Comparison of receptive field properties of neurons in area 17 of normal and bilaterally amblyopic cats

Receptive field properties of extracellularly recorded units in the visual cortex (area 17) of cats made bilaterally amblyopic by a variety of rearing conditions were measured and compared with the properties of units in normal cats. Properties studied included sensitivity to vernier offset, response facilitation to increasing bar length, receptive field size, responsiveness to moving and flashed stimuli, orientation tuning, the relation between mean firing rate and its variance, the amount of overlap of regions of on and off responsiveness in simple and complex cells, and, for flashed stimuli, latency to response onset, time to peak response, and response decay time constant. Behavioural testing of the amblyopic animals showed that spatial resolution was 2–4 times lower and vernier acuity thresholds 10–20 times greater than normal. Despite this, several neuronal response properties did not differ significantly from those in normal animals. These included peak responsiveness to moving stimuli, widths of orientation tuning curves, response variability, and latency to initial response for flashed stimuli. Other properties showed small but significant changes. Sensitivity to vernier offset (impulses per degree of offset) was reduced to nearly half its normal level; receptive field sizes increased by about 24% and an incomplete segregation of regions of on and off responsiveness was found in some cells, which made them hard to classify as simple or complex. Responses to flashed stimuli were smaller and more persistent. Their statistical significance notwithstanding, it seems unlikely that these relatively small response abnormalities in area 17 can fully account for the observed behavioural deficits.     

Simulation Studies of the Role of Intracortical Inhibition in the Formation of Sensitivity to Cross-Shaped Figures

The receptive fields of detector neurons for cross-shaped figures in the visual cortex were modeled in conditions of blockade of intracortical inhibition. The tuning of simulated neurons was compared with and without inhibition in the receptive field. In a simulated detector with convergence from two orientation detectors, acute tuning to the cross widened in the absence of inhibition, becoming invariant to the shape and orientation of the cross. A detector based on the disinhibition mechanism lost cross sensitivity when inhibition was blocked and became a detector for the orientation of a single bar. A model of a receptive field in which the inhibitory zones mask the tuning to a cross-shaped figure and in which blockade of inhibition affects only sensitivity is also proposed. We identified which of the properties of receptive field (configuration, location, zone weightings) allow them to simulate the properties of cat visual cortex field 17 neurons, these being sensitive to the shape and orientation of cross-shaped figures. __________ Translated from Zhurnal Vysshei Nervnoi Deyatel'nosti imeni I. P. Pavlova, Vol. 54, No. 6, pp. 767–775 November–December, 2004.     

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.     

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.     

Response properties of visual cortical neurons in cats reared in stroboscopic illumination

1. The response properties of 182 units were studied in the primary visual cortices (155 in area 18 and 27 in area 17) in eight cats reared from birth in a stroboscopically illuminated environment (frequency, 2/s; duration, 200 microseconds). Multihistogram quantitative testing was carried out in 82 units (64 in area 18 and 18 in area 17). Two hundred three neurons recorded and quantitatively tested in areas 17 and 18 of the normal adult cat were used for comparison. 2. Spatial characteristics of receptive fields investigated using hand-held stimuli were found to be abnormal. The correlation between receptive-field width and eccentricity was lost in area 18 and consequently, receptive fields were significantly wider in area 18 subserving central vision. Cells could be classified according to the spatial characteristics of their receptive fields. There was a much smaller proportion of end-stopped cells in strobe-reared animals. Orientation tuning in the deprived animals was normal except for a small number of cells that showed no selectivity for stimulus orientation. 3. Compilation of velocity-response curves made it possible to classify areas 17 and 18 neurons into four categories: velocity low-pass, velocity broad-band, velocity tuned, and velocity high-pass cells. The proportion of velocity high-pass cells was reduced in area 18 subserving peripheral vision, as was the proportion of velocity-tuned cells in area 18 subserving central vision. 4. In the strobe-reared animal velocity sensitivity was somewhat different from that of the normal animal. Neurons in area 18 subserving the peripheral visual field failed to respond to fast velocities. Neurons in area 17 subserving the central visual field in strobe-reared animals responded to slightly higher velocities than in the normal animal. 5. In the deprived animals the number of neurons that were selective to the direction of motion was strongly reduced. The majority of neurons failed to show a selectivity for direction at all velocities. A number of neurons could be directional at some velocities but were unreliable, since they inverted their preferred direction with velocity changes. 6. Binocular convergence onto visual cortical cells was perturbed. In area 18 the majority of neurons were driven by the contralateral eye. In area 17 most neurons could be driven only by either the ipsilateral or contralateral eye. 7. Quantitative testing (of direction selectivity, sensitivity to high velocities, response latency, and strength) and qualitative testing (receptive-field width, end stopping, and ocular dominance) showed that the normal influence of eccentricity on functional properties was strongly reduced by strobe rearing.     

Response variability and orientation discrimination of single cells in striate cortex of cat

Summary The response of single cells in the striate cortex of cat to a moving light bar of variable orientation was measured by a method providing data on the mean response as well as the standard deviation (SD) at the different stimulus orientations.At the optimal stimulus orientation the SD was about 1/3 of the mean response. Marked differences in this respect were found between simple and complex cells, i.e., the SD for the simple cells was about 1/2 of the mean response and about 1/4 for the complex cells.The variation coefficient (Vc = SD/mean) was minimal at the optimal orientation and increased relatively in the same manner for simple and complex cells as the stimulus orientation was varied away from optimal orientation. The Vc varied with the mean response at optimal orientation in a nonlinear manner. A function is proposed which fits this relationship and which is equally applicable for both simple and complex cells.The mean orientation discrimination (MOD) was defined as that change in orientation angle away from the optimal which produced a response statistically different — on the 1 % level — from the response to the optimal orientation. There were differences in MOD between the two sides of the orientation tuning curve: the mean of the smaller of the two values was 13.5 deg and of the larger 19.7 deg. No significant difference in MOD was found between simple and complex cells despite the fact that the halfwidth of the tuning curves for the two cell types was 19.5 deg and 31.6 deg, respectively.The preciseness in localization of the most sensitive part within the receptive field of single cells was calculated from the variability in time of occurrence of the smallest interspike interval. The degree of preciseness was found to be of the order of 1/4 of the receptive field diameter in both simple and complex cells. When nonoptimal stimulus orientations were presented, the preciseness significantly decreased in complex cells whereas it remained unchanged in simple cells.It is suggested that the same type of intracortical wiring produces orientation selectivity in simple and complex cells, and that the differences in tuning width are mainly due to a larger extension of inhibitory fields in the simple cells. Considering the cortical visual cells as elementary units in a network built for orientation detection and discrimination, the tuning width seems of minor importance for that function.     

Region-specificity of GABAA receptor mediated effects on orientation and direction selectivity in cat visual cortical area 18

The role of GABAergic inhibition in orientation and direction selectivity has been investigated with the GABA_A-Blocker bicuculline in the cat visual cortex, and results indicated a region specific difference of functional contributions of GABAergic inhibition in areas 17 and 18. In area 17 inhibition appeared mainly involved in sculpturing orientation and direction tuning, while in area 18 inhibition seemed more closely associated with temporal receptive field properties. However, different types of stimuli were used to test areas 17 and 18 and further studies performed in area 17 suggested an important influence of the stimulus type (single light bars vs. moving gratings) on the evoked responses (transient vs. sustained) and inhibitory mechanisms (GABA_A vs. GABA_B) which in turn might be more decisive for the specific results than the cortical region. To insert the missing link in this chain of arguments it was necessary to study GABAergic inhibition in area 18 with moving light bars, which has not been done so far. Therefore, in the present study we investigated area 18 cells responding to oriented moving light bars with extracellular recordings and reversible microiontophoretic blockade of GABAergig inhibition with bicuculline methiodide. The majority of neurons was characterized by a pronounced orientation specificity and variable degrees of direction selectivity. GABA_Aergic inhibition significantly influenced preferred orientation and preferred direction in area 18. During the action of bicuculline orientation tuning width increased and orientation and direction selectivity indices decreased. Our results obtained in area 18 with moving bar stimuli, although in the proportion of affected cells similar to those described in area 17, quantitatively matched the findings for direction and orientation specificity obtained with moving gratings in area 18. Accordingly, stimulus type is not decisive in area 18 and the GABA_A dependent, inhibitory intracortical computations involved in orientation specificity are indeed region-specific and in comparison to area 17 less effective in area 18.     

Inhibitory contributions to spatiotemporal receptive-field structure and direction selectivity in simple cells of cat area 17

Intracortical inhibition contributes to direction selectivity in primary visual cortex, but how it acts has been unclear. We investigated this problem in simple cells of cat area 17 by taking advantage of the link between spatiotemporal (S-T) receptive-field structure and direction selectivity. Most cells in layer 4 have S-T-oriented receptive fields in which gradients of response timing across the field confer a preferred direction of motion. Linear summation of responses across the receptive field, followed by a static nonlinear amplification, has been shown previously to account for directional tuning in layer 4. We tested the hypotheses that inhibition acts by altering S-T structure or the static nonlinearity or both. Drifting and counterphasing sine wave gratings were used to measure direction selectivity and S-T structure, respectively, in 17 layer 4 simple cells before and during iontophoresis of bicuculline methiodide (BMI), a GABAA antagonist. S-T orientation was quantified from fits to response temporal phase versus stimulus spatial phase data. Bicuculline reduced direction selectivity and S-T orientation in nearly all cells, and reductions in the two measures were well correlated (r = 0.81) and reversible. Using conventional linear predictions based on response phase and amplitude, we found that BMI-induced changes in S-T structure also accounted well for absolute changes in the amplitude and phase of responses to gratings drifting in the preferred and nonpreferred direction. For each cell we also calculated an exponent used to estimate the static nonlinearity. Bicuculline reduced the exponent in most cells, but the changes were not correlated with reductions in direction selectivity. We conclude that GABAA-mediated inhibition influences directional tuning in layer 4 primarily by sculpting S-T receptive-field structure. The source of the inhibition is likely to be other simple cells with certain spatiotemporal relationships to their target. Despite reductions in the two measures, most receptive fields maintained some directional tuning and S-T orientation during BMI. This suggests that their excitatory inputs, arising from the lateral geniculate nucleus and within area 17, are sufficient to create some S-T orientation and that inhibition accentuates it. Finally, BMI also reduced direction selectivity in 8 of 10 simple cells tested in layer 6, but the reductions were not accompanied by systematic changes in S-T structure. This reflects the fact that S-T orientation, as revealed by our first-order measures of the receptive field, is weak there normally. Inhibition likely affects layer 6 cells via more complex, nonlinear interactions.     

Stimulation of non-classical receptive field enhances orientation selectivity in the cat

We have investigated how the nonclassical receptive field (nCRF) affects dynamic orientation selectivity of cells in the primary visual cortex (V1) in anaesthetized and paralysed cats using the reverse correlation method. We found that tuning to the orientation of the test stimulus depends on the size of the stimulation area. A significant sharpening of orientation tuning was induced by nCRF stimulation, with the magnitude of the effect increasing with the size of stimulation. The effect of the nCRF on the temporal dynamics of orientation tuning was also investigated by examining the tuning over a range of delays from stimulus onset. We found small but detectable changes in both the preferred orientation and the bandwidth of tuning over time when the classical receptive field (CRF) was stimulated alone. Stimulation in nCRF significantly increased the magnitude of these temporal changes. Thus, nCRF stimulation not only enhances the overall orientation selectivity, but also enriches the temporal dynamics of cortical neurones, which may increase the computational power of the visual cortex in information processing.     

Receptive Fields of Bar and Cross-Shaped Figure Detectors in Classical and Combined Mapping

Connections between the excitatory and inhibitory zones of the receptive fields of neurons sensitive to the orientations of single bars and cross-shaped figures in the primary visual cortex were studied by classical and combined mapping. Factor and correlation analysis revealed different relationships between the main characteristics of neurons and their receptive fields for bar and cross detectors. Factor analysis of these connections showed that variables with the greatest weightings, combined into a single factor, were different for different detectors. In bar detectors, there was a direct correlational relationship between background activity and the weighting characteristics of the excitatory and inhibitory zones of their receptive fields. In cross-shaped figure detectors, the indexes of inhibition were positively related to the index of sensitivity to the figure, the characteristics of the excitatory zones of the receptive field, and background activity. In these detectors, increases in the area and weighting of additional receptive field excitatory zones in combined mapping were significantly greater than in bar detectors. The question of the difference in the mechanisms forming the receptive fields of bar and cross-shaped figure detectors, for which direct and recurrent horizontal inhibitory connections with surrounding neurons are more important, is discussed.     

Contrast-dependent changes in spatial frequency tuning of macaque V1 neurons: effects of a changing receptive field size

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.     

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.     

Development of orientation tuning in simple cells of primary visual cortex

Orientation selectivity and its development are basic features of visual cortex. The original model of orientation selectivity proposes that elongated simple cell receptive fields are constructed from convergent input of an array of lateral geniculate nucleus neurons. However, orientation selectivity of simple cells in the visual cortex is generally greater than the linear contributions based on projections from spatial receptive field profiles. This implies that additional selectivity may arise from intracortical mechanisms. The hierarchical processing idea implies mainly linear connections, whereas cortical contributions are generally considered to be nonlinear. We have explored development of orientation selectivity in visual cortex with a focus on linear and nonlinear factors in a population of anesthetized 4-wk postnatal kittens and adult cats. Linear contributions are estimated from receptive field maps by which orientation tuning curves are generated and bandwidth is quantified. Nonlinear components are estimated as the magnitude of the power function relationship between responses measured from drifting sinusoidal gratings and those predicted from the spatial receptive field. Measured bandwidths for kittens are slightly larger than those in adults, whereas predicted bandwidths are substantially broader. These results suggest that relatively strong nonlinearities in early postnatal stages are substantially involved in the development of orientation tuning in visual cortex.     

Axial responses in visual cortical cells: Spatio-temporal mechanisms quantified by Fourier components of cortical tuning curves

Summary The responses of 81 cells from area 17 in paralysed and aneasthetized cats were studied with moving spots and moving bars of different lengths. Tuning curves were measured and plotted as polar-plots. The strongest response of visual cortical cells to a moving bar occurs when the stimulus trajectory crosses the long axis of the receptive field (Hubel and Wiesel 1962). The optimal orientation for a moving and a flashing bar are identical, so that this response-type has been called the orientational component. For a moving spot, however, in most cases the strongest response occurs for motion along the receptive field long axis (axial component). Thus, the axial and orientational components are orthogonal (Wörgötter and Eysel 1989). It is shown that orientational and axial components can display direction selectivity and for short bar stimuli a superposition of the two orthogonal components is demonstrated. Such a superposition in general, resulted in a polar-plot with four peaks 90° apart from each other (four-symmetrical polar-plot). Polar-plots with three or two response peaks were also found; the actual number of response peaks depending on the direction selectivity of the components. In many cells pure axial responses could be elicited with a light spot which stimulates only motion dependent mechanisms. Thus, it was concluded that temporal facilitation is strongly involved in the generation of axial responses. Fourier analysis of polar-plots (SDO-analysis, Wörgötter and Eysel 1987; Wörgötter et al. 1990) was applied to determine the tuning strengths of the different components. In correspondence with the periodicities of a moving oriented stimulus in the visual field, the first harmonic represents directional selectivity, the second orientation selectivity and the fourth harmonic was used to quantify the four-symmetrical superposition effect. It was statistically shown that the strongest superposition (i.e. largest fourth harmonic) occurred for intermediate bar lengths (1–2°). For longer bars only the orientational and for shorter bars predominantly the axial component occurred. In monkey visual cortex (V1, V3) four-symmetrical polar-plots can be obtained even with long stimuli (De Valois et al. 1982; Felleman and Van Essen 1987). Consequentially, we show that a strong fourth order Fourier component occurs. This supports the importance of quantification of higher order symmetries in cortical tuning curves by higher order harmonics in SDO-analysis.     

Spectral receptive field properties explain shape selectivity in area V4

Neurons in cortical area V4 respond selectively to complex visual patterns such as curved contours and non-Cartesian gratings. Most previous experiments in V4 have measured responses to small, idiosyncratic stimulus sets and no single functional model yet accounts for all of the disparate results. We propose that one model, the spectral receptive field (SRF), can explain many observations of selectivity in V4. The SRF describes tuning in terms of the orientation and spatial frequency spectrum and can, in principle, predict the response to any visual stimulus. We estimated SRFs for neurons in V4 of awake primates by linearized reverse correlation of responses to a large set of natural images. We find that V4 neurons have large orientation and spatial frequency bandwidth and often bimodal orientation tuning. For comparison, we estimated SRFs for neurons in primary visual cortex (V1). Consistent with previous observations, we find that V1 neurons have narrower bandwidth than that of V4. To determine whether estimated SRFs can account for previous observations of selectivity, we used them to predict responses to Cartesian gratings, non-Cartesian gratings, natural images, and curved contours. Based on these predictions, we find that the majority of neurons in V1 are selective for Cartesian gratings, whereas the majority of V4 neurons are selective for non-Cartesian gratings or natural images. The SRF describes visual tuning properties with a second-order nonlinear model. These results support the hypothesis that a second-order model is sufficient to describe the general mechanisms mediating shape selectivity in area V4.     

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.     

Dynamics of the orientation-tuned membrane potential response in cat primary visual cortex

Neurons in the primary visual cortex are highly selective for stimulus orientation, whereas their thalamic inputs are not. Much controversy has been focused on the mechanism by which cortical orientation selectivity arises. Although an increasing amount of evidence supports a linear model in which orientation selectivity is conferred upon visual cortical cells by the alignment of the receptive fields of their thalamic inputs, the controversy has recently been rekindled with the suggestion that late cortical input--delayed by multiple synapses--could lead to sharpening of orientation selectivity over time. Here we used intracellular recordings in vivo to examine temporal properties of the orientation-selective response to flashed gratings. Bayesian parameter estimation demonstrated that both preferred orientation and tuning width were stable throughout the response to a single stimulus.     

Correlation of local and global orientation and spatial frequency tuning in macaque V1

Visual cortical neurones display a variety of visual properties. Among those that emerge in the primary visual cortex V1 are sharpening of selectivity for spatial frequency and for orientation. The selectivity for these stimulus attributes can be measured around the peak of the tuning function, usually as bandwidth. Other selectivity measures take into account the response across a broader range of stimulus values. An example of such a global measure is the circular variance of orientation tuning. Here we introduce a similar measure in the spatial frequency domain that takes into account the shape of the tuning curve at frequencies lower than the peak, called the low-spatial frequency variance. Our recent studies with dynamic stimuli suggest that the selectivity for spatial frequency and orientation is strongly correlated with the degree of suppression at low spatial frequencies and off-axis orientations. Here we extend the study of the global tuning to stimulus conditions that measure the response of cells to the presentation of drifting sinusoidal grating stimuli for periods of a few seconds. We find that under such steady-state stimulus conditions there is a strong correlation between the global selectivity measures, orientation circular variance and low spatial frequency variance. Consistent with previous studies, there is a weaker correlation between the local tuning measures, orientation and spatial frequency bandwidth. These results support the idea that there are multiple factors that contribute to tuning and that suppression observed in dynamic experiments is also likely to contribute to the global selectivity for steady-state stimuli.     

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.     

Receptive field size and response latency are correlated within the cat visual thalamus

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.