Functional inhibition in direction-selective retinal ganglion cells: spatiotemporal extent and intralaminar interactions

We recorded from ON-OFF direction-selective ganglion cells (DS cells) in the rabbit retina to investigate in detail the inhibition that contributes to direction selectivity in these cells. Using paired stimuli moving sequentially across the cells' receptive fields in the preferred direction, we directly confirmed the prediction of that a wave of inhibition accompanies any moving excitatory stimulus on its null side, at a fixed spatial offset. Varying the interstimulus distance, stimulus size, luminance, and speed yielded a spatiotemporal map of the strength of inhibition within this region. This "null" inhibition was maximal at an intermediate distance behind a moving stimulus: 1/2 to 11/2 times the width of the receptive field. The strength of inhibition depended more on the distance behind the stimulus than on stimulus speed, and the inhibition often lasted 1-2 s. These spatial and temporal parameters appear to account for the known spatial frequency and velocity tuning of ON-OFF DS cells to drifting contrast gratings. Stimuli that elicit distinct ON and OFF responses to leading and trailing edges revealed that an excitatory response of either polarity could inhibit a subsequent response of either polarity. For example, an OFF response inhibited either an ON or OFF response of a subsequent stimulus. This inhibition apparently is conferred by a neural element or network spanning the ON and OFF sublayers of the inner plexiform layer, such as a multistratified amacrine cell. Trials using a stationary flashing spot as a probe demonstrated that the total amount of inhibition conferred on the DS cell was equivalent for stimuli moving in either the null or preferred direction. Apparently the cell does not act as a classic "integrate and fire" neuron, summing all inputs at the soma. Rather, computation of stimulus direction likely involves interactions between excitatory and inhibitory inputs in local regions of the dendrites.     

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.     

Properties of the surround response mechanism of cat retinal ganglion cells and centre-surround interaction

1. The properties of the surround response mechanism of on-centre cells and its interaction with the centre mechanism were studied by recording from single optic tract fibres. In many of the experiments the spatial distribution of the light within the retinal image of the stimuli was measured.

2. Pure surround responses of on-centre cells were isolated using a centrally located steady light which selectively desensitized (adapted) the centre mechanism. This permitted a peripheral flashing stimulus whose luminance varied over a range as great as 1·38 log units to elicit surround responses which, for any given cell, were of invariant shape. The rate of decay of the firing frequency of the spike burst at `off' varied from cell to cell. The general characteristics of such pure surround responses to squarewave stimuli were described. The plot of the magnitude of pure responses against stimulus luminance, at constant background conditions, was curvilinear.

3. The pure surround response of two off-centre cells was isolated; it was similar in shape to the pure centre response of on-centre cells.

4. Interaction of centre and surround mechanisms of on-centre cells was studied by eliciting a pure central and a pure surround response from the same cell. The electronically obtained algebraic sum of these two pure responses equalled the mixed response of the ganglion cell to simultaneous presentation of the stimuli which evoked the pure responses when presented singly. This is probably best explained by algebraic summation of centre and surround inputs.

5. The pure surround response from two cells to a fixed flashing stimulus was attenuated by a steady field adapting light, both when this was superimposed upon the stimulus and when not superimposed. In the latter case, (i) when the spatial separation between the flashing stimulus and the adapting light was at a minimum, less than 10% of the adapting flux fell inside the boundaries of the stimulating flux, and (ii) the response was attenuated also if the adapting light was in the geometric centre of the receptive field. These results indicate that the adaptation pool of the surround mechanism extends to the central portions of the receptive field.

6. Nearly half the cells tested did not yield pure surround responses. This was probably due to differences, within the ganglion cell population, (i) of the spatial distribution of the ratio of centre to surround signal sensitivity and (ii) of differences in the ratio of centre to surround adaptivity in the receptive field middle. It was not due to excess adaptive flux falling outside the region of maximal centre mechanism adaptivity, nor due to excess stimulus flux falling inside the region of maximal signal sensitivity.

     

Responses of single units in the monkey superior colliculus to stationary flashing stimuli

1. Responses of pan-directional cells in the superficial layers of the superior colliculus in paralysed anaesthetized rhesus monkeys to stationary flashing stimuli have been studied. 2. The receptive field centre response is always of the transient excitatory on-off type, while the surround response is transient inhibitory both at light-on and at light-off. The receptive field centres are circular or slightly elliptical. The average size of the receptive field centres is much larger than that of retinal ganglion cells. All units except those in the far temporal periphery receive binocular input. In each unit the on and off responses have the same latency times. With increasing stimulus area, the latency time at light-on and at light-off first decreases and then remains constant. In most units the number of spikes in the burst at light-on and at light-off first increases, reaches a maximum and then decreases with increasing stimulus area. This decrease demonstrates the presence of an inhibitory surround. 3. A model of spatial and temporal properties of centre and surround mechanisms is tested. Addition of excitatory centre input and inhibitory surround input, which have different spatial and temporal properties, determines the output of the neurone. The centre mechanism gets excitatory input from retinal ganglion cells and shows saturation. The inhibitory surround mechanism is made by an inhibitory interneurone. It could not be decided whether the excitatory input for this interneurone comes from retinal axon collaterals (forward inhibition) or from axon collaterals of "principal" cells in the superior colliculus (backward inhibition).     

Some evidence concerning the physiological basis of the periphery effect in the cat's retina

Summary Peripheral regions of retinal ganglion cell receptive fields were investigated in cerveau isolé cats using automatically presented moving stimuli. By presenting appropriate stimulus configurations it was shown that the excitatory response to peripherally moving objects (the periphery effect) is not due to an extension of receptive field surround properties into the periphery. The differential effects of solid and striped stimuli indicated that the effectiveness of an object in eliciting the periphery effect is related to the amount of moving edge which it presents. Arguments are presented that the periphery effect results from interactions between spatially separated receptive field organizations. Certain difficulties which these findings raise with respect to the current definition of receptive field are discussed.     

Predictive coding for motion stimuli in human early visual cortex

The current study investigates if early visual cortical areas, V1, V2 and V3, use predictive coding to process motion information. Previous studies have reported biased visual motion responses at locations where novel visual information was presented (i.e., the motion trailing edge), which is plausibly linked to the predictability of visual input. Using high-field functional magnetic resonance imaging (fMRI), we measured brain activation during predictable versus unpreceded motion-induced contrast changes during several motion stimuli. We found that unpreceded moving dots appearing at the trailing edge gave rise to enhanced BOLD responses, whereas predictable moving dots at the leading edge resulted in suppressed BOLD responses. Furthermore, we excluded biases in directional sensitivity, shifts in cortical stimulus representation, visuo-spatial attention and classical receptive field effects as viable alternative explanations. The results clearly indicate the presence of predictive coding mechanisms in early visual cortex for visual motion processing, underlying the construction of stable percepts out of highly dynamic visual input.     

Response to motion in extrastriate area MSTl: disparity sensitivity

Many neurons in the lateral-ventral region of the medial superior temporal area (MSTl) have a clear center surround separation in their receptive fields. Either moving or stationary stimuli in the surround modulates the response to moving stimuli in the center, and this modulation could facilitate the perceptual segmentation of a moving object from its background. Another mechanism that could facilitate such segmentation would be sensitivity to binocular disparity in the center and surround regions of the receptive fields of these neurons. We therefore investigated the sensitivity of these MSTl neurons to disparity ranging from three degrees crossed disparity (near) to three degrees uncrossed disparity (far) applied to both the center and the surround regions. Many neurons showed clear disparity sensitivity to stimulus motion in the center of the receptive field. About (1)/(3) of 104 neurons had a clear peak in their response, whereas another (1)/(3) had broader tuning. Monocular stimulation abolished the tuning. The prevalence of cells broadly tuned to near and far disparity and the reversal of preferred directions at different disparities observed in MSTd were not found in MSTl. A stationary surround at zero disparity simply modulated up or down the response to moving stimuli at different disparities in the receptive field (RF) center but did not alter the disparity tuning curve. When the RF center motion was held at zero disparity and the disparity of the stationary surround was varied, some surround disparities produced greater modulation of MSTl neuron response than did others. Some neurons with different disparity preferences in center and surround responded best to the relative disparity differences between center and surround, whereas others were related to the absolute difference between center and surround. The combination of modulatory surrounds and the sensitivity to relative difference between center and surround disparity make these MSTl neurons particularly well suited for the segmentation of a moving object from the background.     

Classification of turtle retinal ganglion cells

1. Receptive fields of 78 retinal ganglion cells were analyzed for their responses to moving and stationary lights that were presented under a variety of stimulus conditions. All cells were sensitive to moving stimuli, and their receptive fields often comprised excitatory and inhibitory sub-regions. 2. Properties used in the classification included responses to stationary flashed stimuli, receptive-field organization, changes in stimulus wavelength and adaptation, movement velocity, and direction of stimulus movement. Eight functional cell classes were derived: simple, ON-sustained, annular, wavelength-sensitive, directionally selective, bar-shaped, large-field, and velocity. 3. Simple cells, representing 21% of the sample, had circular or oval receptive fields of 3-22 degrees that gave transient responses to stationary, flashed lights. Many of these cells, but not all, showed antagonistic center-surround organizations. ON-sustained cells responded for the duration of the stimulus flash or for the duration of a light flash moving through the receptive field. These units comprised 8% of the sample; they had small, circular, non-directional receptive fields and they were most sensitive to red light. Their field sizes did not vary with changes in adaptation level. 4. Annular cells (4% of the sample) gave no responses to any stimulation in the field center, but they responded strongly to stimulation in the surround area, especially to stimuli that moved very slowly through the region. Annular cells were nondirectional, with circular centers of 5-6 degrees diam and annular surround widths of 2-4 degrees. They responded best in light adaptation. 5. Wavelength-sensitive cells, similar to most of the cells sampled, were sensitive to red light when light-adapted. Some cells in addition showed input from rods under dark adaptation. Intensity-response curves for these latter cells showed clear changes from one input to the other as the cells' functional ranges were explored. Some cells responded best to short- or middle-wavelength light, but these were more rarely met. Where multiple receptor inputs could be identified, long-wavelength stimuli evoked transient responses, whereas short-wavelength stimuli favored more sustained spike trains. Wavelength-sensitive cells in this category comprised 5% of the sample.     

Responses to visual contours: spatio-temporal aspects of excitation in the receptive fields of simple striate neurones

1. The properties of the receptive fields of simple cells in the cat striate cortex have been studied by preparing average response histograms both to moving slits of light of different width and to single light-dark edges or contours.2. The movement of a narrow (< 0.3 degrees ) slit across the receptive field gives rise to average response histograms that are either unimodal, bimodal or multimodal. A slit of light has leading (light) and trailing (dark) edges. By increasing the width of the slit it was shown that a discharge peak in the histogram coincides with the passage of one or other of the two edges over a particular region (discharge centre) in the receptive field. Each edge has its own discharge centre which is fired when the edge has the correct orientation and direction of movement.3. The discharge centres in forty-three simple cell receptive fields were located by using one or more of the following stimuli for each cell:(i) slits of different width;(ii) single light and dark edges;(iii) a wide (3 degrees ) slit moved over a range of different velocities.The same locations were obtained when all three procedures were used on the same cell.4. Most cells (79%) discharged to both edges though not necessarily in the same direction of movement. The majority (72%) fired in only one direction and most commonly (51%) the cells responded to both edges in this one direction. In only 16% of cells did both types of edge excite in both directions of movement. When the one type of edge, light or dark, was considered, 84% of the cells were direction selective and, for these cells, the other edge fired only in the same direction (51%), in both directions (7%), only in the opposite direction (5%) or not at all (21%).5. Cells responding in one direction with a unimodal average response histogram may be responding to both edges, the two responses being concealed in the one discharge peak. The two discharge centres are then either nearly coincident or, more usually, slightly offset with respect to one another. Most commonly the dark edge centre is slightly in advance of the light edge centre.6. The discharge peaks in the bimodal and multimodal types come from discharge centres that are spatially separate, each centre firing to only one type of edge. In the case of the bimodal type the light edge centre always lies ahead of the dark edge centre.7. When a cell responds to a single edge in both directions of movement, the type of contrast effective in one direction is always the reverse of that in the other. When the cell responded in both directions, whether to one or both edges, most commonly a light edge discharge centre in one direction occupied approximately the same location in space as the dark edge centre in the reverse direction and vice versa for the other edge.8. Temporal aspects of the discharge of simple cells have been examined by recording the responses to moving slits and single edges over a wide range of velocities.     

Receptive field organization of complex cells in cat striate cortex

Summary The receptive field organization of complex cells was studied by analyzing interaction effects between two stationary flashing light stimuli. One was placed in the most responsive part of the receptive field to produce activity against which effects of the other in different visual field positions could be determined.The receptive field was spatially organized into antagonistic center and flanks just like the fields of simple cells. However, both center and flanks were found within the receptive field area where a single slit evoked discharge. Center and flanks were elongated along the optimal stimulus orientation. The flanks were displaced from the center normal to optimal stimulus orientation.In the center, ON- and OFF-responses were usually about equal in strength and the maximum ON- and OFF-responses occurred in about the same position. This shows that complex cells are activated by input from both ON- and OFF-center cells in the lateral geniculate nucleus (LGN) where the receptive field centers of the LGN cells overlap closely. This explains most of the specific features of complex cells, e.g., the spatially overlapping ON- and OFF-zones, the large response field, the repetitive firing when a slit moves over the receptive field, and the marked non-linear spatial summation.Strong flank suppression occurred with both ON and OFF. The effects were usually stronger on one side of the center. Maximal suppression occurred on the same side with both ON and OFF. This is consistent with the interpretation that complex cells are inhibited by input from both LGN ON- and OFF-center cells with overlapping receptive field centers.A model presuming that complex cells have overlapping but acentric excitatory and inhibitory fields was tested by computer simulation and shown to fit the experimental data. This is the same model as presented for simple cells in the preceding paper (Heggelund 1980), except that the excitatory and inhibitory fields of simple cells have input from either ON- or OFF-center LGN cells, whereas in complex cells they have input from both types.The project was financially supported by the Norwegian Research Council for Science and Humanities     

Receptive-field properties of transcallosal visual cortical neurons in the normal and reeler mouse

The receptive-field properties of neurons in the striate visual cortex of normal and reeler mutant mice were studied with single-unit recording methods in order to determine whether the connections underlying these properties are altered by the developmental abnormality in neuronal position that characterizes reeler neocortex. Neurons with a projection through the corpus callosum were selected for study because they form a physiologically identifiable class of visual cortical neurons with a characteristic distribution of receptive-field properties that can be compared for normal and reeler cortex. Transcallosal cortical neurons in area 17 near its border with area 18a were identified by antidromic stimulation delivered through bipolar electrodes in the contralateral cortex. A computer controlled the visual stimuli, data acquisition, and analysis. Transcallosal neurons were principally found in layers II-III and V in the normal cortex and in a broand band deep in the reeler cortex. These populations had similar distributions of antidromic latencies, indicating that the neurons sampled from normal and reeler cortex were taken from populations with similar axonal diameters and soma sizes. The receptive-field properties of 46 units in 22 normal mice and 28 units in 11 reeler mice were characterized. Transcallosal neurons in both normal and reeler cortex were usually binocularly responsive and dominated by input from the contralateral eye. They exhibited either nonoriented (31 and 48%, respectively) or oriented (69 and 52%) receptive fields. Tuning 10 stimulus velocity was broad, with peak velocity sensitivities ranging from 1 to 1,000 degrees/s. Directional selectivity was present in 41% of normal units ad 32% of reeler units. There was no significant difference between normal and reeler cortex in the distribution of these properties. Transcallosal neurons were also examined for the presence of an inhibitory surround by comparing their responses to moving or stationary stimuli of varying sizes. Of the tested neurons, most (11/17 in normal cortex, 6/9 in reeler) showed evidence of a decrease in response to large moving stimuli. A large proportion (16/20) of normal neurons tested with stationary flashing stimuli had some degree of surround inhibition whereas significantly fewer (5/17) neurons in reeler cortex had this property. Thus, transcallosal neurons in reeler cortex less frequently had an inhibitory surround demonstrable with stationary flashing stimuli, but this difference between normal and reeler was not apparent with a moving stimulus.(ABSTRACT TRUNCATED AT 400 WORDS)     

The inhibitory components in the responses of the lateral suprasylvian area neurons to moving stimuli in cats

The inhibitory components in the neuronal responses of the cat's lateral suprasylvian area (LSA) to moving bright and dark stimuli were investigated. The LSA neurons could be divided into two groups. Neurons of the first group (33%) do not reveal spatial displacement of the inhibitory zones and show displacement of the discharge centers in the receptive field only for one polarity of contrast of moving stimuli, either brighter or darker than the background. The second group (67%) contained the neurons which showed a spatial displacement of the inhibitory components and discharge centers in the receptive field for either polarity of contrasts of the moving stimuli. Tested with stationary flashing stimuli, the majority of neurons in both groups had overlapping ON-OFF discharge regions within their receptive fields. The results obtained with moving stimuli of different speeds and with the masking method suggest the rebound origin of the inhibitory responses in LSA neurons.     

Periodic excitability changes across the receptive fields of complex cells in the striate and parastriate cortex of the cat.

1. Complex cells in cortical areas 17 and 18 of the cat have been studied in response to narrow slits and edges moving across the receptive field in the preferred direction and also to stationary slits of different widths. 2. Average response histograms, recorded as a narrow slit was moved across the receptive field, displayed a periodic series of peaks above a base line level. The response histogram for most area 17 and 18 cells contained five principal peaks; sometimes one or two weaker peaks were present at receptive field borders. The histogram for one cell located at the area 17-18 border showed thirteen distinct peaks. Periodic response patterns were also generated as an extended edge was moved across the receptive field. Plots of cell responses versus slit width for stationary slits of different widths also indicated periodic response pattern. 3. The accuracy of determining the preferred slit orientation was the single most important requirement for demonstrating the periodic response pattern. Significant changes in the appearance of the periodic pattern occurred even upon 5 degrees rotations away from the preferred orientation. 4. Average response histograms were also studied over a wide range of moving slit velocities. The number of peaks across corresponding spacings within the recewptive field remained constant over a range of velocities. Response amplitudes, however, were velocity dependent. Thus the response peaks remain associated with fixed positions within visual space independent of stimulus velocity, even though temporal as well as spatial factors may be involved in response selectivity and the periodic modulation. The most striking periodic response histograms were generated at the velocities which produced the greatest cell firing rates. Area 17 complex cells responded well to velocities of less than 0-5 degrees to 6-0 degrees/sec, but cells in area 18 generally required higher velocities, sometimes as high as 20 degrees--30 degrees/sec, for a good response. 5. Spatial frequencies for the periodic component of the receptive field for area 17 cells in the central visual area covered a range of three octaves up to 5 cycles/degree, and area 18 cells included another octave on the low frequency side. The spatial frequency of a cell was found to be roughly inversely proportional to the receptive field width. Only a small sample of area 18 cells was studied, but these cells tended to represent low spatial frequencies and to respond selectively to high velocity stimuli...     

Ganglion cell discharge and proximal negativity in the pigeon retina.

1. Proximal negativity has been compared with the spike responses and membrane responses of pigeon ganglion cells. 2. Spike discharge peaks develop during the rising phase of proximal negativity. 3. Proximal negativity is never directional, even when recorded concurrently with directional ganglion cells. 4. To moving stimuli, on-centre cells respond in association with the leading edge of the stimulus, off-centre cells with the trailing edge, and on-off cells with the leading and trailing edges. Proximal negativity is produced in association with the leading and trailing edges. 5. Quasi-intracellular records show that discharge peaks occur in association with depolarizing potentials. The synaptic responses of on-off cells bear a closer resemblance to the proximal negative response than do the responses of on-centre cells, off-centre cells, or directional cells.     

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.     

Cutaneous masking. II. Geometry of excitatory andinhibitory receptive fields of single units in somatosensory cortex of the cat.

1. The responses of single neurons in the primary somatosensory cortex of the cat to brief air-pulse stimuli were quantitatively examined. These controlled natural stimuli activated almost exclusively rapidly adapting hair units which, on systematic movement of the stimulus through the receptive field, gave unit-response profiles that showed the classical unimodal tent-shaped distribution. 2. Conditioning stimulus-induced inhibition of a response evoked by a fixed test stimulus was measured by systematically moving the conditioning stimulus through the receptive field. The spatial distribution of in-field inhibitory activity was unimodal and highly covariant with that of the conditioning excitation, the peak inhibition corresponding to the functional center of the excitatory receptive field. 3. Nearly one-half of the units studied evidenced inhibition extending beyond the excitatory receptive field, forming a "surround" inhibitory region; but these were usually restricted areas with rather weak inhibitory effects. 4. Time-course measuring revealed, on the average, inhibition effects measureable from 10 ms before to some 70 ms following conditioning stimulation, with peak inhibition delayed some 10--15 ms from the conditioning stimulus onset. We showed the backward inhibition, occurring with the test stimulus delivered before the onset of the conditioning stimulus, to be a property of the test response duration. Inhibition measured in the surround areas had essentially the same time course as the inhibition calculated from measurements made within the receptive fields. 5. The spatial and temporal profiles of the excitatory and inhibitory cortical unitary activity are thus very similar to the parametric features of psychophysical enhancement and masking. These findings suggest that the excitatory and inhibitory activities related to individual stimuli interact in multipoint stimulus paradigms so that simple unimodal composite profiles are synthesized.     

Neural responses in motor cortex and area 7a to real and apparent motion

The neural activity in area 7a and the arm area of motor cortex was recorded while real or path-guided apparent motion stimuli were presented to behaving monkeys in the absence of a motor response. A smooth stimulus motion was produced in the real motion condition, whereas in the apparent motion condition five stimuli were flashed successively at the vertices of a regular pentagon. The stimuli moved along a low contrast circular path with one of five speeds (180–540 deg/s). We found strong neural responses to real and apparent motion in area 7a and motor cortex. In the motor cortex, a substantial population of neurons showed a selective response to real moving stimuli in the absence of a motor response. This activity was modulated in some cases by the stimulus speed, and some of the neurons showed a response during a particular part of the circular trajectory of the stimulus; the preferred stimulus angular locations were evenly distributed across this neuronal ensemble. It is likely that these neural signals are continuously available to the motor cortex in order to generate responses that demand immediate action. In area 7a, two overlapping populations of neurons were observed. The first comprised cells the activity of which was tuned to the angular location of a circularly moving stimulus in the real motion condition. These cells also responded to apparent motion at high stimulus speeds. A visual receptive field analysis showed that the angular tuning in most of the area 7a neurons did not depend on the spatial location of the stimulus in relation to their receptive field. The second population was selective to apparent moving stimuli and showed a periodic entrainment of activation with the period of the inter-stimulus interval of the flashing dots. Both the angular location and the inter-stimulus interval neural signals can be used to generate precise behavioral responses towards real or apparent moving stimuli.     

Receptive field analysis: responses to moving visual contours by single lateral geniculate neurones in the cat

1. Responses of single geniculate cells to moving light and dark bars and light/dark edges were studied in cats anaesthetized with nitrous oxide/oxygen (70%/30%).2. Over 95% (230 out of 241) of geniculate cells had antagonistic centre-surround receptive fields. Their responses could be characterized as centre-activated or centre-suppressed depending on the receptive field type (ON- or OFF-centre) and the contrast between stimulus and the background (brighter or darker than the background). Moving light and dark edges evoked responses which were very similar to the responses evoked by these stimuli in simple cells of striate cortex.3. A number of cells (45) with antagonistic centre-surround receptive fields were classified according to their X/Y (sustained/transient) properties. Units with sustained properties (X-cells) did not increase their firing rate with an increase of stimulus velocity and some of them showed a clear-cut preference for slow movement (around 1-2 degrees /sec). On the other hand, units with transient properties (Y-cells) showed a clear-cut preference for fast-moving stimuli (50-100 degrees /sec.)4. Elongation of the stimulus beyond the antagonistic surround in both X- and Y-cells produced a clear-cut reduction of amplitude of both centre and surround components of the response. Thus the existence of a suppressive field component beyond the antagonistic surround is confirmed.5. About 5% of cells had receptive fields which did not have an antagonistic centre-surround organization but gave a mixed ON-OFF discharge from the central region of the field. Around the central region there was a silent suppressive zone. These units were not directionally selective, responded preferentially to fast-moving stimuli (25-100 degrees /sec) and had a substantial (spontaneous) maintained activity.     

Interaction effects of visual contours on the discharge frequency of simple striate neurones

1. The discharge frequency of simple neurones in the cat striate cortex responding to the two edges of a slit of light moving over their receptive fields was studied as a function of slit width. While one edge of the slit was discharging the cell, the other edge had a modifying influence on that discharge either by way of facilitation or of inhibition.2. The most common form of the curve relating discharge frequency and slit width had a maximal discharge at narrow slit widths (< 0.5 degrees ) and relative inhibition at medium widths (between 0.5 degrees and 2 degrees ). At greater slit widths there was usually a region of facilitation before the effects of the two edges became independent of one another. Three other response patterns to slits of different width are described.3. The curve relating slit width and response amplitude for a particular cell provides an important clue to the various activity profiles for that cell. An activity profile plots the excitability of a cell along a line through the receptive field in the direction of stimulus movement. Each type of edge, light and dark, has its own set of activity profiles which differ depending upon stimulus parameters such as the direction of the movement of the edge.4. Two other methods were used to provide further data concerning the activity profiles and as a check on the evidence provided by the responses to slits of different width. One of these two methods used the test stimulus against the background of an artificially produced maintained discharge and the other involved the interaction of the two receptive fields of binocularly activated cells.5. A model is put forward to explain the receptive field organization of simple striate neurones which takes into account not only the main features of what is known concerning the synaptology of the visual cortex but also the new data provided by the present paper and the one which precedes it.