The response properties of cells in the middle temporal area (area MT) of owl monkey visual cortex

Recordings from 178 single cells in the middle temporal area (area MT) of owl monkey showed that most cells there are orientation- and direction-selective. They also revealed that a powerful range of binocular interactions occur in area MT, with 20% of the cells being responsive to binocular stimulation only, 5% to monocular stimulation only and about 50% of all cells showing some degree of interocular interaction.     

An inference upon the neural network finding binocular correspondence

Previously, the authors proposed a model of neural network extracting binocular parallax (Hirai and Fukushima, 1975). It is a multilayered network whose final layers consist of neural elements corresponding to binocular depth neurons found in monkey's visual cortex. The binocular depth neuron is selectively sensitive to a binocular stimulus with a specific amount of binocular parallax and does not respond to a monocular one. As described in the last chapter of the previous article (Hirai and Fukushima, 1975), when a binocular pair of input patterns consist of, for example, many vertical bars placed very closely to each other, the binocular depth neurons might respond not only to correct binocular pairs, but also to incorrect ones. Our present study is concentrated upon how the visual system finds correct binocular pairs or binocular correspondence. It is assumed that some neural network is cascaded after the binocular depth neurons and finds out correct binocular correspondence by eliminating the incorrect binocular pairs. In this article a model of such neural network is proposed. The performance of the model has been simulated on a digital computer. The results of the computer simulation show that this model finds binocular correspondence satisfactorily. It has been demonstrated by the computer simulation that this model also explains the mechanism of the hysteresis in the binocular depth perception reported by Fender and Julesz (1967)This work has been done in the NHK Broadcasting Science Research Laboratories     

Stereopsis and the random element in the organization of the striate cortex.

Receptive field position and orientation disparities are both properties of binocularly discharged striate neurons. Receptive field position desparities have been used as a key element in the neural theory for binocular depth discrimination. Since most striate cells in the cat are binocular, these position disparities require that cells immediately adjacent to one another in the cortex should show a random scatter in their monocular receptive field positions. Superimposed on the progressive topographical representation of the visual field on the striate cortex there is experimental evidence for a localized monocular receptive field position scatter. The suggestion is examined that the binocular position disparities are built up out of the two monocular position scatters. An examination of receptive field orientation disparities and their relation to the random variation in the monocular preferred orientations of immediately adjacent striate neurons also leads to the conclusion that binocular orientation disparities are a consequence of the two monocular scatters. As for receptive field position, the local scatter in preferred orientation is superimposed on a progressive representation of orientation over larger areas of the cortex. The representation in the striate cortex of visual field position and of stimulus orientation is examined in relation to the correlation between the disparities in receptive field position and preferred orientation. The role of orientation disparities in binocular vision is reviewed.     

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.     

Pre-attentive segmentation and correspondence in stereo

Traditional stereo grouping models have focused on the problem of stereo correspondence between monocular inputs. Recent physiological data revealed that the disparity selective V2 cells increase their responses when (random-dot stereograms) stimuli within their receptive fields are at or near the boundary of a depth surface. Such highlights to depth (non-luminance) edges are seemingly not computationally required for the correspondence problem. Computationally, these highlights make the boundaries of a depth surface more salient, serving pre-attentive segmentation (between depth planes) and attracting visual attention. In special cases, they enable the psychophysically observed perceptual pop-out of a target from a background of visually identical distractors at a different depth. To achieve the highlights, mutual inhibition between disparity selective cells that are tuned to the same or similar depths is required. However, such mutual inhibition would impede the computation for the correspondence problem, which requires mutual excitation between the same cells. In this work, I introduce a computational model that, I believe, is the first to address both stereo correspondence and pre-attentive stereo segmentation. The computational mechanisms in the model are based on intracortical interactions in V2. I will demonstrate that the model captures the following physiological and psychophysical phenomena: (i) depth-edge highlighting; (ii) disparity capture; (iii) pop-out; and (iv) transparency.     

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.     

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.     

Bistable percepts in the brain: FMRI contrasts monocular pattern rivalry and binocular rivalry

The neural correlates of binocular rivalry have been actively debated in recent years, and are of considerable interest as they may shed light on mechanisms of conscious awareness. In a related phenomenon, monocular rivalry, a composite image is shown to both eyes. The subject experiences perceptual alternations in which the two stimulus components alternate in clarity or salience. The experience is similar to perceptual alternations in binocular rivalry, although the reduction in visibility of the suppressed component is greater for binocular rivalry, especially at higher stimulus contrasts. We used fMRI at 3T to image activity in visual cortex while subjects perceived either monocular or binocular rivalry, or a matched non-rivalrous control condition. The stimulus patterns were left/right oblique gratings with the luminance contrast set at 9%, 18% or 36%. Compared to a blank screen, both binocular and monocular rivalry showed a U-shaped function of activation as a function of stimulus contrast, i.e. higher activity for most areas at 9% and 36%. The sites of cortical activation for monocular rivalry included occipital pole (V1, V2, V3), ventral temporal, and superior parietal cortex. The additional areas for binocular rivalry included lateral occipital regions, as well as inferior parietal cortex close to the temporoparietal junction (TPJ). In particular, higher-tier areas MT+ and V3A were more active for binocular than monocular rivalry for all contrasts. In comparison, activation in V2 and V3 was reduced for binocular compared to monocular rivalry at the higher contrasts that evoked stronger binocular perceptual suppression, indicating that the effects of suppression are not limited to interocular suppression in V1.     

Extrapolating and interpolating surfaces in depth

Random-dot stereograms were generated with a blank area placed in part of the right-hand image so making a patchwork of monocular and binocular areas. The perceived depth and shape of the monocular region, where depth was not explicitly marked, depended in part on the depth and surface orientation of adjacent binocular areas. Thus a monocular rectangle flanked by two binocular rectangles which were placed in different fronto-parallel planes was seen as a sloping surface spanning the depth between the binocular regions, and, under some conditions, the gradient of a sloping binocular plane extended into a neighbouring monocular area. Division of the monocular region into two by textural discontinuities or discontinuities of motion sometimes altered the shape of the extrapolated surface. Often, though, the shape was unchanged by such discontinuities implying that both two- and three-dimensional features are used to segment a scene into separate surfaces. Pictorial cues also contribute to the shape and apparent depth of the monocular surface. For instance, when subjects viewed a display consisting of portions of a cube of which two ends were shown stereoscopically and one side monocularly, the monocular side was seen in three dimensions filling the gap between the ends. When stereo cues were pitted against pictorial cues, sometimes pictorial cues and sometimes stereo cues dominated, and sometimes the surface contained sharp discontinuities enabling both to be accommodated.     

A model of neural network extracting binocular parallax

A model of neural network extracting binocular parallax is proposed. It is a multilayered network composed of analog threshold elements. Three types of binocular neurons are included in this model. They are binocular simple neurons, binocular gate neurons and binocular depth neurons. The final layers of this model consist of elements which correspond to the binocular depth neurons. The performance of the model has been simulated on a digital computer. The results of the computer simulation show that every element of this model acts like neurons found in cat's and monkey's visual system and this model extracts binocular parallax caused by simple line components satisfactorily.     

Fucntional specialization and binocular interaction in the visual areas of rhesus monkey prestriate cortex.

If is is believed that neural mechanisms mediating stereoscopic vision may be localized in specific areas of the visual cortex, then it becomes necessary to be able to define these areas adequately. This is no easy matter in the rhesus monkey, an animal close to man, where the cytoarchitecturally uniform prestriate cortex is folded into deep sulci with secondary gyri. One way around this awkward problem is to use the callosal connections of the prestriate cortex as the anatomical landmarks. Callosal connections are restricted to regions at which the vertical meridian is represented. Since the visual fields, including the vertical meridian, are separately represented in each area, each has its own callosal connections. These are of great help in defining some of the boundaries of these areas, since the boundaries often coincide with the representation of the vertical meridian. With the visual areas thus defined anatomically, it becomes relatively easy to assign recordings to particular areas. Studies of binocular interactions in these areas reveal that most cells in all prestriate areas are binocularly driven. Hence, theoretically, all of the prestriate areas are candidates for stereoscopic mechanisms. The degree of binocular interaction varies from cell to cell. At the two extremes are cells which either respond to monocular stimulation only and are inhibited by binocular stimulation or ones which respond to binocular stimulation only. Changing, as opposed to fixed, disparity is signalled by two types of cells. In one category are cells activated in opposite directions for the two eyes. Such cells are always binocularly driven. In the other category are cells, some of which are monocularly activated, that are capable of responding to changing image size. In the monkey, both these categories of cells have so far been found in the motion area of the superior temporal sulcus only.     

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.     

Does a homogeneous population of elementary movement detectors activate the landing response of blowflies,Calliphora erythrocephala?

The landing response of tethered flying blowflies, Calliphora erythrocephala, was elicited by moving periodic gratings, and by stripes moving apart. The influence of binocular interactions on the landing response was investigated by comparing the responses of intact (“binocular”) animals to the response of flies which had one eye covered with black paint (“monocular” flies) effectively eliminating the input from this eye. Directions of motion eliciting a maximal response (preference direction) were determined in intact animals, and in “molecular” flies for different regions of the visual field. Preference directions determined in “monocular” flies follow the orientation of Z-axes (Fig. 4). Preference directions determined in intact animals and in “monocular” flies differ in the binocular eye region: in intact animals, the preference directions corresponds to vertical directions of motion; whereas the preference direction determined for the same area in “monocular” flies are inclined obliquely against the vertical plane. Sex-specific differences were found for the ventral binocular eye region in which the shift of preference directions is more pronounced in male than in female flies. The experimental data support the hypothesis that elementary movement detectors are aligned along the Z-axes of the eye, and that preference directions deviating from the orientation of elementary movement detectors are caused by binocular interactions.     

Single units and conscious vision.

Figures that can be seen in more than one way are invaluable tools for the study of the neural basis of visual awareness, because such stimuli permit the dissociation of the neural responses that underlie what we perceive at any given time from those forming the sensory representation of a visual pattern. To study the former type of responses, monkeys were subjected to binocular rivalry, and the response of neurons in a number of different visual areas was studied while the animals reported their alternating percepts by pulling levers. Perception-related modulations of neural activity were found to occur to different extents in different cortical visual areas. The cells that were affected by suppression were almost exclusively binocular, and their proportion was found to increase in the higher processing stages of the visual system. The strongest correlations between neural activity and perception were observed in the visual areas of the temporal lobe. A strikingly large number of neurons in the early visual areas remained active during the perceptual suppression of the stimulus, a finding suggesting that conscious visual perception might be mediated by only a subset of the cells exhibiting stimulus selective responses. These physiological findings, together with a number of recent psychophysical studies, offer a new explanation of the phenomenon of binocular rivalry. Indeed, rivalry has long been considered to be closely linked with binocular fusion and stereopsis, and the sequences of dominance and suppression have been viewed as the result of competition between the two monocular channels. The physiological data presented here are incompatible with this interpretation. Rather than reflecting interocular competition, the rivalry is most probably between the two different central neural representations generated by the dichoptically presented stimuli. The mechanisms of rivalry are probably the same as, or very similar to, those underlying multistable perception in general, and further physiological studies might reveal much about the neural mechanisms of our perceptual organization.     

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.     

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.     

Two-dimensional substructure of stereo and motion interactions in macaque visual cortex

The analysis of object motion and stereoscopic depth are important tasks that are begun at early stages of the primate visual system. Using sparse white noise, we mapped the receptive field substructure of motion and disparity interactions in neurons in V1 and MT of alert monkeys. Interactions in both regions revealed subunits similar in structure to V1 simple cells. For both motion and stereo, the scale and shape of the receptive field substructure could be predicted from conventional tuning for bars or dot-field stimuli, indicating that the small-scale interactions were repeated across the receptive fields. We also found neurons in V1 and in MT that were tuned to combinations of spatial and temporal binocular disparities, suggesting a possible neural substrate for the perceptual Pulfrich phenomenon. Our observations constrain computational and developmental models of motion-stereo integration.     

Leopard frog priorities in choosing between prey at different locations

Frogs are able to respond to a prey stimulus throughout their 360° ground-level visual field as well as in the superior visual field. We compared the likelihood of frogs choosing between a more nasally located, ground-level prey versus a more temporally located ground-level prey, when the prey at the nasal location is further away from the frog. Two crickets were presented simultaneously at 9 pairs of angles that included both crickets in the binocular visual field, both crickets in the monocular visual field, or one cricket in the binocular field and one in the monocular field. Frogs chose the more nasally located prey at least 71% of the time when the more temporal prey was in the monocular field; and 64% of the time when both prey were in the binocular field. Frogs tended to choose the more nasally located prey, even though it takes the frog longer to reach the prey. In addition, when given a choice between a prey located at ground level versus a prey located in the superior field, frogs tend to choose the prey at ground-level. These results suggest that there is a neural mechanism that biases frogs’ responses to prey stimuli.     

正常及异常双眼视觉的视动震颤（OKN）反应特性研究

为探讨不同双眼视觉状态下ＯＫＮ反应的特性,对正常人和不同类型的双眼视觉异常者的单眼鼻向及颞向ＯＫＮ反应进行了研究。实验发现：单眼视觉抑制者表现出鼻向与颞向ＯＫＮ反应不对称特性;两眼皆因视剥夺造成双眼视觉异常者主要以ＯＫＮ眼动增益降低为特点;双眼视觉正常者鼻、颞向ＯＫＮ反应是对称的。结果表明：单眼ＯＫＮ眼动反应的不对称特性及增益改变对探讨双眼视觉异常机制有重要意义,为视皮层双眼细胞异常导致单眼ＯＫＮ不对称的假设提供了支持性证据,并对弱视早期诊断及其分类有重要价值。     

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.