Experimental strabismus in the kitten

1. We have examined the relative roles of visual and nonvisual input to striate cortex cells in causing the breakdown of binocularity produced by brief periods of visual-axis misalignment in kittens. 2. In the first study, the binocularity of single neurons recorded from the striate cortex was assessed in kittens reared with either surgical or optical strabismus. Surgical strabismus was induced by performing a unilateral medial rectus tenotomy, and optical strabismus by means of goggles that held prisms of equal power before the two eyes with their bases oriented in opposite directions. The loss of functional binocular connections was of comparable severity in these two groups of kittens. Control kittens, reared wearing goggles containing prisms whose bases were oriented in the same direction, showed normal levels of binocularity. 3. In the second experiment, normal kittens were given a surgical strabismus at around 1 mo of age and kept in total darkness for 2 days, 2 wk, or 4 wk. Cortical binocularity was normal in these kittens. 4. Finally, a group of kittens was reared in the illuminated colony with a symmetric surgical strabismus (bilateral medial rectus tenotomy). These kittens suffered a severe loss in cortical binocularity that was comparable to that seen in control kittens reared with asymmetric (unilateral) strabismus. 5. We conclude that altered visual input caused by misregister of the images falling in the two eyes is necessary and almost certainly sufficient to cause breakdown of cortical binocularity in kittens exposed to brief periods of divergent strabismus and that, when strabismus is induced surgically, this loss of binocularity is not dependent on the symmetry of the surgical manipulation; we thus find no evidence for a special role of afferents from the extraocular muscles in producing this effect.     

Capture of the visual direction of monocular objects by adjacent binocular objects

Investigations of binocular visual direction have concentrated mainly on stationary objects. Eye positions were generally not measured and binocular fixation was assumed to be perfect. During the viewing of stationary objects, vergence errors are not negligible but small. During the viewing of moving objects, however, errors in binocular fixation are much larger. Existing rules for binocular visual direction were examined under the latter, more demanding viewing conditions. Eye movements were measured objectively by the scleral coil technique. Subjects viewed a large stereogram in which the half-images oscillated in counterphase. The stereogram contained two square random-dot patterns placed side by side with a gap in between. A vertical line, visible only to one eye, oscillated in the gap. Subjects were asked to adjust the amplitude of line motion until the line was perceived to be stationary. In so doing, they set amplitudes equal to the amplitudes of half-image motion if the gap between the patterns was narrow. They set amplitudes significantly smaller in wider gaps. Subjects made considerable fixational errors in following the oscillations of the line and the random-dot patterns. The results of the settings and of the retinal errors together refute existing rules for binocular visual direction of monocular objects. Perceived directions of monocular objects cannot be specified by geometrical rules that include only the positions of the objects and of the two eyes. The results suggest that perceived directions of monocular objects are captured by the binocular visual directions of adjacent binocular objects. Capture of binocular visual direction was found to be effective for gaps as wide as 8 deg between the binocular objects. The phenomenon of binocular capture has negative consequences for the general use of nonius lines as indicators of eye position.     

Binocular visual surface perception

Binocular disparity, the differential angular separation between pairs of image points in the two eyes, is the well-recognized basis for binocular distance perception. Without denying disparity's role in perceiving depth, we describe two perceptual phenomena, which indicate that a wider view of binocular vision is warranted. First, we show that disparity can play a critical role in two-dimensional perception by determining whether separate image fragments should be grouped as part of a single surface or segregated as parts of separate surfaces. Second, we show that stereoscopic vision is not limited to the registration and interpretation of binocular disparity but that it relies on half-occluded points, visible to one eye and not the other, to determine the layout and transparency of surfaces. Because these half-visible points are coded by neurons carrying eye-of-origin information, we suggest that the perception of these surface properties depends on neural activity available at visual cortical area V1.     

Synchronizing retinal activity in both eyes disrupts binocular map development in the optic tectum

Spatiotemporal correlations in the pattern of spontaneous and evoked retinal ganglion cell (RGC) activity are believed to influence the topographic organization of connections throughout the developing visual system. We have tested this hypothesis by examining the effects of interfering with these potential activity cues during development on the functional organization of binocular maps in the Xenopus frog optic tectum. Paired recordings combined with cross-correlation analyses demonstrated that exposing normal frogs to a continuous 1 Hz of stroboscopic illumination synchronized the firing of all three classes of RGC projecting to the tectum and induced similar patterns of temporally correlated activity across both lobes of the nucleus. Embryonic and eye-rotated larval animals were reared until early adulthood under equivalent stroboscopic conditions. The maps formed by each RGC class in the contralateral tectum showed normal topography and stratification after strobe rearing, but with consistently enlarged multiunit receptive fields. Maps of the ipsilateral eye, formed by crossed isthmotectal axons, showed significant disorder and misalignment with direct visual input from the retina, and in the eye-rotated animals complete compensatory reorientation of these maps usually induced by this procedure failed to occur. These findings suggest that refinement of retinal arbors in the tectum and the ability of crossed isthmotectal arbors to establish binocular convergence with these retinal afferents are disrupted when they all fire together. Our data thus provide direct experimental evidence that spatiotemporal activity patterns within and between the two eyes regulate the precision of their developing connections.     

Neuronal correlates of amblyopia in the visual cortex of macaque monkeys with experimental strabismus and anisometropia

Amblyopia is a developmental disorder of pattern vision. After surgical creation of esotropic strabismus in the first weeks of life or after wearing -10 diopter contact lenses in one eye to simulate anisometropia during the first months of life, macaques often develop amblyopia. We studied the response properties of visual cortex neurons in six amblyopic macaques; three monkeys were anisometropic, and three were strabismic. In all monkeys, cortical binocularity was reduced. In anisometropes, the amblyopic eye influenced a relatively small proportion of cortical neurons; in strabismics, the influence of the two eyes was more nearly equal. The severity of amblyopia was related to the relative strength of the input of the amblyopic eye to the cortex only for the more seriously affected amblyopes. Measurements of the spatial frequency tuning and contrast sensitivity of cortical neurons showed few differences between the eyes for the three less severe amblyopes (two strabismic and one anisometropic). In the three more severely affected animals (one strabismic and two anisometropic), the optimal spatial frequency and spatial resolution of cortical neurons driven by the amblyopic eye were substantially and significantly lower than for neurons driven by the nonamblyopic eye. There were no reliable differences in neuronal contrast sensitivity between the eyes. A sample of neurons recorded from cortex representing the peripheral visual field showed no interocular differences, suggesting that the effects of amblyopia were more pronounced in portions of the cortex subserving foveal vision. Qualitatively, abnormalities in both the eye dominance and spatial properties of visual cortex neurons were related on a case-by-case basis to the depth of amblyopia. Quantitative analysis suggests, however, that these abnormalities alone do not explain the full range of visual deficits in amblyopia. Studies of extrastriate cortical areas may uncover further abnormalities that explain these deficits.     

Physiological side differences in fluorescein inflow into eyes (author's transl)

Rapid-sequence fluorescein angiography was performed simultaneously on both eyes of each of 241 healthy individuals in order to analyze and compare dye inflow patterns in the eyes of each subject. The many uncertainties of successive investigations resulting from uncontrollable changes in systemic circulation parameters between the two angiographies were thus eliminated. In over 90% of the subjects there was no difference between the two eyes in respect of arm-fundus time nor the filling pattern of choroid and retina. A few subjects did show a difference, but this lasted for only one exposure and was normalized on the second exposure of the sequence. It is therefore concluded that side differences of the fluorescein inflow pattern on simultaneous bilateral angiography are highly suggestive of unilateral circulatory disturbances in the cervical, retinal or choroidal vessel. Such disturbances are practically certain if side differences last for more than just one exposure. In contrast, the filling pattern in cilioretinal arteries was very variable and over one third of the subjects presented with side differences between their two eyes. Therefore the filling pattern of these vessels cannot be used for diagnostic purposes.     

A neural model of contour integration in the primary visual cortex

Experimental observations suggest that contour integration may take place in V1. However, there has yet to be a model of contour integration that uses only known V1 elements, operations, and connection patterns. This article introduces such a model, using orientation selective cells, local cortical circuits, and horizontal intracortical connections. The model is composed of recurrently connected excitatory neurons and inhibitory interneurons, receiving visual input via oriented receptive fields resembling those found in primary visual cortex. Intracortical interactions modify initial activity patterns from input, selectively amplifying the activities of edges that form smooth contours in the image. The neural activities produced by such interactions are oscillatory and edge segments within a contour oscillate in synchrony. It is shown analytically and empirically that the extent of contour enhancement and neural synchrony increases with the smoothness, length, and closure of contours, as observed in experiments on some of these phenomena. In addition, the model incorporates a feedback mechanism that allows higher visual centers selectively to enhance or suppress sensitivities to given contours, effectively segmenting one from another. The model makes the testable prediction that the horizontal cortical connections are more likely to target excitatory (or inhibitory) cells when the two linked cells have their preferred orientation aligned with (or orthogonal to) their relative receptive field center displacements.     

Binocular brightness: A suppression-summation trade off.

Two experiments with 22 undergraduates estimated binocular brightness of targets of large visual extent. On each trial one eye was presented with a fairly intense luminance of 800 cd/m–2, and the other eye with 1 of 12 luminances ranging from zero to 800 cd/m–2. Exp 1, using ganzfeld stimuli, produced a large amount of binocular brightness summation and very little Fechner's paradox, a decrease in binocular brightness that occurs when the luminances to the 2 eyes differ greatly. Exp 2, using a smaller target with very low spatial frequencies, produced greater Fechner's paradox than the ganzfelder, but more binocular summation and less Fechner's paradox than what is usually reported for small targets with abrupt contours. Results suggest a trade-off between suppressive and summative mechanisms involving binocular cells that are spatially tuned. (French abstract) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Visual fields in Short-toed Eagles, Circaetus gallicus (Accipitridae), and the function of binocularity in birds

Visual fields were determined in alert restrained birds using an ophthalmoscopic reflex technique. The region of binocular overlap is relatively small: maximum width of 20 degrees occurs approximately 15 degrees below the horizontal, and the field extends vertically through 80 degrees with the bill tip placed close to the centre. Monocular field width in the horizontal plane is 139 degrees, and the field is asymmetric about the optic axis. The cyclopean field extends through 260 degrees, and the blind area above and behind the head reaches maximum width of 100 degrees close to the horizontal. At the frontal margins of the monocular field the retinal and optical fields do not coincide; the retinal field margin lies approximately 10 degrees inside the optical margin. This gives rise to an apparent binocular field that is twice the width of the functional binocular field. Interspecific comparisons show that the binocular field of Short-toed Eagles is similar in shape and size to those of bird species that differ markedly in phylogeny, ecology, foraging technique, and eye size. This suggests that these relatively narrow binocular fields are a convergent feature of birds whose foraging is guided by visual cues irrespective of whether items are taken directly in the bill or in the feet, as in eagles, and irrespective of the size and shape of the monocular and cyclopean visual fields. It is argued that binocular vision in birds results from the requirement for each monocular field to extend contralaterally to embody a portion of the optical flow field which is radially symmetrical about the direction of travel. This is in contrast to functional explanations of binocularity, such as those concerned with stereopsis, which present it as a means of extracting higher order information through the combination of two monocular images of the same portion of a scene.     

Diagnostic approach in coronary disease in developing countries

The fellow eyes of 13 cases of retinal detachment associated with atopic dermatitis were examined by binocular ophthalmoscope combined with scleral indentation. Breaks in the retina or pars plana were detected in 8 cases (62%); 2 of these eyes had asymptomatic retinal detachment. The most common location of these breaks was at the vitreous base symmetrical to the affected, symptomatic eye. Meticulous ophthalmoscopic examination is recommended for both eyes in patients with retinal detachment associated with atopic dermatitis.     

Binocular matching of dissimilar features in phantom stereopsis

Previously we have demonstrated that quantitative depth perception can be elicited from a stereogram that lacks contrast defined binocular corresponding elements (phantom stereopsis). In this report, we use computer simulation to demonstrate that it is biologically plausible for some known binocular cortical cell types to combine non-conventional matching features. Therefore, binocular matching processes based on the responses of these cells could be a conventional one, namely, looking for similar response patterns in the two eyes. While at cell types we simulated gave identical disparity outputs to the conventional stereogram, they responded differently to the phantom stereogram. Processes other than low-level disparity detectors may have to be invoked in order to achieve a unique depth solution.     

Human oculomotor system accounts for 3-D eye orientation in the visual-motor transformation for saccades

A recent theoretical investigation has demonstrated that three-dimensional (3-D) eye position dependencies in the geometry of retinal stimulation must be accounted for neurally (i.e., in a visuomotor reference frame transformation) if saccades are to be both accurate and obey Listing's law from all initial eye positions. Our goal was to determine whether the human saccade generator correctly implements this eye-to-head reference frame transformation (RFT), or if it approximates this function with a visuomotor look-up table (LT). Six head-fixed subjects participated in three experiments in complete darkness. We recorded 60 degrees horizontal saccades between five parallel pairs of lights, over a vertical range of +/-40 degrees (experiment 1), and 30 degrees radial saccades from a central target, with the head upright or tilted 45 degrees clockwise/counterclockwise to induce torsional ocular counterroll, under both binocular and monocular viewing conditions (experiments 2 and 3). 3-D eye orientation and oculocentric target direction (i.e., retinal error) were computed from search coil signals in the right eye. Experiment 1: as predicted, retinal error was a nontrivial function of both target displacement in space and 3-D eye orientation (e.g., horizontally displaced targets could induce horizontal or oblique retinal errors, depending on eye position). These data were input to a 3-D visuomotor LT model, which implemented Listing's law, but predicted position-dependent errors in final gaze direction of up to 19.8 degrees. Actual saccades obeyed Listing's law but did not show the predicted pattern of inaccuracies in final gaze direction, i.e., the slope of actual error, as a function of predicted error, was only -0. 01 +/- 0.14 (compared with 0 for RFT model and 1.0 for LT model), suggesting near-perfect compensation for eye position. Experiments 2 and 3: actual directional errors from initial torsional eye positions were only a fraction of those predicted by the LT model (e. g., 32% for clockwise and 33% for counterclockwise counterroll during binocular viewing). Furthermore, any residual errors were immediately reduced when visual feedback was provided during saccades. Thus, other than sporadic miscalibrations for torsion, saccades were accurate from all 3-D eye positions. We conclude that 1) the hypothesis of a visuomotor look-up table for saccades fails to account even for saccades made directly toward visual targets, but rather, 2) the oculomotor system takes 3-D eye orientation into account in a visuomotor reference frame transformation. This transformation is probably implemented physiologically between retinotopically organized saccade centers (in cortex and superior colliculus) and the brain stem burst generator.     

Binocular rivalry: ambiguities resolved

For over 100 years, binocular rivalry was seen as the result of competition between the two eyes, involving reciprocal suppression of retinal inputs. Now it emerges that rivalry reflects alternating perceptual interpretations that are represented in the firing patterns of cells in the temporal visual cortex.     

On the binocular summation of chromatic contrast

The binocular summation of chromatic contrast was investigated under a variety of stimulus conditions. Binocular and monocular contrast detection thresholds were measured using 0.5 cpd Gabor patches. It was found that, using stimuli which contained combinations of chromatic and luminance contrast, binocular detection could take place independently in luminance-contrast- and chromatic-contrast-sensitive mechanisms. It was also found that, with chromatic stimuli, levels of binocular summation were above those expected from probability summation between the eyes, and thus showed evidence for binocular neural summation within chromatic detection mechanisms. The implications of these results for (a) the binocularity of chromatic detection mechanisms, and (b) the suggested link between stereopsis and binocular neural summation, are discussed.     

The aftereffect of tracking eye movements

When observers tracked moving stripes across a background either of stationary stripes, or of stripes moving in the opposite direction, they saw a clear motion aftereffect when the stripes stopped moving. The direction of this aftereffect was opposite to that of the previously tracked stripes, and was thus the same as the direction of the retinal movement of the non-tracked stripes. This aftereffect of tracking was shown not to depend upon slippage of the tracked contours on the retina during tracking, or upon the saccadic phase of optokinetic nystagmus. The effect showed storage over a period of time with the eyes shut. It appears that the effect is due to induced movement, and arises originally from stimulation of the retina by background contours in the tracking phase. This was shown by confining the view of the moving target to one eye, while permitting both eyes to be exposed to background stimulation during tracking. After such stimulation the magnitude of the aftereffect was equal in the two eyes.     

Correlation-based development of ocularly matched orientation and ocular dominance maps: determination of required input activities

We extend previous models for separate development of ocular dominance and orientation selectivity in cortical layer 4 by exploring conditions permitting combined organization of both properties. These conditions are expressed in terms of functions describing the degree of correlation in the firing of two inputs from the lateral geniculate nucleus (LGN), as a function of their retinotopic separation and their "type" (ON center or OFF center and left eye or right eye). The development of ocular dominance requires that the correlations of an input with other inputs of the same eye be stronger than or equal to its correlations with inputs of the opposite eye and strictly stronger at small retinotopic separations. This must be true after summing correlations with inputs of both center types. The development of orientation-selective simple cells requires that (1) an input's correlations with other inputs of the same center type be stronger than its correlations with inputs of the opposite center type at small retinotopic separation; and (2) this relationship reverse at larger retinotopic separations within an arbor radius (the radius over which LGN cells can project to a common cortical point). This must be true after summing correlations with inputs serving both eyes. For orientations to become matched in the two eyes, correlated activity within the receptive fields must be maximized by specific between-eye alignments of ON and OFF subregions. Thus the correlations between the eyes must differ depending on center type, and this difference must vary with retinotopic separation within an arbor radius. These principles are satisfied by a wide class of correlation functions. Combined development of ocularly matched orientation maps and ocular dominance maps can be achieved either simultaneously or sequentially. In the latter case, the model can produce a correlation between the locations of orientation map singularities and local ocular dominance peaks similar to that observed physiologically. The model's main prediction is that the above correlations should exist among inputs to cortical layer 4 simple cells before vision. In addition, mature simple cells are predicted to have certain relationships between the locations of the ON and OFF subregions of the left and right eyes' receptive fields.     

Interocular inhibition in strabismus

Many patients with acquired strabismus do not suffer from diplopia and confusion after an individually and age-dependent interval. They inhibit the image of the deviated eye by binocular rilvary and particularly by the physiological ability to disregard visually disturbing stimuli. In strabismus with early onset, binocular rivalry is also demonstrable, even for stimuli that do not normally lead to suppression. On the basis of anomalous retinal correspondence, this rivalry occurs between retinal points onto which the same object projects. The retinal area with the lesser eccentricity receives the dominance. The fovea of the deviated eye is therefore not suppressed. In small-angle strabismus with smaller functional differences between anomalous corresponding retinal points anomalous fusion and even stereopsis can be possible as long as strong suprathreshold stimuli are presented. Strabismic amblyopia as a consequence of interfoveal suppression can only develop before anomalous retinal correspondence dominates in the strabismic child.     

Why do patterned afterimages fluctuate in visibility?

Considers that prolonged patterned afterimages go through sequences of complete and partial visibility and disappearances before they finally cease to be visible. A survey was made of studies examining these fluctuations and the factors involved in them. Theories attributing the fluctuations to retinal processes, eye movements, adaptation of feature detectors, binocular rivalry, binocular interaction, and attention are assessed. It is tentatively concluded that the fluctuations of patterned afterimages are due to two cortical processes: One concerns the interaction between contour detectors, and the other involves inhibition between columns for ocular dominance or disparity detection. (97 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Neural mechanisms underlying stereoscopic vision

The progressive frontalization of both eyes in mammals causes overlap of the left and right visual fields, having as a consequence a region of binocular field with single vision and stereopsis. The horizontal separation of the eyes makes the retinal images of the objects lying in this binocular field have slight horizontal and vertical differences, termed disparities. Horizontal disparities are the main cue for stereopsis. In the past decades numerous physiological studies made on monkeys, which have in many aspects a similar visual system to humans, showed that a population of visual cells are capable of encoding the amplitude and sign of horizontal disparity. Such disparity detectors were found in cortical visual areas V1, V2, V3, V3A, VP, MT (V5) and MST of monkeys and in the superior colliculus of the cat and opossum. According to their disparity tuning function, these cells were first grouped into tuned excitatory, tuned inhibitory, near and far sub-groups. Subsequent studies added two more categories, tuned near and tuned far cells. Asymmetries between left and right receptive field position, on and off regions, and intra-receptive field wiring are believed to be the neural mechanisms of disparity detection. Because horizontal disparity alone is insufficient to compute reliable stereopsis, additional information about fixation distance and angle of gaze is required. Thus, while there is unequivocal evidence of cells capable of detecting horizontal disparities, it is not known how horizontal disparity is calibrated. Sensitivity to vertical disparity and information about the vergence angle or eye position may be the source of this additional information.