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Ralph D Freeman

Publications and source records attributed to Ralph D Freeman.

5 recordsLinked to original sources

Oblique effect: a neural basis in the visual cortex.

The details of oriented visual stimuli are better resolved when they are horizontal or vertical rather than oblique. This "oblique effect" has been confirmed in numerous behavioral studies in humans and to some extent in animals. However, investigations of its neural basis have produced mixed and inconclusive results, presumably due in part to limited sample sizes. We have used a database to analyze a population of 4,418 cells in the cat's striate cortex to determine possible differences as a function of orientation. We find that both the numbers of cells and the widths of orientation tuning vary as a function of preferred orientation. Specifically, more cells prefer horizontal and vertical orientations compared with oblique angles. The largest population of cells is activated by orientations close to horizontal. In addition, orientation tuning widths are most narrow for cells preferring horizontal orientations. These findings are most prominent for simple cells tuned to high spatial frequencies. Complex cells and simple cells tuned to low spatial frequencies do not exhibit these anisotropies. For a subset of simple cells from our population (n = 104), we examined the relative contributions of linear and nonlinear mechanisms in shaping orientation tuning curves. We find that linear contributions alone do not account for the narrower tuning widths at horizontal orientations. By modeling simple cells as linear filters followed by static expansive nonlinearities, our analysis indicates that horizontally tuned cells have a greater nonlinear component than those tuned to other orientations. This suggests that intracortical mechanisms play a major role in shaping the oblique effect.

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Single-neuron activity and tissue oxygenation in the cerebral cortex.

Blood oxygen level-dependent functional magnetic resonance imaging uses alterations in brain hemodynamics to infer changes in neural activity. Are these hemodynamic changes regulated at a spatial scale capable of resolving functional columns within the cerebral cortex? To address this question, we made simultaneous measurements of tissue oxygenation and single-cell neural activity within the visual cortex. Results showed that increases in neuronal spike rate were accompanied by immediate decreases in tissue oxygenation. We used this decrease in tissue oxygenation to predict the orientation selectivity and ocular dominance of neighboring neurons. Our results establish a coupling between neural activity and oxidative metabolism and suggest that high-resolution functional magnetic resonance imaging may be used to localize neural activity at a columnar level.

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Stereoscopic depth processing in the visual cortex: a coarse-to-fine mechanism.

For binocular animals viewing a three-dimensional scene, the left and right eyes receive slightly different information, and the brain uses this 'binocular disparity' to interpret stereoscopic depth. An important theoretical conjecture in this mechanism is that coarse processing precedes and constrains finely detailed processing. We present three types of neurophysiological data from the cat's visual cortex that are consistent with a temporal coarse-to-fine tuning of disparity information. First, the disparity tuning of cortical cells generally sharpened during the time course of response. Second, cells responsive to large and small spatial scale had relatively shorter and longer temporal latencies, respectively. Third, cross-correlation analysis between simultaneously recorded pairs of cortical cells showed that connections between disparity-tuned neurons were generally stronger for coarse-to-fine processing than for fine-to-coarse processing. These results are consistent with theoretical and behavioral studies and suggest that rapid, coarse percepts are refined over time in stereoscopic depth perception.

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Cortical columns: a multi-parameter examination.

Columnar structure in the cerebral cortex has been demonstrated in numerous studies. However, in the visual system, it is not clear from imaging, basic physiological and anatomical approaches how multiple stimulus parameters are related within columns. We have analyzed recordings from pairs of neurons in the striate cortex of the cat using various spatial and temporal parameters. We find that most parameters are clustered within inferred columns with the exception of spatial phase. Diversity of phase could be useful for serial processing in central visual pathways.

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Disinhibition outside receptive fields in the visual cortex.

By definition, the region outside the classical receptive field (CRF) of a neuron in the visual cortex does not directly activate the cell. However, the response of a neuron can be influenced by stimulation of the surrounding area. In previous work, we showed that this influence is mainly suppressive and that it is generally limited to a local region outside the CRF. In the experiments reported here, we investigate the mechanisms of the suppressive effect. Our approach is to find the position of a grating patch that is most effective in suppressing the response of a cell. We then use a masking stimulus at different contrasts over the grating patch in an attempt to disinhibit the response. We find that suppressive effects may be partially or completely reversed by use of the masking stimulus. This disinhibition suggests that effects from outside the CRF may be local. Although they do not necessarily underlie the perceptual analysis of a figure-ground visual scene, they may provide a substrate for this process.

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