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.

scholarly journals V1 orientation plasticity is explained by broadly tuned feedforward inputs and intracortical sharpening

2010 ◽  
Vol 27 (1-2) ◽  
pp. 57-73 ◽  
Author(s):  
ANDREW F. TEICH ◽  
NING QIAN
Keyword(s):  
Tuning Curve ◽  
Orientation Discrimination ◽  
Orientation Tuning ◽  
Plastic Properties ◽  
Simple Cells ◽  
Tuning Curves ◽  
Complex Cells ◽  
Learning And Adaptation ◽  
Mere Presence ◽  
Simple And Complex Cells

AbstractOrientation adaptation and perceptual learning change orientation tuning curves of V1 cells. Adaptation shifts tuning curve peaks away from the adapted orientation, reduces tuning curve slopes near the adapted orientation, and increases the responses on the far flank of tuning curves. Learning an orientation discrimination task increases tuning curve slopes near the trained orientation. These changes have been explained previously in a recurrent model (RM) of orientation selectivity. However, the RM generates only complex cells when they are well tuned, so that there is currently no model of orientation plasticity for simple cells. In addition, some feedforward models, such as the modified feedforward model (MFM), also contain recurrent cortical excitation, and it is unknown whether they can explain plasticity. Here, we compare plasticity in the MFM, which simulates simple cells, and a recent modification of the RM (MRM), which displays a continuum of simple-to-complex characteristics. Both pre- and postsynaptic-based modifications of the recurrent and feedforward connections in the models are investigated. The MRM can account for all the learning- and adaptation-induced plasticity, for both simple and complex cells, while the MFM cannot. The key features from the MRM required for explaining plasticity are broadly tuned feedforward inputs and sharpening by a Mexican hat intracortical interaction profile. The mere presence of recurrent cortical interactions in feedforward models like the MFM is insufficient; such models have more rigid tuning curves. We predict that the plastic properties must be absent for cells whose orientation tuning arises from a feedforward mechanism.

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.

scholarly journals Comparison Among Some Models of Orientation Selectivity

2006 ◽  
Vol 96 (1) ◽  
pp. 404-419 ◽  
Author(s):  
Andrew F. Teich ◽  
Ning Qian
Keyword(s):  
Peak Shift ◽  
Orientation Tuning ◽  
Complex Spectrum ◽  
Complex Cell ◽  
Phase Information ◽  
Simple Complex ◽  
Simple Cells ◽  
Complex Cells ◽  
Spatial Phase ◽  
Opposite Contrast

Several models exist for explaining primary visual cortex (V1) orientation tuning. The modified feedforward model (MFM) and the recurrent model (RM) are major examples. We have implemented these two models, at the same level of detail, alongside a few newer variations, and thoroughly compared their receptive-field structures. We found that antiphase inhibition in the MFM enhances both spatial phase information and orientation tuning, producing well-tuned simple cells. This remains true for a newer version of the MFM that incorporates untuned complex-cell inhibition. In contrast, when the recurrent connections in the RM are strong enough to produce typical V1 orientation tuning, they also eliminate spatial phase information, making the cells complex. Introducing phase specificity into the connections of the RM (as done in an original version of the RM) can make the cells phase sensitive, but the cells show an incorrect 90° peak shift of orientation tuning under opposite contrast signs. An inhibition-dominant version of the RM can generate well-tuned cells across the simple–complex spectrum, but it predicts that the net effect of cortical interactions is to suppress feedforward excitation across all orientations in simple cells. Finally, adding antiphase inhibition used in the MFM into the RM produces a most general model. We call this new model the modified recurrent model (MRM) and show that this model can also produce well-tuned cells throughout the simple–complex spectrum. Unlike the inhibition-dominant RM, the MRM is consistent with data from cat V1, suggesting that the net effect of cortical interactions is to boost simple cell responses at the preferred orientation. These results suggest that the MFM is well suited for explaining orientation tuning in simple cells, whereas the standard RM is for complex cells. The assignment of the RM to complex cells also avoids conflicts between the RM and the experiments of cortical inactivation (done on simple cells) and the spatial-frequency dependency of orientation tuning (found in simple cells). Because orientation-tuned V1 cells show a continuum of simple- to complex-cell behavior, the MRM provides the best description of V1 data.

scholarly journals Local Signals From Beyond the Receptive Fields of Striate Cortical Neurons

2003 ◽  
Vol 90 (2) ◽  
pp. 822-831 ◽  
Author(s):  
James R. Müller ◽  
Andrew B. Metha ◽  
John Krauskopf ◽  
Peter Lennie
Keyword(s):  
Receptive Field ◽  
Cortical Neurons ◽  
Striate Cortex ◽  
Image Representation ◽  
Receptive Fields ◽  
Preferred Orientation ◽  
Orientation Tuning ◽  
Stimulus Space ◽  
Tuning Curves ◽  
Complex Cells

We examined in anesthetized macaque how the responses of a striate cortical neuron to patterns inside the receptive field were altered by surrounding patterns outside it. The changes in a neuron's response brought about by a surround are immediate and transient: they arise with the same latency as the response to a stimulus within the receptive field (this argues for a source locally in striate cortex) and become less effective as soon as 27 ms later. Surround signals appeared to exert their influence through divisive interaction (normalization) with those arising in the receptive field. Surrounding patterns presented at orientations slightly oblique to the preferred orientation consistently deformed orientation tuning curves of complex (but not simple) cells, repelling the preferred orientation but without decreasing the discriminability of the preferred grating and ones at slightly oblique orientations. By reducing responsivity and changing the tuning of complex cells locally in stimulus space, surrounding patterns reduce the correlations among responses of neurons to a particular stimulus, thus reducing the redundancy of image representation.

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.

Functional implications of cross-orientation inhibition of cortical visual cells. I. Neurophysiological evidence

1982 ◽  
Vol 216 (1204) ◽  
pp. 335-354 ◽  
Keyword(s):  
Striate Cortex ◽  
Discharge Rate ◽  
Inhibitory Effect ◽  
Visual Noise ◽  
Simple Cells ◽  
One Dimensional ◽  
Complex Cells ◽  
Visual Cells ◽  
Visual Objects ◽  
Simple And Complex Cells

Simple and complex cells of striate cortex of anaesthetized and paralysed cats were stimulated with two superimposed one-dimensional grating stimuli of different orientations to investigate inhibitory effects of non-optimally oriented stimuli. We confirmed that a stimulus of orientation orthogonal to a cell’s long axis significantly reduces the cell’s discharge rate. Further experiments revealed the following, (i) The inhibition was typically stronger for simple than for complex cells, (ii) It is very broadly tuned for orientation, all orientations outside the cell’s tuning band having a comparable inhibitory effect. (iii) Similarly, it is broadly tuned for spatial frequency. These last two results suggest that the inhibition arises not from a single cell but from a pool of cells, (iv) The pattern of the discharge of the inhibition in response to stimulation by phase-reversed sinusoidal gratings is consistent with the notion that the inhibition arises from complex cells. A second series of recordings of stimulation by visual noise patterns demonstrated how ‘cross-orientation inhibition’ prevents simple cells from responding to two-dimensional visual noise while allowing them to respond to comparable one-dimensional noise patterns. We suggest that this mechanism may serve to render simple cells selectively sensitive to one-dimensional stimuli, such as the contours or borders of visual objects.

Direction-selective adaptation in simple and complex cells in cat striate cortex

1988 ◽  
Vol 59 (4) ◽  
pp. 1314-1330 ◽  
Author(s):  
S. G. Marlin ◽  
S. J. Hasan ◽  
M. S. Cynader
Keyword(s):  
Cortical Neurons ◽  
Striate Cortex ◽  
Selective Adaptation ◽  
Direction Selectivity ◽  
Simple Cells ◽  
Complex Cells ◽  
Preferred Direction ◽  
Directional Preference ◽  
Simple And Complex Cells ◽  
Cat Striate Cortex

1. The selectivity of adaptation to unidirectional motion was examined in neurons of the cat striate cortex. Following prolonged stimulation with a unidirectional high-contrast grating, the responsivity of cortical neurons was reduced. In many units this decrease was restricted to the direction of prior stimulation. This selective adaptation produced changes in the degree of direction selectivity of the cortical units (as measured by the ratio of the response to motion in the preferred direction to that in the nonpreferred direction). 2. The initial strength of the directional preference of a given cortical unit did not determine the degree of direction-selective adaptation. Indeed, even non-direction-selective units could exhibit pronounced direction-selective adaptation. The degree of direction-selective adaptation was also independent of the overall decrease in responsivity during adaptation. 3. There was no difference between simple and complex cells in the total amount of adaptation observed. The selectivity of the adaptation, however, did differ between these two cell types. As a group, simple cells showed significant direction-selective adaptation, whereas complex cells did not. The directional preference of most simple cells decreased following preferred direction adaptation and many highly direction selective simple cells became non-direction selective. In addition, simple cells became significantly more direction selective following nonpreferred direction adaptation. 4. Some complex cells also demonstrated direction-selective adaptation. There was, however, much more variability among complex cells than simple cells. Some complex cells actually increased direction selectivity following preferred direction adaptation. These differences between simple and complex cells suggest that changes in direction selectivity following unidirectional adaptation are not due to simple neuronal fatigue of the unit being recorded, but depend on selective adaptation of afferent inputs to the unit. 5. The spontaneous activity of many cortical neurons decreased following preferred direction adaptation but increased following adaptation in the nonpreferred direction. The response to a stationary grating also decreased following preferred direction adaptation. However, there was very little change in the response to a stationary grating following adaptation in the nonpreferred direction.

scholarly journals Binocular Spatial Phase Tuning Characteristics of Neurons in the Macaque Striate Cortex

1997 ◽  
Vol 78 (1) ◽  
pp. 351-365 ◽  
Author(s):  
Earl L. Smith ◽  
Yuzo M. Chino ◽  
Jinren Ni ◽  
William H. Ridder ◽  
M.L.J. Crawford
Keyword(s):  
Spatial Frequency ◽  
Striate Cortex ◽  
Orientation Tuning ◽  
Absolute Position ◽  
Simple Cells ◽  
Complex Cells ◽  
Spatial Phase ◽  
Binocular Interactions ◽  
Phase Tuning ◽  
Phase Sensitive

Smith, Earl L., III, Yuzo M. Chino, Jinren Ni, William H. Ridder III, and M.L.J. Crawford. Binocular spatial phase tuning characteristics of neurons in the macaque striate cortex. J. Neurophysiol. 78: 351–365, 1997. We employed microelectrode recording techniques to study the sensitivity of individual neurons in the striate cortex of anesthetized and paralyzed monkeys to relative interocular image disparities and to determine the effects of basic stimulus parameters on these cortical binocular interactions. The visual stimuli were drifting sine wave gratings. After the optimal stimulus orientation, spatial frequency, and direction of stimulus movement were found, the cells' disparity tuning characteristics were determined by measuring responses as a function of the relative interocular spatial phase of dichoptic grating pairs. No attempts were made to assess absolute position disparities or horizontal disparities relative to the horopter. The majority (∼70%) of simple cells were highly sensitive to interocular spatial phase disparities, particularly neurons with balanced ocular dominances. Simple cells typically demonstrated binocular facilitation at the optimal phase disparity and binocular suppression for disparities 180° away. Fewer complex cells were phase selective (∼40%); however, the range of disparity selectivity in phase-sensitive complex cells was comparable with that for simple cells. Binocular interactions in non-phase-sensitive complex cells were evidenced by binocular response amplitudes that differed from responses to monocular stimulation. The degree of disparity tuning was independent of a cell's optimal orientation or the degree of direction tuning. However, disparity-sensitive cells tended to have narrow orientation tuning functions and the degree of disparity tuning was greatest for the optimal stimulus orientations. Rotating the stimulus for one eye 90° from the optimal orientation usually eliminated binocular interactions. The effects of phase disparities on the binocular response amplitude were also greatest at the optimal spatial frequency. Thus a cell's sensitivity to absolute position disparities reflects its spatial tuning characteristics, with cells sensitive to high spatial frequencies being capable of signaling very small changes in image disparity. On the other hand, stimulus contrast had relatively little effect on a cell's disparity tuning, because response saturation occurred at the same contrast level for all relative interocular phase disparities. Thus, as with orientation tuning, a cell's optimal disparity and the degree of disparity selectivity were invariant with contrast. Overall, the results show that sensitivity to interocular spatial phase disparities is a common property of striate neurons. A cell's disparity tuning characteristics appear to largely reflect its monocular receptive field properties and the interocular balance between excitatory and inhibitory inputs. However, distinct functional classes of cortical neurons could not be discriminated on the basis of disparity sensitivity alone.

Discrimination of orientation and position disparities by binocularly activated neurons in cat straite cortex

1977 ◽  
Vol 40 (2) ◽  
pp. 260-283 ◽  
Author(s):  
J. I. Nelson ◽  
H. Kato ◽  
P. O. Bishop
Keyword(s):  
Ocular Dominance ◽  
Stimulus Orientation ◽  
Orientation Tuning ◽  
Simple Cells ◽  
Tuning Curves ◽  
Complex Cells ◽  
Eye Rotation ◽  
Striate Neurons ◽  
The Mean ◽  
Orientation Disparity

1. We have examined and compared the ability of binocularly activated striate neurons to make both position disparity and orientation disparity discrimination in the anesthetized (N2O/O2) and paralyzed cat preparation. 2. Accurate knowledge of eye position is essential for disparity studies. Using a retinal projection technique able to detect eye drifts of less than 3' arc per retinal landmark and less than 18' arc cyclorotation disparity, we determined eye drift during the course of 2- to 4-day experiments. After the initial eye rotation due to the anesthesia and the onset of paralysis (see below), rotational drift thereafter was mainly excyclorotatory and, from all causes, rarely totaled more than 4 degrees disparity. All our data have been corrected for this residual cyclorotatory drift. 3. Optimal stimulus orientation disparities were determined from quantitative monocular orientation tuning curves for 74 binocularly activated striate cells (37 simple, 3 hypercomplex I, 31 complex, 3 hypercomplex II) from nine cats. Without exception, the mean optimal stimulus orientation disparity in each of our animals showed a departure from zero disparity equivalent to an incyclorotation of the eyes (mean, 9.2 degrees; range, 2.7 degrees-15.9 degrees). 4. We attribute this mean optimal stimulus orientation disparity shift to ocular cyclorotation as a result of the initial anesthesia and paralysis. Assuming equal intortion, incyclorotation for each eye averages 4.6 degrees. On the assumption that the mean optimal stimulus orientation disparity is zero in normal life, we pooled results from the nine animals about their individual means. For the 74 cells the resulting distribution of the optimal stimulus orientation disparities had a range of about +/-15 degrees (simple cells: SD 4.9 degrees; complex cells: SD 7.4 degrees). 5. We examined the relationship of the sharpness of the orientation tuning curves to ocular dominance, to absolute orientation preference, and to other unit properties. The striking observation was the high correlation between the sharpness of orientation tuning curves for the two eyes of a binocular neuron. For simple cells the mean difference for the half-widths of half-height was only 2.54 degrees, with sharpness showing a high correlation between the two eyes (r=0.915) over half-width at half-heights ranging from 8.5 degrees to 41.8 degrees. Complex cells showed a similar, albeit weaker, correlation. 6. Having shown that, assessed monocularly binocular units show different orientation tunings in the two eyes, we undertook binocular experiments to ascertain if these differences were the optimal disparities of sharply tuned stimulus orientation disparity channels. Using a matrix stimulation paradigm to minimize the effects of spontaneous changes in responsiveness, we have simultaneously extracted bionocular stimulus orientation disparity and position disparity tuning curves from single striate neurons...

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