Effects of fusion between tactile and proprioceptive inputs on tactile perception

Tactile perception is typically considered the result of cortical interpretation of afferent signals from a network of mechanical sensors underneath the skin. Yet, tactile illusion studies suggest that tactile perception can be elicited without afferent signals from mechanoceptors. Therefore, the extent that tactile perception arises from isomorphic mapping of tactile afferents onto the somatosensory cortex remains controversial. We tested whether isomorphic mapping of tactile afferent fibers onto the cortex leads directly to tactile perception by examining whether it is independent from proprioceptive input by evaluating the impact of different hand postures on the perception of a tactile illusion across fingertips. Using the Cutaneous Rabbit Effect, a well studied illusion evoking the perception that a stimulus occurs at a location where none has been delivered, we found that hand posture has a significant effect on the perception of the illusion across the fingertips. This finding emphasizes that tactile perception arises from integration of perceived mechanical and proprioceptive input and not purely from tactile interaction with the external environment.     

Grey matter volume in early human visual cortex predicts proneness to the sound-induced flash illusion

Visual perception can be modulated by sounds. A drastic example of this is the sound-induced flash illusion: when a single flash is accompanied by two bleeps, it is sometimes perceived in an illusory fashion as two consecutive flashes. However, there are strong individual differences in proneness to this illusion. Some participants experience the illusion on almost every trial, whereas others almost never do. We investigated whether such individual differences in proneness to the sound-induced flash illusion were reflected in structural differences in brain regions whose activity is modulated by the illusion. We found that individual differences in proneness to the illusion were strongly and significantly correlated with local grey matter volume in early retinotopic visual cortex. Participants with smaller early visual cortices were more prone to the illusion. We propose that strength of auditory influences on visual perception is determined by individual differences in recurrent connections, cross-modal attention and/or optimal weighting of sensory channels.     

Perception of Body Ownership Is Driven by Bayesian Sensory Inference

Recent studies have shown that human perception of body ownership is highly malleable. A well-known example is the rubber hand illusion (RHI) wherein ownership over a dummy hand is experienced, and is generally believed to require synchronized stroking of real and dummy hands. Our goal was to elucidate the computational principles governing this phenomenon. We adopted the Bayesian causal inference model of multisensory perception and applied it to visual, proprioceptive, and tactile stimuli. The model reproduced the RHI, predicted that it can occur without tactile stimulation, and that synchronous stroking would enhance it. Various measures of ownership across two experiments confirmed the predictions: a large percentage of individuals experienced the illusion in the absence of any tactile stimulation, and synchronous stroking strengthened the illusion. Altogether, these findings suggest that perception of body ownership is governed by Bayesian causal inference—i.e., the same rule that appears to govern the perception of outside world.     

Perception of visual image size by school students of different ages

In psychophysical experiments, 209 high school students with normal vision, 8 to 16 years old, were examined to study the perception of visual image sizes. Observers assessed the length of linear arrow-like figures (the Müller-Lyer illusion) or the length of single lines without arrow-like ends. Distortion of line size perception by 17–21% was obtained in the Müller-Lyer illusion independent of the age of observers. Distortion of size perception was absent in the case of estimation of the length of single line segments. The size differentiation thresholds gradually decreased in both cases with increasing age of observers and were correlated with the acuity of vision. For single lines, they were, on average, 1.3 times lower than the thresholds in the Müller-Lyer illusion. The probable mechanisms of the Müller-Lyer illusion are discussed. The experimental results demonstrate stability of illusion for observers aged 8–16 years, which may be connected with preferential contribution of the lower levels of the visual system to the appearance of the illusion.     

Illusion susceptibility in domestic dogs

Domestic dogs play many vital roles in human lives; however, relatively little is known about how they perceive the world visually. Given dogs’ recent popularity as a subject in cognitive and behavioural studies, it is important to understand how they visually interpret the world around them. One way to evaluate perception is to assess illusion susceptibility; specifically, how visual information is processed, interpreted and modified post-retinally. While illusion susceptibility has been used across a variety of species to comparatively assess the similarities and differences in visual processing and perception, this relatively novel methodological approach has only recently been adapted to evaluate perception in domestic dogs. Here, we present a comprehensive overview of the findings from studies that have evaluated domestic dogs’ illusion susceptibility, highlighting the relevance of these results for those studying illusion susceptibility in animals as well as canine behaviour and cognition. More specifically, the ultimate goal of this review is to answer the questions: (a) Are dogs susceptible to visual illusions? (b) If so, are they susceptible to illusions in a way that parallels humans and/or other animals? (c) Are findings, within dogs, consistent and if not, how might these be interpreted and explained?     

The dissociation between perception and action in the Ebbinghaus illusion: nonillusory effects of pictorial cues on grasp

According to a recently proposed distinction [1] between vision for perception and vision for action, visually guided movements should be largely immune to the perceptually compelling changes in size produced by pictorial illusions. Tests of this prediction that use the Ebbinghaus illusion have revealed only small effects of the illusion on grasp scaling as compared to its effect on perception [2-4]. Nevertheless, some have argued that the small effect on grasp implies that there is a single representation of size for both perception and action [5]. Recent findings, however, suggest that the 2-D pictorial elements, such as those comprising illusory backgrounds, can sometimes be treated as obstacles and thereby influence the programming of grasp [6]. The arrangement of the 2-D elements commonly used in previous studies examining the Ebbinghaus illusion could therefore give rise to an effect on grasp scaling that is independent of its effect on perceptual judgements, even though the two effects are in the same direction. We present evidence demonstrating that when the gap between the target and the illusion-making elements in the Ebbinghaus illusion is equidistant across different perceptual conditions (Figure 1a), the apparent effect of the illusion on grasp scaling is eliminated.     

Perception, action, and Roelofs effect: a mere illusion of dissociation

A prominent and influential hypothesis of vision suggests the existence of two separate visual systems within the brain, one creating our perception of the world and another guiding our actions within it. The induced Roelofs effect has been described as providing strong evidence for this perception/action dissociation: When a small visual target is surrounded by a large frame positioned so that the frame's center is offset from the observer's midline, the perceived location of the target is shifted in the direction opposite the frame's offset. In spite of this perceptual mislocalization, however, the observer can accurately guide movements to the target location. Thus, perception is prone to the illusion while actions seem immune. Here we demonstrate that the Roelofs illusion is caused by a frame-induced transient distortion of the observer's apparent midline. We further demonstrate that actions guided to targets within this same distorted egocentric reference frame are fully expected to be accurate, since the errors of target localization will exactly cancel the errors of motor guidance. These findings provide a mechanistic explanation for the various perceptual and motor effects of the induced Roelofs illusion without requiring the existence of separate neural systems for perception and action. Given this, the behavioral dissociation that accompanies the Roelofs effect cannot be considered evidence of a dissociation of perception and action. This indicates a general need to re-evaluate the broad class of evidence purported to support this hypothesized dissociation.     

The cutaneous rabbit illusion affects human primary sensory cortex somatotopically

We used functional magnetic resonance imaging (fMRI) to study neural correlates of a robust somatosensory illusion that can dissociate tactile perception from physical stimulation. Repeated rapid stimulation at the wrist, then near the elbow, can create the illusion of touches at intervening locations along the arm, as if a rabbit hopped along it. We examined brain activity in humans using fMRI, with improved spatial resolution, during this version of the classic cutaneous rabbit illusion. As compared with control stimulation at the same skin sites (but in a different order that did not induce the illusion), illusory sequences activated contralateral primary somatosensory cortex, at a somatotopic location corresponding to the filled-in illusory perception on the forearm. Moreover, the amplitude of this somatosensory activation was comparable to that for veridical stimulation including the intervening position on the arm. The illusion additionally activated areas of premotor and prefrontal cortex. These results provide direct evidence that illusory somatosensory percepts can affect primary somatosensory cortex in a manner that corresponds somatotopically to the illusory percept.     

Does My Step Look Big In This? A Visual Illusion Leads To Safer Stepping Behaviour

Background

Tripping is a common factor in falls and a typical safety strategy to avoid tripping on steps or stairs is to increase foot clearance over the step edge. In the present study we asked whether the perceived height of a step could be increased using a visual illusion and whether this would lead to the adoption of a safer stepping strategy, in terms of greater foot clearance over the step edge. The study also addressed the controversial question of whether motor actions are dissociated from visual perception.

Methodology/Principal Findings

21 young, healthy subjects perceived the step to be higher in a configuration of the horizontal-vertical illusion compared to a reverse configuration (p = 0.01). During a simple stepping task, maximum toe elevation changed by an amount corresponding to the size of the visual illusion (p<0.001). Linear regression analyses showed highly significant associations between perceived step height and maximum toe elevation for all conditions.

Conclusions/Significance

The perceived height of a step can be manipulated using a simple visual illusion, leading to the adoption of a safer stepping strategy in terms of greater foot clearance over a step edge. In addition, the strong link found between perception of a visual illusion and visuomotor action provides additional support to the view that the original, controversial proposal by Goodale and Milner (1992) of two separate and distinct visual streams for perception and visuomotor action should be re-evaluated.     

Fooling the Eyes: The Influence of a Sound-Induced Visual Motion Illusion on Eye Movements

The question of whether perceptual illusions influence eye movements is critical for the long-standing debate regarding the separation between action and perception. To test the role of auditory context on a visual illusion and on eye movements, we took advantage of the fact that the presence of an auditory cue can successfully modulate illusory motion perception of an otherwise static flickering object (sound-induced visual motion effect). We found that illusory motion perception modulated by an auditory context consistently affected saccadic eye movements. Specifically, the landing positions of saccades performed towards flickering static bars in the periphery were biased in the direction of illusory motion. Moreover, the magnitude of this bias was strongly correlated with the effect size of the perceptual illusion. These results show that both an audio-visual and a purely visual illusion can significantly affect visuo-motor behavior. Our findings are consistent with arguments for a tight link between perception and action in localization tasks.     

The Impact of Spatial Incongruence on an Auditory-Visual Illusion

Background

The sound-induced flash illusion is an auditory-visual illusion – when a single flash is presented along with two or more beeps, observers report seeing two or more flashes. Previous research has shown that the illusion gradually disappears as the temporal delay between auditory and visual stimuli increases, suggesting that the illusion is consistent with existing temporal rules of neural activation in the superior colliculus to multisensory stimuli. However little is known about the effect of spatial incongruence, and whether the illusion follows the corresponding spatial rule. If the illusion occurs less strongly when auditory and visual stimuli are separated, then integrative processes supporting the illusion must be strongly dependant on spatial congruence. In this case, the illusion would be consistent with both the spatial and temporal rules describing response properties of multisensory neurons in the superior colliculus.

Methodology/Principal Findings

The main aim of this study was to investigate the importance of spatial congruence in the flash-beep illusion. Selected combinations of one to four short flashes and zero to four short 3.5 KHz tones were presented. Observers were asked to count the number of flashes they saw. After replication of the basic illusion using centrally-presented stimuli, the auditory and visual components of the illusion stimuli were presented either both 10 degrees to the left or right of fixation (spatially congruent) or on opposite (spatially incongruent) sides, for a total separation of 20 degrees.

Conclusions/Significance

The sound-induced flash fission illusion was successfully replicated. However, when the sources of the auditory and visual stimuli were spatially separated, perception of the illusion was unaffected, suggesting that the “spatial rule” does not extend to describing behavioural responses in this illusion. We also find no evidence for an associated “fusion” illusion reportedly occurring when multiple flashes are accompanied by a single beep.     

Changing the Chevreul illusion by a background luminance ramp: lateral inhibition fails at its traditional stronghold--a psychophysical refutation

The Chevreul illusion is a well-known 19^th century brightness illusion, comprising adjacent homogeneous grey bands of different luminance, which are perceived as inhomogeneous. It is generally explained by lateral inhibition, according to which brighter areas projected to the retina inhibit the sensitivity of neighbouring retinal areas. Lateral inhibition has been considered the foundation-stone of early vision for a century, upon which several computational models of brightness perception are built. One of the last strongholds of lateral inhibition is the Chevreul illusion, which is often illustrated even in current textbooks. Here we prove that lateral inhibition is insufficient to explain the Chevreul illusion. For this aim, we placed the Chevreul staircase in a luminance ramp background, which noticeably changed the illusion. In our psychophysical experiments, all 23 observers reported a strong illusion, when the direction of the ramp was identical to that of the staircase, and all reported homogeneous steps (no illusion) when its direction was the opposite. When the background of the staircase was uniform, 14 saw the illusion, and 9 saw no illusion. To see whether the change of the entire background area or that of the staircase boundary edges were more important, we placed another ramp around the staircase, whose direction was opposite to that of the original, larger ramp. The result is that though the inner ramp is rather narrow (mean = 0.51 deg, SD = 0.48 deg, N = 23), it still dominates perception. Since all conditions of the lateral inhibition account were untouched within the staircase, lateral inhibition fails to model these perceptual changes. Area ratios seem insignificant; the role of boundary edges seems crucial. We suggest that long range interactions between boundary edges and areas enclosed by them, such that diffusion-based models describe, provide a much more plausible account for these brightness phenomena, and local models are insufficient.     

Mass is all that matters in the size-weight illusion

An object in outer space is weightless due to the absence of gravity, but astronauts can still judge whether one object is heavier than another one by accelerating the object. How heavy an object feels depends on the exploration mode: an object is perceived as heavier when holding it against the pull of gravity than when accelerating it. At the same time, perceiving an object’s size influences the percept: small objects feel heavier than large objects with the same mass (size–weight illusion). Does this effect depend on perception of the pull of gravity? To answer this question, objects were suspended from a long wire and participants were asked to push an object and rate its heaviness. This way the contribution of gravitational forces on the percept was minimised. Our results show that weight is not at all necessary for the illusion because the size–weight illusion occurred without perception of weight. The magnitude of the illusion was independent of whether inertial or gravitational forces were perceived. We conclude that the size–weight illusion does not depend on prior knowledge about weights of object, but instead on a more general knowledge about the mass of objects, independent of the contribution of gravity. Consequently, the size–weight illusion will have the same magnitude on Earth as it should have on the Moon or even under conditions of weightlessness.     

A laminar cortical model of stereopsis and 3D surface perception: closure and da Vinci stereopsis

A laminar cortical model of stereopsis and 3D surface perception is developed and simulated. The model describes how monocular and binocular oriented filtering interact with later stages of 3D boundary formation and surface filling-in in the LGN and cortical areas V1, V2, and V4. It proposes how interactions between layers 4, 3B, and 2/3 in V1 and V2 contribute to stereopsis, and how binocular and monocular information combine to form 3D boundary and surface representations. The model includes two main new developments: (1) It clarifies how surface-to-boundary feedback from V2 thin stripes to pale stripes helps to explain data about stereopsis. This feedback has previously been used to explain data about 3D figure-ground perception. (2) It proposes that the binocular false match problem is subsumed under the Gestalt grouping problem. In particular, the disparity filter, which helps to solve the correspondence problem by eliminating false matches, is realized using inhibitory interneurons as part of the perceptual grouping process by horizontal connections in layer 2/3 of cortical area V2. The enhanced model explains all the psychophysical data previously simulated by Grossberg and Howe (2003), such as contrast variations of dichoptic masking and the correspondence problem, the effect of interocular contrast differences on stereoacuity, Panum's limiting case, the Venetian blind illusion, stereopsis with polarity-reversed stereograms, and da Vinci stereopsis. It also explains psychophysical data about perceptual closure and variations of da Vinci stereopsis that previous models cannot yet explain.     

Features of perception of length of segments under conditions of ponzo and Müller-Lyer illusions in schizophrenia

In order to better appreciate the neurophysiologic mechanisms of perception of length under conditions of geometrical visual illusions, we studied sensitivity of mentally healthy subjects and schizophrenic patients to Ponzo and Müller-Lyer illusion. Patients with schizophrenia estimated length of segments of Müller-Lyer figure less precisely. Accuracy of perception of length of segments in Ponzo figure was ambiguously connected with the duration of the disease. Persons suffering from schizophrenia for a short time were less inclined to Ponzo illusion than mentally healthy subjects. On the contrary, patients who suffered from schizophrenia for a long time were more sensitive to this illusion. Ponzo illusion can be used as a marker of schizophrenia which is found out during the specific period of development of the disease. High sensitivity of patients with schizophrenia to Müller-Lyer and Ponzo illusions supports a hypothesis about the role of the global analysis of an image during processing of its low-frequency component in formation of the illusions under study.     

The effects of linear perspective on relative size discrimination in chimpanzees (Pan troglodytes) and humans (Homo sapiens)

In this study, we tested the corridor illusion in three chimpanzees and five humans, applying a relative size discrimination task to assess pictorial depth perception using linear perspective. The subjects were required to choose the physically larger cylinder of two on a background containing drawn linear perspective cues. We manipulated both background and cylinder size in each trial. Our findings suggest that chimpanzees, like humans, exhibit the corridor illusion.     

Subjective Size Perception Depends on Central Visual Cortical Magnification in Human V1

In the Ebbinghaus illusion, the context surrounding an object modulates its subjectively perceived size. Previous work implicates human primary visual cortex (V1) as the neural substrate mediating this contextual effect. Here we studied in healthy adult humans how two different types of context (large or small inducers) in this illusion affected size perception by comparing each to a reference stimulus without any context. We found that individual differences in the magnitudes of the illusion produced by either type of context were correlated with V1 area defined through retinotopic mapping using functional MRI. However, participants'' objective ability to discriminate the size of objects presented in isolation was unrelated to illusion strength and did not correlate with V1 area. Control analyses showed no correlations between behavioral measures and the overall V1 area estimated probabilistically on the basis of neuroanatomy alone. Therefore, subjective size perception correlated with variability in central cortical magnification rather than the anatomical extent of primary visual cortex. We propose that such changes in subjective perception of size are mediated by mechanisms that scale with the extent to which an individual''s V1 selectively represents the central visual field.     

A motion illusion reveals mechanisms of perceptual stabilization

Visual illusions are valuable tools for the scientific examination of the mechanisms underlying perception. In the peripheral drift illusion special drift patterns appear to move although they are static. During fixation small involuntary eye movements generate retinal image slips which need to be suppressed for stable perception. Here we show that the peripheral drift illusion reveals the mechanisms of perceptual stabilization associated with these micromovements. In a series of experiments we found that illusory motion was only observed in the peripheral visual field. The strength of illusory motion varied with the degree of micromovements. However, drift patterns presented in the central (but not the peripheral) visual field modulated the strength of illusory peripheral motion. Moreover, although central drift patterns were not perceived as moving, they elicited illusory motion of neutral peripheral patterns. Central drift patterns modulated illusory peripheral motion even when micromovements remained constant. Interestingly, perceptual stabilization was only affected by static drift patterns, but not by real motion signals. Our findings suggest that perceptual instabilities caused by fixational eye movements are corrected by a mechanism that relies on visual rather than extraretinal (proprioceptive or motor) signals, and that drift patterns systematically bias this compensatory mechanism. These mechanisms may be revealed by utilizing static visual patterns that give rise to the peripheral drift illusion, but remain undetected with other patterns. Accordingly, the peripheral drift illusion is of unique value for examining processes of perceptual stabilization.     

What the 'Moonwalk' illusion reveals about the perception of relative depth from motion

When one visual object moves behind another, the object farther from the viewer is progressively occluded and/or disoccluded by the nearer object. For nearly half a century, this dynamic occlusion cue has been thought to be sufficient by itself for determining the relative depth of the two objects. This view is consistent with the self-evident geometric fact that the surface undergoing dynamic occlusion is always farther from the viewer than the occluding surface. Here we use a contextual manipulation ofa previously known motion illusion, which we refer to as the'Moonwalk' illusion, to demonstrate that the visual system cannot determine relative depth from dynamic occlusion alone. Indeed, in the Moonwalk illusion, human observers perceive a relative depth contrary to the dynamic occlusion cue. However, the perception of the expected relative depth is restored by contextual manipulations unrelated to dynamic occlusion. On the other hand, we show that an Ideal Observer can determine using dynamic occlusion alone in the same Moonwalk stimuli, indicating that the dynamic occlusion cue is, in principle, sufficient for determining relative depth. Our results indicate that in order to correctly perceive relative depth from dynamic occlusion, the human brain, unlike the Ideal Observer, needs additional segmentation information that delineate the occluder from the occluded object. Thus, neural mechanisms of object segmentation must, in addition to motion mechanisms that extract information about relative depth, play a crucial role in the perception of relative depth from motion.     

The role of vergence in the perception of distance: a fair test of Bishop Berkeley's claim

Binocular eye movements were measured while subjects perceived the wallpaper illusion in order to test the claim made by Bishop Berkeley in 1709 that we perceive the distance of nearby objects by evaluating the vergence angles of our eyes. Four subjects looked through a nearby fronto-parallel array of vertical rods (28-35 cm away) as they binocularly fixated a point about 1 meter away. The wallpaper illusion was perceived under these conditions, i.e. the rods appeared farther away than their physical location. We found that although binocular fixation at an appropriate distance was needed to begin perceiving the wallpaper illusion (at least for naive observers), once established, the illusion was quite robust in the sense that it was not affected by changing vergence. No connection between the apparent localization of the rods and vergence was observed. We conclude that it is unlikely that vergence, itself, is responsible for the perceived distance shift in the wallpaper illusion, making it unlikely that vergence contributes to the perception of distance as Bishop Berkeley suggested. We found this to be true even when vergence angles were relatively large (more than 2 deg), the region in which the control of vergence eye movements has been shown to be both fast and effective.