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

1999 ◽  
Vol 16 (6) ◽  
pp. 1115-1121 ◽  
Author(s):  
JUSTIN L. GARDNER ◽  
AKIYUKI ANZAI ◽  
IZUMI OHZAWA ◽  
RALPH D. FREEMAN
Keyword(s):  
Aspect Ratio ◽  
Striate Cortex ◽  
Tuning Curve ◽  
Spatial Summation ◽  
Orientation Selectivity ◽  
Orientation Tuning ◽  
Simple Cells ◽  
Size And Shape ◽  
Nonlinear Contribution ◽  
Linear And Nonlinear

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.

Role of Inhibition in the Specification of Orientation Selectivity of Cells in the Cat Striate Cortex

1989 ◽  
Vol 2 (1) ◽  
pp. 41-55 ◽  
Author(s):  
A. B. Bonds
Keyword(s):  
Striate Cortex ◽  
Temporal Frequency ◽  
Selective Inhibition ◽  
Orientation Selectivity ◽  
Orientation Tuning ◽  
Individual Response ◽  
Cortical Cells ◽  
Simple Cells ◽  
Local Maxima ◽  
Tuning Function

AbstractMechanisms supporting orientation selectivity of cat striate cortical cells were studied by stimulation with two superimposed sine-wave gratings of different orientations. One grating (base) generated a discharge of known amplitude which could be modified by the second grating (mask). Masks presented at nonoptimal orientations usually reduced the base-generated response, but the degree of reduction varied widely between cells. Cells with narrow orientation tuning tended to be more susceptible to mask presence than broadly tuned cells; similarly, simple cells generally showed more response reduction than did complex cells.The base and mask stimuli were drifted at different temporal frequencies which, in simple cells, permitted the identification of individual response components from each stimulus. This revealed that the reduction of the base response by the mask usually did not vary regularly with mask orientation, although response facilitation from the mask was orientation selective. In some sharply tuned simple cells, response reduction had clear local maxima near the limits of the cell's orientation-tuning function.Response reduction resulted from a nearly pure rightward shift of the response versus log contrast function. The lowest mask contrast yielding reduction was within 0.1–0.3 log unit of the lowest contrast effective for excitation.The temporal-frequency bandpass of the response-reduction mechanism resembled that of most cortical cells. The spatial-frequency bandpass was much broader than is typical for single cortical cells, spanning essentially the entire visual range of the cat.These findings are compatible with a model in which weak intrinsic orientation-selective excitation is enhanced in two stages: (1) control of threshold by nonorientation-selective inhibition that is continuously dependent on stimulus contrast; and (2) in the more narrowly tuned cells, orientation-selective inhibition that has local maxima serving to increase the slope of the orientation-tuning function.

Responses of neurons in cat striate cortex to vernier offsets in reverse contrast stimuli

1995 ◽  
Vol 12 (5) ◽  
pp. 805-817 ◽  
Author(s):  
N.v. Swindale
Keyword(s):  
Striate Cortex ◽  
Tuning Curve ◽  
High Sensitivity ◽  
Orientation Tuning ◽  
Vernier Acuity ◽  
Simple Cells ◽  
Tuning Curves ◽  
Complex Cells ◽  
Simple And Complex Cells ◽  
Opposite Contrast

AbstractThis paper examines how the responses of cells in area 17 of the cat vary as a function of the vernier offset between a bright and a dark bar. The study was prompted by the finding that human vernier acuity is reduced for bars or edges of opposite contrast sign (Mather & Morgan, 1986; O'Shea & Mitchell, 1990). Both simple and complex cells showed V-shaped tuning curves for reverse contrast stimuli: i.e. response was minimum at alignment, and increased with increasing vernier offset. For vernier bars with the same contrast sign, γ-shaped tuning curves were found, as reported earlier (Swindale & Cynader, 1986). Sensitivity to offset was inversely correlated in the two paradigms. However, complex cells with high sensitivity to offsets in a normal vernier stimulus were significantly less sensitive to offsets in reverse contrast stimuli. A cell's response to a vernier stimulus in which both bars are bright can be predicted by the shape of its orientation tuning curve, if the vernier stimulus is approximated by a single bar with an orientation equal to that of a line joining the midpoints of the two component bars (Swindale & Cynader, 1986). This approximation did not hold for the reverse contrast condition: orientation tuning curves for compound barswere broad and shallow, rather than bimodal, with peaks up to 40 deg from the preferred orientation. Results from simple cells were compared with predictions made by a linear model of the receptive field. The model predicted the V-shaped tuning curves found for reverse contrast stimuli. It also predicted that absolute values of tuning slopes for vernier offsets in reverse contrast stimuli might sometimes be higher than with normal stimuli. This was observed in some simple cells. The model was unable to explain the shape of orientation tuning curves for compound bars, nor could it explain the breakdown of the equivalent orientation approximation.

Receptive-field characteristics of neurons in cat striate cortex: Changes with visual field eccentricity

1976 ◽  
Vol 39 (3) ◽  
pp. 512-533 ◽  
Author(s):  
J. R. Wilson ◽  
S. M. Sherman
Keyword(s):  
Visual Field ◽  
Receptive Field ◽  
Striate Cortex ◽  
Ocular Dominance ◽  
Receptive Fields ◽  
Direction Selectivity ◽  
Orientation Selectivity ◽  
Simple Cells ◽  
Complex Cells ◽  
Cat Striate Cortex

1. Receptive-field properties of 214 neurons from cat striate cortex were studied with particular emphasis on: a) classification, b) field size, c) orientation selectivity, d) direction selectivity, e) speed selectivity, and f) ocular dominance. We studied receptive fields located throughtout the visual field, including the monocular segment, to determine how receptivefield properties changed with eccentricity in the visual field.2. We classified 98 cells as "simple," 80 as "complex," 21 as "hypercomplex," and 15 in other categories. The proportion of complex cells relative to simple cells increased monotonically with receptive-field eccenticity.3. Direction selectivity and preferred orientation did not measurably change with eccentricity. Through most of the binocular segment, this was also true for ocular dominance; however, at the edge of the binocular segment, there were more fields dominated by the contralateral eye.4. Cells had larger receptive fields, less orientation selectivity, and higher preferred speeds with increasing eccentricity. However, these changes were considerably more pronounced for complex than for simple cells.5. These data suggest that simple and complex cells analyze different aspects of a visual stimulus, and we provide a hypothesis which suggests that simple cells analyze input typically from one (or a few) geniculate neurons, while complex cells receive input from a larger region of geniculate neurons. On average, this region is invariant with eccentricity and, due to a changing magnification factor, complex fields increase in size with eccentricity much more than do simple cells. For complex cells, computations of this geniculate region transformed to cortical space provide a cortical extent equal to the spread of pyramidal cell basal dendrites.

Binocular Combination of Contrast Signals by Striate Cortical Neurons in the Monkey

1997 ◽  
Vol 78 (1) ◽  
pp. 366-382 ◽  
Author(s):  
Earl L. Smith ◽  
Yuzo Chino ◽  
Jinren Ni ◽  
Han Cheng
Keyword(s):  
Cortical Neurons ◽  
Striate Cortex ◽  
Receptive Fields ◽  
Spatial Summation ◽  
Simple Cells ◽  
Complex Cells ◽  
Threshold Response ◽  
Binocular Interactions ◽  
Specific Complex ◽  
Left And Right

Smith, Earl L., III, Yuzo Chino, Jinren Ni, and Han Cheng. Binocular combination of contrast signals by striate cortical neurons in the monkey. J. Neurophysiol. 78: 366–382, 1997. With the use of microelectrode recording techniques, we investigated how the contrast signals from the two eyes are combined in individual cortical neurons in the striate cortex of anesthetized and paralyzed macaque monkeys. For a given neuron, the optimal spatial frequency, orientation, and direction of drift for sine wave grating stimuli were determined for each eye. The cell's disparity tuning characteristics were determined by measuring responses as a function of the relative interocular spatial phase of dichoptic stimuli that consisted of the optimal monocular gratings. Binocular contrast summation was then investigated by measuring contrast response functions for optimal dichoptic grating pairs that had left- to right-eye interocular contrast ratios that varied from 0.1 to 10. The goal was to determine the left- and right-eye contrast components required to produce a criterion threshold response. For all functional classes of cortical neurons and for both cooperative and antagonistic binocular interactions, there was a linear relationship between the left- and right-eye contrast components required to produce a threshold response. Thus, for example for cooperative binocular interactions, a reduction in contrast to one eye was counterbalanced by an equivalent increase in contrast to the other eye. These results showed that in simple cells and phase-specific complex cells, the contrast signals from the two eyes were linearly combined at the subunit level before nonlinear rectification. In non-phase-specific complex cells, the linear binocular convergence of contrast signals could have taken place either before or after the rectification process, but before spike generation. In addition, for simple cells, vector analysis of spatial summation showed that the inputs from the two eyes were also combined in a linear manner before nonlinear spike-generating mechanisms. Thus simple cells showed linear spatial summation not only within and between subregions in a given receptive field, but also between the left- and right-eye receptive fields. Overall, the results show that the effectiveness of a stimulus in producing a response reflects interocular differences in the relative balance of inputs to a given cell, however, the eye of origin of a light-evoked signal has no specific consequence.

The effect of threshold on the relationship between the receptive-field profile and the spatial-frequency tuning cure in simple cells of the cat's striate cortex

1989 ◽  
Vol 3 (5) ◽  
pp. 445-454 ◽  
Author(s):  
Y. Tadmor ◽  
D. J. Tolhurst
Keyword(s):  
Fourier Transform ◽  
Spatial Frequency ◽  
Fourier Transforms ◽  
Frequency Tuning ◽  
Weighting Function ◽  
Tuning Curve ◽  
Spatial Summation ◽  
Response Threshold ◽  
Simple Cells ◽  
Spatial Frequency Tuning

AbstractIt is believed that spatial summation in most simple cells is a linear process. If this were so, then the Fourier transform of a simple cell's line weighting function should predict the cell's spatial frequency tuning curve. We have compared such predictions with experimental measurements and have found a consistent discrepancy: the predicted tuning curve is much too broad. We show qualitatively that this kind of discrepancy is consistent with the well-known threshold nonlinearity shown by most cortical cells. We have tested quantitatively whether a response threshold could explain the observed disagreements between predictions and measurements: a least-squares minimization routine was used to fit the inverse Fourier Transform of the measured frequency tuning curve to the measured line weighting function. The fitting procedure permitted us to introduce a threshold to the reconstructed line weighting function. The results of the analysis show that, for all of the cells tested, the Fourier transforms produced better predictions when a response threshold was included in the model. For some cells, the actual magnitude of the response threshold was measured independently and found to be compatible with that suggested by the model. The effects of nonlinearities of spatial summation are considered.

Comparison of contrast-normalization and threshold models of the responses of simple cells in cat striate cortex

1997 ◽  
Vol 14 (2) ◽  
pp. 293-309 ◽  
Author(s):  
D. J. Tolhurst ◽  
D. J. Heeger
Keyword(s):  
Receptive Field ◽  
Striate Cortex ◽  
Frequency Tuning ◽  
Tuning Curve ◽  
Threshold Model ◽  
Output Stage ◽  
Simple Cells ◽  
Neurophysiological Data ◽  
Contrast Normalization ◽  
The Relationship

AbstractIn almost every study of the linearity of spatiotemporal summation in simple cells of the cat's visual cortex, there have been systematic mismatches between the experimental observations and the predictions of the linear theory. These mismatches have generally been explained by supposing that the initial spatiotemporal summation stage is strictly linear, but that the following output stage of the simple cell is subject to some contrast-dependent nonlinearity. Two main models of the output nonlinearity have been proposed: the threshold model (e.g. Tolhurst & Dean, 1987) and the contrast-normalization model (e.g. Heeger, 1992a, b). In this paper, the two models are fitted rigorously to a variety of previously published neurophysiological data, in order to determine whether one model is a better explanation of the data. We reexamine data on the interaction between two bar stimuli presented in different parts of the receptive field; on the relationship between the receptive-field map and the inverse Fourier transform of the spatial-frequency tuning curve; on the dependence of response amplitude and phase on the spatial phase of stationary gratings; on the relationships between the responses to moving and modulated gratings; and on the suppressive action of gratings moving in a neuron's nonpreferred direction. In many situations, the predictions of the two models are similar, but the contrast-normalization model usually fits the data slightly better than the threshold model, and it is easier to apply the equations of the normalization model. More importantly, the normalization model is naturally able to account very well for the details and subtlety of the results in experiments where the total contrast energy of the stimuli changes; some of these phenomena are completely beyond the scope of the threshold model. Rigorous application of the models' equations has revealed some situations where neither model fits quite well enough, and we must suppose, therefore, that there are some subtle nonlinearities still to be characterized.

Quantitative studies of single-cell properties in monkey striate cortex. II. Orientation specificity and ocular dominance

1976 ◽  
Vol 39 (6) ◽  
pp. 1320-1333 ◽  
Author(s):  
P. H. Schiller ◽  
B. L. Finlay ◽  
S. F. Volman
Keyword(s):  
Striate Cortex ◽  
Ocular Dominance ◽  
Direction Selectivity ◽  
Orientation Selectivity ◽  
Neural Mechanisms ◽  
Orientation Tuning ◽  
Cortical Layers ◽  
Quantitative Studies ◽  
Quantitative Analyses ◽  
Orientation Specificity

1. Quantitative analyses of orientation specificity and ocular dominance were carried out in striate cortex of the rhesus monkey. 2. Sharpness of orientation selectivity was greater for simple (S type) than for complex (CX type) cells. CX-type cells became more broadly tuned in the deeper cortical layers: S-type cells were equally well tuned throughout the cortex. 3. Sharpness of orientation selectivity for S-type cells was similar at all retinal eccentricities studied (0 degrees - 20 degrees from the fovea):in CX-type cells orientation selectivity decreased slightly with increasing eccentricity. 4. The orientation tuning of binocular cells was similar when mapped separately through each eye. 5. Orientation selectivity and direction selectivity are independent of each other, suggesting that separate neural mechanisms give rise to them. 6. More CX-type cells can be binocularly activated than S-type cells (88% versus 49%). The ocular dominance of S-type cells is similar in all cortical layers: for CX-type cells there is an increase in the number of cells in ocular-dominance category 4 in layers 5 and 6.

Oblique Effect: A Neural Basis in the Visual Cortex

2003 ◽  
Vol 90 (1) ◽  
pp. 204-217 ◽  
Author(s):  
Baowang Li ◽  
Matthew R. Peterson ◽  
Ralph D. Freeman
Keyword(s):  
Striate Cortex ◽  
Oblique Effect ◽  
Orientation Tuning ◽  
Neural Basis ◽  
Linear Filters ◽  
Simple Cells ◽  
Tuning Curves ◽  
Complex Cells ◽  
Behavioral Studies ◽  
Spatial Frequencies

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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