Collinearity tolerance of cells in areas 17 and 18 of the cat's visual cortex: Relative sensitivity to straight lines and chevrons

Summary Collinearity tolerance and length dependence of orientation tuning were compared in cells recorded from areas 17 and 18 of the lightly anaesthetised cat's visual cortex. Orientation tuning and interaction between receptive field halves of the same cells are reported in the preceding paper and elsewhere (Hammond and Andrews, 1978a, b).In confirmation of previous work, increase in stimulus length was associated with sharper orientation tuning in all simple and hypercomplex cells, and in most complex cells even in the absence of length summation.Cells in areas 17 and 18 were more sharply tuned for straight lines than for chevrons bent symmetrically about the optimal orientation; tuning for chevrons was noticeably skewed compared with tuning for straight lines. In area 17, the best response was always obtained with a straight line of optimal orientation.The two halves of the receptive fields of some cells in area 18 had dissimilar preferred orientations. Even in cells whose receptive field halves were similarly tuned, broadly tuned, or apparently untuned for orientation, simultaneous stimulation of both halves of the receptive field led to substantial sharpening of tuning. In cells with dissimilarly tuned half fields, the skew in chevron tuning was predictable from the orientation tuning of each half of the receptive field. Two area 18 cells responded consistently better to a chevron stimulus than to a straight line of any orientation.     

Orientation tuning of cells in areas 17 and 18 of the cat's visual cortex

Summary Sharpness and symmetry of orientation tuning were quantitatively investigated and compared in ninety-seven cells from areas 17 and 18 of the lightly-anaesthetised feline visual cortex.Halfwidths of orientation tuning at half-height ranged between 5 ° and 73 ° for long stimuli, with an extreme exception at 111 ° (excluding untuned cells).There was a tendency for cells in area 18 to be more broadly tuned than those in area 17, due largely to the relatively sharp tuning of area 17 simple cells. Confirming previous work, simple cells were more sharply tuned than complex cells in area 17. In area 18, there was no clear distinction in sharpness of tuning between complex type 1 cells (equated with area 17 simple cells), complex type 2 cells (equated with area 17 complex cells), or hypercomplex cells.Approximately 60% of cells in both areas were asymmetrically tuned for orientation: ratios of half-widths to either side of the optimal orientation ranged from 1.0–3.0, exceptionally 5.8. Asymmetry of tuning was more marked in area 18 than in area 17, except that area 18 complex type 2 cells as a group were relatively symmetrically tuned for orientation.Occasional cells with different preferred orientations for opposite directions of motion, for each peak of a bimodal response to a single direction, or for each half of the receptive field were also observed. The latter are described in the following paper (Hammond and Andrews, 1978b).     

Local signals from beyond the receptive fields of striate cortical neurons

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.     

Responses to moving slits by single units in cat striate cortex

Summary A quantitative study has been made of the responses to moving slit stimuli by single units in the cat striate cortex whose receptive fields lay within 5° of the visual axis. Special attention was given to finding the optimal stimulus parameters including slit width, length, orientation and speed. The analysis was largely based on averaged response vs. time histograms. Using the classification of simple and complex responses types, the units were further subdivided on the basis of the number of modes in the response and on the presence or absence of directional selectivity. Simple unimodal units with directional selectivity (SUDS) had the most specific stimulus requirements and nearly always had zero background activity. Complex units usually had a high level of background activity. SUDS units also showed a preference for horizontally- and vertically ****-orientated stimuli. Whenever the response survived reversal of contrast the directional selectivity remained independent of the change. Optimal stimulus speeds varied widely from unit to unit with a mean at 4°/sec: simple bimodal units and complex units tended to have higher optimal stimulus speeds and responded over a wider range of speeds than did simple unimodal units. While SUDS units with very small receptive fields tended to prefer slowly moving stimuli, in general there was no correlation between receptive field size and optimal stimulus speed.Selby Fellow of the Australian Academy of Sciences.     

Classification of receptive field properties in cat visual cortex

Summary The properties of the receptive fields of visual cortex neurons of cats were studied manually and by a computer controlled system using single lines, double lines and multiple lines (gratings). The multiple selectivities of each of the receptive fields studied make it necessary to abandon the concept that each cell functions as a feature detector. Instead, an attempt was made to classify the receptive field properties with the aim to delineate the transfer functions (of the total networks) served by each property. When tested with one-line stimulus, cells with simple receptive field properties diffefed from cells with complex receptive field properties as to their velocity selectivity (simple: 1 ° to 3 °/s; complex: 4 ° to 10 °/s), spontaneous activity (lower for cells with simple properties), optimal firing rate (lower for cells with simple properties) and receptive field size (smaller for cells with simple properties) but not for orientation and direction selectivity. When tested with a 2-lines moving stimulus, the responses of cells with simple properties were facilitated by the progressive separation of the lines whereas the responses of cells with complex receptive field properties were inhibited. When multiple lines, i.e. gratings, were used, an equivalence between simple and X properties and complex and Y properties was shown, while the sustained/transient classification proved to be independent of the simple/complex (X/Y) classification. Thus, receptive field properties can be classified into three categories: one reflects the input to the receptive fields; a second deals with the interactive properties of the fields; while a third appears more related to the overall properties of the network.This research was supported by a postdoctoral fellowship from the Medical Research Council of Canada to Maurice Ptito, a predoctoral fellowship from the National Research Council of Canada to Maryse C. Lassende and NIMH Grant MH12970 and NIMH Career Research Award MH15214 to Karl H. Pribram     

Organization of suppression in receptive fields of neurons in cat visual cortex.

1. The response to an optimally oriented stimulus of both simple and complex cells in the cat's striate visual cortex (area 17) can be suppressed by the superposition of an orthogonally oriented drifting grating. This effect is referred to as cross-orientation suppression. We have examined the spatial organization and tuning characteristics of this suppressive effect with the use of extracellular recording techniques. 2. For a total of 75 neurons, we have measured the size of each cell's excitatory receptive field by use of rectangular patches of drifting sinusoidal gratings presented at the optimal orientation and spatial frequency. The length and width of these grating patches are varied independently. Receptive-field length and width are determined from the dimensions of the smallest grating patch required to elicit a maximal response. 3. The extent of the area from which cross-orientation suppression originates has been measured in an analogous manner. Each neuron is excited by a patch of drifting grating the same size as the receptive field. The response to this stimulus is modulated by a superimposed patch of grating having an orthogonal orientation. After selecting the spatial frequency that produces maximal suppression, the response of each cell is examined as a function of the length and width of the orthogonal (suppressive) grating patch. Results from 29 cells show that the dimensions of the orthogonal grating patch required to elicit maximal suppression are similar to, or smaller than, the dimensions of the excitatory receptive field. Thus cross-orientation suppression originates from within the receptive field. 4. For some cells the spatial frequency tuning of the suppressive effect is much broader than the spatial frequency tuning for excitation. In these cases it is possible to find a spatial frequency that produces suppression but not excitation. With the use of a suppressive stimulus having this spatial frequency, we examined the strength of suppression as a function of orientation for 11 cells. These tests show that suppression occurs at all orientations, including the preferred orientation for excitation. In some cases, suppression is somewhat stronger at the preferred orientation for excitation than at any other orientation. 5. For 12 cells we varied the relative spatial phase between the optimally oriented and orthogonal gratings. In all cases the magnitude of suppression is largely independent of the relative spatial phase. 6. For three binocular cells we examined whether the suppressive effect of a grating oriented orthogonal to the optimum could be mediated dichoptically.(ABSTRACT TRUNCATED AT 400 WORDS)     

Orientation discrimination sensitivity of single units in cat primary visual cortex

Summary Responses of visual cortex (area 17) neurons to moving oriented stimuli were recorded from anesthetized cats. The variance of response (SD²) to repeated identical stimuli was directly proportional to response magnitude (R), (SD² =C²R). The values of C were not found to differ significantly between different types of cortical cells. The relationship predicts that the coefficient of variation (SD/R) will be smallest near the peak of the tuning curve, indicating that the peak response is most reliable for detecting an orientation but not necessarily the most sensitive to a change in orientation. Tuning curves and response variability were then examined to determine the orientation at which the neuron was most sensitive to changes in stimulus orientation using signal detection theory. The discrimination index (d = [R1-R2J/SD) for a 1 degree change in stimulus orientation was greatest along the flanks of the tuning curve. In order to generalize the experimental data, response distributions derived from a model of cells with parameters based on experimental data were examined to determine the minimal discriminable change in stimulus orientation. Changes of stimulus orientation between 0.6 and 5 deg of arc could be detected from single responses of a single cell by an optimal observer with 75% accuracy if the orientation change was centered at the most sensitive part of the tuning curve.Supported in part by grant NS10332     

Local correlation-based circuitry can account for responses to multi-grating stimuli in a model of cat V1

In cortical simple cells of cat striate cortex, the response to a visual stimulus of the preferred orientation is partially suppressed by simultaneous presentation of a stimulus at the orthogonal orientation, an effect known as "cross-orientation inhibition." It has been argued that this is due to the presence of inhibitory connections between cells tuned for different orientations, but intracellular studies suggest that simple cells receive inhibitory input primarily from cells with similar orientation tuning. Furthermore, response suppression can be elicited by a variety of nonpreferred stimuli at all orientations. Here we study a model circuit that was presented previously to address many aspects of simple cell orientation tuning, which is based on local intracortical connectivity between cells of similar orientation tuning. We show that this model circuit can account for many aspects of cross-orientation inhibition and, more generally, of response suppression by nonpreferred stimuli and of other nonlinear properties of responses to stimulation with multiple gratings.     

Orientation tuning of input conductance, excitation, and inhibition in cat primary visual cortex

The input conductance of cells in the cat primary visual cortex (V1) has been shown recently to grow substantially during visual stimulation. Because increasing conductance can have a divisive effect on the synaptic input, theoretical proposals have ascribed to it specific functions. According to the veto model, conductance increases would serve to sharpen orientation tuning by increasing most at off-optimal orientations. According to the normalization model, conductance increases would control the cell's gain, by being independent of stimulus orientation and by growing with stimulus contrast. We set out to test these proposals and to determine the visual properties and possible synaptic origin of the conductance increases. We recorded the membrane potential of cat V1 cells while injecting steady currents and presenting drifting grating patterns of varying contrast and orientation. Input conductance grew with stimulus contrast by 20-300%, generally more in simple cells (40-300%) than in complex cells (20-120%), and in simple cells was strongly modulated in time. Conductance was invariably maximal for stimuli of the preferred orientation. Thus conductance changes contribute to a gain control mechanism, but the strength of this gain control does not depend uniquely on contrast. By assuming that the conductance changes are entirely synaptic, we further derived the excitatory and inhibitory synaptic conductances underlying the visual responses. In simple cells, these conductances were often arranged in push-pull: excitation increased when inhibition decreased and vice versa. Excitation and inhibition had similar preferred orientations and did not appear to differ in tuning width, suggesting that the intracortical synaptic inputs to simple cells of cat V1 originate from cells with similar orientation tuning. This finding is at odds with models where orientation tuning in simple cells is achieved by inhibition at off-optimal orientations or sharpened by inhibition that is more broadly tuned than excitation.     

Cat area 17. III. Response properties and orientation anisotropies of corticotectal cells

The receptive field properties of antidromically identified corticotectal (CT) cells in area 17 were explored in the paralyzed, anesthetized cat. To compare these with another population of infragranular cells, we also examined the receptive field properties of cells in layer 6. Sixty percent of our sample of CT cells showed increased response to increased stimulus length (length summation) and were classified as standard complex cells. The other 40% showed little or no length summation, were generally end stopped, and were classified as special complex cells. Standard and special complex CT cells have complementary orientation anisotropies: the distribution of orientation preferences of standard complex cells is biased toward obliquely oriented stimuli, whereas special complex cells are biased toward horizontally and vertically oriented stimuli. The receptive fields of the cells in our sample were primarily along the horizontal meridian so we cannot determine if these anisotropies are defined relative to the vertical meridian or relative to the meridian passing through the receptive field. The effects of these anisotropies in preferred orientation are minimized by the broad orientation tuning of CT cells. There was no simple relationship between the direction bias of CT cells and the reported direction bias of tectal cells. In contrast to the heterogeneity of corticotectal cells, layer 6 cells uniformly showed strong length summation, tight orientation tuning, and little spontaneous activity.     

Orientation selectivity and the spatial distribution of enhancement and suppression in receptive fields of cat striate cortex cells

The relationship between orientation selectivity and spatial receptive field organization was analyzed. Receptive field maps were made with a dual stimulus technique where an optimally oriented activation slit was presented in the most responsive region to produce activity against which the effect of a test spot in various positions was determined. Both simple and complex cells had receptive fields which were subdivided into adjacent elongated and antagonistic subregions. When the two stimuli were presented in phase (both ON or OFF simultaneously) the fields had a central enhancement region with a strong suppression flank on one or both sides. Optimal slit orientation was related to the location of the suppression flank relative to the location of the central enhancement region, and the degree of orientation selectivity to the shape of the subregions and the distance between them. Estimated orientation tuning curves calculated from the receptive field maps gave satisfactory first approximations to experimental curves. The relative contribution of enhancement and suppression to orientation selectivity was studied by presenting a test slit in different orientations in phase with an optimally oriented activation slit. The orientation selectivity was produced almost exclusively by the flank suppression indicating that orientation selectivity is produced by inhibitory input. The flank suppression lacked any specific orientation selectivity, and it occurred only when both the central region and the flanks were activated in phase. Orientation selectivity in both simple and complex cells is explained by a receptive field organization where the cells have input from partially overlapping excitatory and inhibitory fields which have their centers slightly displaced from each other.     

The contribution of inhibitory mechanisms to the receptive field properties of neurones in the striate cortex of the cat.

1. The iontophoretic application of the GABA antagonist bicuculline to simple and complex cells in the striate cortex of the cat produced extensive modifications of receptive field properties. These modifications appear to relate to a block or reduction of GABA-mediated intracortical inhibitory influences acting on the cells examined. 2. For simple cells the effects of bicuculline on receptive field properties involved a loss of the subdivision of the receptive field into antagonistic "on" and "off" regions, a reduction in orientation specificity and a reduction or elimination of directional specificity. 3. The effect on the "on" and "off" subdivisions of the simple cell receptive field was such that all stationary flashing stimuli, whether covering the whole receptive field, or located within the receptive field over a previously determined "on" or "off" region, resulted in an "on and off" response. 4. The orientation specificity of complex cells was reduced during the application of bicuculline such that in many cases the original specificity of the cell was virtually lost with the response to the orientation at 90 degrees to the optimal being of similar magnitude to the optimal. The directional specificity of complex cells was generally less affected than that of simple cells. Often when large changes in orientation specificity were observed the directional specificity was relatively unaffected. 5. For some cells apparently showing to all visual stimuli only inhibitory responses, the application of bicuculline resulted in the appearance of excitatory responses. 6. In all cases receptive field properties reverted to the original state after termination of the bicuculline application. It was not generally possible to duplicate the effects of bicuculline by raising neuronal excitability with iontophoretically applied glutamate. 7. On the basis of these results it is suggested that the normal subdivision of the simple cell receptive field into separate "on" and "off" regions and its directional specificity are dependent on intracortical inhibitory processes that are blocked by bicuculline. The orientational tuning of simple cells conversely appears to be largely determined by the excitatory input but normally enhanced by lateral type inhibitory processes acting in the orientation domain. 8. It also appears that the excitatory input to some complex cells is not orientation specific. This suggests that for these cells it is extremely unlikely that they receive an orientation specific excitatory input from simple cells.     

A quantitative study of the projection area of the central and the paracentral visual field in area 17 of the cat

Cells in cat's area 17 respond optimally if elongated contrasts are presented at a certain angle or orientation with respect to the retina, or to the visual field, respectively (Hubel and Wiesel, 1962). The preferred orientation and the range of orientation sensitivity of cells in close proximity to one another have been determined in order to investigate the spatial arrangement of the orientation domain in area 17. 1. A slight overrepresentation of vertical and horizontal orientations is seen in cells with complex receptive fields, whereas in cells with simple fields all orientations are represented to an equal degree. The orientation selectivity, defined as the halfwidth of tuning curves constructed from the cells response to a moving stimulus, is less than 60 degrees in more than 80% of all cells investigated, and is on the average 20-30 degrees smaller in cells with simple than in cells with complex receptive fields. 2. In 80% of all cases considered the difference in the preferred orientation between two cells less than 200 mum horizontally distant in area 17 is less than 30 degrees, which is of the order of an individual cells orientation selectivity. Each cell, therefore, will respond to some extent to that orientation which is preferred by the cells in the immediate surroundings. 3. Sequential changes in the preferred orientation between cells successively recorded are observed as the postlateral gyrus is explored from anterior to posterior and from medial to lateral. On these general trends a random variation in the preferred orientation between neighbouring cells of the order of 5-10 degrees is superimposed. One orientation sequence (180 degrees) occupies 700-1200 mum, so that on the average a change in the preferred orientation of the order of 10 degrees is complete after 50 mum distance in the cortex measured parallel to the pial surface. Assuming that 18 different orientations (+/- 5 degrees) functionally represent one complete orientation sequence it is found that 'all' orientations are functionally represented by the cells contained in a cortical cylinder of 300-700 mum in diameter. 4. Cells having the same preferred orientation are grouped together in cortical regions which appear in crossection as a band or a spot. These regions have been termed iso-orientation bands or spots. The diameter of the spots and the small diameter of the bands do not exceed 100 mum. Taking an average orientation selectivity of 40 degrees for cells vertically aligned in area 17 it is calculated that cells situated 100 mum to either side of an iso-orientation band or around an iso-orientation spot still respond with 50% of the discharge to their own optimal orientation ...     

Inactivation of the infragranular striate cortex broadens orientation tuning of supragranular visual neurons in the cat

Intracortical inhibition is believed to enhance the orientation tuning of striate cortical neurons, but the origin of this inhibition is unclear. To examine the possible influence of ascending inhibitory projections from the infragranular layers of striate cortex on the orientation selectivity of neurons in the supragranular layers, we measured the spatiotemporal response properties of 32 supragranular neurons in the cat before, during, and after neural activity in the infragranular layers beneath the recorded cells was inactivated by iontophoretic administration of GABA. During GABA iontophoresis, the orientation tuning bandwidth of 15 (46.9%) supragranular neurons broadened as a result of increases in response amplitude to stimuli oriented about ±20° away from the preferred stimulus angle. The mean (±SD) baseline orientation tuning bandwidth (half width at half height) of these neurons was 13.08±2.3°. Their mean tuning bandwidth during inactivation of the infragranular layers increased to 19.59±2.54°, an increase of 49.7%. The mean percentage increase in orientation tuning bandwidth of the individual neurons was 47.4%. Four neurons exhibited symmetrical changes in their orientation tuning functions, while 11 neurons displayed asymmetrical changes. The change in form of the orientation tuning functions appeared to depend on the relative vertical alignment of the recorded neuron and the infragranular region of inactivation. Neurons located in close vertical register with the inactivated infragranular tissue exhibited symmetric changes in their orientation tuning functions. The neurons exhibiting asymmetric changes in their orientation tuning functions were located just outside the vertical register. Eight of these 11 neurons also demonstrated a mean shift of 6.67±5.77° in their preferred stimulus orientation. The magnitude of change in the orientation tuning functions increased as the delivery of GABA was prolonged. Responses returned to normal approximately 30 min after the delivery of GABA was discontinued. We conclude that inhibitory projections from neurons within the infragranular layers of striate cortex in cats can enhance the orientation selectivity of supragranular striate cortical neurons.     

Effect of body tilt on receptive field orientation of simple visual cortical neurons in unanesthetized cats

Summary The receptive field (RF) orientation of 53 simple visual cortical neurons was determined by recording the activity of single cells during presentation of stationary bars of light. An RF tuning curve was constructed for each cell by averaging the neural discharge resulting from the repeated presentation of a number of slit orientations. RF curves were then determined again, following a 45 ° roll tilt of the entire head and body, and subsequently after the return of the animal to the original horizontal position. RF tuning curves were typical of what others have found to characterize simple cells, and were highly replicable on the return to the starting position. In 73% of the cells studied, the RF orientation after tilt remained unaltered relative to the head axis (±15 °); in the remaining 27% of the cells RF orientations either under- or over-shot the retinal tilt by more than 15 °, and in some cases by as much as 45 °. These results support the hypothesis that the well documented vestibular inputs to visual cortex play a role in modifying the RF orientation selectivity of visual cortical neurons, and suggest that such information may be an important neurophysiological substrate underlying visual spatial constancy mechanisms.This research was supported by NIH grant NS12308, and grants to DLT and NMB from the University of Pittsburgh Medical Alumni Association     

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.     

The velocity tuning of single units in cat striate cortex.

1. The activity of single units was recorded from the striate cortex (area 17) of anaesthetized, paralysed cats. Responses to stimuli moving at different velocities were examined. 2. Peak evoked firing frequency, rather than fotal evoked spikes, is used throughout as a measure of response. The former mea-ure gives curves of response vs. velocity that correlate well with curves of contrast sensitivity vs. velocity, wheras the latter does not. 3. Cortical receptive fields were classified according to the criteria of Hubel & Wiesel. Simple cells were found to prefer lower velocities (mean 2-2 deg sec-1) than complex cells( mean 18-8 deg sec-1). The response of simple cells to stimuli moving faster than 20 deg sec-1 is generally poor; complex cells usually discharge briskly to these speeds. 4. Cells classified as hypercomplex by the end-inhibition criterion were further chara-terized as type I or type II, according to the suggestion of Dreher (1972). Type I units are indistinguishable from simple cells in their velocity tuning, and type II units equally clearly resemble complex cells. These results are therefor consistent with Dreher's sbudivision. 5. Teh selectivity of cells for velocity is variable but can be quite marked. The average selectivities of simple and complex cells are not significantly different. There is an inverse correlation between preferred velocity and the sharpness of velocity selectivity for simple cells; no trend is apparent for other cell types. 6. No clear correlation is observed between the velocity preferances of units and their degree of direction selectivity, or receptive field arrangement. Simple cells with 'sustainef' temporal responses to flashed stimuli tend to prefer slower rates of movement than 'transient' ones, and to be less selective for velocity. 7. The results for different cortical cell-types are compared with the velocity tuning of X- and Y-cells in the lateral geniculate nucleus.     

Modification of response functions of cat visual cortical cells by spatially congruent perturbing stimuli.

Responses of cat striate cortical cells to a drifting sinusoidal grating were modified by the superimposition of a second, perturbing grating (PG) that did not excite the cell when presented alone. One consequence of the presence of a PG was a shift in the tuning curves. The orientation tuning of all 41 cells exposed to a PG and the spatial frequency tuning of 83% of the 23 cells exposed to a PG showed statistically significant dislocations of both the response function peak and center of mass from their single grating values. As found in earlier reports, the presence of PGs suppressed responsiveness. However, reductions measured at the single grating optimum orientation or spatial frequency were on average 1.3 times greater than the suppression found at the peak of the response function modified by the presence of the PG. Much of the loss in response seen at the single grating optimum is thus a result of a shift in the tuning function rather than outright suppression. On average orientation shifts were repulsive and proportional (approximately 0.10 deg/deg) to the angle between the perturbing stimulus and the optimum single grating orientation. Shifts in the spatial frequency response function were both attractive and repulsive, resulting in an overall average of zero. For both simple and complex cells, PGs generally broadened orientation response function bandwidths. Similarly, complex cell spatial frequency response function bandwidths broadened. Simple cell spatial frequency response functions usually did not change, and those that did broadened only 4% on average. These data support the hypothesis that additional sinusoidal components in compound stimuli retune cells' response functions for orientation and spatial frequency.     

Effects of inhibitory gain and conductance fluctuations in a simple model for contrast-invariant orientation tuning in cat V1

The origin of orientation selectivity in primary visual cortex (V1) is a model problem for understanding cerebral cortical circuitry. A key constraint is that orientation tuning width is invariant under changes in stimulus contrast. We have previously shown that this can arise from the combination of feedforward lateral geniculate nucleus (LGN) input and an orientation-untuned component of feedforward inhibition that dominates excitation. However, these models did not include the large background voltage noise observed in vivo. Here, we include this noise and examine a simple model of cat V1 response. Constraining our simulations to fit physiological data, our single model parameter is the strength of feedforward inhibition relative to LGN excitation. With physiological noise, the contrast invariance of orientation tuning depends little on inhibition level, although very weak or very strong inhibition leads to weak broadening or sharpening, respectively, of tuning with contrast. For any inhibition level, an alternative measure of orientation tuning -- the circular variance -- decreases with contrast as observed experimentally. These results arise primarily because the voltage noise causes large inputs to be much more strongly amplified than small ones in evoking spiking responses, relatively suppressing responses to nonpreferred stimuli. However, inhibition comparable to or stronger than excitation appears necessary to suppress spiking responses to nonpreferred orientations to the extent seen in vivo and to allow the emergence of a tuned mean voltage response. These two response properties provide the strongest constraints on model details. Antiphase inhibition from inhibitory simple cells, and not just untuned inhibition from inhibitory complex cells, appears necessary to fully explain these aspects of cortical orientation tuning.     

Directionality of cat striate cortical neurones: contribution of suppression

Summary Direction-selective or direction-biased striate cortical neurones were assessed for absence or incidence of suppression of firing, maximal at 90° or 180° (null suppression) to the optimal direction, in 327 neurones recorded from the striate cortex of cats anaesthetized with N₂O/O₂/halothane. Stimuli were light or dark bars moving over uniform or stationary textured back-grounds; or square-wave gratings of optimal spatial frequency and velocity. Five identified directionality groups were correlated with neuronal class and a range of other receptive field properties. Suppression maximal at 90° to optimum was common amongst direction-biased neurones, rare amongst direction-selective neurones. In the latter group, null suppression (maximal at 180° to optimum) was more prevalent than at 90°. Standard complex cells constituted the majority of complex neurones. They were more commonly direction-biased and less commonly showed suppression than special complex cells. The latter comprised the majority of direction-selective neurones with 180° suppression. Endstopping was seen more frequently in special complex cells, but for each functional class was similarly distributed between the different directionality groups. Based on the mean and mode of partially overlapping distributions, for all neuronal classes direction-selective neurones were more broadly tuned than direction-biased neurones. Special complex neurones were appreciably more broadly tuned than standard complex neurones; those with suppression at 180° were the most broadly tuned neurones in the cortex. Direction-biased neurones with suppression at 90° to optimum were more sharply tuned than those lacking such suppression. Direction-selective neurones had larger receptive fields than direction-biased neurones. In both groups receptive fields decreased in size in the sequence: intermediate complex > standard complex > special complex > simple. Resting discharge was highest amongst direction-selective neurones with 180° suppression, lower in those with 90° suppression or those lacking it, and lowest amongst direction-biased neurones. With the possible exception of the minority of neurones that were silent, low levels of resting discharge have not seriously prejudiced either neuronal categorization or comparisons of tuning selectivity. The pattern which emerges is that suppression maximal in directions orthogonal to the preferred direction/orientation is more commonly associated with sharp tuning and directionbias, whereas null suppression, in the direction opposite that preferred, is associated with broad tuning, direction-selectivity, high resting discharge levels and strong texture sensitivity.