Length and width tuning of neurons in the cat's primary visual cortex

1994 ◽  
Vol 71 (1) ◽  
pp. 347-374 ◽  
Author(s):  
G. C. DeAngelis ◽  
R. D. Freeman ◽  
I. Ohzawa
Keyword(s):  
Receptive Field ◽  
Spatial Frequency ◽  
Spatial Organization ◽  
Striate Cortex ◽  
Frequency Tuning ◽  
Variable Parameter ◽  
Visual Space ◽  
Similar Degree ◽  
Tuning Curves ◽  
Number Of Cycles

1. The classically defined receptive field of a visual neuron is the area of visual space over which the cell responds to visual stimuli. It is well established, however, that the discharge produced by an optimal stimulus can be modulated by the presence of additional stimuli that by themselves do not produce any response. This study examines inhibitory influences that originate from areas located outside of the classical (i.e., excitatory) receptive field. Previous work has shown that for some cells the response to a properly oriented bar of light becomes attenuated when the bar extends beyond the receptive field, a phenomenon known as end-inhibition (or length tuning). Analogously, it has been shown that increasing the number of cycles of a drifting grating stimulus may also inhibit the firing of some cells, an effect known as side-inhibition (or width tuning). Very little information is available, however, about the relationship between end- and side-inhibition. We have examined the spatial organization and tuning characteristics of these inhibitory effects by recording extracellularly from single neurons in the cat's striate cortex (Area 17). 2. For each cortical neuron, length and width tuning curves were obtained with the use of rectangular patches of drifting sinusoidal gratings that have variable length and width. Results from 82 cells show that the strengths of end- and side-inhibition tend to be correlated. Most cells that exhibit clear end-inhibition also show a similar degree of side-inhibition. For these cells, the excitatory receptive field is surrounded on all sides by inhibitory zones. Some cells exhibit only end- or side-inhibition, but not both. Data for 28 binocular cells show that length and width tuning curves for the dominant and nondominant eyes tend to be closely matched. 3. We also measured tuning characteristics of end- and side-inhibition. To obtain these data, the excitatory receptive field was stimulated with a grating patch having optimal orientation, spatial frequency, and size, whereas the end- or side-inhibitory regions were stimulated with patches of gratings that had a variable parameter (such as orientation). Results show that end- and side-inhibition tend to be strongest at the orientation and spatial frequency that yield maximal excitation. However, orientation and spatial frequency tuning curves for inhibition are considerably broader than those for excitation, suggesting that inhibition is mediated by a pool of neurons.(ABSTRACT TRUNCATED AT 400 WORDS)

Effects of rearing kittens with convergent strabismus on development of receptive-field properties in striate cortex neurons

1983 ◽  
Vol 50 (1) ◽  
pp. 265-286 ◽  
Author(s):  
Y. M. Chino ◽  
M. S. Shansky ◽  
W. L. Jankowski ◽  
F. A. Banser
Keyword(s):  
Receptive Field ◽  
Spatial Frequency ◽  
Striate Cortex ◽  
Frequency Tuning ◽  
Tuning Curves ◽  
Spatial Frequency Tuning ◽  
Receptive Field Properties ◽  
Convergent Strabismus ◽  
Striate Neurons ◽  
Contrast Thresholds

Convergent strabismus was induced surgically in seven kittens at 3 wk of age. Recordings were made in 131 cells in the striate cortex of these strabismic kittens after maturation, and the results were compared to the data obtained from 140 striate neurons in normally reared cats. All our samples had receptive fields (RFs) within 5 degrees of the area centralis. The spatial and temporal response properties were quantitatively analyzed by using drifting sinusoidal gratings of various contrasts as well as spatial and temporal frequencies. In contrast to other reports, the receptive-field properties of the striate neurons exclusively driven or dominated by the deviating eye were quite abnormal. The spatial frequency tuning curves in strabismic cats were different from those obtained from normals in that the optimal spatial frequencies and spatial resolutions were shifted to lower values and the bandwidths were significantly broader. The contrast-response functions show that contrast thresholds, measured at the optimal spatial frequency, were significantly higher and the slope of the functions much flatter in strabismic animals. Moreover, receptive-field sizes were much larger and the sharpness of orientation tuning was reduced in strabismic cats. Direction selectivity, however, was normal in those animals. The temporal frequency tuning curves were also abnormal, particularly in that temporal resolution was considerably reduced in strabismic cats compared with normally reared cats. In addition, many cells in strabismic animals exhibited much longer latencies to visual and optic chiasm (OX) stimulations. All these effects, to our great surprise, were also observed in the striate units controlled by the nondeviating eye, although the magnitude of the alteration depends on the receptive-field property. The abnormalities found in spatial frequency tuning, contrast thresholds, and RF sizes were as severe as those revealed in the deviating eye. On the other hand, the effects on other RF properties were minimal or less severe. These physiological findings correspond very well with results from behavioral measurements of spatial contrast sensitivity in the same cats reported elsewhere. It is concluded from these results that the nondeviating eye in convergent strabismic animals, and perhaps humans, should not always be presumed "normal."

Contrast-Dependent Changes in Spatial Frequency Tuning of Macaque V1 Neurons: Effects of a Changing Receptive Field Size

2002 ◽  
Vol 88 (3) ◽  
pp. 1363-1373 ◽  
Author(s):  
Michael P. Sceniak ◽  
Michael J. Hawken ◽  
Robert Shapley
Keyword(s):  
Receptive Field ◽  
Spatial Frequency ◽  
Frequency Tuning ◽  
Fundamental Property ◽  
Spatial Summation ◽  
Field Size ◽  
Stimulus Contrast ◽  
Tuning Curves ◽  
Spatial Frequency Tuning ◽  
Receptive Field Organization

Previous studies on single neurons in primary visual cortex have reported that selectivity for orientation and spatial frequency tuning do not change with stimulus contrast. The prevailing hypothesis is that contrast scales the response magnitude but does not differentially affect particular stimuli. Models where responses are normalized over contrast to maintain constant tuning for parameters such as orientation and spatial frequency have been proposed to explain these results. However, our results indicate that a fundamental property of receptive field organization, spatial summation, is not contrast invariant. We examined the spatial frequency tuning of cells that show contrast-dependent changes in spatial summation and have found that spatial frequency selectivity also depends on stimulus contrast. These results indicate that contrast changes in the spatial frequency tuning curves result from spatial reorganization of the receptive field.

Spatiotemporal organization of simple-cell receptive fields in the cat's striate cortex. II. Linearity of temporal and spatial summation

1993 ◽  
Vol 69 (4) ◽  
pp. 1118-1135 ◽  
Author(s):  
G. C. DeAngelis ◽  
I. Ohzawa ◽  
R. D. Freeman
Keyword(s):  
Receptive Field ◽  
Striate Cortex ◽  
Frequency Tuning ◽  
Temporal Frequency ◽  
Spatial Summation ◽  
Direction Selectivity ◽  
Simple Cells ◽  
Tuning Curves ◽  
Spatiotemporal Receptive Field ◽  
Temporal And Spatial

1. We have tested the hypothesis that simple cells in the cat's visual cortex perform a linear spatiotemporal filtering of the visual image. To conduct this study we note that a visual neuron behaves linearly if the responses to small, brief flashes of light are mathematically related, via the Fourier transform, to the responses elicited by sinusoidal grating stimuli. 2. We have evaluated the linearity of temporal and spatial summation for 118 simple cells recorded from the striate cortex (area 17) of adult cats and kittens at ages 4 and 8 wk postnatal. These neurons represent a subset of the population of cells for which we have described the postnatal development of spatiotemporal receptive-field structure in the preceding paper. Spatiotemporal receptive-field profiles are constructed, with the use of a reverse correlation technique, from the responses to random sequences of small bar stimuli that are brighter or darker than the background. Fourier analysis of spatiotemporal receptive-field profiles yields linear predictions of the cells' spatial and temporal frequency tuning. These predicted responses are compared with spatial and temporal frequency tuning curves measured by the use of drifting, sinusoidal-luminance grating stimuli. 3. For most simple cells, there is good agreement between spatial and temporal frequency tuning curves predicted from the receptive-field profile and those measured by the use of sinusoidal gratings. These results suggest that both spatial and temporal summation within simple cells are approximately linear. There is a tendency for predicted tuning curves to be slightly broader than measured tuning curves, a finding that is consistent with the effects of a threshold nonlinearity at the output of these neurons. In some cases, however, predicted tuning curves deviate from measured responses only at low spatial and temporal frequencies. This cannot be explained by a simple threshold nonlinearity. 4. If linearity is assumed, it should be possible to predict the direction selectivity of simple cells from the structure of their spatiotemporal receptive-field profiles. For virtually all cells, linear predictions correctly determine the preferred direction of motion of a visual stimulus. However, the strength of the directional bias is typically underestimated by a factor of about two on the basis of linear predictions. Consideration of the expansive exponential nonlinearity revealed in the contrast-response function permits a reconciliation of the discrepancy between measured and predicted direction selectivity indexes. 5. Overall, these findings show that spatiotemporal receptive-field profiles obtained with the use of reverse correlation may be used to predict a variety of response properties for simple cells.(ABSTRACT TRUNCATED AT 400 WORDS)

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

1987 ◽  
Vol 57 (3) ◽  
pp. 773-786 ◽  
Author(s):  
B. C. Skottun ◽  
A. Bradley ◽  
G. Sclar ◽  
I. Ohzawa ◽  
R. D. Freeman
Keyword(s):  
Spatial Frequency ◽  
Striate Cortex ◽  
Frequency Tuning ◽  
Single Cells ◽  
Frequency Discrimination ◽  
Frequency Selectivity ◽  
Response Amplitude ◽  
Visual Orientation ◽  
Characteristic Analysis ◽  
Response Variance

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.

Harmonic basis functions for spatial coding in the cat striate cortex

1989 ◽  
Vol 3 (4) ◽  
pp. 351-363 ◽  
Author(s):  
V. D. Glezer ◽  
V. V. Yakovlev ◽  
V. E. Gauzelman
Keyword(s):  
Spatial Frequency ◽  
Striate Cortex ◽  
Weighting Function ◽  
Discrete Distribution ◽  
Basis Functions ◽  
Spatial Coding ◽  
Central Visual Field ◽  
Spatial Frequencies ◽  
The Absolute ◽  
Number Of Cycles

AbstractThe number of subregions in the activity profiles of simple cells varies in different cells from 2–8; that is, the number of cycles in the weighting function varies from 1–4. The distribution of receptive-field (RF) sizes at eccentricities of 0-6 deg are clustered at half-octave intervals and form a discrete distribution with maxima at 0.62, 0.9, 1.24, 1.8, 2.48, and 3.4 deg. The spatial frequencies to which the cells are tuned are also clustered at half-octave intervals, forming a discrete distribution peaking at 0.45, 0.69, 0.9, 1.35, 1.88, 2.7, 3.8, and 5.6 cycles/deg. If we divide the RF sizes by the size of the period of the subregions, then the average indices of complexity (really existing) or the number of cycles in the weighting function form (after normalization) the sequences: 1, 1.41, 2.0, 2.9, 4.15.The relation between the bandwidth of the spatial-frequency characteristic and the optimal spatial frequency is in accordance with predictions of the Fourier hypothesis. The absolute bandwidth does not change with the number of cycles/module. This means that inside the module the absolute bandwidth does not change with the number of the harmonic. The results allow us to suggest the following. A module of the striate cortex, which is a group of cells with RFs of equal size projected onto the same area of central visual field, accounts for the Fourier description of the image. The basis functions of the module are composed of four harmonics only, irrespective of size and position of the module.Besides linear cells (sinusoidal and cosinusoidal elements), the module contains nonlinear cells, performing a nonlinear summation of the responses of sinusoidal and cosinusoidal elements. Such cells are characterized by an index of complexity which is more than the number of cycles in the weighting function and by marked overlap of ON and OFF zones. The analysis of organization suggests that the cells can measure the amplitude and phase of the stimulus.

Receptive Field Size and Response Latency Are Correlated Within the Cat Visual Thalamus

2005 ◽  
Vol 93 (6) ◽  
pp. 3537-3547 ◽  
Author(s):  
Chong Weng ◽  
Chun-I Yeh ◽  
Carl R. Stoelzel ◽  
Jose-Manuel Alonso
Keyword(s):  
Receptive Field ◽  
Spatial Frequency ◽  
Response Latency ◽  
Time Course ◽  
Frequency Tuning ◽  
Receptive Fields ◽  
Cortical Cell ◽  
Field Size ◽  
Receptive Field Size ◽  
Spatial Frequency Tuning

Each point in visual space is encoded at the level of the thalamus by a group of neighboring cells with overlapping receptive fields. Here we show that the receptive fields of these cells differ in size and response latency but not at random. We have found that in the cat lateral geniculate nucleus (LGN) the receptive field size and response latency of neighboring neurons are significantly correlated: the larger the receptive field, the faster the response to visual stimuli. This correlation is widespread in LGN. It is found in groups of cells belonging to the same type (e.g., Y cells), and of different types (i.e., X and Y), within a specific layer or across different layers. These results indicate that the inputs from the multiple geniculate afferents that converge onto a cortical cell (approximately 30) are likely to arrive in a sequence determined by the receptive field size of the geniculate afferents. Recent studies have shown that the peak of the spatial frequency tuning of a cortical cell shifts toward higher frequencies as the response progresses in time. Our results are consistent with the idea that these shifts in spatial frequency tuning arise from differences in the response time course of the thalamic inputs.

Spatial and temporal frequency tuning in striate cortex: functional uniformity and specializations related to receptive field eccentricity

2010 ◽  
Vol 31 (6) ◽  
pp. 1043-1062 ◽  
Author(s):  
Hsin-Hao Yu ◽  
Richa Verma ◽  
Yin Yang ◽  
Heath A. Tibballs ◽  
Leo L. Lui ◽  
...  
Keyword(s):  
Receptive Field ◽  
Striate Cortex ◽  
Frequency Tuning ◽  
Temporal Frequency

The Study of Spatial Frequency Channels for Human Visual System

2019 ◽  
Vol 33 (06) ◽  
pp. 1955007
Author(s):  
Xiangyang Xu ◽  
Qiao Chen ◽  
Ruixin Xu
Keyword(s):  
Receptive Field ◽  
Spatial Frequency ◽  
Visual System ◽  
Human Visual System ◽  
Frequency Tuning ◽  
Tuning Curve ◽  
Vision System ◽  
Human Vision ◽  
Psychophysical Experiment ◽  
Human Vision System

Similar to auditory perception of sound system, color perception of the human visual system also presents a multi-frequency channel property. In order to study the multi-frequency channel mechanism of how the human visual system processes color information, the paper proposed a psychophysical experiment to measure the contrast sensitivities based on 17 color samples of 16 spatial frequencies on CIELAB opponent color space. Correlation analysis was carried out on the psychophysical experiment data, and the results show obvious linear correlations of observations for different spatial frequencies of different observers, which indicates that a linear model can be used to model how human visual system processes spatial frequency information. The results of solving the model based on the experiment data of color samples show that 9 spatial frequency tuning curves can exist in human visual system with each lightness, R–G and Y–B color channel and each channel can be represented by 3 tuning curves, which reflect the “center-around” form of the human visual receptive field. It is concluded that there are 9 spatial frequency channels in human vision system. The low frequency tuning curve of a narrow-frequency bandwidth shows the characteristics of lower level receptive field for human vision system, the medium frequency tuning curve shows a low pass property of the change of medium frequent colors and the high frequency tuning curve of a width-frequency bandwidth, which has a feedback effect on the low and medium frequency channels and shows the characteristics of higher level receptive field for human vision system, which represents the discrimination of details.

Role of suppression in shaping orientation and direction selectivity of complex neurons in cat striate cortex

1996 ◽  
Vol 75 (3) ◽  
pp. 1163-1176 ◽  
Author(s):  
P. Hammond ◽  
J. N. Kim
Keyword(s):  
Spatial Frequency ◽  
Striate Cortex ◽  
Cutoff Frequency ◽  
Direction Selectivity ◽  
Uniform Field ◽  
Directional Tuning ◽  
Tuning Curves ◽  
Spatial Frequencies ◽  
Dominant Eye ◽  
Cat Striate Cortex

1. Single binocularly driven complex neurons in cat striate cortex were recorded extracellularly under nitrous oxide-oxygen-halothane anesthesia and muscle relaxant. Orientational/directional tuning was initially derived for each eye in turn, with sine wave gratings of optimal spatial frequency and velocity, while the other eye viewed a uniform field. 2. For the dominant eye, previously concealed suppression was revealed against elevated levels of firing induced with a conditioning grating, drifting continuously in the preferred direction, simultaneously presented to the nondominant eye. During steady-state binocular conditioning, orientational/directional tuning was reestablished for the dominant eye. In a subset of cells, tuning curves during conditioning were also derived for the reverse configuration, i.e., nondominant eye tuning, dominant eye conditioning: results were qualitatively identical to those for conditioning through the nondominant eye. 3. Neurons were initially segregated into five groups, according to the observed suppression profiles induced at nonoptimal orientations/directions during conditioning: Type 1, suppression centered on orthogonal directions; Type 2, suppression around null directions; Type 3, null suppression combined with orthogonal suppression; Type 4, lateral suppression, maximal for directions immediately flanking those inducing excitation; and Type 5, the residue of cells, totally lacking suppression or showing complex or variable suppression. 4. Sharpness of (excitatory) tuning was correlated with directionality and with class of suppression revealed during binocular conditioning. Direction-biased neurons were more sharply orientation tuned than direction-selective neurons; similarly, neurons exhibiting lateral or orthogonal suppression during conditioning were more sharply tuned than neurons with null suppression. 5. Application of suboptimal directions of conditioning weakened the induced suppression but altered none of its main characteristics. 6. The relationship between excitation, suppression, and spatial frequency was investigated by comparing tuning curves for the dominant eye at several spatial frequencies, without and during conditioning. End-stopped neurons preferred lower spatial frequencies and higher velocities of motion than non-end-stopped neurons. Confirming previous reports, suppression in some neurons was still present for spatial frequencies above the cutoff frequency for excitation, demonstrating the tendency for suppression to be more broadly spatial frequency tuned than excitation. 7. Scatterplots of strength of suppression, in directions orthogonal and opposite maximal excitation, partially segregated neurons of Types 1-3. Clearer segregation of Types 1-4 was obtained by curve-fitting to profiles of suppression, and correlating half-width of tuning for suppression with the angle between the directions of optimal suppression and optimal excitation in each neuron. 8. Two interpretations are advanced-the first, based on three discrete classes of inhibition, orthogonal, null and lateral; the second, based on only two classes, orthogonal and null/lateral--in which null and lateral suppression are manifestations of the same inhibitory mechanism operating, respectively, on broadly tuned direction-selective or on sharply tuned direction-biased neurons. Orthogonal suppression may be untuned for direction, whereas lateral and null suppression are broadly direction tuned. Within each class, suppression is more broadly spatial frequency tuned than excitation. 9. It is concluded that orientational/directional selectivity of complex cells at different spatial frequencies is determined by the balance between tuned excitation and varying combinations of relatively broadly distributed or untuned inhibition.

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.

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