Moving from spatially segregated to transparent motion: A modelling approach

Motion transparency, in which patterns of moving elements group together to give the impression of lacy overlapping surfaces, provides an important challenge to models of motion perception. It has been suggested that we perceive transparent motion when the shape of the velocity histogram of the stimulus is bimodal. To investigate this further, random-dot kinematogram motion sequences were created to simulate segregated (perceptually spatially separated) and transparent (perceptually overlapping) motion. The motion sequences were analysed using the multi-channel gradient model (McGM) to obtain the speed and direction at every pixel of each frame of the motion sequences. The velocity histograms obtained were found to be quantitatively similar and all were bimodal. However, the spatial and temporal properties of the velocity field differed between segregated and transparent stimuli. Transparent stimuli produced patches of rightward and leftward motion that varied in location over time. This demonstrates that we can successfully differentiate between these two types of motion on the basis of the time varying local velocity field. However, the percept of motion transparency cannot be based simply on the presence of a bimodal velocity histogram.     

Integration of Motion Responses Underlying Directional Motion Anisotropy in Human Early Visual Cortical Areas

Recent imaging studies have reported directional motion biases in human visual cortex when perceiving moving random dot patterns. It has been hypothesized that these biases occur as a result of the integration of motion detector activation along the path of motion in visual cortex. In this study we investigate the nature of such motion integration with functional MRI (fMRI) using different motion stimuli. Three types of moving random dot stimuli were presented, showing either coherent motion, motion with spatial decorrelations or motion with temporal decorrelations. The results from the coherent motion stimulus reproduced the centripetal and centrifugal directional motion biases in V1, V2 and V3 as previously reported. The temporally decorrelated motion stimulus resulted in both centripetal and centrifugal biases similar to coherent motion. In contrast, the spatially decorrelated motion stimulus resulted in small directional motion biases that were only present in parts of visual cortex coding for higher eccentricities of the visual field. In combination with previous results, these findings indicate that biased motion responses in early visual cortical areas most likely depend on the spatial integration of a simultaneously activated motion detector chain.     

Acoustic facilitation of object movement detection during self-motion

In humans, as well as most animal species, perception of object motion is critical to successful interaction with the surrounding environment. Yet, as the observer also moves, the retinal projections of the various motion components add to each other and extracting accurate object motion becomes computationally challenging. Recent psychophysical studies have demonstrated that observers use a flow-parsing mechanism to estimate and subtract self-motion from the optic flow field. We investigated whether concurrent acoustic cues for motion can facilitate visual flow parsing, thereby enhancing the detection of moving objects during simulated self-motion. Participants identified an object (the target) that moved either forward or backward within a visual scene containing nine identical textured objects simulating forward observer translation. We found that spatially co-localized, directionally congruent, moving auditory stimuli enhanced object motion detection. Interestingly, subjects who performed poorly on the visual-only task benefited more from the addition of moving auditory stimuli. When auditory stimuli were not co-localized to the visual target, improvements in detection rates were weak. Taken together, these results suggest that parsing object motion from self-motion-induced optic flow can operate on multisensory object representations.     

Multisensory integration in the superior colliculus: a neural network model

Neurons in the superior colliculus (SC) are known to integrate stimuli of different modalities (e.g., visual and auditory) following specific properties. In this work, we present a mathematical model of the integrative response of SC neurons, in order to suggest a possible physiological mechanism underlying multisensory integration in SC. The model includes three distinct neural areas: two unimodal areas (auditory and visual) are devoted to a topological representation of external stimuli, and communicate via synaptic connections with a third downstream area (in the SC) responsible for multisensory integration. The present simulations show that the model, with a single set of parameters, can mimic various responses to different combinations of external stimuli including the inverse effectiveness, both in terms of multisensory enhancement and contrast, the existence of within- and cross-modality suppression between spatially disparate stimuli, a reduction of network settling time in response to cross-modal stimuli compared with individual stimuli. The model suggests that non-linearities in neural responses and synaptic (excitatory and inhibitory) connections can explain several aspects of multisensory integration.     

A Bayesian model of stereopsis depth and motion direction discrimination

 The extraction of stereoscopic depth from retinal disparity, and motion direction from two-frame kinematograms, requires the solution of a correspondence problem. In previous psychophysical work [Read and Eagle (2000) Vision Res 40: 3345–3358], we compared the performance of the human stereopsis and motion systems with correlated and anti-correlated stimuli. We found that, although the two systems performed similarly for narrow-band stimuli, broad-band anti-correlated kinematograms produced a strong perception of reversed motion, whereas the stereograms appeared merely rivalrous. I now model these psychophysical data with a computational model of the correspondence problem based on the known properties of visual cortical cells. Noisy retinal images are filtered through a set of Fourier channels tuned to different spatial frequencies and orientations. Within each channel, a Bayesian analysis incorporating a prior preference for small disparities is used to assess the probability of each possible match. Finally, information from the different channels is combined to arrive at a judgement of stimulus disparity. Each model system – stereopsis and motion – has two free parameters: the amount of noise they are subject to, and the strength of their preference for small disparities. By adjusting these parameters independently for each system, qualitative matches are produced to psychophysical data, for both correlated and anti-correlated stimuli, across a range of spatial frequency and orientation bandwidths. The motion model is found to require much higher noise levels and a weaker preference for small disparities. This makes the motion model more tolerant of poor-quality reverse-direction false matches encountered with anti-correlated stimuli, matching the strong perception of reversed motion that humans experience with these stimuli. In contrast, the lower noise level and tighter prior preference used with the stereopsis model means that it performs close to chance with anti-correlated stimuli, in accordance with human psychophysics. Thus, the key features of the experimental data can be reproduced assuming that the motion system experiences more effective noise than the stereoscopy system and imposes a less stringent preference for small disparities. Received: 2 March 2001 / Accepted in revised form: 5 July 2001     

Characterization of the percepts evoked by discontinuous motion over the perioral skin.

The capacity of human subjects to process information about discontinuous and continuous movement was evaluated. Constant-velocity brushing stimuli were delivered through aperture plates that rested lightly upon the mandibular skin. Each plate consisted of either two spatially separated, slit-like openings or a single continuous, longer opening. It was discovered that percepts of smooth apparent motion were achieved with the split apertures (i.e., from discontinuous movement) for only limited ranges of stimulus velocity. Moreover, the optimal velocity supporting smooth apparent motion increased with the separation between the slit-like openings. In a second series of experiments, subjects' ability to discriminate opposing directions of discontinuous and continuous movement was evaluated. It was found that subjects could derive directional information from percepts elicited by discontinuous movement. However, the capacity to discriminate opposing directions of continuous movement cannot be explained solely in terms of the ability to process information about the change in position of a stimulus from its onset to its offset.     

Auditory aftereffects of approaching and withdrawing sound sources: Dependence on the trajectory and location of adapting stimuli

Perception of approaching and withdrawing sound sources and their action on auditory aftereffects were studied in the free field. Motion of adapting stimuli was mimicked in two ways: (1) simultaneous opposite changes of amplitude of broadband noise impulses at two loudspeakers placed at 1.1 and 4.5 m from the listener; (2) an increase or a decrease of amplitude of broadband noise impulses in only one loudspeaker, the nearer or the remote one. Motion of test stimuli was mimicked in the former way. Listeners determined direction of the test stimuli motion without any adaptation (control) or after adaptation to stationary, slowly moving (with an amplitude change of 2 dB) and rapidly moving (amplitude change of 12 dB) stimuli. Percentages of “withdrawal” reports were used for construction of psychometric curves. Three phenomena of auditory perception were observed. In the absence of adaptation, a growing-louder effect was revealed, i.e., listeners reported more frequently the test sounds as the approaching ones. Once adapted to stationary or slowly moving stimuli, listeners showed a location-dependent aftereffect. Test stimuli were reported as withdrawing more often as compared with control. The effect was associated with the previous one and was weaker when the distance to the loudspeaker producing adapting stimuli was greater. After adaptation to rapidly moving stimuli, a motion aftereffect was revealed. In this case, listeners reported a direction of test stimuli motion as being opposite to that of adapting stimuli. The motion aftereffect was more pronounced when the adapting stimuli motion was mimicked in the former way, as this method allows estimation of their trajectory. There was no relationship between the motion aftereffect and the growing-louder effect, whichever way the adapting stimuli were produced. There was observed a tendency for reduction of aftereffects of approaching and for intensification of aftereffects of withdrawal with growing distance from source of adapting stimuli.     

Responses of neurons in the primary auditory cortex of the cat to the auditory motion stimuli with variable interaural delay

Unit responses in the primary auditory cortex of anesthetized cats to stationary and apparently moving stimuli resulted from a static and dynamically varying interaural delay (ITD) were recorded. The static stimuli consisted of binaurally presented tones and clicks. The dynamic stimuli were produced by in-phase and out-of-phase binaurally presented click trains with time-varying ITD. Sensitivity to ITDs was mostly seen in responses of the neurons with low characteristic frequency (below 2.8 kHz). All cells sampled with static stimuli responded to simulated motion. A motion effect could take the form of a difference in response magnitude depending on the direction of stimulus motion and a shift in the ITD-function opposite the direction of motion. The magnitude of motion effects was influenced by the position of motion trajectory relative to the ITD-function. The greatest motion effect was produced by motion crossing the ITD-function slopes.     

The Intrinsic Electrophysiological Characteristics of Fly Lobula Plate Tangential Cells: III. Visual Response Properties

In this last paper in a series (Borst and Haag, 1996; Haag et al., 1997) about the lobula plate tangential cells of the fly visual system (CH, HS, and VS cells), the visual response properties were examined using intracellular recordings and computer simulations. In response to visual motion stimuli, all cells responded mainly by a graded shift of their axonal membrane potential. While ipsilateral motion resulted in a graded membrane potential shift, contralateral motion led to distinct EPSPs. For HS cells, simultaneous extracellular recorded action potentials of a spiking interneuron, presumably the H2 cell, corresponded to the EPSPs in the HS cell in a one-to-one fashion. When HS cells were hyperpolarized during ipsilateral motion, they mainly produced action potentials, but when they were hyperpolarized during contralateral motion only a slight increase of EPSP amplitude, could be observed. Intracellular application of the sodium channel blocker QX 314 abolished action potentials of HS cells while having little effect on the graded membrane response to ipsilateral motion. HS and CH cells were also studied with respect to their spatial integration properties. For both cell types, their graded membrane response was found to increase less than linearly with the size of the ipsilateral motion pattern. However, while for HS cells various amounts of hyperpolarizing current injected during motion stimulation led to different saturation levels, this was not the case for CH cells. In response to a sinusoidal velocity modulation, CH cells followed pattern motion only up to 10 Hz modulation frequency, but HS cells still revealed significant membrane depolarizations up to about 40 Hz.In the computer simulations, the compartmental models of tangential cells, as derived in the previous papers, were linked to an array of local motion detectors. The model cells revealed the same basic response features as their natural counterparts. They showed a response saturation as a function of stimulus size. In CH-models, however, the saturation was less pronounced than in real CH-cells, indicating spatially nonuniform membrane resistances with higher values in the dendrite. As in the experiments, HS models responded to high-frequency velocity modulation with a higher amplitude than did CH models.     

Properties of individual movement detectors as derived from behavioural experiments on the visual system of the fly

The performance of the fly's movement detection system is analysed using the visually induced yaw torque generated during tethered flight as a behavioural indicator. In earlier studies usually large parts of the visual field were exposed to the movement stimuli; the fly's response, therefore, represented the spatially pooled output signals of a large number of local movement detectors. Here we examined the responses of individual movement detectors. The stimulus pattern was presented to the fly via small vertical slits, thus, nearly avoiding spatial integration of local movement information along the horizontal axis of the eye. The stimulus consisted of a vertically oriented sine-wave grating which was moved with a constant velocity either clockwise or counterclockwise. In agreement with the theory of movement detectors of the correlation type, the time-course of the detector signal is modulated with the spatial phase of the stimulus pattern. It can even assume negative values for some time during the response cycle and thus signal the wrong direction of motion. By spatially integrating the response over sufficiently large arrays of movement detectors these response modulations disappear. Finally, one obtains a signal of the movement detection system which is constant while the pattern moves in one direction and only changes its sign when the pattern reverses its direction of motion. Spatial integration thus represents a simple means to obtain a meaningful representations of motion information.     

Do variables that affect similar bistable apparent-movement displays result in similar changes in perception?

Two bistable apparent-movement displays (i.e. ones that generate two qualitatively different kinds of movement percepts under different conditions) were compared. They were designed to be as similar as possible spatially, and were studied with identical stimulus manipulations to see whether changes in balance between their bistable percepts would be similar. Results show that the two displays had different response characteristics to the same stimulus manipulations. Two models of motion perception that have previously predicted at least one kind of bistable apparent motion were considered in terms of how well they address the current data. As yet, neither model has been shown to predict the motion states and bistable behavior of the two displays studied here. It is concluded that results of the type described here (specifically, differences in the psychophysical functions yielded by two structurally similar but qualitatively different bistable displays) present a challenge for theories of motion perception.     

Visual detection of paradoxical motion in flies

Summary From psychophysics it is known that humans easily perceive motion in Fourier-stimuli in which dots are displaced coherently into one direction. Furthermore, motion can be extracted from Drift-balanced stimuli in which the dots on average have no distinct direction of motion, or even in paradox -motion stimuli where the dots are displaced opposite to the perceived direction of motion. Whereas Fourier-motion can be explained by very basic motion detectors and nonlinear preprocessing of the input can account for the detection of Drift-balanced motion, a hierarchical model with two layers of motion detectors was proposed to explain the perception of -motion. The well described visual system of the fly allows to investigate whether these complex motion stimuli can be detected in a comparatively simple brain.The detection of such motion stimuli was analyzed for various random-dot cinematograms with extracellular recordings from the motion-sensitive Hl-neuron in the third visual ganglion of the blowfly Calliphora erythrocephala. The results were compared to computer-simulations of a hierarchical model of motion detector networks.For Fourier- and Drift-balanced motion stimuli, the Hl-neuron responds directionally selective to the moving object, whereas for -motion stimuli, the preferred direction is given by the dot displacement. Assuming nonlinear preprocessing of the detector input, such as a half-wave rectification, elementary motion detectors of the correlation type can account for these results.Abbreviations EMD elementary motion detector     

Synaptic plasticity can produce and enhance direction selectivity

The discrimination of the direction of movement of sensory images is critical to the control of many animal behaviors. We propose a parsimonious model of motion processing that generates direction selective responses using short-term synaptic depression and can reproduce salient features of direction selectivity found in a population of neurons in the midbrain of the weakly electric fish Eigenmannia virescens. The model achieves direction selectivity with an elementary Reichardt motion detector: information from spatially separated receptive fields converges onto a neuron via dynamically different pathways. In the model, these differences arise from convergence of information through distinct synapses that either exhibit or do not exhibit short-term synaptic depression—short-term depression produces phase-advances relative to nondepressing synapses. Short-term depression is modeled using two state-variables, a fast process with a time constant on the order of tens to hundreds of milliseconds, and a slow process with a time constant on the order of seconds to tens of seconds. These processes correspond to naturally occurring time constants observed at synapses that exhibit short-term depression. Inclusion of the fast process is sufficient for the generation of temporal disparities that are necessary for direction selectivity in the elementary Reichardt circuit. The addition of the slow process can enhance direction selectivity over time for stimuli that are sustained for periods of seconds or more. Transient (i.e., short-duration) stimuli do not evoke the slow process and therefore do not elicit enhanced direction selectivity. The addition of a sustained global, synchronous oscillation in the gamma frequency range can, however, drive the slow process and enhance direction selectivity to transient stimuli. This enhancement effect does not, however, occur for all combinations of model parameters. The ratio of depressing and nondepressing synapses determines the effects of the addition of the global synchronous oscillation on direction selectivity. These ingredients, short-term depression, spatial convergence, and gamma-band oscillations, are ubiquitous in sensory systems and may be used in Reichardt-style circuits for the generation and enhancement of a variety of biologically relevant spatiotemporal computations.     

Dynamic characteristics of spatial mechanisms coding contour structures

Psychophysical thresholds for the detection of luminance targets improve significantly when the targets are presented in a specific context of spatially separated, collinear inducing stimuli defining visual contours. This phenomenon is generally referred to as a special case of detection facilitation called spatial facilitation. Spatial facilitation has been observed with luminance-defined. achromatic stimuli on achromatic backgrounds as well as with targets and inducers defined by colour contrast. This paper reviews psychophysical results from detection experiments with human observers showing the conditions under which spatially separated contour inducers facilitate the detection of simultaneously presented target stimuli. The findings point towards two types of spatial mechanisms: (i) Short-range mechanisms that are sensitive to narrowly spaced stimuli of small size and, at distinct target locations, selective to the contrast polarity of targets and inducers. (ii) Long-range mechanisms that are triggered by longer stimuli, generate facilitation across wider spatial gaps between targets and inducers, and are insensitive to their contrast polarity. Spatial facilitation with chromatic stimuli requires a longer inducer exposure than spatial facilitation with achromatic stimuli, which is already fully effective at inducer exposures of 30 ms. This difference in temporal dynamics indicates some functional segregation between mechanisms for colour and luminance contrast in spatial coding. In general, spatially induced detection facilitation can to a large extent be explained by mechanisms involving from-short-to-long-range interactions between cortical detectors.     

Traveling waves in a simple population model involving growth and death

The structure of solutions to a simple spatially dependent population model involving growth and death is investigated. Two forms of motility of the population are considered: (1) random motion only modeled by a Fickian law, and (2) a directed component of motion (chemotaxis), included in addition to the random motion. Under certain growth conditions a traveling wave of constant speed is approached. This speed can be increased by the addition of the chemotaxis with a corresponding increase in the asymptotic population. Development of initial conditions into a wave is illustrated numerically.     

In vivo noninvasive measurement of spatially resolved corneal elasticity in human eyes using Lamb wave optical coherence elastography

Current elastography techniques are limited in application to accurately assess spatially resolved corneal elasticity in vivo for human eyes. The air‐puff optical coherence elastography (OCE) with an eye motion artifacts correction algorithm is developed to distinguish the in vivo cornea vibration from the eye motion and visualize the Lamb wave propagation clearly in healthy subjects. Based on the Lamb wave model, the phase velocity dispersion curve in the high‐frequency is calculated to obtain spatially resolved corneal elasticity accurately with high repeatability. It is found that the corneal elasticity has regional variations and is correlated with intraocular pressure, which suggests that the method has the potential to provide noninvasive measurement of spatially resolved corneal elasticity in clinical practice.     

Sensitivity to colour- and to orientation-carried motion respectively improves and deteriorates under equiluminant background conditions.

This study presents two distinct effects produced by manipulation of the background illumination on the directional sensitivity to colour- and orientation-carried motion. The two motion percepts were produced with two of a class of stimuli extensively used by the first and last authors in apparent-motion studies. The stimuli were designed to produce motion perception by virtue of spatiotemporal matching of (a) colour with orientation systematically mismatched (Colour across Orientation, CxO) and of (b) orientation with colour systematically mismatched (OxC). An increase in background illumination from dark to the equiluminance point (relative to the luminance of the discrete stimulus microelements) entails a significant increase and decrease of directional performances with CxO and OxC stimuli, respectively. It is proposed that these anti-symmetrical background effects have distinct neurophysiological origins. For CxO stimuli, improvement of directional performances at the equiluminant point is presumably due to the inactivation of the inhibitory effect of the luminance-motion pathway on the chromatic-motion pathway. The opposite effect obtained with OxC stimuli, previously referred to as the veto effect (Gorea and Papathomas, 1988 Invest. Ophthal. Vis. Sci. Suppl., 29, 265), is supposed to be entailed by the inactivation of the luminance-oriented mechanism, the only motion sensitive mechanism activated by this stimulus configuration.     

A Common Cortical Circuit Mechanism for Perceptual Categorical Discrimination and Veridical Judgment

Perception involves two types of decisions about the sensory world: identification of stimulus features as analog quantities, or discrimination of the same stimulus features among a set of discrete alternatives. Veridical judgment and categorical discrimination have traditionally been conceptualized as two distinct computational problems. Here, we found that these two types of decision making can be subserved by a shared cortical circuit mechanism. We used a continuous recurrent network model to simulate two monkey experiments in which subjects were required to make either a two-alternative forced choice or a veridical judgment about the direction of random-dot motion. The model network is endowed with a continuum of bell-shaped population activity patterns, each representing a possible motion direction. Slow recurrent excitation underlies accumulation of sensory evidence, and its interplay with strong recurrent inhibition leads to decision behaviors. The model reproduced the monkey's performance as well as single-neuron activity in the categorical discrimination task. Furthermore, we examined how direction identification is determined by a combination of sensory stimulation and microstimulation. Using a population-vector measure, we found that direction judgments instantiate winner-take-all (with the population vector coinciding with either the coherent motion direction or the electrically elicited motion direction) when two stimuli are far apart, or vector averaging (with the population vector falling between the two directions) when two stimuli are close to each other. Interestingly, for a broad range of intermediate angular distances between the two stimuli, the network displays a mixed strategy in the sense that direction estimates are stochastically produced by winner-take-all on some trials and by vector averaging on the other trials, a model prediction that is experimentally testable. This work thus lends support to a common neurodynamic framework for both veridical judgment and categorical discrimination in perceptual decision making.     

Visual Benefits in Apparent Motion Displays: Automatically Driven Spatial and Temporal Anticipation Are Partially Dissociated

Many behaviourally relevant sensory events such as motion stimuli and speech have an intrinsic spatio-temporal structure. This will engage intentional and most likely unintentional (automatic) prediction mechanisms enhancing the perception of upcoming stimuli in the event stream. Here we sought to probe the anticipatory processes that are automatically driven by rhythmic input streams in terms of their spatial and temporal components. To this end, we employed an apparent visual motion paradigm testing the effects of pre-target motion on lateralized visual target discrimination. The motion stimuli either moved towards or away from peripheral target positions (valid vs. invalid spatial motion cueing) at a rhythmic or arrhythmic pace (valid vs. invalid temporal motion cueing). Crucially, we emphasized automatic motion-induced anticipatory processes by rendering the motion stimuli non-predictive of upcoming target position (by design) and task-irrelevant (by instruction), and by creating instead endogenous (orthogonal) expectations using symbolic cueing. Our data revealed that the apparent motion cues automatically engaged both spatial and temporal anticipatory processes, but that these processes were dissociated. We further found evidence for lateralisation of anticipatory temporal but not spatial processes. This indicates that distinct mechanisms may drive automatic spatial and temporal extrapolation of upcoming events from rhythmic event streams. This contrasts with previous findings that instead suggest an interaction between spatial and temporal attention processes when endogenously driven. Our results further highlight the need for isolating intentional from unintentional processes for better understanding the various anticipatory mechanisms engaged in processing behaviourally relevant stimuli with predictable spatio-temporal structure such as motion and speech.