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R J Snowden

Publications and source records attributed to R J Snowden.

At least 37 records · Page 2Linked to original sources

Spatial frequency adaptation: threshold elevation and perceived contrast.

We have measured the spread of contrast adaptation across the dimension of spatial frequency. Threshold elevation was tightly tuned to the adapting spatial frequency but became much broader as test contrast was increased. This means that, for a given test frequency, there are some frequencies which do not raise threshold but do result in a loss of perceived contrast. The contrast dependence, retinal specificity and interocular transfer of adaptation effects elicited from same-and remote-frequency adaptation were compared. While we were able to show some distinct differences between threshold and suprathreshold tests, we were unable to demonstrate any reliable differences in the retinal specificity and interocular transfer between same- and remote-frequency adaptation.

Adaptation, Ocular↗

The processing of temporal modulation at different levels of retinal illuminance.

How does our temporal vision change as the mean illuminance reduces? We have examined the processing of near-threshold temporal information for a range of illuminance values (2850--0.15 phot td). At high illuminance, the modulation transfer function can be shown to be mediated via three underlying temporal filters that vary in sensitivity with spatial frequency. As the mean illuminance decreases these channels appear to change their sensitivity. Even at the lowest (scotopic) illuminance levels we were able to find evidence for at least two channels mediating detection threshold. There are also changes in the tuning properties of these channels such that the processing of high temporal frequencies is differentially compromised, resulting in a reduction in the flicker fusion limit of each channel, and a shift in the peak of the band-pass channel. The slope of the fall-off in sensitivity at high temporal frequencies is unaffected by test spatial frequency at each illuminance level, suggesting its limiting factor is one that is insensitive to spatial frequency. We propose that the changes in the tuning of the temporal filters occur because of an early (e.g. photoreceptor) change in the response dynamics, or by interactions between photoreceptors, rather than changes at or beyond the level of the channel response.

Adult↗

The effect of contrast adaptation on briefly presented stimuli.

Wilson and Humanski (1993) have recently reported evidence that adapting to low temporal frequency sinewave gratings yields little threshold elevation for briefly presented test stimuli. We postulated that brief stimuli may be detected by a transient channel which would be minimally affected by a low temporal frequency adapting pattern. We therefore measured the effect of adaptation on briefly presented test stimuli for a wider range of adapting temporal frequencies. The results indicate that adaptation may yield threshold elevation for briefly presented stimuli and that threshold elevation is greater for high than low temporal frequency adapting patterns. These results are consistent with the hypothesis that briefly presented stimuli are detected by a transient channel.

Adaptation, Ocular↗

Isolation and characteristics of a steady-state visually-evoked potential in humans related to the motion of a stimulus.

We have examined the visual potential evoked by two motion stimuli. In the first stimulus (termed coherent motion) a random-dot pattern oscillated between phases of coherent and incoherent ("snowstorm") motion, and in the second a random-dot pattern alternated in direction of motion (termed direction change). We found that the response to the coherent motion stimulus is low-pass with respect to speed, has low contrast sensitivity and increases steadily with the contrast of the stimuli. The direction change visually-evoked potential (VEP) is band-pass with respect to speed, has high contrast sensitivity but then saturates and even reduces as the stimulus contrast is raised above 0.1. The behaviour of the direction change VEP is similar in nature to results from psychophysical experiments of motion perception and to the known properties of directionally selective cells of the cortex. On the other hand the behaviour of the coherent motion VEP suggests this may not be mediated by a mechanism specific to motion.

Contrast Sensitivity↗

Perceived contrast as a function of adaptation duration.

We measured how the perceived contrast of a sinusoidal grating fades as a function of time. Measurements were made for a range of temporal and spatial frequencies and eccentricities. Patterns of high temporal and low spatial frequency exhibited a greater and more rapid loss of apparent contrast (fade) than those of medium frequencies. The rate and amount of fading for a subgroup of moderate frequencies increased when presented peripherally rather than foveally. Further measurements revealed that gratings of disparate spatial frequencies, but with the same threshold sensitivity, exhibit very different fading characteristics but equal threshold elevation. We conclude that the differential loss of apparent contrast is not an artefact of differing proximities to threshold, nor can it be accounted for by differences in the adaptability of underlying spatio-temporal mechanisms at threshold. The differences in fading may thus reflect either a difference in the adaptability of underlying channels above threshold or a differential contribution of such channels to perceived contrast.

Adaptation, Ocular↗

Is global motion really based on spatial integration of local motion signals?

Previous studies have shown that a random-dot kinematogram (RDK) comprising dots, each of which takes a random walk in direction or speed over time, can appear to flow in a single direction. This has been interpreted as evidence for the existence of a co-operative network linking neurons sensitive to different directions/speeds and different spatial locations. We have investigated the possibility that global motion perception in such patterns might simply reflect motion energy detection at a coarse spatial scale (such that many dots fall in the receptive field of one energy detector) without the need to encode local dot motions on a fine spatial scale and then integrate their motions over space. We created random-walk RDKs and then spatially high-pass filtered them to remove low spatial frequencies. Perception of global motion was unimpaired for both direction and speed random walks, showing that the phenomenon is not reliant on low spatial frequencies and must, therefore, involve integration of local motion signals across space, as originally postulated.

Discrimination, Psychological↗

Adaptability of the visual system is inversely related to its sensitivity.

Prolonged viewing of a high-contrast pattern (the adapter) makes similar patterns harder to detect. This threshold-elevation effect was measured as a function of the contrast of the adapter, with use of sinusoidal grating patterns. Increases in the spatial frequency, temporal frequency, or eccentricity of the stimuli had two major consequences. First, the minimum contrast required for detection of the pattern rose; and second, the function that related threshold elevation to adapting contrast became steeper. It is suggested that this increase in the slope of the function reflects the increased gain of these mechanisms, which might occur as compensation for their relatively poor sensitivity. Changing the subject's ability to detect the adapting pattern by means of masks of an orthogonal orientation had markedly different effects on the threshold-elevation-versus-adapting-contrast function. Under these conditions the slope was decreased. The results support the idea that the human visual system tries to compensate for differences in sensitivity by having different gains be associated with different mechanisms.

Adaptation, Ocular↗

Tuning of MST neurons to spiral motions.

Cells in the dorsal division of the medial superior temporal area (MSTd) have large receptive fields and respond to expansion/contraction, rotation, and translation motions. These same motions are generated as we move through the environment, leading investigators to suggest that area MSTd analyzes the optical flow. One influential idea suggests that navigation is achieved by decomposing the optical flow into the separate and discrete channels mentioned above, that is, expansion/contraction, rotation, and translation. We directly tested whether MSTd neurons perform such a decomposition by examining whether there are cells that are preferentially tuned to intermediate spiral motions, which combine both expansion/contraction and rotation components. The finding that many cells in MSTd are preferentially selective for spiral motions indicates that this simple three-channel decomposition hypothesis for MSTd does not appear to be correct. Instead, there is a continuum of patterns to which MSTd cells are selective. In addition, we find that MSTd cells maintain their selectivity when stimuli are moved to different locations in their large receptive fields. This position invariance indicates that MSTd cells selective for expansion cannot give precise information about the retinal location of the focus of expansion. Thus, individual MSTd neurons cannot code, in a precise fashion, the direction of heading by using the location of the focus of expansion. The only way this navigational information could be accurately derived from MSTd is through the use of a coarse, population encoding. Positional invariance and selectivity for a wide array of stimuli suggest that MSTd neurons encode patterns of motion per se, regardless of whether these motions are generated by moving objects or by motion induced by observer locomotion.

Animals↗

Subtractive and divisive adaptation in the human visual system.

Sensory systems can adapt to the conditions imposed on them. In the visual system, adapting to a pattern increases the threshold of the ability to see that pattern, and reduces the perceived contrast of the pattern above threshold. Most neurons of the striate cortex reduce their responsiveness after being stimulated for some time by a high-contrast pattern. Such an effect may lie behind these psychophysical adaptation phenomena. These adaptation effects have been reported to be confined to patterns of similar orientation, which is understandable in that the visual neurons that adapt are only excited by a small range of orientations. Neurophysiological evidence suggests that neurons with different orientation preferences have inhibitory interconnections. It is therefore of interest to explore the possible effects of these connections on perception. Here we show that adapting to a horizontal pattern can reduce the perceived contrast of a vertical test pattern more than a horizontal test pattern. These 'cross-orientation' effects are modelled by a division-like process, whereas the more normal 'similar-orientation' effects are modelled by a subtractive process.

Adaptation, Ocular↗

The response of neurons in areas V1 and MT of the alert rhesus monkey to moving random dot patterns.

We studied the response of single units to moving random dot patterns in areas V1 and MT of the alert macaque monkey. Most cells could be driven by such patterns; however, many cells in V1 did not give a consistent response but fired at a particular point during stimulus presentation. Thus different dot patterns can produce a markedly different response at any particular time, though the time averaged response is similar. A comparison of the directionality of cells in both V1 and MT using random dot patterns shows the cells of MT to be far more directional. In addition our estimates of the percentage of directional cells in both areas are consistent with previous reports using other stimuli. However, we failed to find a bimodality of directionality in V1 which has been reported in some other studies. The variance associated with response was determined for individual cells. In both areas the variance was found to be approximately equal to the mean response, indicating little difference between extrastriate and striate cortex. These estimates are in broad agreement (though the variance appears a little lower) with those of V1 cells of the anesthetized cat. The response of MT cells was simulated on a computer from the estimates derived from the single unit recordings. While the direction tuning of MT cells is quite wide (mean half-width at half-height approximately 50 degrees) it is shown that the cells can reliably discriminate much smaller changes in direction, and the performance of the cells with the smallest discriminanda were comparable to thresholds measured with human subjects using the same stimuli (approximately 1.1 degrees). Minimum discriminanda for individual cells occurred not at the preferred direction, that is, the peak of their tuning curves, but rather on the steep flanks of their tuning curves. This result suggests that the cells which may mediate the discrimination of motion direction may not be the cells most sensitive to that direction.

Animals↗

Orientation bandwidth: the effect of spatial and temporal frequency.

The orientation bandwidths of psychophysically defined channels of human vision were estimated by two techniques for a wide range of spatial and temporal frequencies. The first technique was an adaptation paradigm, where the subjects' ability to see patterns of various orientations was measured before and after adapting to a high contrast pattern. The second technique evaluated subjects' ability to discriminate between two gratings of different orientations in relation to their ability to detect the patterns. Bandwidths were unaffected by temporal frequency at high spatial frequencies but increased with temporal frequency at low spatial frequencies. Bandwidths increased modestly with decreasing spatial frequency at low temporal frequencies but more drastically at high temporal frequencies. Both techniques gave similar results except for patterns with very low spatial and high temporal frequencies. In this region the stimulus appears "spatial-frequency doubled" which may be used as a cue for the orientation discrimination task.

Adaptation, Ocular↗

Temporal properties of human visual filters: number, shapes and spatial covariation.

The temporal properties of the foveal visual filters were revealed using a method which is a variant on previously used noise masking paradigms. This enables the temporal properties of the mechanisms underlying threshold detection of a spatio-temporal probe to be measured. In accord with recent suggestions these results support the existence of three temporal mechanisms. The evidence for the third, higher temporal mechanism is only persuasive at low spatial frequencies. Furthermore, the results suggest that although there is some degree of spatio-temporal covariation in the filtering properties either of individual filters or across the filter population, the well known spatio-temporal covariation in human detection sensitivity is adequately explained by a sensitivity scaling of individual temporal filters with approximately invariant temporal properties.

Contrast Sensitivity↗

Temporal frequency filters in the human peripheral visual field.

The temporal filtering properties of the human peripheral field were investigated by means of measuring: (1) modulation transfer functions for a range of spatial frequencies at four visual field locations (0, 10, 30 and 50 degrees), (2) the contrast of a masking stimulus required to extinguish the visibility of just suprathreshold probes. Results suggest that the number of temporal filters governing detection threshold is dependent upon both eccentricity and spatial frequency. For near-foveal viewing three temporal filters were found (one low-pass and two band-pass), whereas at far eccentricities only one was found (band-pass). A similar result was obtained by modeling the modulation transfer function by simply scaling the sensitivities of three independently derived filters. Our data suggest that (1) changes in the modulation transfer function with respect to spatial frequency and eccentricity can be adequately explained by the changes in sensitivity of a small number of spatio-temporal separable filters; (2) the peripheral field is not merely a coarser version of the fovea but has qualitative differences which may be thought to emphasize the transient properties of the stimulus.

Contrast Sensitivity↗

The perception of visual motion.

Recent developments have led to a greater insight into the complex processes of perception of visual motion. A better understanding of the neuronal circuitry involved and advances in electrophysiological techniques have allowed researchers to alter the perception of an animal with a stimulating electrode. In addition, studies have further elucidated the processes by which signals are combined and compared, allowing a greater understanding of the effects of selective brain damage.

Animals↗

Sensitivity to relative and absolute motion.

The threshold of sensitivity to movement could be governed by mechanisms that are sensitive either to change in spatial position, or directly to the movement itself. The use of spatially complex patterns (random-dot patterns) has been suggested to eliminate the former strategy allowing examination of the movement detecting mechanisms in isolation. By means of such a technique, thresholds for directional judgements were determined for patterns which underwent either a simple displacement or a shearing displacement. Thresholds for shearing motion were found to be around one half of those for simple motion, suggesting that relative, rather than absolute, motion governs performance for small displacements. This contrasts with previous experiments which showed that absolute motion governs performance for much larger displacements.

Attention↗

Measurement of visual channels by contrast adaptation.

Inspection of a high-contrast grating pattern affects our ability to detect patterns that are similar. This technique can be used to infer the underlying mechanisms of the visual system. By using this technique, measurements of the bandwidth of orientation channels are taken for different levels of adapting contrast and adapting duration. If the threshold elevation is plotted as the difference between the unadapted and adapted threshold in decibels, then the orientation bandwidth is invariant if taken at some fraction of the maximum elevation. This results from the fact that, as the orientation difference between the adapting and test patterns increases, the function relating threshold elevation to adapting contrast reduces in slope. These data contradict the often-used 'equivalent contrast transformation' (in which the fall off in the adaptation effect with respect to orientation is expressed in terms of an equivalent reduction in adapting contrast) as this would produce quite different bandwidths at different adapting contrasts. The data also address the issue of the neuronal mechanisms of adaptation.

Adaptation, Ocular↗

The temporal integration and resolution of velocity signals.

The temporal properties of human visual motion detection were explored. Experiment 1 measured thresholds for speed discrimination as a function of stimulus duration. Thresholds fell asymptotically to a Weber fraction around 0.06 over a period of approx. 100 msec, with faster speeds asymptoting at slightly shorter stimulus durations. A second experiment required subjects to discriminate a pattern that was modulated between two speeds from one which remained at a constant speed. The minimum depth of the modulation required to make this judgement was found to be equivalent to a Weber fraction of 0.3 at low modulation rates, around five times greater than when the velocities were presented in isolation (expt 1). At some higher modulation rate performance dramatically declined. The modulation rate at which this occurred decreased with stimulus speed, and increased with stimulus size. The results of expt 1 seem consistent with the known properties of primary motion sensors, while the results of the latter experiments may arise from a later stage integrating the output of these primary motion sensors.

Discrimination, Psychological↗

The response of area MT and V1 neurons to transparent motion.

An important use of motion information is to segment a complex visual scene into surfaces and objects. Transparent motions present a particularly difficult problem for segmentation because more than one velocity vector occurs at each local region in the image, and current machine vision systems fail in these circumstances. The fact that motion transparency is prevalent in natural scenes, and yet artificial systems display an inability to analyze it, suggests that the primate visual system has developed specialized methods for perceiving transparent motion. Also, the currently prevalent model of physiological mechanisms for motion-direction selectivity employs inhibitory interactions between neurons; such interactions would silence neurons under transparent conditions and render the visual system blind to transparent motion. To examine how the primate visual system solves this transparency problem, we recorded the activity of direction-selective cells in the first (area V1) and in a later (area MT) stage in the cortical motion-processing pathway in behaving monkeys. The visual stimuli consisted of random dot patterns forming single moving surfaces, transparent surfaces, and motion discontinuities. We found that area V1 cells responded to their preferred direction of movement even under transparent conditions, whereas area MT cells were suppressed under the transparent condition. These data suggest a simple solution to the transparency problem at the level of area V1. More than one motion vector can be represented at a single retinal location by different subpopulations of neurons tuned to different directions of motion; these subpopulations may represent the early stage for segmenting different, transparent surfaces. The results also suggest that facilitatory mechanisms, which unlike inhibitory interactions are largely unaffected by transparent conditions, play an important role in direction selectivity in area V1. The inhibitory interactions for different motion directions for area MT neurons may contribute to a mechanism for smoothing or averaging the velocity field, computations thought to be necessary for reducing noise and interpolating moving surfaces from sparse information.

Animals↗