Dichoptic Presentation Research Articles

Serial spatial filters in vision

Observers viewing two superimposed laser interference fringes of nearly equal spatial frequency see an illusory grating of low spatial frequency, even when the spatial frequency of the fringes exceeds the resolution limit. This grating is a product of nonlinear distortion within the visual system ( MacLeod, Williams and Makous (1992) Vision Research, 32, 347–363]. By separately manipulating the spatial frequencies of the interference fringes and the distortion gratings, we decomposed the contrast sensitivity function into two serial components separated by the nonlinear process. Losses in the optics of the eye were avoided by use of laser interferometry. Spatial summation preceding the nonlinear stage was restricted to the light-collecting area of individual cones and was directly proportional to the diameters of cone inner segments at three retinal eccentricities; this suggests that light is trapped within cones at the level of their inner segments. Even 30° from the fovea, the nonlinear stage precedes the site where separate signals from individual cones are no longer maintained; this leads us to suggest that the nonlinear process lies within the retina. In addition, spatial antagonism precedes the nonlinear stage; this places the nonlinear process at a site following the outer segments of the cones. Dichoptic presentation of the interference fringes failed to produce illusory gratings; that is, the nonlinearities within the binocular pathway do not produce distortions like those produced by the monocular nonlinearity.

Oscillatory discharge in the visual system: does it have a functional role?

1. The discharge of individual neurons in the visual cortex and lateral geniculate nucleus (LGN) of anesthetized and paralyzed cats and kittens was examined for the presence of oscillatory activity. Neural firing was evoked through the monoptic or dichoptic presentation of drifting gratings and random sequences of flashed bars. The degree to which different oscillatory frequencies were present in neural discharge was quantified by computation of the power spectra of impulse train responses. 2. Action potentials from single cells were recorded extracellularly and isolated on the basis of amplitude. Receptive-field properties of the neurons under study were characterized initially by their discharge in response to gratings of sinusoidal luminance. By varying orientation and spatial frequency, optimal stimulus characteristics were determined. Oscillation analysis was performed on spike trains acquired during repeated presentations of the optimal stimulus by identification of power spectra peaks in the frequency range of rhythmic potentials observed in electroencephalograph studies (30-80 Hz). The amplitude and frequency of the largest peak in this range was used to characterize oscillatory strength and frequency. All discharge in which the peak amplitude exceeded the high-frequency noise by a factor > 1.5 was classified as oscillatory. 3. Of the 342 cortical cells examined, 147 cells displayed oscillatory activity in the 30 to 80-Hz range during portions of their visual response. Sixty out of 169 simple cells, 82 out of 166 complex cells, and 5 out of 7 special complex cells exhibited oscillations. There was no laminar bias in the distribution of oscillatory cells; the proportions of oscillatory cells were similar in all layers. All oscillatory discharge was variable with respect to frequency and strength between successive presentations of the same optimal stimulus. In as little as 10 s, for example, peak frequencies shifted by a factor of two. For many cells, these trial-to-trial variations obscured detectable oscillations when all trials were averaged together. 4. The potential role of neuronal maturation in the generation of oscillatory activity was investigated by studying neuronal responses from kittens at 4 wk postnatal. Of the 80 kitten cells studied, 27 exhibited oscillatory discharge. Although oscillations in the kitten visual cortex spanned the same frequency range as that seen in the adult, oscillations in the midfrequency range (36-44 Hz) are more common in the adult cortex. 5. To explore the possibility that oscillations might play a functional role in vision, we investigated the dependence of oscillations on different stimulus parameters.(ABSTRACT TRUNCATED AT 400 WORDS)

両眼網膜非対応をもたないステレオグラムに対する奥行反応： “Ｌａｕ効果”について

We examined the idea that, when the Zöllner figure in one eye and the principal lines in the figure in the other eye were presented, the perceived depth is due to "phenomenal" disparity. The phenomenal disparity is defined as apparent displacement between two illusory "oblique" principal lines in one eye and two "parallel" principal lines in the other. Observes were asked whether depth was seen or not at upper or lower part of the two perceived lines and, if seen, which line was closer. Seventeen observers showed that only 9% of their responses was consistent with the idea, although they reported depths for disparity stimuli. This suggests that the dichoptic presentation of such figures should not be considered as that requires the processing of binocular disparity.

Visual evoked potentials elicited by subjective contour figures

In order to investigate the relationship between the appearance of illusory figures and the wave form of visual evoked potentials (VEPs), 8 different visual pattern stimuli were presented to 8 normal subjects. Four of the stimuli (experimental stimuli) produced subjective figures and contours (squares and discs). The 4 other stimuli (reference stimuli), although equal to the experimental stimuli in the amount of physical energy, did not produce the illusion of squares or discs. Electrodes were placed on the scalp at central and occipital locations. Three prominent peaks in the occipital record were observed in all subjects. An amplitude difference of VEP N180 (N2) between the subjective figures and the reference stimuli was found in the values for each subject. Enhancement of the VEP of the illusory figure stimuli was observed for a specific component (N2), whereas the amplitude values at the central components and the occipital P120 (P2) and P280 (P3) were almost the same as the reference values. The VEP (N2 component) amplitude enhancement at the occipital area for subjective figure stimuli suggests that illusory contour formation takes place at higher levels in the visual system. This was known from experiments using dichoptic presentation.

Brightness perception and filling-in

Three experiments were performed in which a stimulus with homogeneous color and luminance was masked by a second stimulus containing contours. In the first experiment the target was a large white disk and the mask was a white circle concentric with the disk but of smaller radius. We found that the mask had a large (up to 2 log unit) suppressive effect on the brightness of the target, but only inside the radius of the mask. With monoptic presentation of target and mask, the greatest suppression was observed with an SOA of 50–100 msec. With dichoptic presentation the strongest suppression was obtained with simultaneous stimuli. The second experiment demonstrated that the latest time at which masking was effective was correlated with the distance between the edge of the target stimulus and the contour in the mask. One possible explanation of the results from these two experiments is that the masking contour is interfering with the propagation of a brightness signal from the target's border. In the third experiment gaps were introduced into the masking circle. Surprisingly, even with rather large gaps there was significant suppression of brightness in the center of the target. We have encountered difficulties attempting to account for these findings with known physiological mechanisms such as lateral inhibition. A qualitative explanation of the results that looks promising is a two-component process involving brightness filling-in and smoothing to satisfy fixed boundary conditions at contours.

Spatial sensitization in the short wavelength sensitive pathways under dichoptic viewing conditions

Spatial sensitization (Westheimer) functions were measured under conditions that isolated the short wavelength sensitive pathways. The variable diameter pedestal and the test probe were either presented to the same eye (monoptic presentation) or to different eyes (dichoptic presentation). The most significant new finding was that a dichoptically presented, small, blue pedestal caused threshold elevations of about 1–2 long units for an S cone detected probe. However, a large pedestal caused little or no change in threshold. This result contrasts with previous results using white light stimuli, which showed that steadily presented dichoptic pedestals caused little or no threshold change. Furthermore, we show there is little masking when the probe is detected by the isolated M or L cone pathways. These data thus demonstrate a binocular, size dependent interaction revealed only when S cones detect the probe.

Simultaneous orientation contrast for lines in the human fovea

When it is surrounded by lines of a differing orientation, a test line changes its apparent orientation in a direction away from that of the surround lines. Using a nulling technique to arrive at numerical values, the properties of this simultaneous orientation contrast have been analyzed: it diminishes with distance of the surround lines; rises and then falls off as a function of surround line orientation; decreases with exposure duration; is sharply tuned (± 100msec) for synchrony of test and surround line presentation; is robust to differences between test and surround line disparity but not intensity; and is reduced with dichoptic presentation of test and surround lines. Orientation contrast can be induced in a variety of oriented features, including illusory contours, an ellipse, a moving dot and a row of dots or lines, but two dots alone don't suffice. The results are taken as evidence that orientation is a domain sui generis, in which simultaneous contrast is exhibited in the same manner as in the domains of color, brightness and disparity.

Adaptation of suprathreshold contrast and luminance stimuli.

The temporal and spatial properties of the difference in perceived contrast and brightness of two suprathreshold stimuli presented successively in different retinal locations were determined. Stimulus onset asynchrony (SOA) was varied and the perceived contrast or brightness of the first stimulus (S1) was measured as a function of SOA by matching the contrast or luminance of the second stimulus (S2) to that of S1. The two stimuli overlapped in time for 200 ms to allow the comparison to be made. The adjusted values for S2 could well be fitted with an exponential decay function of SOA. For luminance increments and decrements the time constant for this function was 253 ms; for checkerboards with checks of size 16 min square the time constant was 164 ms. The difference in perceived contrast was dependent on initial contrast in a nonlinear fashion. It increased with increasing check size and was independent of the mean luminance and spatial proximity of the two stimuli. The phenomenon was observed with different pattern types and with dichoptic presentation, but could only be seen when direct comparison of the two stimuli was possible.

Changes in luminance affect dichoptic unique yellow

An extension of the Benzschawel–Guth ATDN model [ Color Res. Appl.9, 133 ( 1984)] predicts that, for dichoptic presentation, the green-to-red (G/R) ratio required to obtain a unique yellow percept should decrease with increasing luminance, because the green postreceptor hue response increases faster than the red postreceptor hue response with increasing intensity. However, for monoptic presentation the model predicts that the G/R ratio should remain invariant, because the neural responses from the receptors increase at the same rate with increases in luminance. Data from four subjects confirmed these predictions.

Monocular and binocular response properties of cells in the striate-recipient zone of the cat's lateral posterior-pulvinar complex.

1. We have studied response properties of single cells in the striate-recipient zone of the cat's lateral posterior-pulvinar (LP-P) complex. This zone is in the lateral section of the lateral posterior nucleus (LP1). Our purpose was to determine basic response characteristics of these cells and to investigate the possibility that the LP-P complex is a center of integration that is dominated by input from visual cortex. 2. The majority (72%) of cells in the striate-recipient zone respond to drifting sinusoidal gratings with unmodulated discharge. 3. Cells in the LP1 are selective to the orientation of gratings, and tuning functions have a mean bandwidth of 31 degrees. More than one-half of these units are direction-selective. The preferred orientation and the tuning widths for the two eyes are generally well matched. However, a few cells exhibited the interesting property of opposite preferred directions for the two eyes. Orientation tuning for a small group of cells was different for the mean discharge and first harmonic components, suggesting a convergence from different inputs to these cells. 4. Two-thirds of LP1 cells are tuned to low spatial frequencies (less than 0.5 c/deg). The tuning is broad with a mean bandwidth of 2.2 octaves. The remaining one-third of the units are low-pass because they show no attenuation of their responses to low spatial frequencies. Both eyes exhibit the same spatial frequency preference and the same spatial frequency tuning. There is a high correlation between spatial frequency and orientation selectivities. 5. All cells tested are tuned for temporal frequency with a sharp attenuation for low frequencies. The optimal values range between 4 and 8 Hz, and the mean bandwidth is 2.2 octaves. 6. Cells in LP1 are mostly binocular. When monocular, cells are almost always contralaterally driven. Dichoptic presentation of gratings reveals the presence of strong binocular interaction. In almost all cases, these interactions are phase specific. The cell's discharge is facilitated at particular phases and inhibited at phases 180 degrees away. These binocular interactions are orientation dependent. 7. Twenty-five percent of the cells with phase-specific binocular facilitation appear to be monocular when each eye is tested separately. For three cells, we observed a non-phase-specific inhibitory effect of the silent eye. 8. Our findings indicate that LP1 cells form a relatively homogeneous group, suggesting a high degree of integration of multiple cortical inputs.(ABSTRACT TRUNCATED AT 400 WORDS)

Early Motion Processes and the Kinetic Depth Effect

It has been known for over 30 years that motion information alone is sufficient to yield a vivid impression of three-dimensional object structure. For example, a computer simulation of a transparent sphere, the surface of which is randomly speckled with dots, gives no impression of depth when presented as a stationary pattern on a visual display. As soon as the sphere is made to rotate in a series of discrete steps or frames, its 3-D structure becomes apparent. Three experiments are described which use this stimulus, and find that depth perception in these conditions depends crucially on the spatial and temporal properties of the display: 1. Depth is seen reliably only for between-frame rotations of less than 15°, using two-frame and four-frame sequences. 2. Parametric observations using a wide range of frame durations and inter-frame intervals reveal that depth is seen only for inter-frame intervals below 80 msec and is optimal when the stimulus can be sampled at intervals of about 40–60 msec. 3. Monoptic presentation of two frames of the stimulus is sufficient to yield depth, but the impression is destroyed by dichoptic presentation. These data are in close agreement with the observed limits of direction perception in experiments using “short-range” stimuli. It is concluded that depth perception in the motion display used in these experiments depends on the outputs of low-level or “short-range” motion detectors.

Spatial properties of mechanisms for detection of moving dot targets in dynamic visual noise.

The maximum displacement threshold for direction discrimination (dmax) was determined for single or paired dot targets moving discretely against a background of dynamic visual noise. dmax rose as the spatial density of noise was reduced, or when the interframe interval was decreased. dmax was greater for dot pairs than for single dots, and rose progressively as the distance between the dots was reduced. dmax was also greater if the orientation of the target dot pairs differed from the orientation of paired dots in the background noise. Dichoptic presentation of the target and background noise allowed the target to be detected with an accuracy that did not depend on displacement.

Monocular motion sensing, binocular motion perception

The two-process account of motion perception and its binocular organization were addressed in experiments on apparent movement (AM) with three types of grating: sinusoidal; random bar width; and square-wave with missing fundamental (MF). Monocular MF gratings sampled four times per cycle of drift always appeared to move backwards. Here AM was unrelated to the spatial appearance of the pattern, and followed the motion of the dominant spatial frequency component (the third harmonic). We take this reversed AM to be characteristic of “short-range” motion sensors. It did not occur dichoptically, implying that the direction-selective mechanism of motion sensors is purely monocular. AM was seen with dichoptic presentation for all three types of grating. Performance improved with the length of the stimulus sequence, as predicted by probability summation. This result reconciles previous positive and negative findings on dichoptic AM. The perceived direction of dichoptic AM was consistent with polarity-selective matching of features over time (the “long-range process”). The most telling effect supporting featurematching in dichoptic motion was that dichoptic MF motion reversed direction with a change in the visible features of the pattern (induced by changes in contrast and pulse duration); monocular apparent motion did not. Two routes from spatial frequency channels to the perception of object motion are discussed.

Semantic priming without awareness: some methodological considerations and replications.

Previous studies of subliminal semantic priming effects are considered. The distinction between subjective and objective detection thresholds proposed by Cheesman and Merikle (1984) is accepted, and it is concluded that no convincing evidence has been adduced to support or deny the existence of semantic priming effects at or below objective threshold. New criteria for the assessment of objective thresholds are proposed, and four experiments are reported in each of which pattern-masked primes were presented 10% below objective threshold. The first three experiments presented primes, mask, and targets binocularly; the fourth used dichoptic presentation. Data were analysed using both conventional F ratios and the quasi F ratio advocated by Clark (1973). Results showed that significant subliminal semantic priming effects occurred in each experiment when conventional F ratios were used, and significant effects occurred in all but the first experiment when quasi F was used. Binocular and dichoptic presentation of stimuli gave semantic priming effects whose magnitudes did not differ significantly. It is concluded that semantic priming effects can be obtained below objective detection threshold. It is also concluded that replication with different participants and different stimuli is preferable to the use of quasi F ratios in investigations where verbal stimuli are used.

The effect of area, separation, and dichoptic presentation on the perception of achromatic color.

In a center-surround arrangement of two luminances, luminance ratio, area of the surrounding higher luminances, and separation of the two luminances were varied. Decreasing the area of the surrounding luminance caused the achromatic color in the center to be lighter, and a similar but smaller effect was caused by increasing separation up to approximately 30′ of visual angle; increasing separation beyond 30′ did not lighten the color further. At higher luminance ratios, which resulted in darker colors, the effect of decreasing the area of the surround became more pronounced. Dichoptic presentation of the surround and center luminances resulted in perception of achromatic colors in the center that depended on luminance ratios, in a manner similar to perception of binocularly presented luminances. Dichoptically produced colors were some-what lighter than those that resulted when center and surround were given to each eye

Spatial phase dependence and the role of motion detection in monocular and dichoptic forward masking

Contrast thresholds for briefly flashed gratings were measured by the QUEST procedure, under conditions of forward masking by gratings of the same spatial frequency (usually 1 c/deg). Low-contrast masks reduced threshold at short onset asynchronies (0 to 50 msec), while higher contrasts raised threshold over a broader temporal range (0 to 100–140 msec). Both effects depended on the spatial phase relation, but in different ways. Threshold reduction at 0–±90deg phases probably arises from spatio-temporal filtering by direction-selective mechanisms. This conclusion was supported by computer simulation of a motion detector model. The direction-selective stage of motion analysis may be entirely monocular, since facilitation at 90 deg was abolished by dichoptic presentation. Threshold elevation was phase-dependent at short SOA's (20–50 msec), with a minimum at ±90 deg, but was not phase-dependent at longer SOA's (70–140 msec). In-phase masking (0 deg) was about equally strong monocularly and dichoptically, but dichoptic threshold elevation showed no phase-dependence at any SOA. Threshold elevation at longer SOA's, and with dichoptic presentation, may reflect a purely suppressive binocular masking effect, unselective for spatial phase, and its basis may be the same as contrast adaptation. At short SOA's, monocular and binocular masking data apparently reflect a mixture of this phase-independent suppression and phase-selective facilitation.

Duration of visual afterimages on modulated backgrounds: Postreceptoral processes

A foveal afterimage produced by a small photoflash increases in duration when the luminance of a 5.8° diameter background on which it is seen is temporally modulated. At a modulation frequency of 1 Hz and a depth of modulation of 52%, the duration of the afterimage is prolonged by 335% compared to the duration obtained on a steady background. This increase has been attributed to the functional border resulting from the difference in excitability between bleached and unbleached photoreceptors. Afterimage duration is also prolonged, although only by 20%, when the luminance of the background is kept constant, while the luminance of an annular surround is modulated. This finding suggests a weak effect of neural lateral interaction (via area contrast). If the background luminance is modulated only in the contralateral eye (dichoptic presentation), afterimage duration increases by as much as 54% compared to the unmodulated state. This result indicates that afterimages, in part, are sustained by processes mediated by the visual cortex.

Binocular suppression in cortical neurons.

Dichoptic presentation of patterns similar in shape but of very different contrast results in the perception of only the high contrast pattern (binocular suppression). When recording from binocular neurons of the cat visual cortex, we have found an effect which is strikingly similar to this perceptual phenomenon. If a high and a low contrast grating are presented simultaneously, one to each eye, the cell's response to the low contrast stimulus is suppressed.

The contribution of blue-sensitive cones to spatial responses of post-receptoral visual channels in man.

Psychophysical methods developed for the investigation of spatial and temporal pathways in human vision have been applied in combination with the two-colour increment threshold technique of W. S. Stiles to study the way in which signals from blue-sensitive cones are transmitted along the visual pathways. A flicker sensitive spatio-temporal filter, designated 'ST2', has been examined by background modulation methods, and spatial filters sensitive to bars of a specific width by grating adaptation methods employing dichoptic presentation of stimuli. It is shown that the blue-sensitive (pi 3) spectral mechanism contributes to both classes of filter response, in a manner similar to that observed for the red-sensitive spectral mechanism. The binocularly driven, bar-sensitive filters have broad-band spectral response characteristics, thus the data demonstrate that signals arising in blue-sensitive cones converge onto a luminance channel. The results of this investigation, together with those previously published for a second (ST1) spatio-temporal filter, describe a variety of post-receptoral responses involving the pi 3 spectral mechanism.

Frequency of Fusion and of Loss of Fusion, and Binocular Depth Perception with Alternating Stimulus Presentation

Stereoscopic vision was investigated with an experimental design allowing dichoptic stimulus presentation at different frequencies of image alternation. For twenty subjects the frequency of binocular fusion and the frequency of loss of fusion to one stereoscopic image was measured as a function of the convergence angle. In thirteen subjects no dependence of the fusion frequency was found, while seven subjects showed a marked increase of the fusion frequency with increasing angle of convergence. In all cases the frequency of fusion was higher than the frequency of loss of fusion. Both frequencies, however, are lower than the flicker fusion frequency. Under conditions where no monocular cues and no references for stereoptic depth comparisons were presented, the apparent distance of the image from the observer could not be assessed, but perception of relative motion in depth was possible. All subjects assessed the direction of motion accurately down to changes of the convergence angle of 0.2 deg s-1.