes without nameable gist) and thus corroborates the shift from more detailed processing of images in greyscale to more gist-based processing of coloured images.
Abstract: After viewing directional motion, one is likely to perceive a subsequently presented directionally ambiguous motion as being in the same direction as the prior motion. The perceptual bias towards the most recent percept gradually develops as the interval between the prior stimulus and a subsequent test becomes longer. This form of positive bias, or priming, is created in an automatic fashion. It remain unclear how such perceptual bias could be eliminated by a stimulus manipulation. Here we examine whether presentation of a stimulus, which was neutral as to the competing perceptual interpretations, during the interval between prior and test stimuli, disrupts the development of the priming effect. In experiments with ambiguous motion, we used stationary gratings as the neutral stimuli, and in an experiment with binocular rivalry between orthogonal gratings, we used a plaid pattern consisting of the two rival gratings. In both cases, presenting the neutral stimuli reduced the perceptual bias. These findings show that the visual system dynamically calibrates its internal bias using a recent percept and that this internal bias can be nullified by presenting neutral stimuli.
Abstract: Perceptual priming is generally regarded as a passive and automatic process, as it is obtained even without awareness of the prime. Recent studies have introduced a more active form of perceptual priming in which priming for a subsequent ambiguous stimulus is triggered by the subjective percept, that is, interpretation of a previous ambiguous stimulus. This phenomenon known as stabilization does not require a conscious effort to actively maintain one perceptual interpretation. In this study, we show that distraction of attention, during and even after the prime presentation, interferes with the build-up of perceptual memory for stabilization. This implies that despite the apparent automaticity, stabilization involves an active attentional process for encoding and retention. The disruption during the encoding can be attributed to the reduction in sensory signals for the prime. However, the disruption during the retention suggests that the implicit memory trace of the prime necessitates the attentional resource to fully develop. The active nature of the build-up of perceptual memory for stabilization is consistent with the idea that perceptual memory increases its strength gradually over a few seconds. These findings suggest that seemingly automatic and effortless cognitive processes can compete with online perceptual processing for common attentional resources.
Abstract: We investigated the effects of different viewpoints on remapping visuo-motor space, and whether remapping happens differently during the planning and the online control phase of goal-directed movements. Participants tapped targets on a monitor that was placed horizontally flat and flush with the table in front of them. They viewed the layout of the scene, including the monitor, and their hand, through video goggles attached to a camera. The camera could be moved along a semi-circle with the monitor as the circle's center. On each trial, the camera was randomly positioned at one of seven locations on the semi-circle (-90 to +90 degrees in 30 degrees steps), always at eye height. The time needed to tap the target was quickest when the camera was approximately facing the participant and progressively increased when the camera was located more to the sides (Experiment 1). There was no effect of camera location on performance when participants only saw the static layout of the scene and were not allowed to see the scene or their hand during the movement (Experiment 2). By comparison, the dependency of performance on camera location increased when participants did not have information about the layout of the scene at the start of the trial, and could only perform remapping when their hand was visible during the movement (Experiment 3). These results indicate that visuo-motor remapping happens differently during the planning phase when only static information about the layout is available, and during the control phase when dynamic information about the moving hand is also available.
Abstract: The position of a moving object is often mislocalised in the direction of movement. At the input stage of visual processing, the position of a moving object should still be represented veridically, whereas it should become closer to the mislocalised position at a later processing stage responsible for positional judgment. Here, we show that visual transients expose the veridical position of a moving object represented in early visual areas. For example, when a ring is flashed on a moving bar, the part of the bar within the ring is perceived at the veridical position, whereas the part outside the ring is perceived to be ahead of the ring as in the flash-lag effect. Our observations suggest that a filling-in process is triggered at the edges of the flash. This indicates that, in early cortical areas, moving objects are still represented at their veridical positions, and the perceived location is determined by the higher visual areas.
Abstract: How does the brain estimate time? This old question has led to many biological and psychological models of time perception (R. A. Block, 1989; P. Fraisse, 1963; J. Gibbon, 1977; D. L. I. Zakay, 1989). Because time cannot be directly measured at a given moment, it has been proposed that the brain estimates time based on the number of changes in an event (S. W. Brown, 1995; P. Fraisse, 1963; W. D. Poynter, 1989). Consistent with this idea, dynamic visual stimuli are known to lengthen perceived time (J. F. Brown, 1931; S. Goldstone & W. T. Lhamon, 1974; W. T. Lhamon & S. Goldstone, 1974, C. O. Z. Roelofs & W. P. C. Zeeman, 1951). However, the kind of information that constitutes the basis for time perception remains unresolved. Here, we show that the temporal frequency of a stimulus serves as the "clock" for perceived duration. Other aspects of changes, such as speed or coherence, were found to be inconsequential. Time dilation saturated at a temporal frequency of 4-8 Hz. These results suggest that the clock governing perceived time has its basis at early processing stages. The possible links between models of time perception and neurophysiological functions of early visual areas are discussed.
Abstract: Attentional selection plays a critical role in conscious perception. When attention is diverted, even salient stimuli fail to reach visual awareness. Attention can be voluntarily directed to a spatial location or a visual feature for facilitating the processing of information relevant to current goals. In everyday situations, attention and awareness are tightly coupled. This has led some to suggest that attention and awareness might be based on a common neural foundation, whereas others argue that they are mediated by distinct mechanisms. A body of evidence shows that visual stimuli can be processed at multiple stages of the visual-processing streams without evoking visual awareness. To illuminate the relationship between visual attention and conscious perception, we investigated whether top-down attention can target and modulate the neural representations of unconsciously processed visual stimuli. Our experiments show that spatial attention can target only consciously perceived stimuli, whereas feature-based attention can modulate the processing of invisible stimuli. The attentional modulation of unconscious signals implies that attention and awareness can be dissociated, challenging a simplistic view of the boundary between conscious and unconscious visual processing.
Abstract: Perceived color at a point in space is not determined simply by the color directly stimulating the corresponding retinal position. Surface color is informed by flanking edge signals, which also serve to inhibit the intrusion of signals from neighboring surfaces. Spatially continuous local interactions among color and luminance signals have been implicated in a propagation process often referred to as filling-in. Here, we report a phenomenon of discrete color filling whereby color jumps over luminance gaps filling into disconnected regions of the stimulus. This color filling is found to be blocked at boundaries defined by texture. The color filling is also highly specific to the elements belonging to a common perceptual surface, even when multiple surfaces are transparently overlaid. Our results indicate that color filling can be governed by a host of visual cues outside the realm of first-order color and brightness, via their impact on perceptual surface segmentation and segregation.
Abstract: We report a distortion of subjective time perception in which the duration of a first interval is perceived to be longer than the succeeding interval of the same duration. The amount of time expansion depends on the onset type defining the first interval. When a stimulus appears abruptly, its duration is perceived to be longer than when it appears following a stationary array. The difference in the processing time for the stimulus onset and motion onset, measured as reaction times, agrees with the difference in time expansion. Our results suggest that initial transient responses for a visual onset serve as a temporal marker for time estimation, and a systematic change in the processing time for onsets affects perceived time.
Abstract: Visual neurons show fast adaptive behavior in response to brief visual input. However, the perceptual consequences of this rapid neural adaptation are less known. Here, we show that brief exposure to a moving adaptation stimulus-ranging from tens to hundreds of milliseconds-influences the perception of a subsequently presented ambiguous motion test stimulus. Whether the ambiguous motion is perceived to move in the same direction (priming), or in the opposite direction (rapid motion aftereffect) varies systematically with the duration of the adaptation stimulus and the adaptation-test blank interval. These biases appear and decay rapidly. Moreover, when the adapting stimulus is itself ambiguous, these effects are not produced. Instead, the percept for the subsequent test stimulus is biased to the perceived direction of the adaptation stimulus. This effect (perceptual sensitization) builds gradually over the time between the adaptation and test stimuli. Our results indicate that rapid adaptation plays a role mainly within early motion processing, whereas a slow potentiation controls the sensitivity at a later stage.
Abstract: When our visual system is confronted with ambiguous stimuli, the perceptual interpretation spontaneously alternates between the competing incompatible interpretations. The timing of such perceptual alternations is highly stochastic and the underlying neural mechanisms are poorly understood. We show that perceptual alternations can be triggered by a transient stimulus presented nearby. The induction was tested for four types of bistable stimuli: structure-from-motion, binocular rivalry, Necker cube, and ambiguous apparent motion. While underlying mechanisms may vary among them, a transient flash induced time-locked perceptual alternations in all cases. The effect showed a dependence on the adaptation to the dominant percept prior to the presentation of a flash. These perceptual alternations show many similarities to perceptual disappearances induced by transient stimuli (Kanai and Kamitani, 2003 Journal of Cognitive Neuroscience 15 664-672; Moradi and Shimojo, 2004 Vision Research 44 449-460). Mechanisms linking these two transient-induced phenomena are discussed.
Abstract: When each eye is confronted with a dissimilar stimulus, the percept will generally alternate between the two. This phenomenon is known as binocular rivalry. Although binocular rivalry occurs at locations where targets overlap spatially, the area surrounding rivalrous targets can modulate their dominance. Here we show that during binocular rivalry of oppositely moving gratings, a surrounding grating moving in the same direction as one of the two leads to increased dominance of the opposite direction of motion in the center. This increased dominance of the opposite direction in the center was observed irrespective of the eye to which the surround was presented. Inspection of the results for different conditions reveals that the preference for the opposite direction of motion cannot be explained by a single mechanism operating beyond binocular fusion. We therefore suggest that this phenomenon is the outcome of center-surround interactions at multiple levels along the pathway of visual motion processing.
Abstract: When you see a red ball rolling across the floor, the ball's redness, roundness and motion appear to be unified and inseparably bound together as features of the ball. But neurophysiological evidence indicates that visual features such as colour, shape and motion are processed in separate regions of the brain. Here we describe an illusion that exploits this separation, causing colour and motion to be recombined incorrectly while a stable stimulus is being viewed continuously.
Abstract: Visual processing involves hierarchical stages in which local features are initially analyzed and subsequently grouped into objects and surfaces. In the domain of motion perception, transparent motion has been used as a powerful tool to investigate the mechanisms underlying the grouping of local features. Here, we report a novel way of creating motion transparency from oscillating dots (MTOD). In this stimulus, individual dots move back and forth over a small distance. When the dots are oscillating in synchrony, global surfaces are also perceived as moving back and forth. However, when the oscillation desynchronizes, the percept turns into two moving surfaces that are sliding over each other continuously (streaming motion). The percept of MTOD is similar to conventional transparent motion, where individual dots move only in one direction. Also, when streaming motion is perceived, the detection of oscillation is impaired. This blindness to the oscillation becomes stronger, as the signal strength for the streaming motion is increased. These findings suggest that when global visual representations are constructed, weak and inconsistent local signals are discarded.
Abstract: A moving object is perceived to lie beyond a static object presented at the same time at the same retinal location (flash-lag effect or FLE). Some studies report that if the moving stimulus stops moving (flash-terminated condition or FTC) the instant the flash occurs, a FLE does not occur. Other studies, using different stimuli, report that the FLE does, in fact, occur in the FTC. The FTC is thus a crucial turning point in theories of flash-lag. Unraveling the mystery of the FLE in the FTC will help unravel the mechanisms underpinning flash-lag and perhaps even perceptual localization in general. Our experiments show that eccentricity of the moving stimulus was a contributing factor, as were eccentricity of the flashed stimulus and spatial separation between the two stimuli. Other factors, such as contrast and offset of moving stimulus, also modulate the magnitude of the FLE in the FTC. We surmise that uncertainty in determining the position in space of a moving stimulus is a key requirement for the lag-effect. A lag-effect in the FTC challenges influential models, such as differential latency, motion extrapolation, and postdiction. Based partly on the notion of an asymmetric spread of activity that arises because of the sheer nature of motion and from a combination of established physiological mechanisms, we propose a schematic account of the present findings that subsumes previous psychological models and scaffolds past experimental findings.
Abstract: A visual transient due to a sudden visual change is generally considered to draw our attention to a location of interest. In a series of experiments we investigated how visual transients facilitate change detection in a scene. In line with earlier reports, we found that a transient sensation has its roots in a temporal interaction at a monocular processing level. Interestingly, we also show that visual transients make it possible to detect a change in the eye of origin, despite the fact that observers have no clue as to which eye is stimulated. That is, visual transients are detected even when there is no perceptual change in the visual content after binocular fusion. More importantly, we show that observers cannot distinguish the transient due to a change in eye of origin from a feature change (the orientation of a Gabor). Both are perceived as actual changes. We conclude that a transient signal is sufficient for the visual system to judge whether something has changed over time.
Abstract: In this study, we investigated the influence of smooth-pursuit eye movements on saccade initiation in response to a sudden jump of a continuously moving target. We replicated the finding by Tanaka et al. (1998) that saccadic eye movements in the direction opposite to preceding pursuit have longer latencies than those in the same direction. We confirmed that this asymmetry is indeed due to an inhibitory effect of smooth pursuit on saccade initiation in the opposite direction rather than facilitation of saccade initiation in the same direction. The inhibitory effect decreased strongly when subjects knew the jump direction in advance. This supports the notion that the prolonged latencies of backward saccades are not due to orbital mechanics or low-level motor processing. Furthermore, we found that the range of saccade directions inhibited by a pursuit movement is broad, covering all directions that did not have the same horizontal component as the pursuit direction. This is in contrast with the predictions of "Inhibition of Saccade Return" (ISR, Hooge and Frens 2000), which is restricted to a smaller confined area.
Abstract: After prolonged fixation, a stationary object placed in the peripheral visual field fades and disappears from our visual awareness, especially at low luminance contrast (the Troxler effect). Here, we report that similar fading can be triggered by visual transients, such as additional visual stimuli flashed near the object, apparent motion, or a brief removal of the object itself (blinking). The fading occurs even without prolonged adaptation and is time-locked to the presentation of the visual transients. Experiments show that the effect of a flashed object decreased monotonically as a function of the distance from the target object. Consistent with this result, when apparent motion, consisting of a sequence of flashes was presented between stationary disks, these target disks perceptually disappeared as if erased by the moving object. Blinking the target disk, instead of flashing an additional visual object, turned out to be sufficient to induce the fading. The effect of blinking peaked around a blink duration of 80 msec. Our findings reveal a unique mechanism that controls the visibility of visual objects in a spatially selective and time-locked manner in response to transient visual inputs. Possible mechanisms underlying this phenomenon will be discussed.
Abstract: Dynamic properties of horizontal vestibulo-ocular reflex (VOR) and optokinetic response (OKR) were studied in mice. The VOR was examined in the dark (VORD), in the light (VORL) and in the condition in which most of the visual field moves synchronously with the head motion (VORF). A mouse and/or a surrounding screen with vertical stripes was rotated sinusoidally, and the gain and phase of eye movements were measured in wide dynamic stimulation ranges. The working conditions of VOR and OKR were supplementary; OKR worked at low speeds of head turn and VOR at high speeds. Examination of VORL and VORF revealed non-linear interaction of VOR and OKR. The continuous sinusoidal head oscillation coupled with the in-phase or the out-of-phase oscillation of the surrounding screen, decreased or increased the VORD gain, and increased or decreased the VORD phase lead, respectively. Continuous oscillation of the surrounding screen increased the OKR gain and decreased the phase delay. These changes of VOR and OKR work to reduce the retinal slip. The present study provides fundamental information concerning the dynamic properties of VOR and OKR and the nature of their adaptive modifications in mice, which have been extensively used in genetic manipulation recently.
