Binocular single vision and perceptual processing.

Stimuli with small binocular disparities are seen as single, despite their differing visual directions for the two eyes. Such stimuli also yield stereopsis, but stereopsis and single vision can be dissociated. The occurrence of binocular single vision depends not only on the disparities of individual stimulus elements, but also on the geometrical relation of different parts of the pattern presented to each eye. A pair of vertical bars with opposite binocular disparities is seen as single if the pair is moderately widely spaced but not if it is narrow. Vertical alignment and identity in length of such bars also increase the occurrence of double vision. It is argued that these effects reflect the extraction of features of the monocular patterns, with these detected monocular features determining the binocular percept. Single and double vision of bars differing in orientation can be similarly analysed. The occurrence of relatively elaborate processing of monocular signals does not exclude the possibility that binocular interaction can occur between signals that have not been so processed. Multiple sites or types of binocular interaction are likely.     

Binocular cues and the control of prehension

The present study was designed to assess the importance of binocular information (i.e. binocular disparity and angle of convergence) in the control of prehension. Previous studies which have addressed this question have typically used the same experimental manipulation: comparing prehensile movements executed either under binocular conditions to those executed when one eye was occluded (monocular). However this may not be the correct comparison as in addition to depriving the subject of binocular depth cues. it also deprives the subject of any visual information in one eye. Therefore we determined the prehensile performance when the subject viewed the target object and scene with either (i) two different views (binocular), (ii) two identical views (bi-ocular), or (iii) one view only (monocular). Overall, the qualitative and quantitative performance in the bi-ocular and monocular control conditions was very similar on all the main measures (and different from the performance in the binocular condition). We conclude that the deficits in performance observed found for 'monocular' reaches should be attributed to the lack of local depth information specified by the binocular cues. In addition we speculate that convergence angle and binocular disparity, although involved in both the pre-movement and movement-execution phases of the reach, the cues may be weighted differently in both phases of a prehension movement depending on the behavioural strategy involved.     

The Role of Binocular Disparity in Stereoscopic Images of Objects in the Macaque Anterior Intraparietal Area

Neurons in the macaque Anterior Intraparietal area (AIP) encode depth structure in random-dot stimuli defined by gradients of binocular disparity, but the importance of binocular disparity in real-world objects for AIP neurons is unknown. We investigated the effect of binocular disparity on the responses of AIP neurons to images of real-world objects during passive fixation. We presented stereoscopic images of natural and man-made objects in which the disparity information was congruent or incongruent with disparity gradients present in the real-world objects, and images of the same objects where such gradients were absent. Although more than half of the AIP neurons were significantly affected by binocular disparity, the great majority of AIP neurons remained image selective even in the absence of binocular disparity. AIP neurons tended to prefer stimuli in which the depth information derived from binocular disparity was congruent with the depth information signaled by monocular depth cues, indicating that these monocular depth cues have an influence upon AIP neurons. Finally, in contrast to neurons in the inferior temporal cortex, AIP neurons do not represent images of objects in terms of categories such as animate-inanimate, but utilize representations based upon simple shape features including aspect ratio.     

Binocular coordination: reading stereoscopic sentences in depth

The present study employs a stereoscopic manipulation to present sentences in three dimensions to subjects as they read for comprehension. Subjects read sentences with (a) no depth cues, (b) a monocular depth cue that implied the sentence loomed out of the screen (i.e., increasing retinal size), (c) congruent monocular and binocular (retinal disparity) depth cues (i.e., both implied the sentence loomed out of the screen) and (d) incongruent monocular and binocular depth cues (i.e., the monocular cue implied the sentence loomed out of the screen and the binocular cue implied it receded behind the screen). Reading efficiency was mostly unaffected, suggesting that reading in three dimensions is similar to reading in two dimensions. Importantly, fixation disparity was driven by retinal disparity; fixations were significantly more crossed as readers progressed through the sentence in the congruent condition and significantly more uncrossed in the incongruent condition. We conclude that disparity depth cues are used on-line to drive binocular coordination during reading.     

Binocular visual processing in the owl's telencephalon.

Single neurons recorded from the owl's visual Wulst are surprisingly similar to those found in mammalian striate cortex. The receptive fields of Wulst neurons are elaborated, in an apparently hierarchical fashion, from those of their monocular, concentrically organized inputs to produce binocular interneurons with increasingly sophisticated requirements for stimulus orientation, movement and binocular disparity. Output neurons located in the superficial laminae of the Wulst are the most sophisticated of all, with absolute requirements for a combination of stimuli, which include binocular presentation at a particular horizontal binocular disparity, and with no response unless all of the stimulus conditions are satisfied simultaneously. Such neurons have the properties required for 'global stereopsis', including a receptive field size many times larger than their optimal stimulus, which is more closely matched to the receptive fields of the simpler, disparity-selective interneurons. These marked similarities in functional organization between the avian and mammalian systems exist in spite of a number of structural differences which reflect their separate evoluntionary origins. Discussion therefore includes the possibility that there may exist for nervous systems only a very small number of possible solutions, perhaps a unique one, to the problem of stereopsis.     

Depth generalization from stereo to motion parallax in the owl

Although many sources of three-dimensional information have been isolated and demonstrated to contribute independently, to depth vision in animal studies, it is not clear whether these distinct cues are perceived to be perceptually equivalent. Such ability is observed in humans and would seem to be advantageous for animals as well in coping with the often co-varying (or ambiguous) information about the layout of physical space. We introduce the expression primary-depth-cue equivalence to refer to the ability to perceive mutually consistent information about differences in depth from either stereopsis or motion-parallax. We found that owls trained to detect relative depth as a perceptual category (objects versus holes) when specified by binocular disparity alone (stereopsis), immediately transferred this discrimination to novel stimuli where the equivalent depth categories were available only through differences in motion information produced by head movements (observer-produced motion-parallax). Motion-parallax discrimination did occur under monocular viewing conditions and reliable performance depended heavily on the amplitude of side-to-side head movements. The presence of primary-depth-cue equivalence in the visual system of the owl provides further conformation of the hypothesis that neural systems evolved to detect differences in either disparity or motion information are likely to share similar processing mechanisms.     

Visual cortical responses to the input from the amblyopic eye are suppressed during binocular viewing

Amblyopia is a visual disorder caused by an anomalous early visual experience. It has been suggested that suppression of the visual input from the weaker eye might be a primary underlying mechanism of the amblyopic syndrome. However, it is still an unresolved question to what extent neural responses to the visual information coming from the amblyopic eye are suppressed during binocular viewing. To address this question we measured event-related potentials (ERP) to foveal face stimuli in amblyopic patients, both in monocular and binocular viewing conditions. The results revealed no difference in the amplitude and latency of early components of the ERP responses between the binocular and fellow eye stimulation. On the other hand, early ERP components were reduced and delayed in the case of monocular stimulation of the amblyopic eye as compared to the fellow eye stimulation or to binocular viewing. The magnitude of the amblyopic effect measured on the ERP amplitudes was comparable to that found on the fMRI responses in the fusiform face area using the same face stimuli and task conditions. Our findings showing that the amblyopic effects present on the early ERP components in the case of monocular stimulation are not manifested in the ERP responses during binocular viewing suggest that input from the amblyopic eye is completely suppressed already at the earliest stages of visual cortical processing when stimuli are viewed by both eyes.     

Panum's phenomenon and the confluence of signals from the two eyes in stereoscopy

The signals from the two eyes must be routed to allow either eye to have access to the processing mechanisms for position, shape, colour, etc.; at the same time, information as to the eye of origin must be retained for the purposes of stereoscopy. The study of this confluence of signals from the two eyes was approached psychophysically by studying induced position and depth changes of adjacent binocular and monocular stimuli in the human fovea. It was demonstrated that a monocular visual stimulus located near a binocular one acquires a depth signal, according to a scheme originally proposed by Panum. The effect is unspecific as regards feature shape and brightness, and falls off with a length constant of about 15 minutes of arc in the fovea. A monocular stimulus also affects the apparent depth of its binocular neighbour in a centre-surround manner; disparity pooling changes to disparity repulsion when features are separated by distances of about 3 minutes of arc in the fovea. The findings led to the development of a scheme of uniocular connectivity to a matrix of depth units. Excitation patterns here would depend on the state of the input lines, the intrinsic neuronal interaction properties, and contextural configuring influences from other parts of the nervous system. Experiments showing the spatial extent of pooling and repulsive interaction within the disparity domain help to characterize the stimulus processing in this neural ensemble.     

Predicting Visual Consciousness Electrophysiologically from Intermittent Binocular Rivalry

Purpose

We sought brain activity that predicts visual consciousness.

Methods

We used electroencephalography (EEG) to measure brain activity to a 1000-ms display of sine-wave gratings, oriented vertically in one eye and horizontally in the other. This display yields binocular rivalry: irregular alternations in visual consciousness between the images viewed by the eyes. We replaced both gratings with 200 ms of darkness, the gap, before showing a second display of the same rival gratings for another 1000 ms. We followed this by a 1000-ms mask then a 2000-ms inter-trial interval (ITI). Eleven participants pressed keys after the second display in numerous trials to say whether the orientation of the visible grating changed from before to after the gap or not. Each participant also responded to numerous non-rivalry trials in which the gratings had identical orientations for the two eyes and for which the orientation of both either changed physically after the gap or did not.

Results

We found that greater activity from lateral occipital-parietal-temporal areas about 180 ms after initial onset of rival stimuli predicted a change in visual consciousness more than 1000 ms later, on re-presentation of the rival stimuli. We also found that less activity from parietal, central, and frontal electrodes about 400 ms after initial onset of rival stimuli predicted a change in visual consciousness about 800 ms later, on re-presentation of the rival stimuli. There was no such predictive activity when the change in visual consciousness occurred because the stimuli changed physically.

Conclusion

We found early EEG activity that predicted later visual consciousness. Predictive activity 180 ms after onset of the first display may reflect adaption of the neurons mediating visual consciousness in our displays. Predictive activity 400 ms after onset of the first display may reflect a less-reliable brain state mediating visual consciousness.     

A laminar cortical model of stereopsis and 3D surface perception: closure and da Vinci stereopsis

A laminar cortical model of stereopsis and 3D surface perception is developed and simulated. The model describes how monocular and binocular oriented filtering interact with later stages of 3D boundary formation and surface filling-in in the LGN and cortical areas V1, V2, and V4. It proposes how interactions between layers 4, 3B, and 2/3 in V1 and V2 contribute to stereopsis, and how binocular and monocular information combine to form 3D boundary and surface representations. The model includes two main new developments: (1) It clarifies how surface-to-boundary feedback from V2 thin stripes to pale stripes helps to explain data about stereopsis. This feedback has previously been used to explain data about 3D figure-ground perception. (2) It proposes that the binocular false match problem is subsumed under the Gestalt grouping problem. In particular, the disparity filter, which helps to solve the correspondence problem by eliminating false matches, is realized using inhibitory interneurons as part of the perceptual grouping process by horizontal connections in layer 2/3 of cortical area V2. The enhanced model explains all the psychophysical data previously simulated by Grossberg and Howe (2003), such as contrast variations of dichoptic masking and the correspondence problem, the effect of interocular contrast differences on stereoacuity, Panum's limiting case, the Venetian blind illusion, stereopsis with polarity-reversed stereograms, and da Vinci stereopsis. It also explains psychophysical data about perceptual closure and variations of da Vinci stereopsis that previous models cannot yet explain.     

A scheme for binocular depth perception suggested by neurophysiological evidence

A scheme suggested by neurophysiological evidence is proposed to account for the perceptual phenomena related to binocular stereopsis, especially those observed with Julesz' random stereograms. In the scheme, monocular local features are extracted first. Then the correspondence between the left and right local features is searched for. The correspondence is not one-to-one in general. Thus a sort of direction column due to Blakemore is formed. Each unit in the column is binocular and the receptive field belonging to one eye is located in the same part of the visual field as long as the unit belongs to the same column. However, the receptive fields belonging to the other eye are horizontally displaced to one another. That is, each unit is characterized by binocular disparity. If the correspondence is not one-to-one, then several units belonging to the same column respond simultaneously. Binocular stereopsis can be established if one-to-one correspondence is determined to yield global three dimensional regions. The determination of one-to-one correspondence is carried out through a sort of laterally interacting circuitry in the disparity domain. After the determination of local correspondence, three dimensional global regions are formed by detecting the boundary and by filling-in occluded regions. The results of computer simulation are presented regarding Julesz' stereograms with various types of perturbation. Furthermore, the case of random-dot stereogram in which there is a size difference between the left and right images is simulated. Finally, the computer simulation related to the hysteresis in binocular depth perception is carried out.     

Oscillatory binocular system and temporal segmentation of stereoscopic depth surfaces

A dynamical neural network model of binocular stereopsis is proposed to solve the problem of segmentation which remains ambiguous even when the problem of binocular correspondence is solved. Being compatible with the recent neurophysiological findings (Engel et al. 1991), the model assumes that neural cells show oscillatory activities and that segmentation into a coherent depth surface is coded by synchronization of activities. Employing appropriate constraints for segmentation, the present model shows proper segmentation of depth surfaces and also solves segmentational ambiguity caused by a gap. It is newly shown that binocularly-unmatched monocular cells are discriminated in temporal segmentation of monocular cells caused by recurrent interactions between monocular and binocular cells. Integrative interactions with the other visual components through temporal segmentation are also discussed.     

A computational model of stereoscopic prey capture in praying mantises

We present a simple model which can account for the stereoscopic sensitivity of praying mantis predatory strikes. The model consists of a single “disparity sensor”: a binocular neuron sensitive to stereoscopic disparity and thus to distance from the animal. The model is based closely on the known behavioural and neurophysiological properties of mantis stereopsis. The monocular inputs to the neuron reflect temporal change and are insensitive to contrast sign, making the sensor insensitive to interocular correlation. The monocular receptive fields have a excitatory centre and inhibitory surround, making them tuned to size. The disparity sensor combines inputs from the two eyes linearly, applies a threshold and then an exponent output nonlinearity. The activity of the sensor represents the model mantis’s instantaneous probability of striking. We integrate this over the stimulus duration to obtain the expected number of strikes in response to moving targets with different stereoscopic disparity, size and vertical disparity. We optimised the parameters of the model so as to bring its predictions into agreement with our empirical data on mean strike rate as a function of stimulus size and disparity. The model proves capable of reproducing the relatively broad tuning to size and narrow tuning to stereoscopic disparity seen in mantis striking behaviour. Although the model has only a single centre-surround receptive field in each eye, it displays qualitatively the same interaction between size and disparity as we observed in real mantids: the preferred size increases as simulated prey distance increases beyond the preferred distance. We show that this occurs because of a stereoscopic “false match” between the leading edge of the stimulus in one eye and its trailing edge in the other; further work will be required to find whether such false matches occur in real mantises. Importantly, the model also displays realistic responses to stimuli with vertical disparity and to pairs of identical stimuli offering a “ghost match”, despite not being fitted to these data. This is the first image-computable model of insect stereopsis, and reproduces key features of both neurophysiology and striking behaviour.     

Cognitive Functions and Stereopsis in Patients with Parkinson’s Disease and Alzheimer’s Disease Using 3-Dimensional Television: A Case Controlled Trial

Stereopsis or depth perception is an awareness of the distances of objects from the observer, and binocular disparity is a necessary component of recognizing objects through stereopsis. In the past studies, patients with neurodegenerative disease (Alzheimer dementia, AD; Parkinson’s disease IPD) have problems of stereopsis but they did not have actual stimulation of stereopsis. Therefore in this study, we used a 3-dimensional (3D) movie on 3D television (TV) for actual stereopsis stimulation. We propose research through analyzing differences between the three groups (AD, IPD, and Controls), and identified relations between the results from the Titmus Stereo Fly Test, and the 3D TV test. The study also looked into factors that affect the 3D TV test. Before allowing the patients to watch TV, we examined Titmus stereo Fly Test and cognitive test. We used the 3D version of a movie, of 17 minutes 1 second duration, and carried out a questionnaire about stereopsis. The scores of the stereopsis questionnaire were decreased in AD patients, compared with in IPD and controls, although they did not have any difference of Titmus Stereo Fly Test scores. In IPD patients, cognitive function (Montreal cognitive assessment, MoCA) scores were correlated with the scores of the stereopsis questionnaire. We could conclude that Titmus fly test could not distinguish between the three groups and cognitive dysfunction contributes to actual stereopsis perception in IPD patients. Therefore the 3D TV test of AD and IPD patients was more effective than Titmus fly test.     

Binocular Combination of Second-Order Stimuli

Phase information is a fundamental aspect of visual stimuli. However, the nature of the binocular combination of stimuli defined by modulations in contrast, so-called second-order stimuli, is presently not clear. To address this issue, we measured binocular combination for first- (luminance modulated) and second-order (contrast modulated) stimuli using a binocular phase combination paradigm in seven normal adults. We found that the binocular perceived phase of second-order gratings depends on the interocular signal ratio as has been previously shown for their first order counterparts; the interocular signal ratios when the two eyes were balanced was close to 1 in both first- and second-order phase combinations. However, second-order combination is more linear than previously found for first-order combination. Furthermore, binocular combination of second-order stimuli was similar regardless of whether the carriers in the two eyes were correlated, anti-correlated, or uncorrelated. This suggests that, in normal adults, the binocular phase combination of second-order stimuli occurs after the monocular extracting of the second-order modulations. The sensory balance associated with this second-order combination can be obtained from binocular phase combination measurements.     

Brain damage and global stereopsis.

When a single object lies in front of or beyond the plane of fixation its retinal image lies on disparate positions in the two eyes. This 'local' retinal disparity is an excellent cue to depth, and retinal disparties of a few seconds of arc are detectable by people and monkeys. However, most visual scenes produce a complex array of contours in each eye and we can detect the disparity in the arrays despite the ambiguous nature of the disparities, i.e. each contour in one eye could be related to any of several similar contours in the other eye. This ability, known as 'global' stereopsis, may be selectively impaired following brain damage in man. Global stereopsis was measured in rhesus monkeys before and after removing a different cortical visual area in different groups of animals. Only removal of the inferotemporal cortex impaired global stereopsis. The result is related to the findings with human patients and to receptive field properties of neurons in the inferotemporal cortex of monkeys.     

Stereoscopic mechanisms: binocular responses of the striate cells of cats to moving light and dark bars

New knowledge concerning the internal structure and response properties of the receptive fields of striate cells calls for a fresh appraisal of their binocular interactions in the interest of a better understanding of the neural mechanisms underlying binocular depth discrimination. Binocular position-disparity response profiles were recorded from 71 simple and B-cells in response to moving light and dark bars. Predominantly excitatory (PE) cells (N = 48) had disparity response profiles that were spatially closely similar to their respective monocular responses. In addition, the centrally located excitatory subregions were flanked on one or both sides by non-specific inhibitory regions. PE cells with a preferred stimulus orientation within 30 degrees of the vertical (N = 17) showed binocular facilitations with maximal values that were always more than twice (mean 3.3) the sum of the two monocular responses to the same stimuli and generally greater than the facilitations shown by cells with orientations more than 30 degrees from the vertical (N = 29; mean 2.2 times the sum of the respective monocular responses). The strength of the binocular facilitation depended on the stimulus contrast, the facilitation decreasing with increasing contrast. The receptive-field disparity distribution of the 31 PE cells capable of making significant horizontal disparity discriminations has standard deviations of 0.37 degrees and 0.40 degrees, respectively. Predominantly inhibitory cells (PI) (N = 23) showed two basic types of disparity response profile: symmetric (N = 17) and asymmetric (N = 6). Uncertainty regarding the precise location of the binocular fixation point in the anaesthetized and paralysed preparation made it difficult to categorize PI cells adequately.     

Depth resolution in the pigeon

Summary Pigeons possess a binocular visual field and a retinal region of higher cellular density pointing to the center of this overlap. These features and the precision of pecking behavior suggest that in this lateral-eyed bird cues other than monocular ones might participate in depth judgements.Pigeons were trained with an operant procedure to discriminate between luminous points differing in depth which appeared to the observer as floating in the dark. The accuracy of depth judgements was found to be a function of the ratio between the interstimulus distance and the mean eyes-to-stimulus distance. In a first test (experiment I) no external binocular disparity cues were available, the animal only seeing one luminous point at a time (near or far). In a second test (experiment II) where binocular disparity cues were available, the animal having this time to discriminate a pair of points placed at equal depth from a pair placed at unequal depths, only one pair being visible at a time, depth resolution did not improve. This suggests that, at least within the range of distances explored, the pigeon has no stereoscopic vision. Notwithstanding this, binocular cues do play a role, since when tests were done comparing binocular with monocular viewing (experiment III), monocular depth resolution was significantly worse.     

Binocular visual integration in the crustacean nervous system

Although the behavioral repertoire of crustaceans is largely guided by visual information their visual nervous system has been little explored. In search for central mechanisms of visual integration, this study was aimed at identifying and characterizing brain neurons in the crab involved in binocular visual processing. The study was performed in the intact animal, by recording intracellularly the response to visual stimuli of neurons from one of the two optic lobes. Identified neurons recorded from the medulla (second optic neuropil), which include sustaining neurons, dimming neurons, depolarizing and hyperpolarizing tonic neurons and on-off neurons, all presented exclusively monocular (ipsilateral) responses. In contrast, all wide field movement detector neurons recorded from the lobula (third optic neuropil) responded to moving stimuli presented to the ipsilateral and to the contralateral eye. In these cells, the responses evoked by ipsilateral or contralateral stimulation were almost identical, as revealed by analysing the number and amplitude of the elicited postsynaptic potentials and spikes, and the ability to habituate upon repeated visual stimulation. The results demonstrate that in crustaceans important binocular processing takes place at the level of the lobula.