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A Anzai

Publications and source records attributed to A Anzai.

11 recordsLinked to original sources

Joint-encoding of motion and depth by visual cortical neurons: neural basis of the Pulfrich effect.

Motion and stereoscopic depth are fundamental parameters of the structural analysis of visual scenes. Because they are defined by a difference in object position, either over time or across the eyes, a common neural machinery may be used for encoding these attributes. To examine this idea, we analyzed responses of binocular complex cells in the cat striate cortex to stimuli of various intra- and interocular spatial and temporal shifts. We found that most neurons exhibit space-time-oriented response profiles in both monocular and binocular domains. This indicates that these neurons encode motion and depth jointly, and it explains phenomena such as the Pulfrich effect. We also found that the relationship between neuronal tuning of motion and depth conforms to that predicted by the use of motion parallax as a depth cue. These results demonstrate a joint-encoding of motion and depth at an early cortical stage.

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Neural mechanisms for encoding binocular disparity: receptive field position versus phase.

The visual system uses binocular disparity to discriminate the relative depth of objects in space. Because the striate cortex is the first site along the central visual pathways at which signals from the left and right eyes converge onto a single neuron, encoding of binocular disparity is thought to begin in this region. There are two possible mechanisms for encoding binocular disparity through simple cells in the striate cortex: a difference in receptive field (RF) position between the two eyes (RF position disparity) and a difference in RF profiles between the two eyes (RF phase disparity). Although there is evidence that supports each of these schemes, both mechanisms have not been examined in a single study to determine their relative roles. In this study, we have measured RF position and phase disparities of individual simple cells in the cat's striate cortex to address this issue. Using a sophisticated RF mapping technique that employs binary m-sequences, we have obtained left and right eye RF profiles of two or more cells recorded simultaneously. A version of the reference-cell method was used to estimate RF position disparity. We find that RF position disparities generally are limited to values that are not sufficient to encode large binocular disparities. In contrast, RF phase disparities cover a wide range of binocular disparities and exhibit dependencies on RF orientation and spatial frequency in a manner expected for a mechanism that encodes binocular disparity. These results suggest that binocular disparity is encoded mainly through RF phase disparity. However, RF position disparity may play a significant role for cells with high spatial frequency selectivity that are constrained to have only small RF phase disparities.

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Neural mechanisms for processing binocular information I. Simple cells.

The visual system integrates information from the left and right eyes and constructs a visual world that is perceived as single and three dimensional. To understand neural mechanisms underlying this process, it is important to learn about how signals from the two eyes interact at the level of single neurons. Using a sophisticated receptive field (RF) mapping technique that employs binary m-sequences, we have determined the rules of binocular interactions exhibited by simple cells in the cat's striate cortex in relation to the structure of their monocular RFs. We find that binocular interaction RFs of most simple cells are well described as the product of left and right eye RFs. Therefore the binocular interactions depend not only on binocular disparity but also on monocular stimulus position or phase. The binocular interaction RF is consistent with that predicted by a model of a linear binocular filter followed by a static nonlinearity. The static nonlinearity is shown to have a shape of a half-power function with an average exponent of approximately 2. Although the initial binocular convergence of signals is linear, the static nonlinearity makes binocular interaction multiplicative at the output of simple cells. This multiplicative binocular interaction is a key ingredient for the computation of interocular cross-correlation, an algorithm for solving the stereo correspondence problem. Therefore simple cells may perform initial computations necessary to solve this problem.

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Neural mechanisms for processing binocular information II. Complex cells.

Complex cells in the striate cortex exhibit extensive spatiotemporal nonlinearities, presumably due to a convergence of various subunits. Because these subunits essentially determine many aspects of a complex cell receptive field (RF), such as tuning for orientation, spatial frequency, and binocular disparity, examination of the RF properties of subunits is important for understanding functional roles of complex cells. Although monocular aspects of these subunits have been studied, little is known about their binocular properties. Using a sophisticated RF mapping technique that employs binary m-sequences, we have examined binocular interactions exhibited by complex cells in the cat's striate cortex and the binocular RF properties of their underlying functional subunits. We find that binocular interaction RFs of complex cells exhibit subregions that are elongated along the frontoparallel axis at different binocular disparities. Therefore responses of complex cells are largely independent of monocular stimulus position or phase as long as the binocular disparity of the stimulus is kept constant. The binocular interaction RF is well described by a sum of binocular interaction RFs of underlying functional subunits, which exhibit simple cell-like RFs and a preference for different monocular phases but the same binocular disparity. For more than half of the complex cells examined, subunits of each cell are consistent with the characteristics specified by an energy model, with respect to the number of subunits as well as relationships between the subunit properties. Subunits exhibit RF binocular disparities that are largely consistent with a phase mechanism for encoding binocular disparity. These results indicate that binocular interactions of complex cells are derived from simple cell-like subunits, which exhibit multiplicative binocular interactions. Therefore binocular interactions of complex cells are also multiplicative. This suggests that complex cells compute something analogous to an interocular cross-correlation of images for a local region of visual space. The result of this computation can be used for solving the stereo correspondence problem.

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Neural mechanisms underlying binocular fusion and stereopsis: position vs. phase.

The visual system utilizes binocular disparity to discriminate the relative depth of objects in space. Since the striate cortex is the first site along the central visual pathways at which signals from the left and right eyes converge onto a single neuron, encoding of binocular disparity is thought to begin in this region. There are two possible mechanisms for encoding binocular disparity through simple cells in the striate cortex: a difference in receptive field (RF) position between the two eyes (RF position disparity) and a difference in RF profile between the two eyes (RF phase disparity). Although there have been studies supporting each of the two encoding mechanisms, both mechanisms have not been examined in a single study. Therefore, the relative roles of the two mechanisms have not been determined. To address this issue, we have mapped left and right eye RFs of simple cells in the cat's striate cortex using binary m-sequence noise, and then we have estimated RF position and phase disparities. We find that RF position disparities are generally limited to small values that are not sufficient to encode large binocular disparities. In contrast, RF phase disparities cover a wide range of binocular disparities and exhibit dependencies on orientation and spatial frequency in a manner expected for a mechanism that encodes binocular disparity. These results indicate that binocular disparity is mainly encoded through RF phase disparity. However, RF position disparity may play a significant role for cells with high spatial frequency selectivity, which are constrained to small RF phase disparities.

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Receptive field structure in the visual cortex: does selective stimulation induce plasticity?

Sensory areas of adult cerebral cortex can reorganize in response to long-term alterations in patterns of afferent signals. This long-term plasticity is thought to play a crucial role in recovery from injury and in some forms of learning. However, the degree to which sensory representations in primary cortical areas depend on short-term (i.e., minute to minute) stimulus variations remains unclear. A traditional view is that each neuron in the mature cortex has a fixed receptive field structure. An alternative view, with fundamentally different implications for understanding cortical function, is that each cell's receptive field is highly malleable, changing according to the recent history of the sensory environment. Consistent with the latter view, it has been reported that selective stimulation of regions surrounding the receptive field induces a dramatic short-term increase in receptive field size for neurons in the visual cortex [Pettet, M. W. & Gilbert, C. D. (1992) Proc. Natl. Acad. Sci. USA 89, 8366-8370]. In contrast, we report here that there is no change in either the size or the internal structure of the receptive field following several minutes of surround stimulation. However, for some cells, overall responsiveness increases. These results suggest that dynamic alterations of receptive field structure do not underlie short-term plasticity in the mature primary visual cortex. However, some degree of short-term adaptability could be mediated by changes in responsiveness.

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Contrast coding by cells in the cat's striate cortex: monocular vs. binocular detection.

Many psychophysical studies of various visual tasks show that performance is generally better for binocular than for monocular observation. To investigate the physiological basis of this binocular advantage, we have recorded, under monocular and binocular stimulation, contrast response functions for single cells in the striate cortex of anesthetized and paralyzed cats. We applied receiver operating characteristic analysis to our data to obtain monocular and binocular neurometric functions for each cell. A contrast threshold and a slope were extracted from each neurometric function and were compared for monocular and binocular stimulation. We found that contrast thresholds and slopes varied from cell to cell but, in general, binocular contrast thresholds were lower, and binocular slopes were steeper, than their monocular counterparts. The binocular advantage ratio, the ratio of monocular to binocular thresholds for individual cells, was, on average, slightly higher than the typical ratios reported in human psychophysics. No single rule appeared to account for the various degrees of binocular summation seen in individual cells. We also found that the proportion of cells likely to contribute to contrast detection increased with stimulus contrast. Less contrast was required under binocular than under monocular stimulation to obtain the same proportion of cells that contribute to contrast detection. Based on these results, we suggest that behavioral contrast detection is carried out by a small proportion of cells that are relatively sensitive to near-threshold contrasts. Contrast sensitivity functions (CSFs) for the cell population, estimated from this hypothesis, agree well with behavioral data in both the shape of the CSF and the ratio of binocular to monocular sensitivities. We conclude that binocular summation in behavioral contrast detection may be attributed to the binocular superiority in contrast sensitivity of a small proportion of cells which are responsible for threshold contrast detection.

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Linear and nonlinear contributions to orientation tuning of simple cells in the cat's striate cortex.

Orientation selectivity is one of the most conspicuous receptive-field (RF) properties that distinguishes neurons in the striate cortex from those in the lateral geniculate nucleus (LGN). It has been suggested that orientation selectivity arises from an elongated array of feedforward LGN inputs (Hubel & Wiesel, 1962). Others have argued that cortical mechanisms underlie orientation selectivity (e.g. Sillito, 1975; Somers et al., 1995). However, isolation of each mechanism is experimentally difficult and no single study has analyzed both processes simultaneously to address their relative roles. An alternative approach, which we have employed in this study, is to examine the relative contributions of linear and nonlinear mechanisms in sharpening orientation tuning. Since the input stage of simple cells is remarkably linear, the nonlinear contribution can be attributed solely to cortical factors. Therefore, if the nonlinear component is substantial compared to the linear contribution, it can be concluded that cortical factors play a prominent role in sharpening orientation tuning. To obtain the linear contribution, we first measure RF profiles of simple cells in the cat's striate cortex using a binary m-sequence noise stimulus. Then, based on linear spatial summation of the RF profile, we obtain a predicted orientation-tuning curve, which represents the linear contribution. The nonlinear contribution is estimated as the difference between the predicted tuning curve and that measured with drifting sinusoidal gratings. We find that measured tuning curves are generally more sharply tuned for orientation than predicted curves, which indicates that the linear mechanism is not enough to account for the sharpness of orientation-tuning. Therefore, cortical factors must play an important role in sharpening orientation tuning of simple cells. We also examine the relationship of RF shape (subregion aspect ratio) and size (subregion length and width) to orientation-tuning halfwidth. As expected, predicted tuning halfwidths are found to depend strongly on both subregion length and subregion aspect ratio. However, we find that measured tuning halfwidths show only a weak correlation with subregion aspect ratio, and no significant correlation with RF length and width. These results suggest that cortical mechanisms not only serve to sharpen orientation tuning, but also serve to make orientation tuning less dependent on the size and shape of the RF. This ensures that orientation is represented equally well regardless of RF size and shape.

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