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Robert M Shapley

Publications and source records attributed to Robert M Shapley.

7 recordsLinked to original sources

Effect of stimulus size on the dynamics of orientation selectivity in Macaque V1.

Previous research has established that orientation selectivity depends to a great extent on suppressive mechanisms in the visual cortex. In this study, we investigated the spatial organization and the time-course of these mechanisms. Stimuli were presented in circular windows of "optimal" and "large" radii. The two stimulus sizes were chosen based on an area-response function measured with drifting gratings at high contrast. The "optimal" size was defined as the smallest radius that elicited the peak response (average value of 0.45 degrees ), whereas "large" was defined as two to five times the optimal size. We found that the peak amplitude of tuned enhancement and untuned suppression varied <10% on average with stimulus radius, indicating that they are mainly concentrated in the classical receptive field. However, tuned suppression--in those cells that showed it--was significantly stronger with large stimuli, indicating that this component has a contribution from beyond the classical receptive field. These results imply that spatial context (in large stimuli) enhances orientation selectivity by increasing tuned suppression. We also characterized the time evolution of enhancement, of untuned suppression, and of tuned suppression. The time-course of tuned suppression was markedly slower in time-to-peak and longer in its persistence than untuned suppression. Therefore tuned suppression is likely to be generated by long-range recurrent connections or cortico-cortical feedback, whereas untuned suppression is mainly generated locally in V1.

Animals↗

Differences in real and illusory shape perception revealed by backward masking.

Illusory contours (ICs) are thought to be a result of processes involved in the perceptual recovery of occluded surfaces. Here, we investigate the relationship between real and illusory contour perception using a shape discrimination task and backward masking paradigm. ICs can mask other ICs when times between mask onset and stimulus onset, or SOAs, are very long ( approximately 300 ms), but real contours (RCs) are not similarly effective. Masking is absent for RC masks at perceptually salient contrasts, as well as for those with contrast lowered to match the perceived brightness of the illusory surface. We also find that RCs are not masked at long SOAs, either by ICs or by other RCs. Finally, the masking seen between ICs can occur for different sizes of target and mask. The cross-size masking would not be expected if the masking were at a level sensitive to retinal contour location. The late masking therefore may be related to a higher level of processing of shape categories and surfaces, the level at which shapes defined by ICs and RCs are differentially represented.

Form Perception↗

Information tuning of populations of neurons in primary visual cortex.

Neurons in macaque primary visual cortex (V1) show a diversity of orientation tuning properties, exhibiting a broad distribution of tuning width, baseline activity, peak response, and circular variance (CV). Here, we studied how the different tuning features affect the performance of these cells in discriminating between stimuli with different orientations. Previous studies of the orientation discrimination power of neurons in V1 focused on resolving two nearby orientations close to the psychophysical threshold of orientation discrimination. Here, we developed a theoretical framework, the information tuning curve, that measures the discrimination power of cells as a function of the orientation difference, deltatheta, of the two stimuli. This tuning curve also represents the mutual information between the neuronal responses and the stimulus orientation. We studied theoretically the dependence of the information tuning curve on the orientation tuning width, baseline, and peak responses. Of main interest is the finding that narrow orientation tuning is not necessarily optimal for all angular discrimination tasks. Instead, the optimal tuning width depends linearly on deltatheta. We applied our theory to study the discrimination performance of a population of 490 neurons in macaque V1. We found that a significant fraction of the neuronal population exhibits favorable tuning properties for large deltatheta. We also studied how the discrimination capability of neurons is distributed and compared several other measures of the orientation tuning such as CV with Chernoff distances for normalized tuning curves.

Action Potentials↗

Effects of contrast and size on orientation discrimination.

Motivated by the recent physiological finding that a neuron's receptive field can increase in size by a factor of 2-4-fold at low contrast [Nat. Neurosci. 2 (1999) 733, Proc. Natl. Acad. Sci. USA 96 (1999) 12073], we sought to examine whether a psychophysical task might reflect the contrast dependent changes in the size/structure of a receptive field. We postulate that since spatial summation is not contrast invariant, a task that relies on the spatial structure of a receptive field, such as orientation discrimination, should also be affected by changes in contrast. Previously, orientation discrimination thresholds have been reported to be roughly independent of the contrast of a stimulus for most of the visible range of contrasts [i.e. J. Neurophysiol. 57 (1987) 773, J. Opt. Soc. Am. 6 (1989) 713, Vis. Res. 30 (1990) 449, Vis. Res. 39 (1999) 1631]. Here, we found large improvements in orientation discrimination with contrast that were dependent on stimulus area. Furthermore, the apparent constancy of orientation discrimination for large area stimuli is possibly a result of a floor effect on the threshold. Therefore we conclude that there is not strong evidence for contrast invariant orientation discrimination. We interpret these results in the context of recent neurophysiological results about the expansion of cortical cells' receptive fields at low contrast.

Contrast Sensitivity↗

Space and time maps of cone photoreceptor signals in macaque lateral geniculate nucleus.

We studied neurons in the central visual field representation of the lateral geniculate nucleus (LGN) in macaque monkeys by mapping their receptive fields in space and time. The mapping was performed by reverse correlation of a spike train of a neuron with pseudorandom, binary level stimuli (m-sequence grids). Black and white m-sequence grids were used to map the receptive field for luminance. The locations of receptive field center and surround were determined from this luminance map. To map the contribution of each cone class to the receptive field, we designed red-green or blue-yellow m-sequence grids to isolate the influence of that cone (long, middle, or short wavelength-sensitive: L, M, or S). Magnocellular neurons generally received synergistic input from L and M cones in both the center and the surround. A minority had cone-antagonistic (M-L) input to the surround. Red-green opponent parvocellular neurons received opponent cone input (L+M- or M+L-) that overlapped in space, as sampled by our stimulus grid, but that had somewhat different extents. For example, an L+ center parvocellular neuron would be L+/M- in both center and surround, but the L+ signal would be stronger in the center and the M- signal stronger in the surround. Accordingly, the luminance receptive field would be spatially antagonistic: on-center/off-surround. The space-time maps also characterized LGN dynamics. For example, magnocellular responses were transient, red-green parvocellular responses were more sustained, and blue-on responses were the most sustained for both luminance and cone-isolating stimuli. For all cell types the surround response peaked 8-10 msec later than the center response.

Action Potentials↗

Orientation selectivity in macaque V1: diversity and laminar dependence.

We studied the steady-state orientation selectivity of single neurons in macaque primary visual cortex (V1). To analyze the data, two measures of orientation tuning selectivity, circular variance and orientation bandwidth, were computed from the tuning curves. Circular variance is a global measure of the shape of the tuning curve, whereas orientation bandwidth is a local measure of the sharpness of the tuning curve around its peak. Circular variance in V1 was distributed broadly, indicating a great diversity of orientation selectivity. This diversity was also reflected in the individual cortical layers. However, there was a tendency for neurons with high circular variance, meaning low selectivity for orientation, to be concentrated in layers 4C, 3B, and 5. The relative variation of orientation bandwidth across the cortical layers was less than for circular variance, but it showed a similar laminar dependence. Neurons with large orientation bandwidth were found predominantly in layers 4C and 3B. There was a weak correlation between orientation selectivity and the level of spontaneous activity of the neurons. We also assigned a response modulation ratio for each cell, which is a measure of the linearity of spatial summation. Cells with low modulation ratios tended to have higher circular variance and bandwidth than those with high modulation ratios. These findings suggest a revision to the classical view that nonoriented receptive fields are principally found in layer 4C and the cytochrome oxidase-rich blobs in layer 2/3. Instead, a broad distribution of tuning selectivity is found in all cortical layers, and neurons that are weakly tuned for orientation are ubiquitous in V1 cortex.

Action Potentials↗

A psychophysical correlate of contrast dependent changes in receptive field properties.

Recent physiological investigations have demonstrated that a neuron's area of spatial summation can vary depending on stimulus contrast. Specifically, when the same stimulus is presented to a neuron at a low contrast, the area of summation (or neuron's receptive field) can increase by at least a factor of two, compared to that estimated with a high contrast stimulus. We sought to examine this phenomenon psychophysically by using an orientation discrimination task carried out in the presence of contextual stimuli. We have found previously that orientation discrimination thresholds for a sine-wave grating are elevated by the presence of a surround pattern of similar orientation (with an offset) and spatial frequency. However, when these patterns were separated by a gap of mean luminance exceeding roughly 1 deg, thresholds dropped to the level measured using the center pattern alone. Here, we examined the surround pattern's effect on orientation thresholds as a function of the contrast of the center and surround. We find that when both are presented at a low contrast, the detrimental influence of the surround on orientation thresholds is maintained over larger gap separations. We also find that the spatial frequency and orientation selectivity of the surround's masking effect on orientation thresholds is broader at low contrast than at high contrast. Although the results support the idea of a spatial reorganization of the mechanisms involved in the task at low contrast, these changes are insufficient, in and of themselves, to account for the data. We suggest that additional influences possibly reflecting image segmentation also affect performance.

Contrast Sensitivity↗