The interpretation of a moving retinal image

It is shown that from a monocular view of a rigid, textured, curved surface it is possible, in principle, to determine the gradient of the surface at any point, and the motion of the eye relative to it, from the velocity field of the changing retinal image, and its first and second spatial derivatives. The relevant equations are redundant, thus providing a test of the rigidity assumption. They involve, among other observable quantities, the components of shear of the retinal velocity field, suggesting that the visual system may possess specialized channels for computing these components.     

Sensitivity of the Goldfish Motion Detection System Revealed by Incoherent Random Dot Stimuli: Comparison of Behavioural and Neuronal Data

Background

Global motion detection is one of the most important abilities in the animal kingdom to navigate through a 3-dimensional environment. In the visual system of teleost fish direction-selective neurons in the pretectal area (APT) are most important for global motion detection. As in all other vertebrates these neurons are involved in the control of slow phase eye movements during gaze stabilization. In contrast to mammals cortical pathways that might influence motion detection abilities of the optokinetic system are missing in teleost fish.

Results

To test global motion detection in goldfish we first measured the coherence threshold of random dot patterns to elicit horizontal slow phase eye movements. In addition, the coherence threshold of the optomotor response was determined by the same random dot patterns. In a second approach the coherence threshold to elicit a direction selective response in neurons of the APT was assessed from a neurometric function. Behavioural thresholds and neuronal thresholds to elicit slow phase eye movements were very similar, and ranged between 10% and 20% coherence. In contrast to these low thresholds for the optokinetic reaction and APT neurons the optomotor response could only be elicited by random dot patterns with coherences above 40%.

Conclusion

Our findings suggest a high sensitivity for global motion in the goldfish optokinetic system. Comparison of neuronal and behavioural thresholds implies a nearly one-to-one transformation of visual neuron performance to the visuo-motor output. In addition, we assume that the optomotor response is not mediated by the optokinetic system, but instead by other motion detection systems with higher coherence thresholds.     

The effect of target orientation on the visual acuity and the spatial frequency response of the locust eye

The eye of the locust was shown to distinguish between vertical and horizontal alignments of narrow slit targets, and responded more strongly with horizontal alignments. In all cases the response increased as slit width decreased, to a limit of about 0.1 degrees. Similarly the eye responded more strongly to gratings in horizontal alignment, with a peak in response close to a spatial frequency of 3 cyc/deg. These results are discussed and comparisons are made with previous findings on insect and other eyes.     

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.     

Landscape structure and locust swarming: a satellite's eye view

Desert locust Schistocerca gregaria outbreaks consistently start in the same places, suggesting that certain landscapes are particularly favourable for outbreaking. Outbreaks are generated by multiplication, concentration and gregarisation of locust populations. Previous research has shown how small-scale vegetation patterns in desert ecosystems influence locust gregarisation; the present study examines the effects of large-scale landscape structure on locust multiplication and concentration. NOAA/AVHRR satellite imagery was used to relate abundance and spatial distribution of resources at the landscape scale to the historical record of locust outbreaks. Threshold NDVI values were investigated to define what constitutes 'resource' for locusts. The first part of the study showed that abundance and spatial distribution of resource were not sufficient to distinguish between outbreak and non-outbreak areas in the western part of the locust distribution area. Thus, outbreak danger zones cannot be identified by landscape structure at this spatial resolution. The second analysis investigated spatio-temporal patterns of vegetation growth in two locust breeding areas with very different landscape structure; in both cases, the patterns differed significantly between outbreaking and non-outbreaking years. In Mauritania, a flat homogeneous desert landscape, both resource abundance and fragmentation were higher in outbreaking years. On the Red Sea coast, a fragmented landscape, resource spatial distribution was consistent between years, and abundance alone was a significant predictor of outbreaking. High resource abundance promotes locust multiplication, and contraction of resource into small patches increases locust concentration; these two mechanisms explain how landscape structure influences locust outbreaking.     

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.     

Motion detection in the presence and absence of background motion in anAnolis lizard

Anolis lizards respond to a moving object viewed in the periphery of their visual field by turning their eye to fixate the object with their central fovea. This paper describes the relative effectiveness of different patterns of motion of a small black lure in eliciting these eye movements and the way motion of a backdrop of vegetation affects the response. The stimulus was positioned 45 degrees from the animal's line of gaze and oscillated in the vertical axis at different frequencies between 0.5 and 10 Hz. At each frequency, the amplitude of the oscillation was increased until the lizard flicked its eye towards the stimulus. The minimum amplitude needed for response (0.22 degrees of visual angle) was independent of frequency and waveform. The probability of any response occurring was, however, lower at higher frequencies (7 and 10 Hz) and a 1.5 Hz square wave evoked the greatest proportion of responses. Sinusoidal oscillation of a background of vegetation at 1.6 Hz during or before motion of the stimulus lure reduced the probability of an eye flick but did not raise the minimum amplitude needed for a response. The suppressive effect was greatest when the lure was oscillated at frequencies close to that of the background. It is concluded that Anolis, which rely upon motion to detect objects in the periphery of the visual field, filter out irrelevant motion such as that of windblown vegetation by responding preferentially to particular patterns of motion and short term habituation to commonly present patterns of motion.     

Knowing with which eye we see: utrocular discrimination and eye-specific signals in human visual cortex

Neurophysiological and behavioral reports converge to suggest that monocular neurons in the primary visual cortex are biased toward low spatial frequencies, while binocular neurons favor high spatial frequencies. Here we tested this hypothesis with functional magnetic resonance imaging (fMRI). Human participants viewed flickering gratings at one of two spatial frequencies presented to either the left or the right eye, and judged which of the two eyes was being stimulated (utrocular discrimination). Using multivoxel pattern analysis we found that local spatial patterns of signals in primary visual cortex (V1) allowed successful decoding of the eye-of-origin. Decoding was above chance for low but not high spatial frequencies, confirming the presence of a bias reported by animal studies in human visual cortex. Behaviorally, we found that reliable judgment of the eye-of-origin did not depend on spatial frequency. We further analyzed the mean response in visual cortex to our stimuli and revealed a weak difference between left and right eye stimulation. Our results are thus consistent with the interpretation that participants use overall levels of neural activity in visual cortex, perhaps arising due to local luminance differences, to judge the eye-of-origin. Taken together, we show that it is possible to decode eye-specific voxel pattern information in visual cortex but, at least in healthy participants with normal binocular vision, these patterns are unrelated to awareness of which eye is being stimulated.     

Tonic effects of stationary luminous slits on the discharge rates of some locust visual interneurones

The presence of an illuminated slit in the visual field of a locust compound eye produced changes in the tonic discharge rate of the DCMD and three other visual interneurones, recorded in a connective. The DCMD discharge peaked initially in the range of low slit subtenses, but over a period of minutes of exposure its character changed so that there was a rise at high subtenses also. When the luminance of a slit of fixed subtense was increased in steps, there was an initial rise then a sharp fall in discharge, indicating an abrupt onset of inhibition. Lateral spread of inhibition could account for the peak in response to slits, at a subtense falling well within the acceptance angle of a single ommatidium. The results show the ability of some visual interneurones to maintain a changed level of discharge in the presence of a stationary object in the visual field of the eye.     

Effect of refractive error in visual tests with sinusoidal gratings

While sinusoidal grating stimuli are finding increasing application in clinical tests for visual disturbance, the possible effects on the test results of simple defocus of the retinal image, due to refractive or accommodative error, are not always acknowledged. Theoretical considerations and experimental measurements emphasize that the degradation in visual performance due to such defocus effects is minimal when the eye pupil is small and the spatial frequency of the grating test objects is low.     

Neurobiology of polarization vision in the locust Schistocerca gregaria

The polarization pattern of the blue sky serves as an important reference for spatial orientation in insects. To understand the neural mechanisms involved in sky compass orientation we have analyzed the polarization vision system in the locust Schistocerca gregaria. As in other insects, photoreceptors adapted for the detection of sky polarization are concentrated in a dorsal rim area (DRA) of the compound eye. Stationary flying locusts show polarotactic yaw-torque responses when illuminated through a rotating polarizer from above. This response is abolished after painting the DRAs. Central stages of the polarization vision system, revealed through tracing studies, include dorsal areas in the lamina and medulla, the anterior lobe of the lobula, the anterior optic tubercle, the lateral accessory lobe and the central complex. Physiological analysis of polarization-sensitive (POL) neurons has focussed on the optic tubercle and on the central complex. Each POL neuron was maximally excited at a certain e-vector (phimax) and was maximally inhibited at an e-vector perpendicular to phimax. The neurons had large visual fields, and many neurons received input from both eyes. The neuronal organization of the central complex suggests a role as a spatial compass within the locust brain.     

Some results on translation invariance in the human visual system

Four experiments were conducted to study the nature of visual translation invariance in humans. In all the experiments, subjects were trained to discriminate between a previously unknown target and two non-target distractors presented at a fixed retinal location to one side of the fixation point. In a subsequent test phase, this performance was compared with the performance when the patterns were presented either centrally at the fixation point or at a location on the other side of the fixation point, opposite to the location where the patterns were learned, but where acuity was identical to what it was at the learned location. Two different experimental paradigms were used. One used an eye movement control device (Experiment 1) to ensure the eye could not move relative to the patterns to be learned. In the other three experiments, presentation duration of the patterns was restricted to a short enough period to preclude eye movements. During the training period in Experiments 1 and 2, presentation location of the patterns was centered at 2.4 deg in the periphery, whereas in Experiments 3 and 4 presentation eccentricity was reduced to 0.86 and 0.49 deg. In all four experiments performance dropped when the pattern had to be recognized at new test positions. This result suggests that the visual system does not apply a global transposition transformation to the retinal image to compensate for translations. We propose that, instead, it decomposes the image into simple features which themselves are more-or-less translation invariant. If in a given task, patterns can be discriminated using these simple features, then translation invariance will occur. If not, then translation invariance will fail or be incomplete.     

Relating Lateralization of Eye Use to Body Motion in the Avoidance Behavior of the Chameleon (Chamaeleo chameleon)

Lateralization is mostly analyzed for single traits, but seldom for two or more traits while performing a given task (e.g. object manipulation). We examined lateralization in eye use and in body motion that co-occur during avoidance behaviour of the common chameleon, Chamaeleo chameleon. A chameleon facing a moving threat smoothly repositions its body on the side of its perch distal to the threat, to minimize its visual exposure. We previously demonstrated that during the response (i) eye use and body motion were, each, lateralized at the tested group level (N = 26), (ii) in body motion, we observed two similar-sized sub-groups, one exhibiting a greater reduction in body exposure to threat approaching from the left and one – to threat approaching from the right (left- and right-biased subgroups), (iii) the left-biased sub-group exhibited weak lateralization of body exposure under binocular threat viewing and none under monocular viewing while the right-biased sub-group exhibited strong lateralization under both monocular and binocular threat viewing. In avoidance, how is eye use related to body motion at the entire group and at the sub-group levels? We demonstrate that (i) in the left-biased sub-group, eye use is not lateralized, (ii) in the right-biased sub-group, eye use is lateralized under binocular, but not monocular viewing of the threat, (iii) the dominance of the right-biased sub-group determines the lateralization of the entire group tested. We conclude that in chameleons, patterns of lateralization of visual function and body motion are inter-related at a subtle level. Presently, the patterns cannot be compared with humans'' or related to the unique visual system of chameleons, with highly independent eye movements, complete optic nerve decussation and relatively few inter-hemispheric commissures. We present a model to explain the possible inter-hemispheric differences in dominance in chameleons'' visual control of body motion during avoidance.     

Orientation sensitive properties of visually driven neurons in extrastriate area 21a of cat cortex

Orientation sensitive properties of extrastriate area 21a neurons were investigated. Special attention was paid to the qualitative characteristics of neuron responses to the different orientations of visual stimulus motion across neuron classical receptive fields (CRF). The results of experiments have shown that a group of neurons (31%) in area 21a with specialized responses to moving visual stimuli changed their direction selective (DS) characteristics depending on the orientation of the stimulus movement. Some neurons reveal an abrupt drop of the direction sensitivity index (DI) to certain orientation (58%), and some show significant increase of DI at one of applied orientations of stimulus motion (22%). Detailed investigation of response patterns of non-directional neurons to different orientations of stimulus motion have revealed clear-cut qualitative differences, such as different regularities in the distribution of inter-peak inhibitory intervals in the response pattern in dependence of the orientation of stimulus motion. The investigation of neuron CRF stationary functional organization did not reveal correlations between RF's spatial functional organization, and that of qualitative modulations of neuron response patterns. A suggestion was put forward, that visual information central processing of orientation discrimination is a complex integrative process that includes quantitative as well as qualitative transformations of neuron activity.     

VEP Responses to Op-Art Stimuli

Several types of striped patterns have been reported to cause adverse sensations described as visual discomfort. Previous research using op-art-based stimuli has demonstrated that spurious eye movement signals can cause the experience of illusory motion, or shimmering effects, which might be perceived as uncomfortable. Whilst the shimmering effects are one cause of discomfort, another possible contributor to discomfort is excessive neural responses: As striped patterns do not have the statistical redundancy typical of natural images, they are perhaps unable to be encoded efficiently. If this is the case, then this should be seen in the amplitude of the EEG response. This study found that stimuli that were judged to be most comfortable were also those with the lowest EEG amplitude. This provides some support for the idea that excessive neural responses might also contribute to discomfort judgements in normal populations, in stimuli controlled for perceived contrast.     

Motion-based analysis of spatial patterns by the human visual system

BACKGROUND: It is known that the visibility of patterns presented through stationary multiple slits is significantly improved by pattern movements. This study investigated whether this spatiotemporal pattern interpolation is supported by motion mechanisms, as opposed to the general belief that the human visual cortex initially analyses spatial patterns independent of their movements. RESULTS: Psychophysical experiments showed that multislit viewing could not be ascribed to such motion-irrelevant factors as retinal painting by tracking eye movements or an increase in the number of views by pattern movements. Pattern perception was more strongly impaired by the masking noise moving in the same direction than by the noise moving in the opposite direction, which indicates the direction selectivity of the pattern interpolation mechanism. A direction-selective impairment of pattern perception by motion adaptation also indicates the direction selectivity of the interpolation mechanism. Finally, the map of effective spatial frequencies, estimated by a reverse-correlation technique, indicates observers' perception of higher spatial frequencies, the recovery of which is theoretically impossible without the aid of motion information. CONCLUSIONS: These results provide clear evidence against the notion of separate analysis of pattern and motion. The visual system uses motion mechanisms to integrate spatial pattern information along the trajectory of pattern movement in order to obtain clear perception of moving patterns. The pattern integration mechanism is likely to be direction-selective filtering by V1 simple cells, but the integration of the local pattern information into a global figure should be guided by a higher-order motion mechanism such as MT pattern cells.     

Individual Pause-and-Go Motion Is Instrumental to the Formation and Maintenance of Swarms of Marching Locust Nymphs

The principal interactions leading to the emergence of order in swarms of marching locust nymphs was studied both experimentally, using small groups of marching locusts in the lab, and using computer simulations. We utilized a custom tracking algorithm to reveal fundamental animal-animal interactions leading to collective motion. Uncovering this behavior introduced a new agent-based modeling approach in which pause-and-go motion is pivotal. The behavioral and modeling findings are largely based on motion-related visual sensory inputs obtained by the individual locust. Results suggest a generic principle, in which intermittent animal motion can be considered as a sequence of individual decisions as animals repeatedly reassess their situation and decide whether or not to swarm. This interpretation implies, among other things, some generic characteristics regarding the build-up and emergence of collective order in swarms: in particular, that order and disorder are generic meta-stable states of the system, suggesting that the emergence of order is kinetic and does not necessarily require external environmental changes. This work calls for further experimental as well as theoretical investigation of the neural mechanisms underlying locust coordinative behavior.     

Sensitivity variations in the visual system,contrast resolution and eye movements

Attention is drawn to the fact that under normal visual conditions the sensitivity of the receptor units of the visual system are subject to spatial and temporal variations, and that consequently in performing pattern recognition the visual cortex has to discriminate between external luminance structure and internal sensitivity structure. It is suggested that eye movements are the method by which this discrimination is performed. In a simplified model analysis it is shown that eye movements are a suitable mechanism for this discrimination. Implications of this model for detection threshold and stabilized retinal images are discussed. A new interpretation of the adaptation to sine wave grids is given.     

Collision avoidance and a looming sensitive neuron: size matters but biggest is not necessarily best

Locusts possess visual neurons that can be uniquely identified in each locust and that respond selectively to looming stimuli, giving the animal a warning of impending collision. It has been suggested that one such neuron, the lobula giant movement detector (LGMD), issues this warning by generating a peak in its response that occurs ca. 25 ms after a looming object reaches a subtense of 17 degrees on the eye. This peak is proposed to be a trigger for escape behaviour. We use both modelling and electrophysiological techniques to show that this early peak in LGMD response is not the 'essential functional variable' used naturally by the locust to trigger escape, but rather results from the unnaturally large stimulus used in the previous experimental work. The natural predators of Locusta in Africa, where the locust evolved, are small birds such as the fiscal shrike Lanius collaris humeralis and the carmine bee-eater Merops nubicus, with pectoral diameters of 40-45 mm (measurements from museum specimens). Locusta in flight are less than 100 mm wing tip to wing tip. When a locust views small approaching objects, the response of the LGMD continues to increase throughout the object's approach and the locust is able to trigger escape behaviours without the LGMD response peaking prior to collision.