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.
We analyzed the relationship between eye movements and neuronal responses recorded from area MT in alert monkeys trained to maintain visual fixation during the presentation of moving patterns. The monkeys made small saccades which moved the eyes with velocities that spanned the sensitivity range of MT neurons. The saccades evoked changes in the neuronal response that depended upon (1) the level of stimulus-evoked activity amidst which the saccade occurred and (2) the direction of the saccade relative to the preferred direction of the neuron. Most notably, saccades were able to suppress stimulus-evoked activity when they caused retinal image flow that opposed the neuron's preference and were able to elicit a response or enhance weak activity when they caused flow in the neuron's preferred direction. On average, the disturbance lasted 40 ms beginning about 40 ms following saccade onset. Using these parameters, we simulated synthetic spike trains from an imaginary pair of similarly tuned neurons and determined that the interneuronal correlation due to saccades should be negligible at all but the lowest ongoing firing rates. This conclusion was supported from our data by the observation that response variance for single MT spike trains was not measurably reduced during periods of stable gaze compared to periods when eye movement exceeded a stability criterion (0.1 deg during 0.5 s). While the intrusions caused by saccades are too short-lived and infrequent to account for the variability of MT neuronal response (counter to the finding in V1 of Gur et al., 1997), the clear directional signal that they carry in area MT suggests that motion perception is not blocked during saccades by suppression at early stages in the visual pathway.
In humans, esotropia of early onset is associated with a profound asymmetry in smooth pursuit eye movements. When viewing is monocular, targets are tracked well only when they are moving nasally with respect to the viewing eye. To determine whether this pursuit abnormality reflects an anomaly in cortical visual motion processing, we recorded eye movements and cortical neural responses in nonamblyopic monkeys made strabismic by surgery at the age of 10-60 d. Eye movement recordings revealed the same asymmetry in the monkeys' pursuit eye movements as in humans with early-onset esotropia. With monocular viewing, pursuit was much stronger for nasalward motion than for temporalward motion, especially for targets presented in the nasal visual field. However, for targets presented during ongoing pursuit, temporalward and nasalward image motion was equally effective in modulating eye movement. Single-unit recordings made from the same monkeys, under anesthesia, revealed that MT neurons were rarely driven binocularly, but otherwise had normal response properties. Most were directionally selective, and their direction preferences were uniformly distributed. Our neurophysiological and oculomotor measurements both suggest that the pursuit defect in these monkeys is not due to altered cortical visual motion processing. Rather, the asymmetry in pursuit may be a consequence of imbalances in the two eyes' inputs to the "downstream" areas responsible for the initiation of pursuit.
Extrastriate cortical area MT is thought to process behaviorally important visual motion signals. Psychophysical studies suggest that visual motion signals may be analyzed by multiple mechanisms, a "first-order" one based on luminance, and a "second-order" one based upon higher level cues (e.g. contrast, flicker). Second-order motion is visible to human observers, but should be invisible to first-order motion sensors. To learn if area MT is involved in the analysis of second-order motion, we measured responses to first-and second-order gratings of single neurons in area MT (and in one experiment, in area V1) in anesthetized, paralyzed macaque monkeys. For each neuron, we measured directional and spatio-temporal tuning with conventional first-order gratings and with second-order gratings created by spatial modulation of the flicker rate of a random texture. A minority of MT and V1 neurons exhibited significant selectivity for direction or orientation of second-order gratings. In nearly all cells, response to second-order motion was weaker than response to first-order motion. MT cells with significant selectivity for second-order motion tended to be more responsive and more sensitive to luminance contrast, but were in other respects similar to the remaining MT neurons; they did not appear to represent a distinct subpopulation. For those cells selective for second-order motion, we found a correlation between the preferred directions of first-and second-order motion, and weak correlations in preferred spatial frequency. These cells preferred lower temporal frequencies for second-order motion than for first-order motion. A small proportion of MT cells seemed to remain selective and responsive for second-order motion. None of our small sample of V1 cells did. Cells in this small population, but not others, may perform "form-cue invariant" motion processing (Albright, 1992).
O'Keefe, Lawrence P., Jonathan B. Levitt, Daniel C. Kiper, Robert M. Shapley, and J. Anthony Movshon. Functional organization of owl monkey lateral geniculate nucleus and visual cortex. J. Neurophysiol. 80: 594–609, 1998. The nocturnal, New World owl monkey ( Aotus trivirgatus) has a rod-dominated retina containing only a single cone type, supporting only the most rudimentary color vision. However, it does have well-developed magnocellular (M) and parvocellular (P) retinostriate pathways and striate cortical architecture [as defined by the pattern of staining for the activity-dependent marker cytochrome oxidase (CO)] similar to that seen in diurnal primates. We recorded from single neurons in anesthetized, paralyzed owl monkeys using drifting, luminance-modulated sinusoidal gratings, comparing receptive field properties of M and P neurons in the lateral geniculate nucleus and in V1 neurons assigned to CO “blob,” “edge,” and “interblob” regions and across layers. Tested with achromatic stimuli, the receptive field properties of M and P neurons resembled those reported for other primates. The contrast sensitivity of P cells in the owl monkey was similar to that of P cells in the macaque, but the contrast sensitivities of M cells in the owl monkey were markedly lower than those in the macaque. We found no differences in eye dominance, orientation, or spatial frequency tuning, temporal frequency tuning, or contrast response for V1 neurons assigned to different CO compartments; we did find fewer direction-selective cells in blobs than in other compartments. We noticed laminar differences in some receptive field properties. Cells in the supragranular layers preferred higher spatial and lower temporal frequencies and had lower contrast sensitivity than did cells in the granular and infragranular layers. Our data suggest that the receptive field properties across functional compartments in V1 are quite homogeneous, inconsistent with the notion that CO blobs anatomically segregate signals from different functional “streams.”
Schematic and reduced eyes for the horse, cow, sheep, pig, cat, and dog were calculated from optical parameter values obtained from the literature. Calculations were performed with the aid of a computer program using standard Gaussian equations and a homogeneous lens model. Calculated schematic eyes had refractive states within 1.5 D of emmetropia; retinal image sizes ranged from 0.22 (cat and dog) to 0.44 (horse) mm/deg. The cat eye presented here is more nearly emmetropic than a previously published cat schematic eye. The effects of altering refractive indices, radii of curvature, and thicknesses of refractive components upon refractive state and retinal image size are examined. The largest differences resulted from changes made to refractive indices of the lens and vitreous and to the vitreous depth.
We tested the hypothesis that neurons in the primary visual cortex (V1) adapt selectively to contingencies in the attributes of visual stimuli. We recorded from single neurons in macaque V1 and measured the e¡ects of adaptation either to the sum of two gratings (compound stimulus) or to the individual gratings. According to our hypothesis, there would be a component of adaptation that is speci¢c to the compound stimulus. In a ¢rst series of experiments, the two gratings di¡ered in orientation. One grating had optimal orientation and the other was orthogonal to it, and therefore did not activate the neuron under study. These experiments provided evidence in favour of our hypothesis. In most cells adaptation to the compound stimulus reduced responses to the compound stimulus more than it reduced responses to the optimal grating, and the responses to the compound stimulus were reduced more by adaptation to the compound stimulus than by adaptation to the individual gratings. This suggests that a component of adaptation was speci¢c to (and caused by) the simultaneous presence of the two orientations in the compound stimulus. To test whether V1 neurons could adapt to other contingencies in the stimulus attributes, we performed a second series of experiments, in which the component gratings were parallel but di¡ered in spatial frequency, and were both e¡ective in activating the neuron under study. These experiments failed to reveal convincing contingent e¡ects of adaptation, suggesting that neurons cannot adapt equally well to all types of contingency.
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