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Responses of primary visual cortical neurons to binocular disparity without depth perception.

The identification of brain regions that are associated with the conscious perception of visual stimuli is a major goal in neuroscience. Here we present a test of whether the signals on neurons in cortical area V1 correspond directly to our conscious perception of binocular stereoscopic depth. Depth perception requires that image features on one retina are first matched with appropriate features on the other retina. The mechanisms that perform this matching can be examined by using random-dot stereograms, in which the left and right eyes view randomly positioned but binocularly correlated dots. We exploit the fact that anticorrelated random-dot stereograms (in which dots in one eye are matched geometrically to dots of the opposite contrast in the other eye) do not give rise to the perception of depth because the matching process does not find a consistent solution. Anti-correlated random-dot stereograms contain binocular features that could excite neurons that have not solved the correspondence problem. We demonstrate that disparity-selective neurons in V1 signal the disparity of anticorrelated random-dot stereograms, indicating that they do not unambiguously signal stereoscopic depth. Hence single V1 neurons cannot account for the conscious perception of stereopsis, although combining the outputs of many V1 neurons could solve the matching problem. The accompanying paper suggests an additional function for disparity signals from V1: they may be important for the rapid involuntary control of vergence eye movements (eye movements that bring the images on the two foveae into register).

Animals↗

Stimulus mislocalization depends on spatial frequency.

It was previously reported that briefly presented peripheral stimuli are perceived closer to fixation than continuously presented stimuli at the same eccentricity; this effect has, however, not proved consistently replicable. In this study it was investigated whether the misperception of location might depend upon the spatial frequency content of the stimulus. Spatial-frequency-filtered vertical bars were displayed briefly and their locations were judged relative to continuously visible comparison spots. For monocular stimuli, a significant foveopetal mislocalization of the bar was obtained that increased in size as spatial frequency was lowered. Even larger mislocalizations were obtained for dichoptically presented horizontally disparate pairs of bars, and this effect also increased at low spatial frequencies. Possible underlying mechanisms are discussed, and spatial frequency is suggested to have been the confounding factor in previous studies.

Eye Movements↗

Evidence for the correcting-mechanism explanation of the Kanizsa amodal shrinkage.

An object phenomenally shrinks in its horizontal dimension when shown on a 2-D plane as if the central portion of the object were partially occluded by another vertical one in 3-D space (the Kanizsa amodal shrinkage). We examined the predictions of the correcting-mechanism hypothesis proposed by Ohtsuka and Ono (2002, Proceedings of SPIE 4864 167-174), which states that an inappropriate operation of the mechanism that corrects a phenomenal increase in monocularly visible areas accompanied by a stereoscopic occluder gives rise to the illusion. In this study we measured the perceived width (or height in experiment 3) of a square seen behind a rectangle, while controlling other factors which potentially influence the illusion, such as the division of space or depth stratification. The results of five experiments showed that (a) the perceived width was not influenced when the occluder had a relatively large binocular disparity, but was underestimated when the occluder did not have disparity, and (b) the shrinkage diminished when the foreground rectangle was transparent, was horizontally oriented, or contained no pictorial occlusion cues. These results support the hypothesis that the correcting mechanism, triggered by pictorial occlusion cues, contributes to the Kanizsa shrinkage.

Form Perception↗

[Synthesis of binocular disparity with motion parallax in depth slant perception: statistical efficiency approach].

Statistical efficiency approach is used in order to investigate sampling properties of both binocular disparity and motion parallax processes in depth slant perception and to examine the independent decisions model (Mulligan & Shaw, 1980) consisting of these processes. We carried out two experiments in which each cue was displayed solely (i.e., single condition) and both cues were displayed simultaneously (i.e., multiple condition). A two-alternative forced-choice (2AFC) task was used under the situation where a Gaussian noise was added to the stimulus value of depth slant. Statistical efficiencies were calculated in each experiment. The results showed that sampling size from binocular disparity was at least comparable or larger by a factor of 2.5-3.5 with that from motion parallax and that efficiencies of the multiple condition considerably exceeded those of the single condition. This suggests invalidity of the independent decisions model.

Cues↗

Impaired vertical phoria adaptation in patients with cerebellar dysfunction.

PURPOSE: To determine whether phoria adaptation to a vertical prism disparity is altered in patients with cerebellar dysfunction. METHODS: With a computer-aided haploscope, adaptive responses of fusion-free eye position to a 10- or 30-minute period was measured in subjects wearing a 3-prism diopter vertical prism over one eye. Thirteen patients with well-documented cerebellar diseases who did not have manifest ocular misalignment or limited versional eye movement and age-matched healthy subjects participated. RESULTS: The mean +/- SD percentage of vertical phoria adaptation was 13% +/- 22% and 20% +/- 16% for the 10- and 30-minute adaptations, respectively. These levels were significantly smaller than the respective ones in the age-matched control group (P < 0.001, repeated measures MANOVA). Seven (54%) of 13 patients, including two with genetically confirmed pure cerebellar lesions (spinocerebellar ataxia type 6), showed markedly reduced responses to both the 10- and 30-minute adaptations. In all three patients with acute cerebellar ataxia, the adaptive response was improved at the same time as remission of cerebellum-associated neurologic deficits. CONCLUSIONS: Phoria adaptation to vertical binocular disparity is frequently impaired in patients with cerebellar dysfunction. These results bolster the hypothesis that phoria adaptation is a cerebellar-dependent response.

Adaptation, Ocular↗

Organization of disparity-selective neurons in macaque area MT.

Neurons selective for binocular disparity are found in a number of visual cortical areas in primates, but there is little evidence that any of these areas are specialized for disparity processing. We have examined the organization of disparity-selective neurons in the middle temporal visual area (MT), an area shown previously to contain an abundance of disparity-sensitive neurons. We recorded extracellularly from MT neurons at regularly spaced intervals along electrode penetrations that passed through MT either normal to the cortical surface or at a shallow oblique angle. Comparison of multiunit and single-unit recordings shows that neurons are clustered in MT according to their disparity selectivity. Across the surface of MT, disparity-selective neurons are found in discrete patches that are separated by regions of MT that exhibit poor disparity tuning. Within disparity-selective patches of MT, we typically observe a smooth progression of preferred disparities (e.g. , near to far) as our electrode travels parallel to the cortical surface. In electrode penetrations normal to the cortical surface, on the other hand, MT neurons generally have similar disparity tuning, with little variation from one recording site to the next. Thus disparity-tuned neurons are organized into cortical columns by preferred disparity, and preferred disparity is mapped systematically within larger, disparity-tuned patches of MT. Combined with other recent findings, the data suggest that MT plays an important role in stereoscopic depth perception in addition to its well known role in motion perception.

Animals↗

Monocular transparency generates quantitative depth.

Monocular zones adjacent to depth steps can create an impression of depth in the absence of binocular disparity. However, the magnitude of depth is not specified. We designed a stereogram that provides information about depth magnitude but which has no disparity. The effect depends on transparency rather than occlusion. For most subjects, depth magnitude produced by monocular transparency was similar to that created by a disparity-defined depth probe. Addition of disparity to monocular transparency did not improve the accuracy of depth settings. The magnitude of depth created by monocular occlusion fell short of that created by monocular transparency.

Adult↗

Aging and the perception of 3-D shape from dynamic patterns of binocular disparity.

In two experiments, we investigated the ability of younger and older observers to perceive and discriminate 3-D shape from static and dynamic patterns of binocular disparity. In both experiments, the younger observers' discrimination accuracies were 20% higher than those of the older observers. Despite this quantitative difference, in all other respects the older observers performed similarly to the younger observers. Both age groups were similarly affected by changes in the magnitude of binocular disparity, by reductions in binocular correspondence, and by increases in the speed of stereoscopic motion. In addition, observers in both age groups exhibited an advantage in performance for dynamic stereograms when the patterns of binocular disparity contained significant amounts of correspondence "noise." The process of aging does affect stereopsis, but the effects are quantitative rather than qualitative.

Adult↗

Binocular interactions and disparity coding in area 21a of cat extrastriate visual cortex.

We have examined, using both qualitative and quantitative techniques, binocular interactions of extracellularly recorded single neurons in the extrastriate cortical area 21a of anaesthetized and paralysed cats. Consistent with previous reports we have found that: (a) all area 21a neurons were orientation-selective, with about 65% of them preferring orientations within 30 degrees of the vertical; and (b) over 75% of area 21a cells could be activated through either eye. Furthermore, a significant minority (4 cells; about 10%) of a subpopulation of 39 neurons in which binocular interactions were examined quantitatively, were "obligatory binocular neurons", that is, they responded very weakly, if at all, to the monocular stimuli presented through either eye but responded vigorously to simultaneous stimulation through both eyes. Almost 70% (27/39) of neurons tested quantitatively for binocular interaction have shown significant modulation (over 50%) of their peak responses in relation to binocular positional retinal disparities. The majority of neurons sensitive to binocular positional disparities resembled either "tuned excitatory" (22 cells; 56.5% of the sample) or "tuned inhibitory" (2 cells; 5% of our sample) cells. In particular, they gave, respectively, maximal or minimal responses to optimally oriented, moving photic stimuli when the receptive fields plotted through each eye completely or partially overlapped. Although neurons recorded in area 21a have relatively large receptive fields (mean width 3.3 +/- 1.1 degrees; range 2.0-5.6 degrees), the mean width of the disparity tuning curve (2.8 +/- 1.0 degrees; range 1.3-4.8 degrees) for our sample of area 21a neurons was similar to those of neurons with significantly smaller receptive fields, recorded in areas 17 and 18 of cat's primary visual cortex. We conclude that area 21a of the cat, like areas 17 and 18 of primary visual cortex, is likely to play an important role in binocular depth discrimination and might constitute a "higher order" area for stereoscopic binocular vision.

Animals↗

[Effects on depth perception and pattern recognition in random dot stereograms by changing the matrix dot density].

INTRODUCTION: Julesz named the ability of depth perception in random dot stereograms as "global stereopsis". At present there are only high-density random dot stereograms with different disparities in use in the ophthalmological practice. The aim of our exploration is to measure the effects on pattern recognition and depth perception by thinning out the dot density in random dot stereograms down to very low levels (< 1%). METHODS: Experiments were performed by 43 volunteer observers using two sequences of random dot stereograms. Each sequence has its own constant disparity and within a sequence the dot density decreased to values less than 1%. Additionally we compared the performance in these tests with the performance in the conventional haploscopic stereoscopic tests (Titmus, TNO, Randot). RESULTS: Of the observers 48.8% (n = 21) were able to establish a complete depth perception and pattern recognition ("global stereopsis") at a matrix dot density of 0.3% and a disparity of 730" (12, 17'). Another group of subjects (48.8%, n = 21) saw the sub-matrix at 0.3% dot density in depth as a star field ("local stereopsis") but was not able to recognise the form of the sub-matrix. A significant correlation does not exist between these results and those from the conventional stereoscopic test. DISCUSSION: We assume from these results that the very low density random dot stereograms measure a new quality of stereoscopic vision, from which we can obtain information about the density of cortical binocular activated elements in the human brain.

Adolescent↗

Shape analysis and stereopsis for human depth perception.

The perceived relative depth of two isolated short parallel lines in the center of a scene is known to depend on the disparities and positions of other items in the scene, as well as on their own disparities. We demonstrate here that the shapes of these other items also contribute significantly to the perceived depth, and that these non-disparity influences on depth judgements may already be evident when only three dots are presented as stimuli. When two short vertical test lines are surrounded by a trapezoidal "picture frame", the perceived relative depth of the test lines is affected by the shape of the trapezoid as well as by the disparities assigned to its vertical parallel sides. The influence of the trapezoidal frame can be interpreted as an effect of perspective. The induced relative depth of the test lines is measured by recording the amount of "compensating disparity" that must be given to one of the lines in order for observers to judge the two test lines to be equidistant from the observer's viewing position. Surprisingly, for fixed disparities of the vertical edges of the surrounding picture frame, the induced depth of the test lines increases as the difference in the lengths of the vertical sides increases, regardless of whether the perspective interpretation of the difference in the lengths is consistent with or in conflict with the disparity-defined slant. Shape-related apparent depth changes are especially sensitive to the shape of the trapezoid if it is nearly rectangular, and are comparable in magnitude to those resulting from changes in disparity of the surrounding frame. When a pair of short vertical parallel test lines is presented alone, without a surrounding frame or any other items in the scene, excellent relative depth discrimination is displayed by most subjects. However, if the lines are replaced by squares, trapezoids, triangles, single horizontal lines, or other figures of about the same size as the original test lines, the slant discrimination threshold for these plane figures for naïve observers become poorer by a factor of 20-100. By the use of a feedback signal, observers can be trained to use only disparity cues and ignore shape effects. Some observers have difficulty ignoring the shapes of some figures, the "difficult" figures being different for each observer. After training, the relative depth thresholds for most figures approach those of the original unconnected parallel test lines.

Depth Perception↗

[Can fixation disparity be detected reliably by measurement and correctional techniques according H.J. Haase (MKH)?].

BACKGROUND: The theory of the "Measuring and Correction Methods of H.-J. Haase" (MCH) states that a small misalignment of one eye, called fixation disparity, indicates a difficulty in overcoming a "vergence position of rest" that is different from ortho position. This difficulty, so the theory, can cause asthenopic complaints, such as headaches, and these complaints can be relieved by prisms. The theory further claims that fixation disparity can be ascertained by a series of tests which depend on the subject's perception. The tests most decisive for the diagnosis of a so-called fixation disparity type 2 consist of stereo displays. The magnitude of the prism that allows the subject to see the test configurations in symmetry is thought to be the one that corrects the "vergence position of rest". METHODS: Nine subjects with healthy eyes in whom a "fixation disparity type 2" had been diagnosed were selected for the study. Misalignment of the eyes was determined according to the principle of the unilateral cover test. Targets identical for both eyes were presented on the screen of the Polatest E. Then, the target was deleted for one eye and the ensuing position change of the other eye was measured, using the search coil technique. This test was performed both with and without the MCH prism. RESULTS: In all 9 subjects the misalignment was less than 10 minutes of arc, i.e. in the range of normal fixation instability. Averaging across the 9 subjects, the deviation of the eye (misaligned according to MCH) was 0.79 +/- 3.45 minutes of arc in the direction opposed to that predicted by the MCH, a value not significantly different from zero. The MCH prism elicited a fusional vergence movement the magnitude of which corresponded to the magnitude of the MCH prism. CONCLUSION: Ascertaining fixation disparity with the MCH is unreliable. Accordingly, it appears dubious to correct a "vergence position of rest" on the basis of the MCH.

Adult↗

Disparity and shading cues cooperate for surface interpolation.

In two experiments, we tested whether disparity and shading cues cooperated for surface interpolation. Observers adjusted a probe dot to/lie on a surface specified either by a sparse disparity field, a continuous stereo shading or monocular shading gradient, or both cues. Observers' adjustments were very consistent with disparity information but their adjustments were much more variable with shading information. However, observers significantly improved their precision when both cues were present, relative to when only disparity information was present. These results cannot be explained by assuming that separate modules analyze disparity and shading information, even if observers optimally combined these cues. Rather, we attribute this improvement to a process through which the shading gradient constrains the disparity field in regions where disparities cannot be directly measured. This cooperative process may be based on the natural covariation existing between these cues produced by the retinal projection of smooth surfaces.

Adult↗

Relationship between phase and energy methods for disparity computation.

The phase and energy methods for computing binocular disparity maps from stereograms are motivated differently, have different physiological relevances, and involve different computational steps. Nevertheless, we demonstrate that at the final stages where disparity values are made explicit, the simplest versions of the two methods are exactly equivalent. The equivalence also holds when the quadrature-pair construction in the energy method is replaced with a more physiologically plausible phase-averaging step. The equivalence fails, however, when the phase-difference receptive field model is replaced by the position-shift model. Additionally, intermediate results from the two methods are always quite distinct. In particular, the energy method generates a distributed disparity representation similar to that found in the visual cortex, while the phase method does not. Finally, more elaborate versions of the two methods are in general not equivalent. We also briefly compare these two methods with some other stereo models in the literature.

Animals↗

The interaction of binocular disparity and motion parallax in the computation of depth.

Depth from binocular disparity and motion parallax has traditionally been assumed to be the product of separate and independent processes. We report two experiments which used classical psychophysical paradigms to test this assumption. The first tested whether there was an elevation in the thresholds for detecting the 3D structure of corrugated surfaces defined by either binocular disparity or motion parallax following prolonged viewing (adaptation) of supra-threshold surfaces defined by either the same or different cue (threshold elevation). The second experiment tested whether the depth detection thresholds for a compound stimulus, containing both binocular disparity and motion parallax, were lower than the thresholds determined for each of the components separately (sub-threshold summation). Experiment 1 showed a substantial amount of within- and between-cue threshold elevation and experiment 2 revealed the presence of sub-threshold summation. Together, these results support the view that the combination of binocular disparity and motion parallax information is not limited to a linear, weighted addition of their individual depth estimates but that the cues can interact non-linearly in the computation of depth.

Adaptation, Ocular↗

Remote operation: a selective review of research into visual depth perception.

Some perceptual motor operations are performed remotely; examples include the handling of life-threatening materials and surgical procedures. A camera conveys the site of operation to a TV monitor, so depth perception relies mainly on pictorial information, perhaps with enhancement of the occlusion cue by motion. However, motion information such as motion parallax is not likely to be important. The effectiveness of pictorial information is diminished by monocular and binocular information conveying flatness of the screen and by difficulties in scaling: Only a degree of relative depth can be conveyed. Furthermore, pictorial information can mislead. Depth perception is probably adequate in remote operation, if target objects are well separated, with well-defined edges and familiar shapes. Stereoscopic viewing systems are being developed to introduce binocular information to remote operation. However, stereoscopic viewing is problematic because binocular disparity conflicts with convergence and monocular information. An alternative strategy to improve precision in remote operation may be to rely on individuals who lack binocular function: There is redundancy in depth information, and such individuals seem to compensate for the lack of binocular function.

Depth Perception↗

Binocular coordination of the eyes during reading: word frequency and case alternation affect fixation duration but not fixation disparity.

This experiment investigated whether properties of the text being read affect binocular coordination of the eyes during reading. Readers' binocular eye movements were recorded while they read sentences that contained high- and low-frequency words. In addition, half of the sentences were presented in normal case, and half were presented in alternating case (i.e., AlTeRnAtInG cAsE). Past research has suggested that the visual system tolerates less binocular fixation disparity with alternating than with normal case (Heller & Radach, 1999). While both word frequency and alternating case produced large effects on fixation durations on the target word, neither manipulation affected the magnitude of fixation disparity. It is concluded that linguistic and visual properties of the text being read do not influence binocular coordination of the eyes during reading. Additional analyses also showed no difference in fixation disparity between reading and a nonlinguistic task. Implications of these results for split-fovea models of reading are discussed.

Analysis of Variance↗