Changes in reactivity of visual cortical neurons after local photic stimulation of their receptive fields in cats

Recovery cycles of unit responses in the primary visual cortex to local photic stimulation of their receptive fields were studied in unanesthetized, immobilized cats by the paired stimulus method. In most cases the process of recovery of neuronal reactivity did not follow a steady course. Recovery from depression evoked by the first stimulus took place more suddenly in neurons in the central part of the visual field, and initial recovery of activity was more complete than in peripheral neurons. Differences in the synchronization of inhibitory and excitatory inputs to neurons responsible for central and peripheral vision are discussed.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 13, No. 3, pp. 233–240, May–June, 1981.     

Visual cortical unit responses to photic stimulation of different zones of the receptive fields in cats

In acute experiments on unanesthetized curarized cats the intensity functions, response thresholds, inhibition thresholds, and differential sensitivity of 96 neurons in the primary visual projection cortex were investigated by extracellular recording of unit activity during central and peripheral stimulation of their receptive fields. In darkness the neurons had wide threshold and above-threshold reliefs (3–30°). The threshold reliefs of the receptive fields of some cells were found to be V-shaped, whereas others were marked by alternation of zones of increased and reduced excitability. Sensitivity of both excitatory and inhibitory inputs of the receptive field as a rule was greatest in the center. Inhibitory inputs of different cortical neurons were much more standard and less sensitive to light, and they were mainly activated within the intermediate (mesoptic) range of brightnesses. During light adaptation the threshold contour of the receptive field narrows sharply, mainly because of the fall in sensitivity of its peripheral inputs. Compared with the lateral geniculate body and retina, the relative number of low-threshold elements, sensitivity in the system of inhibitory elements, and differential brightness sensitivity are greater in the cortex. The mechanisms of formation of receptive fields of cortical neurons and their modification during changes in the level of adaptation, and also the role of excitatory and inhibitory inputs of the cell in these effects are discussed.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 11, No. 3, pp. 227–235, May–June, 1979.     

Receptive field dynamics of neurons in the cat visual cortex and lateral geniculate body

Spike responses of single neurons in the primary visual cortex and lateral geniculate body to random presentation of local photic stimuli in different parts of the receptive field of the cell were studied in acute experiments on curarized cats. Series of maps of receptive fields with time interval of 20 msec obtained by computer enabled the dynamics of the excitatory and inhibitory zones of the field to be assessed during development of on- and off-responses to flashes. Receptive fields of all cortical and lateral geniculate body neurons tested were found to undergo regular dynamic reorganization both after the beginning and after the end of action of the photic stimulus. During the latent period of the response no receptive field was found in the part of the visual field tested, but later a small zone of weak responses appeared only in the center of the field. Gradually (most commonly toward 60–100 msec after application of the stimulus) the zone of the responses widened to its limit, after which the recorded field began to shrink, ending with complete disappearance or disintegration into separate fragments. If two bursts of spikes were generated in response to stimulation, during the second burst the receptive field of the neuron changed in the same way. The effects described were clearly exhibited if the level of background illumination, the intensity of the test bars, their contrast with the background, duration, angles subtended, and orientation were varied, although the rate and degree of reorganization of the receptive field in this case changed significantly. The functional importance of the effect for coding of information about the features of a signal by visual cortical neurons is discussed.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 14, No. 6, pp. 622–630, November–December, 1982.     

Temporal summation in cat visual cortical neurons

Characteristics of temporal summation in neurons of area 17 of the visual cortex in acute experiments on unanesthetized, immobilized cats. During light adaptation, extracellular spike responses of these neurons to optimal local photic stimuli of varied duration — from 5 to 1000 msec — were studied. The critical duration of temporal summation of excitation, determined by the supraliminal method using the criterion of maximal discharge frequency in the first volley of the spike response, varied in different cells from 5 to 100 msec; neurons with summation lasting 15–100 msec (mean 31.45±5.67 msec) were found most frequently. Neurons with central receptive fields differed significantly from cellswith peripheral fields in the shorter critical duration of temporal summation, the lower frequency of spontaneous discharges, and the shorter duration of the first volley of the response. Summation time in neurons with simple receptive fields was significantly shorter than in neurons with complex receptive fields. The results of these experiments are compared with data in the literature obtained by the study of retinal and lateral geniculate neurons in cats and are discussed from the stand-point of division of ascending afferent projections in the visual system into X-and Y-groups (Ia and Ib).Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 13, No. 4, pp. 345–352, July–August, 1981.     

Receptive fields of neurons in the lateral suprasylvian area of the cat

The structure of receptive fields of single neurons in the lateral suprasylvian area of the cat's cortex was studied. Receptive fields of neurons in this area are larger (up to 2000 deg² or more) than those of the visual projection cortex. A difference was found in the sizes of these fields of the same neuron when measured by presentation of a black object and spot of light. Experimental results showed that most neurons of the area (104 of 148) that are sensitive to visual stimulation respond clearly to flashes of a stationary spot of light. Because of this feature the structure of the receptive fields of the neurons were studied by point by point testing of their whole surface. Intensities of on- and off-components of on-off neurons were found to differ. Only 16% of receptive fields had equal numbers of discharges in on- and off-components of the on-off response. Dominance of one component was observed in 84% of on-off neurons. Receptive fields with several discharge centers are a characteristic feature of neurons in this area. A concentric organization of the receptive fields was found in 11% of neurons.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan, Translated from Neirofiziologiya, Vol. 14, No. 3, pp. 278–283, May–June, 1982.     

Responses of nervous elements of the visual cortex of the chipmunk Eutamias sibiricus to moving and stationary stimuli]

The receptive field organization of cortical units has been studied in experiments with testing by moving and stationary light spots. The size of the receptive fields varied from 3 degrees to 10 degrees. Receptive fields which were tested by a stationary light spot exhibited various types of organization. Some of the neurons produced extensive excitatory on- and off-responses to stimulation by a light spot. Neuronal excitation evoked by light decreased if the stimulus was near the field boundary. Some of the neurons produced either on- or off-responses in any point of the receptive field. A small part of neurons had receptive fields with on- and off-reactions in the center, and either on- or off-responses at the peripheral zones. Most of the neurons exhibited specialization with respect to high-speed motion.     

Responses of receptive fields of frog retinal ganglion cells to the leading and trailing edges of moving stimuli

The position of on- and off-discharge centers in class 1 and 3 receptive fields of the frog retina was determined with the aid of moving bars of different lengths. On- and off-centers of receptive fields of the first group coincide, those of the second are spatially separate, and in fields of the 3rd group the discharge center of one contrast sign occupies the central position and discharge centers of the opposite sign are located at the periphery, to its right and left. Receptive fields of the frog retina thus have features which approximate them to the concentric receptive fields of geniculate neurons and the fields of the cat visual cortex. Asymmetry in the responses was found: during movement in opposite directions the distance between the discharge centers changed, during movement to one side only one of the peripheral centers was revealed, whereas during movement to the other side the second center was revealed on the opposite side of the receptive field. This asymmetry of spatiotemporal relations in the receptive fields is similar to that found in the fields of cortical neurons and is connected with their directional properties.Research Institute of Applied Mathematics and Cybernetics, N. I. Lobachevskii State University, Gor'kii. Translated from Neirofiziologiya, Vol. 12, No. 1, pp. 75–85, January–February, 1980.     

Receptive fields of visual cortical neurons during changes in parameters of photic stimulation in cats

Spatial excitability contours in receptive fields of visual cortical neurons during changes in the physical and physiological parameters of photic stimulation were investigated in acute experiments on immobilized cats under conditions of dark, mesopic, and low photopic adaptation. With the change from dark to low mesopic adaptation the shape and size of the receptive fields detected by testing with flashes of constant intensity are unchanged, but with the transition to low photopic adaptation the receptive field becomes long and very narrow in 72% of cases, and the acuity of its orientational and directional tuning becomes much sharper. Against an unchanged background illumination, loss of brightness of the test light slit leads to narrowing of the measurable receptive field. Excitability contours of the receptive field estimated on the basis of absolute threshold of the cell response and level of intensity necessary to obtain the same number of spikes in the response become much narrower as the threshold criterion rises and during dark adaptation. Reactivity contours of the receptive field in response to stimulation of physiologically equal intensities (equal to the increase in threshold) under conditions of photopic adaptation also are much narrower than reactivity contours under conditions of dark adaptation. Evaluation of receptive fields with allowance for the possible contribution of effects of light scatter on the screen and in the ocular media showed that in most cases their size cannot be explained by these phenomena.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 12, No. 2, pp. 115–123, March–April, 1980.     

Effects of local adaptation of receptive field on sensitivity of the cat visual cortex neurons to cruciform images

The responses to flashing single light bars of different orientation and to cruciform images (CI) were compared in 9 neurons of the cat striate cortex possessing high specific sensitivity to CI, during local adaptation of various receptive field (RF) zones. In most neurons, a two- to threefold reduction in the response to CI with a constantly present bar of optimum or orthogonal orientation, if compared with a response to the figure consisting of two flashing bars, was found. Responses to the CI including an adaptation bar were often increased, if compared with those observed at usual orientation tuning. The role of a cross-orientation inhibition in the formation of a selective sensitivity to CI in the neurons of the visual cortex is discussed.Neirofiziologiya/Neurophysiology, Vol. 27, No. 2, pp. 134–139, March–April, 1995.     

Thresholds and latencies of retinal ganglion cell responses in cats to local stimulation of various parts of their receptive field

Depending on their responses to separate stimulation of the center and periphery of the receptive field, all ganglion cells of the cat retina can be subdivided into two types: ON-center (OFF-periphery) and OFF-center (ON-periphery). By all the parameters studied these ON- and OFF-systems were symmetrical. This apparently reflects, first, the equality of informativeness of illumination and darkening of individual areas of the visual field and, second, adaptation in order to widen the dynamic range of the visual channel of information transmission. Thresholds of unit responses to stimulation of the periphery and center of their receptive fields were identical. The latent periods of the unit responses were much longer in the first case than in the second. This is regarded as providing the functional basis for discrimination between "central" and "peripheral" unit responses by higher structures.Institute of Control Problems, Academy of Sciences of the USSR. Translated from Neirofiziologiya, Vol. 3, No. 6, pp. 644–649, November–December, 1971.     

Investigation of functional organization of receptive fields in the cat visual cortex

Receptive fields of neurons in Area 17 of the visual cortex were investigated in cats. Concentrically shaped fields, fields responding selectively to orientation of a strip or edge, and fields which can be regarded as intermediate between the first two types are described. The boundary between zones of summation and of lateral inhibition coincides in some receptive fields with the boundary between central and peripheral zones with opposite forms of response, while in other fields they do not coincide. For some cells there is no peripheral zone or it may disappear with worsening of the state of function. Cells were observed for which an increase in area of the stimulus in the central zone inhibits the response reaction. Analysis of these data suggests that several cells of the geniculate ganglion converge on some cortical neurons, and several cortical cells on others. An effect of adaptive inhibition was found in which constant illumination of an area in the center of the receptive field inhibits the response in another part. It is shown that this effect is unconnected with the action of scattered light. Constant illumination of the peripheral part of the receptive field deinhibits adaptive inhibition. The boundary between the zones of summation and of lateral inhibition coincides with the boundary between the zones of adaptive inhibition and deinhibition.I. V. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 1, No. 1, pp. 90–100, July–August, 1969.     

Rabbit visual cortical neurons with simple and complex receptive fields

Receptive fields of neurons of the rabbit visual cortex selective for stimulus orientation were investigated. These receptive fields were less well differentiated than those of the analogous neurons of the cat visual cortex (large in size and circular in shape). Two mechanisms of selectivity for stimulus orientation were observed: inhibition between on and off zones of the receptive field (sample type) and oriented lateral inhibition within the same zone of the receptive field (complex type). Lateral inhibition within the same zone of the receptive field also took place in unselective neurons; "complex" selective neurons differed from them in the orientation of this inhibition. A combination of both mechanisms was possible in the receptive field of the same neuron. It is suggested that both simple and complex receptive fields are derivatives of unselective receptive fields and that "complex" neurons are not the basis for a higher level of analysis of visual information than in "simple" neurons.A. N. Severtsov Institute of Evolutionary Morphology and Ecology of Animals, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 10, No. 1, pp. 13–21, January–February, 1978.     

Structure of receptive fields of cat pulvinar neurons sensitive to photic stimulation

Receptive fields of 262 pulvinar neurons were studied. Receptive fields of 142 of these neurons were studied in detail with the aid of a stationary spot of light, flashing in different parts of the receptive field. Depending on responses to presentation of the stationary stimulus the neurons were divided into six groups. The first group included neurons with on—off responses to photic stimulation (44 of 142), the second group neurons with off responses only (42 of 142). In cells of the third group (19 of 142) an on response only was recorded in all structures of the receptive field tested. Neurons of the fourth group (eight of 142) had a receptive field of similar structure to that of the simple receptive fields of neurons in cortical area 17. The fifth group (10 of 142) included neurons with a receptive field of concentric structure, the sixth (19 of 142) consisted of neurons with receptive fields with multiple discharge centers. The structure of the receptive field of these neurons was mosaic, with an irregular distribution of exciting and "silent" zones. The mean response latency of the pulvinar neurons was 40–70 msec. Responses of neurons with shorter (20 msec) and longer (130–160 msec) latent periods also were recorded.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 11, No. 1, pp. 3–10, January–February, 1979.     

Inhibitory zones of receptive fields of the lateral geniculate body and visual cortex of the cat

Unit responses to moving strips were investigated. The organization of the inhibitory zones in the receptive fields of the lateral geniculate body and visual cortex of the cat was compared. The response in the receptive field of the lateral geniculate body was inhibited only during simultaneous stimulation of the excitatory and inhibitory zones of the field. Stimulation of the inhibitory zone in the receptive field of the visual cortex was effective for a long time (several hundreds of milliseconds) after stimulation of the excitatory zone. The inhibitory zones of the simple and complex receptive fields of the visual cortex differed significantly. An increase in the width of the strip above the optimal size reduced the inhibitory effect in the complex fields. This was not observed in the simple receptive fields. The functional and structural models of the receptive field of the visual cortex are discussed.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 5, No. 2, pp. 201–209, March–April, 1973.     

Functional characteristics of visual cortical neurons during local photic stimulation of their receptive fields in cats

A group of functional characteristics of 103 neurons in visual cortical area 17 was investigated in acute experiments on curarized, light-adapted cats during a change in various parameters of the local photic stimuli. The average threshold sensitivity of the neuron population was 32 dB (0.052 nit), the sharpness of orientation tuning was 37°, the critical summation time was 57 msec, and the reactivity recovery time 190 msec. Photic sensitivity was lower during light adaptation than during dark adaptation, orientation selectivity of the neurons was increased, temporal summation was lengthened, and the time required by the neuron to recovery from after-inhibition was shortened. Several properties of the cortical neurons depended on the accentricity of their receptive fields: Cells with centrally localized receptive fields on average had lower thresholds and shorter summation time and they recovered their reactivity more quickly; their activity was of a higher frequency and they more often generated short phasic discharges than neurons with receptive fields in the peripheral part of the visual field. The mechanisms responsible for changes in the properties of neurons in the central and peripheral visual channels during dark and light adaptation are discussed. The presence of several inhibitory subsystems in the cortex regulating unit activity in the primary visual projection area is postulated.     

Characteristics of photic stimulus intensity coding by visual cortical neurons of light-adapted cats

Dependence of response characteristics of 70 visual cortical neurons on intensity of photic stimuli (bars) of optimal shape, size, orientation, location, and duration, presented under conditions of photopic light adaptation, were studied in curarized cats. Intensity functions were used to estimate response thresholds of the neuron, its differential sensitivity, the optimal intensity and the band width of brightnesses which the neuron could effectively code. Most of the neurons tested (70%) had nonmonotonous intensity functions with marked inhibitory distortion of gradual dependence in the middle part of the brightness range studied. Threshold of neuronal responses to light differed by 5 or 6 orders of magnitude. The threshold of the response and its minimal latent period were directly connected: Neurons with the lowest response threshold mainly responded after the shortest latent period. The maximal discharge frequency and differential sensitivity of the cell also correlated directly. The range of intensities within which the neuron exhibited maximal differential sensitivity, i.e., was able to code the strength of the light most effectively, lay between 0 and –20 dB. A sharp increase in differential sensitivity of the cortical neurons was found under light adaptation conditions compared with dark adaptation. Differences in characteristics of cortical neurons with their receptive field in the central and peripheral parts of the visual field were found.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 15, No. 3, pp. 211–217, May–June, 1983.     

Excitability cycles and rhythmic responses of visual cortical neurons to diffuse and punctiform light stimuli

The responses of the rabbit's visual cortical neurons to paired and rhythmic punctiform stimulation of their receptive fields were compared with responses to diffuse photic stimuli and the electric stimulation of the optic nerve. Diffuse photic and electric stimuli evoke a short-lasting initial activation of a neuron, followed by an inhibitory phase, during which the response to repetitive stimulation is suppressed. By contrast, localized stimulation of the neuron's receptive field with a spot of light produces an intensive and longer-lasting activation which is not followed by profound inhibition. Fusion of the responses to paired and rhythmic localized stimuli is therefore possible when the intervals between stimuli are brief enough. Rhythmic stimulation is capable of evoking sustained activation of a neuron during the entire duration of light flicker. During stimulation of a single point of the receptive field such prolonged activation can take place only within a limited range of stimulation frequencies (up to 15/sec), while higher frequencies evoke responses to the onset and offset of sequences of light flashes only. If, however, rhythmic stimuli alternate between the various points of a receptive field, prolonged neuronal activation is observed with any frequency of stimulation.A. N. Severtsov Institute of Evolutionary Morphology and Ecology of Animals, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 3, No. 3, pp. 252–259, May–June, 1971.     

Somatosensory cortical unit responses to stimulation of the vibrissae in cats

Characteristics of extra- and intracellular responses of 57 neurons in the vibrissal projection zone of the first somatosensory area of the cat cortex were investigated. The intensity of both excitatory and inhibitory unit responses was found to diminish during successive stimulation of different parts of the receptive fields in the direction from center toward periphery. Usually, when central parts of receptive fields were stimulated, inhibition in the unit responses was postexcitatory, whereas when peripheral parts were stimulated inhibition could precede excitation. The possibility of an increase in the role of interaction between excitatory and inhibitory processes arising in neurons in response to vibrissal stimulation with an increase in the distance from center to periphery of receptive fields of single cortical cells is discussed. Neurons found during one insertion of the microelectrode were seen to have common center for their receptive fields, but the diameters of the receptive fields of individual neurons could differ significantly. Moreover, during such vertical insertions responses of neurons with primary inhibition to the stimuli presented were recorded.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 12, No. 2, pp. 124–130, March–April, 1980.     

Neurons with visual receptive fields independent of eye position in the caudal section of the ventral bank of cat cruciate slucus

Neurons responding to tactile and visual stimulation were found in the caudal section of the cruciate slucus ventral bank in awake cats. Tactile receptive fields were located on the face, mainly around the mouth. Visual stimuli evoked a response when presented close to the tactile receptive field. It was found that the visual responses of these bimodal neurons located in layer VI of the cortex display spatial consistency. The position of these visual receptive fields remained constant through saccadic eye movements, while still linked to the tactile receptive field.Institute for Research into Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neifofiziologiya, Vol. 18, No. 6, pp. 800–805, November–December, 1986.     

Dynamic and static receptive field characteristics of single neurons in the lateral suprasylvian area in cats

Static and dynamic properties of receptive fields of neurons in the lateral suprasylvian area of the cat cerebral cortex were studied. Neurons with different dynamic characteristics may have an identical static organization of their receptive fields; strict correlation is thus not found between these two characteristics of neurons in this area. Most black-sensitive neurons were found to have a receptive field with off-response. Stimulus contrast reversal tests showed that generation of responses to presentation of both black and light stimuli takes place as a result of excitation of the same area of the receptive field and is not due to spatially different on- and off-zones.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 16, No. 1, pp. 116–123, January–February, 1984.