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B C Skottun

Publications and source records attributed to B C Skottun.

At least 19 recordsLinked to original sources
PubMed1994

Illusory contours and linear filters.

The border between two abutting gratings is sometimes referred to as an illusory contour. In the Fourier spectrum of such a pattern there is no energy corresponding to the precise orientation of this contour. Neuronal responses elicited by such stimuli are therefore thought to be incompatible with linear filtering. The present investigation tested this notion. Abutting gratings were Fourier analyzed and passed through spatial and temporal filters with properties comparable with those of cortical neurons. It was found that the distribution of amplitudes in the Fourier spectrum is such that these stimuli may indeed stimulate neurons based on linear filtering. In the case of drifting stimuli the amount of stimulation may be maximal when the illusory contour is at the optimal orientation for bars with less stimulation on either side of this orientation.

Cerebral Cortex↗
PubMed1991

Classifying simple and complex cells on the basis of response modulation.

Hubel and Wiesel (1962; Journal of Physiology, London, 160, 106-154) introduced the classification of cortical neurons as simple and complex on the basis of four tests of their receptive field structure. These tests are partly subjective and no one of them unequivocally places neurons into distinct classes. A simple, objective classification criterion based on the form of the response to drifting sinusoidal gratings has been used by several laboratories, although it has been criticized by others. We review published and unpublished evidence which indicates that this simple and objective criterion reliability divides neurons of the striate cortex in both cats and monkeys into two groups that correspond closely to the classically-described simple and complex classes.

Animals↗
PubMed1988

Responses of simple and complex cells to random dot patterns: a quantitative comparison.

1. There are several reports that random dot patterns are potent stimuli for cortical complex cells but not for simple cells. This finding is regarded as evidence against Hubel and Wiesel's hierarchical model of cortical circuitry, in which simple cells are the principal input to complex cells. We have reinvestigated the question quantitatively by recording responses to dot patterns from 106 cells in area 17 and the 17/18 border region of normal adult cats. 2. The cells were classified as simple (n = 62) or complex (n = 40) (4 were end stopped or hypercomplex) on the basis of whether they gave modulated (AC) or unmodulated (DC) responses to drifting sine gratings. 3. Although there are large within-group differences, we found both simple and complex cells that respond to bright random dots on a dark background, drifted across the receptive field at 3 degrees/s. The responses at the optimal direction averaged 6.2 and 18.1 spikes/s (spontaneous activity subtracted) for simple and complex cells, respectively. 4. We also recorded responses to drifting sine gratings. Complex cells were also found to respond more than simple cells to these stimuli. For each cell, we calculated a dot index expressing the dot response relative to grating response. The dot index averaged 0.43 for simple cells and 0.55 for complex cells. It therefore appears that much of the difference in response to dot patterns reflects a difference in general responsivity. 5. In subsamples of cells, we examined the effects of varying dot density, dot size, and drift velocity. These variables affect different cells in a manner largely independent of cell class. Most simple cells in our sample responded well to random dot patterns at several velocities, at two different dot sizes and at both 3 and 50% dot densities. 6. Our results agree with previous studies in showing that complex cells respond more vigorously than simple cells to dot patterns, but the fact that many simple cells also respond to these stimuli makes our results consistent with a hierarchical model of cortical circuitry.

Animals↗
PubMed1987

Effects of contrast and spatial frequency on vernier acuity.

We have examined vernier acuity using sinusoidal luminance gratings. Vernier thresholds were affected by both grating contrast and spatial frequency. With fixed (50%) contrast gratings, vernier thresholds reached minimum values of approximately 10 sec of arc at spatial frequencies between 6 and 16 c/deg. Vernier thresholds for all spatial frequencies are related to contrast by a power law with exponents of approximately -0.8. Thresholds approach half a grating period (180 deg phase shift) as grating contrast approaches detection thresholds. We discuss our results in relation to three models for vernier detection. Most of our data are consistent with the predictions of Wilson's [(1986) Vision Res. 26, 453-469] model. Detection of vernier off-sets at low spatial frequencies may depend on detection of the horizontal border formed between the two halves of the grating.

Form Perception↗
PubMed1987

Visual orientation and spatial frequency discrimination: a comparison of single neurons and behavior.

Neurons in the visual cortex respond selectively to stimulus orientation and spatial frequency. Changes in response amplitudes of these neurons could be the neurophysiological basis of orientation and spatial frequency discrimination. We have estimated the minimum differences in stimulus orientation and spatial frequency that can produce reliable changes in the responses of individual neurons in cat visual cortex. We compare these values with orientation and spatial frequency discrimination thresholds determined behaviorally. Slopes of the tuning functions and response variability determine the minimum orientation and spatial frequency differences that can elicit a reliable response change. These minimum values were obtained from single cells using receiver operating characteristic (ROC) analysis. The average minimum orientation and spatial frequency differences that could be signaled reliably by cells from our sample were 6.4 degrees (n = 22) and 21.3% (n = 18), respectively. These values are approximately 0.20 of the average full tuning width at one-half height of the cells. Although these average values are well above the behaviorally determined thresholds, the most selective cells signaled orientation and frequency differences of 1.84 degrees and 5.25%, respectively. These values are of the same order of magnitude as the behavioral thresholds. We show that, because of slow fluctuations in a cell's responsivity, ROC analysis overestimates response variability. We estimate that these slow response fluctuations elevated our estimates of single cell "thresholds" by, on average, 30%. Our data point to an approximate correspondence between orientation and spatial frequency discrimination "thresholds" determined behaviorally and those estimated from the most selective single cortical cells. Interpretation of this quantitative correspondence is considered in the discussion.

Animals↗
PubMed1987

The effects of contrast on visual orientation and spatial frequency discrimination: a comparison of single cells and behavior.

We have compared the effects of contrast on human psychophysical orientation and spatial frequency discrimination thresholds and on the responses of individual neurons in the cat's striate cortex. Contrast has similar effects on orientation and spatial frequency discrimination: as contrast is increased above detection threshold, orientation and spatial frequency discrimination performance improves but reaches maximum levels at quite low contrasts. Further increases in contrast produce no further improvements in discrimination. We measured the effects of contrast on response amplitude, orientation and spatial frequency selectivity, and response variance of neurons in the cat's striate cortex. Orientation and spatial frequency selectivity vary little with contrast. Also, the ratio of response variance to response mean is unaffected by contrast. Although, in many cells, response amplitude increases approximately linearly with log contrast over most of the visible range, some cells show complete or partial saturation of response amplitude at medium contrasts. Therefore, some cells show a clear increase in slope of the orientation and spatial frequency tuning functions with increasing contrast, whereas in others the slopes reach maximum values at medium contrasts. Using receiver operating characteristic analysis, we estimated the minimum orientation and spatial frequency differences that can be signaled reliably as a response change by an individual cell. This analysis shows that, on average, the discrimination of orientation or spatial frequency improves with contrast at low contrasts more than at higher contrasts. Using the optimal stimulus for each cell, we estimated the contrast threshold of 48 neurons. Most cells had contrast thresholds below 5%. Thresholds were only slightly higher for nonoptimal stimuli. Therefore, increasing the contrast of sinusoidal gratings above approximately 10% will not produce large increases in the number of responding cells. The observed effects of contrast on the response characteristics of nonsaturating cortical cells do not appear consistent with the psychophysical results. Cells that reach their maximum response at low-to-medium contrasts may account for the contrast independence of psychophysical orientation and spatial frequency discrimination thresholds at medium and high contrasts.

Animals↗
PubMed1986

Effect of contrast on spatial frequency tuning of neurones in area 17 of cat's visual cortex.

Previous investigations have revealed that perceived spatial frequency of gratings rises as contrast is lowered. In order to account for this finding it has been postulated that the spatial frequency which produces the maximum response from cortical neurones decreases with contrast. We have examined this hypothesis by determining optimal spatial frequency at 3-5 different contrast levels for 37 neurones in the cat striate cortex. For the complete sample no systematic changes in optimal spatial frequency was observed. However, a shift in the predicted direction was found for cells tuned to high spatial frequencies.

Animals↗
PubMed1986

Orientation discrimination in amblyopia.

Using extended sinusoidal gratings to avoid potential problems of eccentric fixation, the authors have studied orientation discrimination in amblyopia. For all subjects, elevated orientation discrimination thresholds at high spatial frequencies were found. However, raised thresholds decrease with decreasing spatial frequency, and can be normal at low frequencies. Orientation discrimination thresholds for both amblyopic and non-amblyopic eyes are independent of contrast over most of the visible range. Therefore, amblyopic orientation discrimination thresholds cannot be mimicked in non-amblyopic eyes by reducing contrast. Control experiments show that the orientation discrimination deficits are not restricted to vertical stimuli and that they are not a result of exaggerated cyclotorsional eye movements.

Amblyopia↗
PubMed1985

Neurophysiological evaluation of the differential response model for orientation and spatial-frequency discrimination.

Recent models have attempted to reconcile low psychophysical orientation and spatial-frequency discrimination thresholds with relatively broad orientation and spatial-frequency tuning of cortical neurons. These models have relied on the ability of the neurons to convert small stimulus changes into reliable response changes. We have examined this ability in a sample of neurons from the cat's striate cortex. We present here data from two cells that reliably signaled the smallest orientation and spatial-frequency differences. Using receiver operating characteristic analysis, we find that these cells could reliably signal orientation differences of 1.84 deg and spatial-frequency differences of 0.073 octave. We compare these single-cell results to cat and human behavioral discrimination thresholds.

Animals↗
PubMed1984

Stimulus specificity of binocular cells in the cat's visual cortex: ocular dominance and the matching of left and right eyes.

Most cells in the striate cortex respond to visual stimulation through either eye. We have examined quantitatively the matching of response specificity for the two eyes. Our intention was to determine the degree to which this matching depends on ocular dominance. We used standard single cell recording techniques and studied responses to sinusoidal gratings of different spatial frequencies, orientations, and contrasts. For all tests, stimuli were randomly interleaved both with respect to the value of each parameter, and the eye which was stimulated. After estimating ocular dominance qualitatively and quantitatively, we measured: response modulation (to help identify whether a cell was simple or complex), orientation and spatial frequency tuning, and contrast response functions (to estimate contrast thresholds). Results show that: (1) Response modulation is well matched between the two eyes, but there is a slight tendency for the dominant eye to respond with less modulation. (2) Optimal orientation and spatial frequency and their respective tuning widths were similar for the two eyes. In general, tuning functions for the two eyes differed mainly in slope. However, in each case, there was a tendency for the dominant eye to have broader tuning widths. (3) In most cases, contrast response functions for the two eyes differed mainly in their slopes. Extrapolation to spontaneous levels suggests that estimated contrast thresholds are relatively independent of ocular dominance although, again, there ws a tendency for the dominant eye to exhibit slightly lower estimated thresholds. These findings demonstrate that response characteristics between the two eyes are generally well matched regardless of relative response strength. There are, however, small but clear differences between the two eyes for all parameters we measured which are related to and demonstrate that ocular dominance influences the degree of matching between the two eyes.

Animals↗
PubMed1984

The effects of large orientation and spatial frequency differences on spatial discriminations.

We have examined two questions: (1) can the finest orientation discrimination be achieved only between stimuli with similar spatial frequency content? and likewise, (2) can the lowest spatial frequency discrimination thresholds be achieved only with parallel gratings? In 2 AFC tests we found that neither type of discrimination was affected by stimulus differences along the other dimension. However, some small decreases in method of adjustment matching accuracy were associated with large differences along the secondary dimensions. Considering the neurophysiological implications, these data suggest that fine orientation and spatial frequency discrimination can occur even though separate populations of neurones in the primary visual cortex may be activated by the two stimuli to be discriminated.

Discrimination, Psychological↗
PubMed1984

Spatial summation in subregions of simple-cell receptive fields in cat striate cortex as a function of slit length.

Spatial summation along the optimum stimulus orientation in subregions of simple-cell receptive fields in cat striate cortex was studied quantitatively by measuring the response to stationary light slits of variable length. Before summation analysis, the cell's discharge field was mapped by flashing a test slit on and off in a sequence of positions through the receptive field. A static activation procedure was used to determine the extension of subregions of the receptive field where light stimulation increased (enhancement) or decreased (suppression) the firing rate. An activation slit in the optimum orientation was positioned in the most responsive position of the discharge field and the effects of a parallel test slit, in a series of broadside positions, were assessed from the changes induced in the discharge elicited by the activation slit. Length-response curves for on and off responses were made by positioning a test slit in the respective subregions of the discharge field. The activation procedure was used to make length-response curves for suppression. A test slit of variable length was positioned in a suppression region defined by the activation profiles and an activation slit of fixed length was centred in the most responsive discharge field position. Length summation was found for all cells, both with respect to on and off responses, and suppression. The curves for on and off responses had a maximum value beyond which the response declined or levelled off, but some cells had a secondary, more shallow increase beyond an initial, steeply rising part. Similar properties were found for summation of suppression except that the effects were opposite in sign. Curves made for both on and off regions in the same cell often differed in shape. Such differences were also found when length-response curves made in different suppression regions of the same cell were compared. The various length-summation functions were explained by a model presuming that simple-cell receptive fields consist of partially overlapping non-concentric excitatory and inhibitory fields. This arrangement would also explain why length-response curves in various subregions often had different shapes.

Animals↗
PubMed1983

Spatial summation in the receptive field of simple cells in the cat striate cortex.

Spatial summation was studied quantitatively through width response curves made with an optimally oriented test slit of variable width, and by comparing the response to combined presentation of several parallel slits with the response to each slit alone. Prior to summation analysis, the cell's discharge field (DF) was mapped by presenting a test slit ON and OFF across the receptive field. Activation profiles, showing the extension of subregions where light stimulation increased (enhancement) or decreased the firing rate (suppression), were made by presenting an optimally oriented activation slit in the most responsive DF-position. Against this activity the effects of a parallel test slit were determined in a series of broadside positions. Width response curves were made over the subregions of the DF and the activation profiles. Spatial summation was found in all cells, but the width of the summation region was smaller than the width of the subregions in the respective profiles. The width of the summation region was related to the degree of activation rather than to specific locations within the receptive field. The effect produced by several slits presented together deviated from the algebraic sum of the effects produced by each slit alone. Linear summation was rarely found. Accumulated response curves obtained by integration of DF or activation profiles were compared with width response curves to test linearity of summation. Linear summation throughout the whole receptive field was never found. A satisfactory fit was found only over a narrow region showing that summation was linear within a small part of the summation region. Linearity ended near response maxima or minima in the response profiles. The results indicate that the receptive field of simple cells consists of overlapping excitatory and inhibitory fields, and that the exact location and width of enhancement and suppression zones are determined by an activity-dependent balance between excitatory and inhibitory inputs.

Animals↗