Involvement of the Extrageniculate System in the Perception of Optical Illusions: A Functional Magnetic Resonance Imaging Study

Research on the neural processing of optical illusions can provide clues for understanding the neural mechanisms underlying visual perception. Previous studies have shown that some visual areas contribute to the perception of optical illusions such as the Kanizsa triangle and Müller-Lyer figure; however, the neural mechanisms underlying the processing of these and other optical illusions have not been clearly identified. Using functional magnetic resonance imaging (fMRI), we determined which brain regions are active during the perception of optical illusions. For our study, we enrolled 18 participants. The illusory optical stimuli consisted of many kana letters, which are Japanese phonograms. During the shape task, participants stated aloud whether they perceived the shapes of two optical illusions as being the same or not. During the word task, participants read aloud the kana letters in the stimuli. A direct comparison between the shape and word tasks showed activation of the right inferior frontal gyrus, left medial frontal gyrus, and right pulvinar. It is well known that there are two visual pathways, the geniculate and extrageniculate systems, which belong to the higher-level and primary visual systems, respectively. The pulvinar belongs to the latter system, and the findings of the present study suggest that the extrageniculate system is involved in the cognitive processing of optical illusions.     

A new method to elicit and measure movement illusions in stroke by means of muscle tendon vibration: the Standardized Kinesthetic Illusion Procedure (SKIP)

Abstract

Purpose: Muscle tendon vibration (MTV) strongly activates muscle spindles and can evoke kinaesthetic illusions. Although potentially relevant for sensorimotor rehabilitation in stroke, MTV is scarcely used in clinical practice, likely because of the absence of standardised procedures to elicit and characterise movement illusions. This work developed and validated a Standardised Kinaesthetic Illusion Procedure (SKIP) to favour the use of MTV-induced illusions in clinical settings.

Materials and methods: SKIP scores were obtained in 15 individuals with chronic stroke and 18 age- and gender-matched healthy counterparts. A further 13 healthy subjects were tested to provide more data with the general population. MTV was applied over the Achilles tendon and SKIP scoring system characterised the clearness and direction of the illusions of ankle dorsiflexion movements.

Results: All healthy and stroke participants perceived movement illusions. SKIP scores on the paretic side were significantly lower compared to the non paretic and healthy. Illusions were less clear and sometimes in unexpected directions with the impaired ankle, but still possible to elicit in the presence of sensorimotor deficits.

Conclusions: SKIP represents an ancillary and potentially useful clinical method to elicit and characterise illusions of movements induced by MTV. SKIP could be relevant to further assess the processing of proprioceptive afferents in stroke and their potential impact on motor control and recovery. It may be used to guide therapy and improve sensorimotor recovery. Future work is needed to investigate the metrological properties of our method (reliability, responsiveness, etc.), and also the neurophysiological underpinnings of MTV-induced illusions.     

A robust optical respiratory trigger for small rodents in clinical whole-body MR systems.

An increasing number of animal experiments are currently conducted on clinical MR systems. Motion artefacts due to breathing can become quite apparent, in particular with abdominal examinations. These artefacts can be reduced by using a triggered acquisition. However, the built-in detectors in human whole-body scanners are usually not sensitive enough to detect the tiny movements of small rodents. Therefore, a sensitive optical motion detector was developed together with a simple, robust analogue circuit. This circuit converts the original optical signal into an electrical one, compensates slow drifts and offsets, and finally generates a transistor-transistor logic trigger signal as input for the clinical whole-body magnetic resonance scanner. The trigger was successfully applied in mouse experiments.     

La grotte de Niaux,théâtre des illusions

Niaux Cavern, one of the great decorated caves, has been continuously open to the public. Despite its apparent accessibility, there is much that remains beyond our comprehension. One of the most inaccessible images is in the much visited Salon Noir. It is panel number four which contains a powerful but puzzling figure sometimes referred to as the Crossed Bison. This paper is an attempt to reach into the mind of the individual who produced that image to show how that artist's evident understanding of optical illusions, as well as an intuitive understanding of the theory of mind, propelled its creation for a specific graphic purpose.     

Looking at Op Art from a computational viewpoint

Arts history tells an exciting story about repeated attempts to represent features that are crucial for the understanding of our environment and which, at the same time, go beyond the inherently two-dimensional nature of a flat painting surface: depth and motion. In the twentieth century, Op artists such as Bridget Riley began to experiment with simple black and white patterns that do not represent motion in an artistic way but actually create vivid dynamic illusions in static pictures. The cause of motion illusions in such paintings is still a matter of debate. The role of involuntary eye movements in this phenomenon is studied here with a computational approach. The possible consequences of shifting the retinal image of synthetic wave gratings, dubbed as 'riloids', were analysed by a two-dimensional array of motion detectors (2DMD model), which generates response maps representing the spatial distribution of motion signals generated by such a stimulus. For a two-frame sequence reflecting a saccadic displacement, these motion signal maps contain extended patches in which local directions change only little. These directions, however, do not usually precisely correspond to the direction of pattern displacement that can be expected from the geometry of the curved gratings as an instance of the so-called 'aperture problem'. The patchy structure of the simulated motion detector response to the displacement of riloids resembles the motion illusion, which is not perceived as a coherent shift of the whole pattern but as a wobbling and jazzing of ill-defined regions. Although other explanations are not excluded, this might support the view that the puzzle of Op Art motion illusions could potentially have an almost trivial solution in terms of small involuntary eye movement leading to image shifts that are picked up by well-known motion detectors in the early visual system. This view can have further consequences for our understanding of how the human visual system usually compensates for eye movements, in order to let us perceive a stable world despite continuous image shifts generated by gaze instability.     

Adelson's tile and snake illusions: a Helmholtzian type of simultaneous lightness contrast

Adelson's tile, snake, and some other lightness illusions of the same type were measured with the Munsell neutral scale for twenty observers. It was shown that theories based on low-level luminance contrast processing could hardly explain these illusions. Neither can those based on luminance X-junctions. On the other hand, Helmholtz's idea, that simultaneous lightness contrast originates from an error in judgement of apparent illumination, has been elaborated so as to account for the tile and snake illusions as well as other demonstrations presented in this report.     

Apparent Time Interval of Visual Stimuli Is Compressed during Fast Hand Movement

The influence of body movements on visual time perception is receiving increased attention. Past studies showed apparent expansion of visual time before and after the execution of hand movements and apparent compression of visual time during the execution of eye movements. Here we examined whether the estimation of sub-second time intervals between visual events is expanded, compressed, or unaffected during the execution of hand movements. The results show that hand movements, at least the fast ones, reduced the apparent time interval between visual events. A control experiment indicated that the apparent time compression was not produced by the participants’ involuntary eye movements during the hand movements. These results, together with earlier findings, suggest hand movement can change apparent visual time either in a compressive way or in an expansive way, depending on the relative timing between the hand movement and visual stimulus.     

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 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.     

On the contours of apparent motion: A new perspective on visual space-time

Previous results on the perception of motion indicate that perceived motion paths cannot be explained solely in terms of simple feature-specific analyzers. This is particularly true of apparent (phi) motion. In this paper we develop a dynamic network, with simple filtering and summation properties, which can predict the geometric paths of apparent motion in various spatio-temporal configurations. The network assumptions predict a non-Euclidean metric for the visual space-time of motion perception and we consider the implications of such distortions for various visual displays, including illusions.     

Visual illusions induced by electrical stimulation of the labyrinth

In normal subjects, electrical stimulation of the labyrinth with surface electrodes located on the mastoid process induced illusions of shifting of a fixed point of light in darkness similar to the oculogyral illusion induced by rotatory vestibular stimulation. Monoaural anodal stimulation of the right labyrinth induced apparent shift of the target to the left; with cathodal stimulation, it shifted to the right; threshold current was 0.35–0.6 mA. When the current strength increased, the amplitude and rate of apparent movement of the target increased approximately linearly. With binaural, bipolar stimulation, the illusory movement of the target was toward the site of the cathode and the threshold decreased by 1.5–2.5 times. With binaural, monopolar stimulation, the target seemed to shift along the vertical and the threshold current was 1.4–3.0 mA. Eye movement appeared at substantially higher currents than those resulting in apparent movement of the target. It is suggested that visual illusions are linked not to vestibular eye-movement reactions, but to the effect vestibular signals have on the spatial perception system.Institute of Problems of Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 23, No. 3, pp. 321–327, May–June, 1991.     

Families of stationary patterns producing illusory movement: insights into the visual system.

A computational explanation of the illusory movement experienced upon extended viewing of Enigma, a static figure painted by Leviant, is presented. The explanation relies on a model for the interpretation of three-dimensional motion information contained in retinal motion measurements. This model shows that the Enigma figure is a special case of a larger class of figures exhibiting the same illusory movement and these figures are introduced here. Our explanation suggests that eye movements and/or accommodation changes cause weak retinal motion signals, which are interpreted by higher-level processes in a way that gives rise to these illusions, and proposes a number of new experiments to unravel the functional structure of the motion pathway.     

What are lightness illusions and why do we see them?

Lightness illusions are fundamental to human perception, and yet why we see them is still the focus of much research. Here we address the question by modelling not human physiology or perception directly as is typically the case but our natural visual world and the need for robust behaviour. Artificial neural networks were trained to predict the reflectance of surfaces in a synthetic ecology consisting of 3-D “dead-leaves” scenes under non-uniform illumination. The networks learned to solve this task accurately and robustly given only ambiguous sense data. In addition—and as a direct consequence of their experience—the networks also made systematic “errors” in their behaviour commensurate with human illusions, which includes brightness contrast and assimilation—although assimilation (specifically White's illusion) only emerged when the virtual ecology included 3-D, as opposed to 2-D scenes. Subtle variations in these illusions, also found in human perception, were observed, such as the asymmetry of brightness contrast. These data suggest that “illusions” arise in humans because (i) natural stimuli are ambiguous, and (ii) this ambiguity is resolved empirically by encoding the statistical relationship between images and scenes in past visual experience. Since resolving stimulus ambiguity is a challenge faced by all visual systems, a corollary of these findings is that human illusions must be experienced by all visual animals regardless of their particular neural machinery. The data also provide a more formal definition of illusion: the condition in which the true source of a stimulus differs from what is its most likely (and thus perceived) source. As such, illusions are not fundamentally different from non-illusory percepts, all being direct manifestations of the statistical relationship between images and scenes.     

Influence of fingertip contact on illusory arm movements.

Recent evidence suggests that reaching movements are more accurate when end point contact occurs, suggesting that fingertip contact contributes to a final estimation of arm position. In the present study we tested two hypotheses: 1). that fingertip contact influences illusions of arm movement produced by muscle vibration and 2). that this influence depends on the a priori context of the stability of the contact surface. Subjects sat with their elbows on a table and eyes closed. They demonstrated the perceived orientation of the left (cue) arm by mirroring the location with the right (report) arm. We manipulated deep proprioceptive cues by vibrating the left biceps brachia, causing illusions of elbow extension, and tested whether these illusions were altered when the fingertip remained in contact with a stable external surface. The context at this point represents a prior assumption that the external contact surface is stable. Midway through the experiment, the context was changed by challenging the prior assumption that the contact surface was stable by demonstrating that it could move. Unbeknownst to the subject, the external contact surface remained stable during data collection throughout the experiment. As expected, without tactile cues, biceps vibration caused illusory elbow extension. Conditions with fingertip contact and biceps vibration in the stable context demonstrated that contact largely eliminated the overestimation of cue arm elbow angle. However, in the context of a possibly unstable (movable) contact surface, the reports of elbow extension returned. Thus a priori notions about the stability context of an external contact surface influence how this tactile cue is integrated with proprioceptive sensory modalities to generate an estimate of arm location in space. These findings support the notion that tactile cues are used to calibrate proprioception against external spatial frameworks.     

Head-centred motion perception in the ageing visual system

Stationary objects appear to move in the opposite direction to a pursuit eye movement (Filehne illusion) and moving objects appear slower when pursued (Aubert-Fleischl phenomenon). Both illusions imply that extra-retinal, eye-velocity signals lead to lower estimates of speed than corresponding retinal motion signals. Intriguingly, the velocity (i.e. speed and direction) of the Filehne illusion depends on the age of the observer, especially for brief display durations (Wertheim and Bekkering, 1992). This suggests relative signal size changes as the visual system matures. To test the signal-size hypothesis, we compared the Filehne illusion and Aubert-Fleischl phenomenon in young and old observers using short and long display durations. The trends in the Filehne data were similar to those reported by Wertheim and Bekkering. However, we found no evidence for an effect of age or duration in the Aubert-Fleischl phenomenon. The differences between the two illusions could not be reconciled on the basis of actual eye movements made. The findings suggest a more complicated explanation of the combined influence of age and duration on head-centred motion perception than that described by the signal-size hypothesis.     

Model of a predatory stealth behaviour camouflaging motion

A computational model of a stealth strategy inspired by the apparent mating tactics of male hoverflies is presented. The stealth strategy (motion camouflage) paradoxically allows a predator to approach a moving prey in such a way that it appears to be a stationary object. In the model, the predators are controlled by neural sensorimotor systems that base their decisions on realistic levels of input information. They are shown to be able to employ motion camouflage to approach prey that move along both real hoverfly flight paths and artificially generated flight paths. The camouflaged approaches made demonstrate that the control systems have an ability to predict future prey movements. This is illustrated using two- and three-dimensional simulations.     

Non-linear rate-equilibrium free energy relationships and Hammond behavior in protein folding

Non-linear rate-equilibrium relationships upon mutation or changes in solvent conditions are frequently observed in protein folding reactions and are usually interpreted in terms of Hammond behavior. Here we first give a general overview over the concept of transition state movements in chemical reactions and discuss its application to protein folding. We then show examples for genuine Hammond behavior and for apparent transition state movements caused by other effects like changes in the rate-limiting step of the folding reaction or ground state effects, i.e. structural changes in either the native state or the unfolded state. These examples show that apparent transition state movements can easily be mistaken for Hammond behavior. We describe experimental tests using self- and cross-interaction parameters to distinguish between structural changes in a single transition state following Hammond behavior and apparent transition state movements caused by other effects.     

Geometrical illusions: study and modelling

The phenomena of geometrical illusions of extent suggest that the metric of a perceived field is different from the metric of a physical stimulus. The present study investigated the Müller-Lyer and Oppel-Kundt illusions as functions of spatial parameters of the figures, and constructed a neurophysiological model. The main idea of the modelling is based on the uncertainty principle, according to which distortions of size relations of certain parts of the stimulus, so-called geometrical illusions, are determined by processes of spatial filtering in the visual system. Qualitative and quantitative agreement was obtained between psychophysical measurement of the strength value of the illusions and the predictions of our model. Received: 18 June 1996 / Accepted in revised form: 24 June 1997     

A possible explanation of the low-level brightness–contrast illusions in the light of an extended classical receptive field model of retinal ganglion cells

The low-level brightness–contrast illusions constitute a special class within visual illusions. Speculations exist that these illusions may be processed through the filtering action of the retinal ganglion cells without necessitating much intervention from higher order processes of visual perception. Concept of the classical receptive field of the ganglion cell, derived from early physiological studies, prompted the idea that a Difference of Gaussian (DoG) model might explain the low-level illusions. In spite of its many successes, the DoG model fails to explain some of these illusions. It has been shown in this paper that it is possible to simulate those illusions with a model that takes into cognizance the role of the extended classical receptive field     

Propriomuscular coding of kinaesthetic sensation

The role of propriomuscular information in kinaesthetic sensation was studied. Experiments were carried out on human subjects in whom kinaesthetic illusions were induced by applying tendon vibration with a variable frequency. Six patterns of frequency modulation were used, four of which had an arbitrary form and the other two mimicked natural Ia discharges. The results show that the shape of the illusory movements recorded depended on the type of vibratory pattern used. A mathematical model for the propriomuscular information decoding process is proposed. It takes into account both the agonist and antagonist muscle spindle populations as sources of kinaesthetic information and is based on the assumption that position and velocity information are additively combined. The experimental data show a good fit with the theoretical data obtained by means of model simulation, thus validating our initial hypothesis. Various aspects of the experimental results and the hypotheses involved in the model are discussed.