Spatial arrangements of responses by cells in the cat visual cortex to light and dark bars and edges

Summary Detailed examination is made of the responses of visual cortical cells (area 17, border 17–18 and adjacent area 18) in the anaesthetized cat to stationary flashing bars and to bars (lines) and edges moving at their optimal velocities. Particular attention is given to the receptive field organization of cells in the simple family. While there is good general agreement between the main receptive field subregions revealed by stationary and moving stimuli, the responses to moving light and dark bars, supplemented by the responses to moving light and dark edges, provide a much more rapid, accurate and complete guide to the spatial organization of the receptive fields than do the response profiles to a stationary flashing bar. Moving light and dark bars between them generally reveal more subregions in the receptive fields of simple cells than is evident from the response profiles to a stationary flashing bar, particularly when the receptive fields have many subregions. In addition the responses to moving edges provide a rapid guide to spatial summation across the width of a subregion and the possible antagonistic effects of the next subregion in sequence.Two subclasses of cells in the simple family have been recognized: ordinary simple and fast simple cells. Two cell classes (A-cells and silent periodic cells) having properties intermediate between simple and complex types are discriminated and their properties described.     

Modeling V1 disparity tuning to time-varying stimuli.

Most models of disparity selectivity consider only the spatial properties of binocular cells. However, the temporal response is an integral component of real neurons' activities, and time-varying stimuli are often used in the experiments of disparity tuning. To understand the temporal dimension of V1 disparity representation, we incorporate a specific temporal response function into the disparity energy model and demonstrate that the binocular interaction of complex cells is separable into a Gabor disparity function and a positive time function. We then investigate how the model simple and complex cells respond to widely used time-varying stimuli, including motion-in-depth patterns, drifting gratings, moving bars, moving random-dot stereograms, and dynamic random-dot stereograms. It is found that both model simple and complex cells show more reliable disparity tuning to time-varying stimuli than to static stimuli, but similarities in the disparity tuning between simple and complex cells depend on the stimulus. Specifically, the disparity tuning curves of the two cell types are similar to each other for either drifting sinusoidal gratings or moving bars. In contrast, when the stimuli are dynamic random-dot stereograms, the disparity tuning of simple cells is highly variable, whereas the tuning of complex cells remains reliable. Moreover, cells with similar motion preferences in the two eyes cannot be truly tuned to motion in depth regardless of the stimulus types. These simulation results are consistent with a large body of extant physiological data, and provide some specific, testable predictions.     

Spatio-temporal interactions and the spatial phase preferences of visual neurons

Summary We recorded single neuron responses in the cat's lateral geniculate nucleus (LGN) and visual cortex to compound stimuli composed of two sinusoidal gratings in a 21 frequency ratio. To probe visual receptive field symmetry, we varied the relative spatial phase of the two components and measured the effect on neuronal responses. We expected that on-center LGN neurons would respond best to gratings combined in positive cosine (bright bar) phase, while off-center LGN neurons would respond best to gratings combined in negative cosine (dark bar) phase. When drifting stimuli were used, cells' phase preferences were roughly 90 deg away from the expected values; when stationary, contrast-modulated stimuli were used, phase preferences were as originally predicted. Computer simulations showed that this discrepancy could be explained by taking into account the cells' temporal properties. Thus, tests using drifting stimuli confound the spatial structure of visual neural receptive fields with their temporal response characteristics. A small sample of data from cortical neurons reveals the same confound.     

Space and spatial frequency: analysis and representation in the macaque striate cortex

Summary Simple cells in the macaque striate cortex were tested with bars, edges and gratings. Spatial frequency tuning curves could be predicted from the spatial profiles plotted with bars and edges and the bandwidth could be evaluated more accurately by computing the mean from measured and predicted tuning curves. The results suggest that the mean relative spatial frequency bandwidth (f/fo) is nearly constant and of a moderate value. But at each optimal spatial frequency, cells with different bandwidths (about a factor of two) were recorded. The shapes of spatial response profiles resemble the corresponding spatial and spatial frequency characteristics of line and edge detectors evaluated psychophysically. Among the remaining cell types, concentric cells tend to be tuned to lower spatial frequencies and have broader bandwidths, whereas periodic cells prefer higher spatial frequencies and have narrower bandwidths. Thus the mean relative bandwidth tends to decrease significantly with spatial frequency (as required by a system of patch-by-patch Fourier analysis) only when cells with poor orientation selectivity and the non-linear silent periodic cells are included along with the simple cells. Simple cells, on their own, seem to form a quasi-linear contrast processing system which is more biased towards spatial accuracy than spatial frequency selectivity.Supported by the Wellcome Trust and the Alexander von Humboldt Foundation     

Spatial frequency and orientation tuning curves of visual neurones in the cat: Effects of mean luminance

Summary Experiments have been performed on unanaesthetized and paralysed cats. The tuning curves for spatial frequency of retinal, lateral geniculate and simple and complex cells of the cortex have been determined in response to sinusoidal gratings of various spatial frequencies at different levels of mean luminance. For all neurones, decreasing the mean luminance leads to a progressive loss of spatial resolution and contrast sensitivity. Retinal ganglion cells of type X show, for scotopic levels of luminance, a flattening of their spatial frequency tuning curves in the low spatial frequency range. For geniculate and cortical neurones, on the contrary, the spatial frequency characteristics at the various levels of luminance remain practically invariant in their bandwidth. On the average, complex cells still respond to mean luminances ten times lower than simple cells. The tuning curves for orientation of cortical cells maintain, to a first approximation, the same shape at the various levels of mean luminance. The results are discussed, comparing the electrophysiological with psychophysical data.     

Spatial summation of responses in receptive fields of single cells in cat striate cortex

Summary Spatial summation of responses in striate neurons in cats under N₂O/O₂ anaesthesia was examined quantitatively both along the line of the optimal stimulus orientation (length summation) using moving light bars and single light and dark edge stimuli, and at right angles to the optimal orientation (width summation) using stationary flashing bars. Activity profiles and length-response curves were prepared from simple, complex and hypercomplex I and II cells. An activity profile indicates the responsiveness of a cell at locations along the length of its receptive field. The activity profiles from all cell types were usually well fitted by Gaussian functions. Length summation occurs both in end-free (simple and complex) and, to a lesser extent, in end-stopped (hypercomplex I and II) cells over a wide range of stimulus contrasts (0.13 to 0.95). The linearity of length summation was tested either by comparing the recorded length-response curves with the curves predicted from the linear integration of the activity profiles or by comparing the response to the activation of two regions of the receptive field with the sum of the responses to each region activated separately. Although length summation was usually non-linear (either greater than or less than direct proportionality) it was more nearly linear in complex than it was in simple and hypercomplex I cells. Mechanisms responsible for non-linear length summation were studied, including a threshold for discharge, response saturation and summation of end-zone inhibition. Complex cells show little width summation for bars wider than 0.3 °. In simple and hypercomplex I cells there was also relatively little width summation either in an ON or an OFF discharge region at contrasts above about 0.4 but at lower contrasts width summation may be approximately linear. Spatial summation of responses does not appear to be a useful characteristic for distinguishing one striate cell type from another.     

Spatial frequency tuning and contrast threshold of striate neurons in Siamese cats

Summary The spatial frequency tuning and the contrast-response function of striate neurons in Siamese cats were investigated with drifting sinusoidal gratings of high contrast, and the results were compared to the data obtained in normally pigmented cats. The optimal spatial frequency of the tuning curves obtained from Siamese cats was shifted toward lower values, and the mean optimal spatial frequency was significantly lower as compared to that measured in normal controls. Furthermore, the spatial resolution was severely reduced in Siamese cats, and many tuning curves in these animals showed unusually broad band width. The contrast response functions are characterized by higher contrast thresholds and shallower slopes in experimental animals. The units in Siamese cats had much larger receptive fields. Finally, these abnormalities were found in both simple and complex striate neurons. The present findings are discussed in terms of anomalies in pre-cortical visual neurons and their possible relation to the visual behavior of Siamese cats.     

Directional selectivity and spatiotemporal structure of receptive fields of simple cells in cat striate cortex.

1. Simple cells in cat striate cortex were studied with a number of stimulation paradigms to explore the extent to which linear mechanisms determine direction selectivity. For each paradigm, our aim was to predict the selectivity for the direction of moving stimuli given only the responses to stationary stimuli. We have found that the prediction robustly determines the direction and magnitude of the preferred response but overestimates the nonpreferred response. 2. The main paradigm consisted of comparing the responses of simple cells to contrast reversal sinusoidal gratings with their responses to drifting gratings (of the same orientation, contrast, and spatial and temporal frequencies) in both directions of motion. Although it is known that simple cells display spatiotemporally inseparable responses to contrast reversal gratings, this spatiotemporal inseparability is demonstrated here to predict a certain amount of direction selectivity under the assumption that simple cells sum their inputs linearly. 3. The linear prediction of the directional index (DI), a quantitative measure of the degree of direction selectivity, was compared with the measured DI obtained from the responses to drifting gratings. The median value of the ratio of the two was 0.30, indicating that there is a significant nonlinear component to direction selectivity. 4. The absolute magnitudes of the responses to gratings moving in both directions of motion were compared with the linear predictions as well. Whereas the preferred direction response showed only a slight amount of facilitation compared with the linear prediction, there was a significant amount of nonlinear suppression in the nonpreferred direction. 5. Spatiotemporal inseparability was demonstrated also with stationary temporally modulated bars. The time course of response to these bars was different for different positions in the receptive field. The degree of spatiotemporal inseparability measured with sinusoidally modulated bars agreed quantitatively with that measured in experiments with stationary gratings. 6. A linear prediction of the responses to drifting luminance borders was compared with the actual responses. As with the grating experiments, the prediction was qualitatively accurate, giving the correct preferred direction but underestimating the magnitude of direction selectivity observed.(ABSTRACT TRUNCATED AT 400 WORDS)     

Development of spatial frequency selectivity in striate cortex of vision-deprived cats

Summary Single unit activity was recorded in the striate cortex of vision-deprived cats aged between 3 and 8 weeks. Contrast sensitivity or response measurements made using moving sinusoidal gratings were used to construct spatial frequency tuning curves. At 3 weeks sensitivity, selectivity (assessed both as the narrowness of the tuning curve bandwidth and as the proportion of selective cells), and optimal spatial frequency, are all better than in 2 week old normally reared cats, and comparable with those of 3 week old normally reared cats. After 3 weeks of age no further improvements take place, although distributions of sensitivity, best spatial frequency and bandwidth overlap with those of normal cortical cells, and selectivity is clearly better than in adult LGN cells. These results are consistent with the idea that the spatial properties of cortical cells are at least partly predetermined, but that many cells require visual experience to develop normally.     

The two-dimensional spectral structure of simple receptive fields in cat striate cortex

1. A quantitative, general purpose method was developed for measuring the responses of visual neurons to stimuli distributed with high resolution over the two-dimensional (2D) spatial frequency domain. The stimuli consisted of drifting sinusoidal gratings of nonsaturating contrasts whose spatial frequency and orientation were drawn in random order from a 16 X 16 array of coordinates covering each neuron's responsive area. This method was applied to a population of 36 simple cells in area 17 of cat. 2. The response of each simple cell to drifting sinusoidal gratings appeared as a rectified sinusoidal modulation of the spike frequency. The degree of rectification varied from cell to cell, but for each cell, the form of the response was constant irrespective of stimulus spatial frequency, orientation, or contrast. The amplitude of the average response at the stimulus temporal frequency was used as the response metric at all spectral coordinates. Variations in this amplitude over two spectral dimensions forms a surface that we call the 2D spectral response profile. 3. For each cell, the 2D spectral response profile was localized to a limited region of the complete 2D spatial frequency domain. In bidirectionally responsive cells, there were two lobes in the surface disposed with mirror symmetry about the origin. In all cells, each lobe exhibited a single maximum and the response decayed smoothly in every direction away from the maximum. Isoresponse amplitude contours were elliptical and often, but not always, elongated about an axis of symmetry passing through the origin. 4. We tested the hypothesis that orientation and spatial frequency tuning are independent by forming scaled radial and angular sections through 2D spectral response profiles. In virtually every case polar separability did not obtain, that is, orientation selectivity depended on spatial frequency and vice versa. 5. In contrast, more than half the cells had 2D spectral response profiles that were Cartesian separable. The 2D spectral response profiles of most of the remaining cells were neither polar nor Cartesian separable, because the response profiles were elongated about an axis of symmetry that did not pass through the origin. 6. These results are discussed in terms of the constraints they place on models of the contributions simple cells make toward the neural representation of images.     

Direction selectivity of simple cells in cat striate cortex to moving light bars

The response properties of 84 simple striate cells in anaesthetized (N2O/O2 supplemented with sodium pentobarbital) and paralyzed cats were examined quantitatively using narrow optimally-oriented light and dark bars moving at optimal velocities. Different cells gave two to five spatially-offset response peaks, the light bar and the dark bar response peaks alternating with one another. With only 5 exceptions, the cells had the same preferred direction for movement of the dark bar as for the light bar. Static-field plots were prepared from 32 of the 84 cells using stationary flashing bars. The receptive fields of different cells had from two to four subregions responding either at light on (ON subregion) or at light off (OFF subregion) although one cell had only a single subregion. In the preferred direction of stimulus movement cells gave either the same number of response peaks to moving bars as there were subregions or one additional response peak. The additional response peak, termed a boundary response, always occurred at the end of the sequence of response peaks and was always completely direction selective. The direction selectivities of the individual response peaks in the responses from 49 of the 84 cells were analyzed. To ensure that each response peak and the corresponding peak in the opposite direction both came from the same subregion, the 49 cells were selected on the basis of having a response in the nonpreferred direction sufficient for analysis and of having a stimulus velocity less than 2.5 degrees/s so as to avoid significant spatial shifts of the peaks due to response latencies. For all but two of the 49 cells, the response peaks in any given profile always showed a progressively greater degree of direction selectivity as the stimulus advanced from one subregion to the next, the first subregion giving the least directionally-selective response peak and the last subregion the most directionally-selective peak. This observation was independent of the direction of stimulus motion and of the particular sequence in which the ON and the OFF subregions were traversed by the stimulus. The response patterns observed experimentally have been correlated with theoretical response patterns based on the responses of lateral geniculate neurons.     

Cortical maps of separable tuning properties predict population responses to complex visual stimuli

In the earliest cortical stages of visual processing, a scene is represented in different functional domains selective for specific features. Maps of orientation and spatial frequency preference have been described in the primary visual cortex using simple sinusoidal grating stimuli. However, recent imaging experiments suggest that the maps of these two spatial parameters are not sufficient to describe patterns of activity in different orientation domains generated in response to complex, moving stimuli. A model of cortical organization is presented in which cortical temporal frequency tuning is superimposed on the maps of orientation and spatial frequency tuning. The maps of these three tuning properties are sufficient to describe the activity in orientation domains that have been measured in response to drifting complex images. The model also makes specific predictions about how moving images are represented in different spatial frequency domains. These results suggest that the tangential organization of primary visual cortex can be described by a set of maps of separable neuronal receptive field features including maps of orientation, spatial frequency, and temporal frequency tuning properties.     

Accuracy of subspace mapping of spatiotemporal frequency domain visual receptive fields

Orientation and spatial frequency selectivities are fundamental properties of cells in the early visual cortex. Although they are customarily tested with drifting sinusoidal gratings, a recently developed subspace reverse correlation method may be a better replacement for obtaining a selectivity map in a joint orientation and spatial frequency domain at higher resolution efficiently. These two methods are examined for their accuracy and data compatibility for cells in areas 17 and 18 of anesthetized and paralyzed cats. Peaks and bandwidths of tuning curves from these two methods are highly correlated. However, spatial frequency bandwidths obtained by reverse correlation tend to be slightly narrower for the subspace reverse correlation than those from the drifting grating tests. Consistency between the two methods is improved if the entire duration of data containing signal are taken into account for the subspace reverse correlation rather than using the map only at the optimal correlation delay. Examination of convergence of the subspace mapping process shows that reliable 2-day profiles can be obtained within 5-10 min. for the majority of cells. Temporal dynamics of tuning properties are also examined more directly with the subspace mapping than with the drifting gratings. For many cells, the optimal spatial frequency shifts substantially, measured as a fraction of tuning bandwidth, over the time course of response. In comparison, the optimal orientation remains highly stable throughout the duration of response. Overall, these results suggest that the subspace reverse correlation is a better substitute for the conventional method.     

Effects of stimulus orientation on spatial frequency function of the visual evoked potential

Visual performance is better in response to vertical and horizontal stimuli than oblique ones in many visual tasks; this is called the orientation effect. In order to elucidate the electrophysiological basis of this psychophysical effect, we studied the effects of stimulus orientation on the amplitudes and latencies of visual evoked potentials (VEPs) over different spatial frequencies of the visual stimulation. VEPs to sinusoidal gratings at four orientations (vertical, horizontal, and oblique at 45 degrees and 135 degrees) with eight spatial frequencies (0.5-10.7 cycles/deg) at reversal rates of 1 Hz and 4 Hz were recorded in nine subjects. At 1-Hz stimulation, the amplitude and latency of P100 were measured. At 4-Hz stimulation, VEPs were Fourier-analyzed to obtain phase and amplitude of the second harmonic response (2F). At 1-Hz stimulation, P100 latencies were decreased for oblique stimuli compared with those for horizontal and vertical stimuli at lower spatial frequencies. Conversely, those for oblique stimuli were increased compared with those for horizontal and vertical stimuli at higher spatial frequencies. At 4-Hz stimulation, spatial tuning observed in 2F amplitude of the oblique gratings shifted to lower spatial frequencies when compared with those of vertical stimulation. The alteration of the VEP spatial frequency function caused by the oblique stimuli was in good agreement with the orientation effect observed in psychophysical studies. Our study may have a clinical implication in that VEP testing with stimuli in more than one orientation at slow and fast temporal modulations can be useful in evaluating neurological disease affecting the visual system.     

Direction selectivity of simple cells in cat striate cortex to moving light bars

Summary Quantitative estimates of the direction selectivities of 118 simple cells in response to moving light bars were expressed as a percentage calculated from the ratio of the response peaks: (preferred minus nonpreferred)/preferred. Virtually all simple cells were direction selective to some degree (mean direction selectivity 73.6%). Static-field plots to a stationary flashing bar were prepared from 74 of the 118 cells. Particular attention was given to the 42 cells with only two subregions in their static-field plot, one subregion responding at light on and the other at light off. It was concluded that interactive effects between subregions, whether synergistic or antagonistic, have little if any influence on the direction selective mechanism when the stimulus is a narrow light bar. Eighty two of the 118 cells were also tested with moving light and dark edges and of these 53 had response profiles with only two response peaks, one to the light edge and the other to the dark edge. Forty one of the 53 cells were each not only direction selective for both a light edge and a dark edge but also had a preferred direction for both edges that was the same as that for a light bar. Only two cells had preferred directions for both light and dark edges that were opposite to the direction preferred by the light bar. With one possible exception, every cell with two response peaks to moving edges and two subregions in the static-field plot showed a one-to-one correspondence between the ordinal sequence of the response peaks and the ordinal sequence of the subregions. Depending upon the polarity of the moving edge and the ordinal sequence of the subregions, the mean level of the direction selectivity to a moving edge was significantly below that to a narrow moving light bar. This reduction in the degree of the direction selectivity appears to be due to an interaction between the subregions leading to a reduction in the amplitude of the response in the preferred direction rather than a suppression of the direction selective mechanism that operates in the nonpreferred direction. Moving edges cause a weak interactive effect between the subregions that seems always to reduce the degree of the direction selectivity, never increasing it.     

Areal influences on complex cells in cat striate cortex: stimulus-specificity of width and length summation

Summary In single neurones recorded from the striate cortex of cats anaesthetized with N₂O/O₂/halothane, receptive field dimensions, length specificity and areal extent of drive were assessed for different classes of visual stimuli. Receptive fields were mapped as rectangular minimum response fields (MRFs). Spatial summation along the axis of preferred orientation was assessed: for moving bars whose length was varied (length summation); and for height variation of a square-wave grating patch against a uniform grey background, or a patch of moving texture against a stationary background of similar texture. In complementary tests a moving square-wave grating background was progressively occluded by a uniform grey foreground mask of variable height; or a mask of stationary texture of variable height progressively occluded a background of moving texture. In parallel measurements, the width of grating or textured patches or masks was varied whilst maintaining height constant. Broadly speaking, the areal influence of each class of stimulus was comparable, and distinct from extra-receptive field phenomena in evoking responses from within the receptive field, but not from surrounding areas. The masking paradigm provided the most sensitive measure of receptive field height and width. However, in some neurones length summation, the degree of endstopping, and the directional bias depended critically on the stimulus configuration used. Length summation tended to be more dramatic for short bars than for gratings. Length summation for texture was significantly more pronounced than for an oriented bar in special and in intermediate complex neurones. By contrast, endstopping was typically less intense for gratings than for bars, and least pronounced for texture. Because of stimulus specificity, complex neurones assigned to particular functional subgroups on the basis of their response to oriented bars may exhibit quite different patterns of behaviour for other classes of stimuli.     

The nature of V1 neural responses to 2D moving patterns depends on receptive-field structure in the marmoset monkey

A plaid pattern is formed when two sinusoidal gratings of different orientations are added together. Previous work has shown that V1 neurons selectively encode the direction and orientation of the component gratings in a moving plaid but not the direction of the plaid itself (Movshon et al. 1985). We recorded the responses of 49 direction-selective neurons to moving gratings and plaid patterns in area V1 of the anesthetized marmoset monkey (Callithrix jacchus). The responses of V1 neurons to rectangular patches of varying lengths and widths containing gratings of optimal spatial frequency were used to measure size and aspect ratio of the receptive-field subunits. We measured responses to plaid patterns moving in different directions and graded the magnitude of the response to the direction of motion of the plaid and the response to the direction of motion of the component gratings. We found significant correlations between receptive-field structure and the type and strength of its response to moving plaid patterns. The strength of pattern and component responses was significantly correlated with the interrelated properties of direction tuning width (Spearman's r = 0.82, P < 0.001), and receptive-field subunit aspect ratio (Spearman's r = -0.79, P < 0.001). Neurons with broad direction tuning and short, wide receptive-field subunits gave their greatest response when the plaid moved in their preferred direction. Conversely, neurons with narrow direction tuning and long, narrow receptive-field subunits gave their greatest responses when the plaid moved in a direction such that one of its components moved in the preferred direction.     

Spatial and temporal properties of neurons of the lateral suprasylvian cortex of the cat

Neurons in the posteromedial lateral suprasylvian cortex (PMLS) of cats were recorded extracellularly to investigate their response to stimulation by bars and by sinusoidal gratings. Two general types of cells were identified: those that modulated in synchrony with the passage of drifting bars and gratings and those that responded with an unmodulated increase in discharge. Both types responded to contrast reversed gratings with a modulation of activity: the cells that modulated to drifting gratings modulated to the first harmonic of contrast reversed gratings (at appropriate spatial phase and frequency), whereas those that did not modulate to drifting gratings always modulated to the second harmonic of contrast reversed gratings. No cell had a clear null point. Nearly all cells were selective for spatial frequency. The preferred frequency ranged from 0.1 to 1 cycles per degree (cpd), and selectivity bandwidths (full width at half height) were around two octaves. Preferred spatial frequency was not correlated with receptive field size, but bandwidth and receptive field size were positively correlated. Preferred spatial frequency decreased with eccentricity, at about 0.05 octaves/deg. The response of all cells increased as a function of grating contrast up to a saturation level. The contrast threshold for response to a grating of optimal parameters was approximately 1% for most cells and the saturation contrast approximately 10%. The contrast gain was approximately 25 spikes/s per log unit of contrast. All cells were tuned for temporal frequency, preferring frequencies from approximately 3 to 10 Hz, with a selectivity bandwidth approximately 2 octaves. For some cells, the spatial selectivity did not depend on the temporal frequency and vice versa. Others were spatiotemporally coupled, with the preferred temporal frequency being lower at high than at low spatial frequencies, and the preferred spatial frequency lower at high than at low temporal frequencies. Previous results showing broad velocity tuning to a bar were replicated and found to be predictable from the combined spatial and temporal tuning of PMLS cells and the Fourier spectrum of a bar. Preferred temporal frequency steadily decreased with eccentricity, at 0.025 octaves/deg. The results for PMLS cells are compared with those of other visual areas. Acuity and spatial preference and selectivity bandwidth is comparable to all areas except area 17, where they are a factor of about two higher. Temporal selectivity in PMLS is as fine as observed in other areas. The possibility that PMLS cells may be involved with motion detection and detection of motion in depth is discussed.     

Spatial frequency characteristics of nearby neurons in cats’ visual cortex

Various methods have allowed mapping of responses to several stimulus features on the cortical surface, particularly edge orientation and motion direction. The cortical mapping of spatial frequencies (SF), which is the basic property that leads to perception of spatial details of visual objects, is still controversial. We recorded simultaneously extracellular action potentials from neighboring cells in superficial layers of the area 17–18 border region of anesthetized cats. Responses of nearby cells to sine-wave gratings of varying SF were analyzed. Spatial frequency tuning curves were cross-correlated to establish the degree of similarity between the curves and optimal SFs were compared for each pair of neurons. The investigation showed that only about a half of nearby neurons exhibited close optimal SFs and similar tuning curves. The results suggest that SF channels do not show a clear clustering within a small pool of neurons. Such organization may contribute to the perception of spatial details at all orientations and motion directions.     

The role of GABAergic inhibition in the response properties of neurones in cat visual area 18

Bicuculline methiodide was iontophoretically applied to single neurones in cat area 18 to investigate how removal of gamma-aminobutyrate mediated inhibition affects the visual response properties. Moving sinusoidal gratings were used to study spatial and temporal response characteristics. Orientation sensitivity and spatial and temporal frequency tuning curves were determined with and without iontophoretically applied bicuculline. In most neurones, orientation sensitivity and spatial frequency tuning remained largely unaffected, whereas temporal frequency tuning was very much broadened. It is suggested that the dominant excitatory input to area 18 cells is a spatially organized input from area 17 and local inhibition in area 18 sharpens primarily temporal selectivity. An alternative explanation of our results would be that the distribution of synapses mediating temporal tuning in area 18 is fundamentally different from that mediating spatial frequency and orientation tuning, which may be located at sites distant from the cell body and relatively inaccessible to the drug application.