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.

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.

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 Learning and Adaptation in a Recurrent Model of V1 Orientation Selectivity

2003 ◽  
Vol 89 (4) ◽  
pp. 2086-2100 ◽  
Author(s):  
Andrew F. Teich ◽  
Ning Qian
Keyword(s):  
Tuning Curve ◽  
Human Subjects ◽  
Peak Shift ◽  
Orientation Selectivity ◽  
Orientation Tuning ◽  
Physiological Data ◽  
Tuning Curves ◽  
Psychophysical Data ◽  
Learning And Adaptation ◽  
Experimental Findings

Learning and adaptation in the domain of orientation processing are among the most studied topics in the literature. However, little effort has been devoted to explaining the diverse array of experimental findings via a physiologically based model. We have started to address this issue in the framework of the recurrent model of V1 orientation selectivity and found that reported changes in V1 orientation tuning curves after learning and adaptation can both be explained with the model. Specifically, the sharpening of orientation tuning curves near the trained orientation after learning can be accounted for by slightly reducing net excitatory connections to cells around the trained orientation, while the broadening and peak shift of the tuning curves after adaptation can be reproduced by appropriately scaling down both excitation and inhibition around the adapted orientation. In addition, we investigated the perceptual consequences of the tuning curve changes induced by learning and adaptation using signal detection theory. We found that in the case of learning, the physiological changes can account for the psychophysical data well. In the case of adaptation, however, there is a clear discrepancy between the psychophysical data from alert human subjects and the physiological data from anesthetized animals. Instead, human adaptation studies can be better accounted for by the learning data from behaving animals. Our work suggests that adaptation in behaving subjects may be viewed as a short-term form of learning.

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...

scholarly journals Spatial and Temporal Features of Synaptic to Discharge Receptive Field Transformation in Cat Area 17

2010 ◽  
Vol 103 (2) ◽  
pp. 677-697 ◽  
Author(s):  
Lionel G. Nowak ◽  
Maria V. Sanchez-Vives ◽  
David A. McCormick
Keyword(s):  
Spatial Frequency ◽  
Response Latency ◽  
Frequency Tuning ◽  
Receptive Fields ◽  
Width Ratio ◽  
Area 17 ◽  
Simple Cells ◽  
Complex Cells ◽  
Temporal Features ◽  
Simple And Complex Cells

The aim of the present study was to characterize the spatial and temporal features of synaptic and discharge receptive fields (RFs), and to quantify their relationships, in cat area 17. For this purpose, neurons were recorded intracellularly while high-frequency flashing bars were used to generate RFs maps for synaptic and spiking responses. Comparison of the maps shows that some features of the discharge RFs depended strongly on those of the synaptic RFs, whereas others were less dependent. Spiking RF duration depended poorly and spiking RF amplitude depended moderately on those of the underlying synaptic RFs. At the other extreme, the optimal spatial frequency and phase of the discharge RFs in simple cells were almost entirely inherited from those of the synaptic RFs. Subfield width, in both simple and complex cells, was less for spiking responses compared with synaptic responses, but synaptic to discharge width ratio was relatively variable from cell to cell. When considering the whole RF of simple cells, additional variability in width ratio resulted from the presence of additional synaptic subfields that remained subthreshold. Due to these additional, subthreshold subfields, spatial frequency tuning predicted from synaptic RFs appears sharper than that predicted from spiking RFs. Excitatory subfield overlap in spiking RFs was well predicted by subfield overlap at the synaptic level. When examined in different regions of the RF, latencies appeared to be quite variable, but this variability showed negligible dependence on distance from the RF center. Nevertheless, spiking response latency faithfully reflected synaptic response latency.

scholarly journals Substructure of Direction-Selective Receptive Fields in Macaque V1

2003 ◽  
Vol 89 (5) ◽  
pp. 2743-2759 ◽  
Author(s):  
Margaret S. Livingstone ◽  
Bevil R. Conway
Keyword(s):  
Receptive Fields ◽  
Direction Selectivity ◽  
Positive Interactions ◽  
Null Direction ◽  
Simple Cells ◽  
Complex Cells ◽  
Preferred Direction ◽  
Orientation Axis ◽  
Interaction Regions ◽  
Simple And Complex Cells

We used two-dimensional (2-D) sparse noise to map simultaneous and sequential two-spot interactions in simple and complex direction-selective cells in macaque V1. Sequential-interaction maps for both simple and complex cells showed preferred-direction facilitation and null-direction suppression for same-contrast stimulus sequences and the reverse for inverting-contrast sequences, although the magnitudes of the interactions were weaker for the simple cells. Contrast-sign selectivity in complex cells indicates that direction-selective interactions in these cells must occur in antecedent simple cells or in simple-cell-like dendritic compartments. Our maps suggest that direction selectivity, and on andoff segregation perpendicular to the orientation axis, can occur prior to receptive-field elongation along the orientation axis. 2-D interaction maps for some complex cells showed elongated alternating facilitatory and suppressive interactions as predicted if their inputs were orientation-selective simple cells. The negative interactions, however, were less elongated than the positive interactions, and there was an inflection at the origin in the positive interactions, so the interactions were chevron-shaped rather than band-like. Other complex cells showed only two round interaction regions, one negative and one positive. Several explanations for the map shapes are considered, including the possibility that directional interactions are generated directly from unoriented inputs.

Space-time spectra of complex cell filters in the macaque monkey: A comparison of results obtained with pseudowhite noise and grating stimuli

1994 ◽  
Vol 11 (4) ◽  
pp. 805-821 ◽  
Author(s):  
James P. Gaska ◽  
Lowell D. Jacobson ◽  
Hai-Wen Chen ◽  
Daniel A. Pollen
Keyword(s):  
Spatial Frequency ◽  
Macaque Monkey ◽  
Spike Rate ◽  
Second Order ◽  
Orientation Tuning ◽  
Complex Cell ◽  
Interaction Function ◽  
Tuning Curves ◽  
Complex Cells ◽  
Order Kernel

AbstractWhite noise stimuli were used to estimate second-order kernels for complex cells in cortical area VI of the macaque monkey, and drifting grating stimuli were presented to the same sample of neurons to obtain orientation and spatial-frequency tuning curves. Using these data, we quantified how well second-order kernels predict the normalized tuning of the average response of complex cells to drifting gratings.The estimated second-order kernel of each complex cell was transformed into an interaction function defined over all spatial and temporal lags without regard to absolute position or delay. The Fourier transform of each interaction function was then computed to obtain an interaction spectrum. For a cell that is well modeled by a second-order system, the cell’s interaction spectrum is proportional to the tuning of its average spike rate to drifting gratings. This result was used to obtain spatial-frequency and orientation tuning predictions for each cell based on its second-order kernel. From the spatial-frequency and orientation tuning curves, we computed peaks and bandwidths, and an index for directional selectivity.We found that the predictions derived from second-order kernels provide an accurate description of the change in the average spike rate of complex cells to single drifting sine–wave gratings. These findings are consistent with a model for complex cells that has a quadratic spectral energy operator at its core but are inconsistent with a spectral amplitude model.

Cone Inputs to Simple and Complex Cells in V1 of Awake Macaque

2007 ◽  
Vol 97 (4) ◽  
pp. 3070-3081 ◽  
Author(s):  
Gregory D. Horwitz ◽  
E. J. Chichilnisky ◽  
Thomas D. Albright
Keyword(s):  
Linear Models ◽  
Noise Analysis ◽  
Receptive Fields ◽  
Light Responses ◽  
Simple Cells ◽  
Complex Cells ◽  
V1 Neurons ◽  
Spike Triggered Averaging ◽  
Simple And Complex Cells ◽  
Simple And Complex Cell

Rules by which V1 neurons combine signals originating in the cone photoreceptors are poorly understood. We measured cone inputs to V1 neurons in awake, fixating monkeys with white-noise analysis techniques that reveal properties of light responses not revealed by purely linear models used in previous studies. Simple cells were studied by spike-triggered averaging that is robust to static nonlinearities in spike generation. This analysis revealed, among heterogeneously tuned neurons, two relatively discrete categories: one with opponent L- and M-cone weights and another with nonopponent cone weights. Complex cells were studied by spike-triggered covariance, which identifies features in the stimulus sequence that trigger spikes in neurons with receptive fields containing multiple linear subunits that combine nonlinearly. All complex cells responded to nonopponent stimulus modulations. Although some complex cells responded to cone-opponent stimulus modulations too, none exhibited the pure opponent sensitivity observed in many simple cells. These results extend the findings on distinctions between simple and complex cell chromatic tuning observed in previous studies in anesthetized monkeys.

scholarly journals Influence of Contrast on Orientation and Temporal Frequency Tuning in Ferret Primary Visual Cortex

2004 ◽  
Vol 91 (6) ◽  
pp. 2797-2808 ◽  
Author(s):  
Henry J. Alitto ◽  
W. Martin Usrey
Keyword(s):  
Visual Cortex ◽  
Primary Visual Cortex ◽  
Cortical Neurons ◽  
Frequency Tuning ◽  
Temporal Frequency ◽  
Orientation Tuning ◽  
Visual Response ◽  
Stimulus Contrast ◽  
Tuning Curves ◽  
Simple And Complex Cells

Neurons in primary visual cortex are highly sensitive to the contrast, orientation, and temporal frequency of a visual stimulus. These three stimulus properties can be varied independently of one another, raising the question of how they interact to influence neuronal responses. We recorded from individual neurons in ferret primary visual cortex to determine the influence of stimulus contrast on orientation tuning, temporal-frequency tuning, and latency to visual response. Results show that orientation-tuning bandwidth is not affected by contrast level. Thus neurons in ferret visual cortex display contrast-invariant orientation tuning. Stimulus contrast does, however, influence the structure of orientation-tuning curves as measures of circular variance vary inversely with contrast for both simple and complex cells. This change in circular variance depends, in part, on a contrast-dependent change in the ratio of null to preferred orientation responses. Stimulus contrast also has an influence on the temporal-frequency tuning of cortical neurons. Both simple and complex cells display a contrast-dependent rightward shift in their temporal frequency-tuning curves that results in an increase in the highest temporal frequency needed to produce a half-maximum response (TF50). Results show that the degree of the contrast-dependent increase in TF50 is similar for cortical neurons and neurons in the lateral geniculate nucleus (LGN) and indicate that subcortical mechanisms likely play a major role in establishing the degree of effect displayed by downstream neurons. Finally, results show that LGN and cortical neurons experience a contrast-dependent phase advance in their visual response. This phase advance is most pronounced for cortical neurons indicating a role for both subcortical and cortical mechanisms.

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.

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