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R M Shapley

Publications and source records attributed to R M Shapley.

At least 19 recordsLinked to original sources
PubMed2002

Suppression of neural responses to nonoptimal stimuli correlates with tuning selectivity in macaque V1.

Neural responses in primary visual cortex (area V1) are selective for the orientation and spatial frequency of luminance-modulated sinusoidal gratings. Selectivity could arise from enhancement of the cell's response by preferred stimuli, suppression by nonoptimal stimuli, or both. Here, we report that the majority of V1 neurons do not only elevate their activity in response to preferred stimuli, but their firing rates are also suppressed by nonoptimal stimuli. The magnitude of suppression is similar to that of enhancement. There is a tendency for net response suppression to peak at orientations near orthogonal to the optimal for the cell, but cases where suppression peaks at oblique orientations are observed as well. Interestingly, selectivity and suppression correlate in V1: orientation and spatial frequency selectivity are higher for neurons that are suppressed by nonoptimal stimuli than for cells that are not. This finding is consistent with the idea that suppression plays an important role in the generation of sharp cortical selectivity. We show that nonlinear suppression is required to account for the data. However, the precise structure of the neural circuitry generating the suppressive signal remains unresolved. Our results are consistent with both feedback and (nonlinear) feed-forward inhibition.

Animals↗
PubMed2001

The relatively small decline in orientation acuity as stimulus size decreases.

Orientation acuity was measured with circular patches of sinusoidal gratings of various sizes. Threshold estimates were lowest (acuity highest) for the largest size patch, and increased as the stimulus size was reduced, consistent with the results of many researchers using line stimuli. These results are compared with the predictions of a simple and widely accepted model of spatial vision whereby the output of independent feed-forward filters are combined to produce threshold estimates. Specifically, the rectified output of a number of independent filters (i.e. Gabors) spanning the stimulus space (i.e. orientation) are combined via Bayesian decision theory. This model cannot account quantitatively for the relatively low thresholds estimated for the small sized stimuli when compared to the thresholds measured with larger patches. Application of a comparable analysis, with preliminary measurements of neuronal responses from primary visual cortex replacing the response rectified Gabor filter's responses, provides a more reasonable account of behavioral acuity. This indicates a fundamental inadequacy of the feed-forward filter model in accounting for V1 neurons' role in perception.

Adult↗
PubMed2001

Contextual influences on orientation discrimination: binding local and global cues.

We sought to determine how local and global features within an image interact by examining whether orientation discrimination thresholds could be modified by contextual information. In particular, we investigated how local orientation signals within an image are pooled together, and whether this pooling process is dependent on the global orientation content present in the image. We find that observers' orientation judgments depend on surround contextual information, with performance being optimal when the center and surround stimuli are clearly distinct. In cases where the center and surround were not clearly segregated, we report two sets of results. If there was an ambiguity regarding the perception of a global structure (i.e. a small mismatch between local cues), observers' performance was impaired. If there was no mismatch and local and global cues were consistent with the perception of a single surface, observers performed as well as in the distinct surfaces case. Although some of our results can be largely accounted for by interactions between differently oriented filters, other aspects are more difficult to reconcile with this explanation. We suggest that low level filtering constrains observers' performance, and that influences arising from image segmentation modify how local orientation signals are pooled together.

Contrast Sensitivity↗
PubMed2000

Visual function and brain organization in non-decussating retinal-fugal fibre syndrome.

Functional neuroimaging, psychophysical and electrophysiological investigations were performed in a patient with non-decussating retinal-fugal fibre syndrome, an inborn achiasmatic state in which the retinal projections of each eye map entirely to the ipsilateral primary visual cortex. Functional magnetic resonance imaging (fMRI) studies showed that for monocularly presented simple visual stimuli, only the ipsilateral striate cortex was activated. Within each hemisphere's striate cortex, the representation of the two hemifields overlapped extensively. Despite this gross miswiring, visual functions that require precise geometrical information (such as vernier acuity) were normal, and there was no evidence for the confounding of visual information between the overlapping ipsi-lateral and contralateral representations. Contrast sensitivity and velocity judgments were abnormal, but their dependence on the orientation and velocity of the targets suggests that this deficit was due to ocular instabilities, rather than the miswiring per se. There were no asymmetries in performance observed in visual search, visual naming or illusory contour perception. fMRI analysis of the latter two tasks under monocular viewing conditions indicated extensive bilateral activation of striate and prestriate areas. Thus, the remarkably normal visual behavior achieved by this patient is a result of both the plasticity of visual pathways, and efficient transfer of information between the hemispheres.

Adolescent↗
PubMed1999

Illusory contour perception and amodal boundary completion: evidence of a dissociation following callosotomy.

A fundamental problem in form perception is how the visual system can link together spatially separated contour fragments to form the percept of a unitary shape. Illusory contours and amodal completion are two phenomena that demonstrate this linking process. In the present study we investigate these phenomena in the divided hemispheres of two callosotomy ("split-brain") patients. The data suggest that dissociable neural mechanisms are responsible for the generation of illusory contours and amodal completion. Although both cerebral hemispheres appear to be equally capable of perceiving illusory contours, amodal completion is more readily utilized by the right hemisphere. These results suggest that illusory contours may be attributable to low-level visual processes common to both hemispheres, whereas amodal completion reflects a higher-level, lateralized process.

Adult↗
PubMed1998

Functional organization of owl monkey lateral geniculate nucleus and visual cortex.

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."

Animals↗
PubMed1994

Linearity and non-linearity in cortical receptive fields.

Visual neurons in striate (V1) cortex have been studied as feature detectors or as spatiotemporal filters. A useful way to distinguish between these two conceptual approaches is by studying the way in which visual signals are pooled across space and time. Many neurons in layer IV of striate cortex exhibit linear spatial summation and their response time course is consistent with linear temporal summation. Neurons in supragranular and infragranular layers sum signals in a non-linear manner. A particularly important non-linearity seen in many cortical complex cells is non-linear summation along an axis parallel to their preferred orientation. This leads to responsiveness to 'illusory contours', borders defined by texture differences only. These and other results on non-linear summation of chromatic and achromatic signals imply that V1 cortex performs sophisticated and complex image processing and is not simply an array of spatiotemporal filters.

Animals↗
PubMed1994

On temporal codes and the spatiotemporal response of neurons in the lateral geniculate nucleus.

1. The present work relates recent experimental studies of the temporal coding of visual stimuli (McClurkin, Optican, Richmond, and Gawne, Science 253: 675, 1991) to the measurements of the spatiotemporal receptive fields of neurons within the lateral geniculate of primate. 2. We analyze both new and previously described magnocellular and parvocellular single units. The spatiotemporal impulse response function of the unit, defined as the time-resolved average firing rate in response to a weak stimulus flashed at a given location and time, is characterized by the singular value decomposition. This analysis allows one to represent the impulse response by a small number, two to three, of spatial and temporal modes. Both magnocellular and parvocellular units are weakly nonseparable, with major and minor modes that account, respectively, for approximately 78 and 22% of the response. The major temporal mode for both types is essentially identical for the first 100 ms. At later times the response of magnocellular units changes sign and decays slowly, whereas the response of parvocellular units decays relatively rapidly. 3. The spatiotemporal impulse response function completely determines the response of a unit to an arbitrary stimulus when linear response theory is valid. Using the measured impulse response, combined with a rectifying neuronal input-output relation, we calculate the responses to a complete set of spatial luminance patterns constructed of "Walsh" functions. Our predicted temporal responses are in qualitative agreement with those reported for parvocellular units (McClurkin, Optican, Richmond, and Gawne, J. Neurophysiol. 66: 794, 1991). Under the additional assumptions of Poisson statistics for the probability of spiking and a plausible background firing rate, we predict the performance of a unit in the Walsh pattern discrimination task as quantified by mutual information. Our prediction is again consistent with the reported results. 4. Last, we consider the issue of temporal coding within linear response. For stimuli presented for fixed time intervals, the singular value decomposition provides a natural relation between the temporal modes of the neuronal response and the spatial pattern of the stimulus. Although it is tempting to interpret each temporal mode as an independent channel that encodes orthogonal features of the stimulus, successively higher order modes are increasingly unreliable and do not significantly increase the discrimination capabilities of the unit.

Animals↗
PubMed1993

Macaque V1 neurons can signal 'illusory' contours.

We describe here a new view of primary visual cortex (V1) based on measurements of neural responses in V1 to patterns called 'illusory contours' (Fig. 1a, b). Detection of an object's boundary contours is a fundamental visual task. Boundary contours are defined by discontinuities not only in luminance and colour, but also in texture, disparity and motion. Two theoretical approaches can account for illusory contour perception. The cognitive approach emphasizes top-down processes. An alternative emphasizes bottom-up processing. This latter view is supported by (1) stimulus constraints for illusory contour perception and (2) the discovery by von der Heydt and Peterhans of neurons in extrastriate visual area V2 (but not in V1) of macaque monkeys that respond to illusory contours. Using stimuli different from those used previously, we found illusory contour responses in about half the neurons studied in V1 of macaque monkeys. Therefore, there are neurons as early as V1 with the computational power to detect illusory contours and to help distinguish figure from ground.

Animals↗
PubMed1992

Spatial structure of cone inputs to receptive fields in primate lateral geniculate nucleus.

Human colour vision depends on three classes of cone photoreceptors, those sensitive to short (S), medium (M) or long (L) wavelengths, and on how signals from these cones are combined by neurons in the retina and brain. Macaque monkey colour vision is similar to human, and the receptive fields of macaque visual neurons have been used as an animal model of human colour processing. P retinal ganglion cells and parvocellular neurons are colour-selective neurons in macaque retina and lateral geniculate nucleus. Interactions between cone signals feeding into these neurons are still unclear. On the basis of experimental results with chromatic adaptation, excitatory and inhibitory inputs from L and M cones onto P cells (and parvocellular neurons) were thought to be quite specific (Fig. 1a). But these experiments with spatially diffuse adaptation did not rule out the 'mixed-surround' hypothesis: that there might be one cone-specific mechanism, the receptive field centre, and a surround mechanism connected to all cone types indiscriminately (Fig. 1e). Recent work has tended to support the mixed-surround hypothesis. We report here the development of new stimuli to measure spatial maps of the linear L-, M- and S-cone inputs to test the hypothesis definitively. Our measurements contradict the mixed-surround hypothesis and imply cone specificity in both centre and surround.

Animals↗
PubMed1992

Broadband temporal stimuli decrease the integration time of neurons in cat striate cortex.

We have studied the responses of striate cortical neurons to stimuli whose contrast is modulated in time by either a single sinusoid or by the sum of eight sinusoids. The sum-of-sinusoids stimulus resembles white noise and has been used to study the linear and nonlinear dynamics of retinal ganglion cells (Victor et al., 1977). In cortical neurons, we have found different linear and second-order responses to single-sinusoid and sum-of-sinusoids inputs. Specifically, while the responsivity near the optimal temporal frequency is lower for the sum-of-sinusoids stimulus, the responsivity at higher temporal frequencies is relatively greater. Along with this change in the response amplitudes, there is a systematic change in the time course of responses. For complex cells, the integration time, the effective delay due to a combination of actual delays and low-pass filter stages, changes from a median of 85 ms with single sinusoids to 57 ms with a sum of sinusoids. For simple cells, the integration times for single sinusoids range from 44-100 ms, but cluster tightly around 40 ms for the sum-of-sinusoids stimulus. The change in time constant would argue that the increased sensitivity to high frequencies cannot be explained by a static threshold, but must be caused by a fundamental alteration in the response dynamics. These effects are not seen in the retina (Shapley & Victor, 1981) and are most likely cortical in origin.

Animals↗
PubMed1991

Fine structure of receptive-field centers of X and Y cells of the cat.

We investigated the fine structure of receptive field centers of X and Y cells of the retina and lateral geniculate nucleus of the cat using sinusoidal grating stimuli of high spatial frequency. By measuring orientation tuning and spatial-frequency tuning at multiple orientations, the two-dimensional sensitivity distribution was examined. We found that receptive-field centers typically have multiple sensitivity peaks that can be modeled as several spatially offset subunits. A subunit structure was found in both X and Y cells, with an average number of subunits per receptive-field center of approximately 2.9 in X cells and approximately 4.6 in Y cells. In X cells these subunits may correspond to individual cone bipolar inputs. In Y cells, the subunits may reflect the structure of the dendritic tree. The observation of the subunit structure of the receptive-field center, in conjunction with manipulation of the retinal wiring through pharmacological intervention, may provide a new tool for probing the circuitry of the retina.

Animals↗
PubMed1991

Gender differences in apparent motion perception.

Distance disparity is a strong cue to element correspondence in apparent motion. Using a 2-AFC paradigm we have previously shown that shape similarity also plays a role. We now demonstrate a small gender difference in these effects: women are more sensitive to distance disparity, whereas men are more sensitive to differences in shape. Furthermore, in the competing presence of a shape cue, women's sensitivity to distance decreases while men's sensitivity is unaffected. These observations may be related to putative gender differences in the 'form' and 'motion-spatial relations' cortical pathways.

Adult↗
PubMed1991

Directional selectivity and spatiotemporal structure of receptive fields of simple cells in cat striate cortex.

1. Simple cells in cat striate cortex were studied with a number of stimulation paradigms to explore the extent to which linear mechanisms determine direction selectivity. For each paradigm, our aim was to predict the selectivity for the direction of moving stimuli given only the responses to stationary stimuli. We have found that the prediction robustly determines the direction and magnitude of the preferred response but overestimates the nonpreferred response. 2. The main paradigm consisted of comparing the responses of simple cells to contrast reversal sinusoidal gratings with their responses to drifting gratings (of the same orientation, contrast, and spatial and temporal frequencies) in both directions of motion. Although it is known that simple cells display spatiotemporally inseparable responses to contrast reversal gratings, this spatiotemporal inseparability is demonstrated here to predict a certain amount of direction selectivity under the assumption that simple cells sum their inputs linearly. 3. The linear prediction of the directional index (DI), a quantitative measure of the degree of direction selectivity, was compared with the measured DI obtained from the responses to drifting gratings. The median value of the ratio of the two was 0.30, indicating that there is a significant nonlinear component to direction selectivity. 4. The absolute magnitudes of the responses to gratings moving in both directions of motion were compared with the linear predictions as well. Whereas the preferred direction response showed only a slight amount of facilitation compared with the linear prediction, there was a significant amount of nonlinear suppression in the nonpreferred direction. 5. Spatiotemporal inseparability was demonstrated also with stationary temporally modulated bars. The time course of response to these bars was different for different positions in the receptive field. The degree of spatiotemporal inseparability measured with sinusoidally modulated bars agreed quantitatively with that measured in experiments with stationary gratings. 6. A linear prediction of the responses to drifting luminance borders was compared with the actual responses. As with the grating experiments, the prediction was qualitatively accurate, giving the correct preferred direction but underestimating the magnitude of direction selectivity observed.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1990

Light adaptation in the primate retina: analysis of changes in gain and dynamics of monkey retinal ganglion cells.

The responses of monkey retinal ganglion cells to sinusoidal stimuli of various temporal frequencies were measured and analyzed at a number of mean light levels. Temporal modulation tuning functions (TMTFs) were measured at each mean level by varying the drift rate of a sine-wave grating of fixed spatial frequency and contrast. The changes seen in ganglion cell temporal responses with changes in adaptation state were similar to those observed in human subjects and in turtle horizontal cells and cones tested with sinusoidally flickering stimuli; "Weber's Law" behavior was seen at low temporal frequencies but not at higher temporal frequencies. Temporal responses were analyzed in two ways: (1) at each light level, the TMTFs were fit by a model consisting of a cascade of low- and high-pass filters; (2) the family of TMTFs collected over a range of light levels for a given cell was fit by a linear negative feedback model in which the gain of the feedback was proportional to the mean light level. Analysis (1) revealed that the temporal responses of one class of monkey ganglion cells (M cells) were more phasic at both photopic and mesopic light levels than the responses of P ganglion cells. In analysis (2), the linear negative feedback model accounted reasonably well for changes in gain and dynamics seen in three P cells and one M cell. From the feedback model, it was possible to estimate the light level at which the dark-adapted gain of the cone pathways in the primate retina fell by a factor of two. This value was two to three orders of magnitude lower than the value estimated from recordings of isolated monkey cones. Thus, while a model which includes a single stage of negative feedback can account for the changes in gain and dynamics associated with light adaptation in the photopic and mesopic ranges of vision, the underlying physical mechanisms are unknown and may involve elements in the primate retina other than the cone.

Adaptation, Ocular↗
PubMed1988

Background light and the contrast gain of primate P and M retinal ganglion cells.

Retinal ganglion cells projecting to the monkey lateral geniculate nucleus fall into two classes: those projecting to the magnocellular layers of the nucleus (M cells) have a higher contrast gain to luminance patterns at photopic levels of retinal illumination than those projecting to the parvocellular layers (P cells). We report here that this difference in luminance contrast gain between M and P cells is maintained at low levels of mean retinal illumination. In fact, our results suggest that in the mesopic and scotopic ranges of mean illumination, the M-cell/magnocellular pathway is the predominant conveyor of information about spatial contrast to the visual cortex.

Animals↗
PubMed1987

Linear mechanisms of directional selectivity in simple cells of cat striate cortex.

The role of linear spatial summation in the directional selectivity of simple cells in cat striate cortex was investigated. The experimental paradigm consisted of comparing the response to drifting grating stimuli with linear predictions based on the response to stationary contrast-reversing gratings. The spatial phase dependence of the response to contrast-reversing gratings was consistent with a high degree of linearity of spatial summation within the receptive fields. Furthermore, the preferred direction predicted from the response to stationary gratings generally agreed with the measurements made with drifting gratings. The amount of directional selectivity predicted was, on average, about half the measured value, indicating that nonlinear mechanisms act in concert with linear mechanisms in determining the overall directional selectivity.

Action Potentials↗