Structural organization of receptive fields of lateral suprasylvian cortical neurons in cats

The substructural organization of receptive fields of lateral suprasylvian cortical neurons, sensitive to movement of visual stimuli, was investigated in cats. The experimental results showed that receptive fields of neurons in this cortical area, judging by responses to movement, consist mainly of cells with qualitatively different characteristics. With the unmasked method of presentation of a moving stimulus, a reduction in the amplitude of movement as a rule evoked a directional response of the cell, whereas with the masked method, and with the same amplitudes of movement, a nondirectional response appeared. The receptive fields of some neurons were particularly sensitive to movement of borders but did not respond to the body of the stimulus like receptive fields of neurons described in other visual structures. Heterogeneity of the substructural organization of receptive fields of lateral suprasylvian cortical neurons can be explained by convergence of inputs on the neuron and it is regarded as the basis of integrative mechanisms in this structure.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 17, No. 3, pp. 293–300, May–June, 1985.     

‘Vector white noise’: a technique for mapping the motion receptive fields of direction-selective visual neurons

A technique is described and tested for mapping the sensitivities and preferred directions of motion at different locations within the receptive fields of direction-selective motion-detecting visual neurons. The procedure is to record the responses to a number of visual stimuli, each stimulus presentation consisting of a set of short, randomly-oriented, moving bars arranged in a square grid. Each bar moves perpendicularly to its long axis. The vector describing the sensitivity and preferred direction of motion at each grid location is obtained as a sum of the unit vectors defining the directions of motion of the bars in each of the stimuli at that location, weighted by the strengths of the corresponding responses. The resulting vector field specifies the optimum flow field for the neuron. The advantage of this technique over the conventional approach of probing the receptive field sequentially at each grid location is that the parallel nature of the stimulus is sensitive to nonlinear interactions (such as shunting inhibition for mutual facilitation) between different regions of the visual field. The technique is used to determine accurately the motion receptive fields of direction-selective motion detecting neurons in the optic lobes of insects. It is potentially applicable to motion-sensitive neurons with highly structured receptive fields, such as those in the optic tectum of the pigeon or in area MST of the monkey.     

A system of insect neurons sensitive to horizontal and vertical image motion connects the medulla and midbrain

Summary The response properties and gross morphologies of neurons that connect the medulla and midbrain in the butterfly Papilio aegeus are described. The neurons presented give direction-selective responses, i.e. they are excited by motion in the preferred direction and the background activity of the cells is inhibited by motion in the opposite, null, direction. The neurons are either maximally sensitive to horizontal motion or to slightly off-axis vertical upward or vertical downward motion, when tested in the frontal visual field. The responses of the cells are dependent on the contrast frequency of the stimulus with peak values at 5–10 Hz. The receptive fields of the medulla neurons are large and are most sensitive in the frontal visual field. Examination of the local and global properties of the receptive fields of the medulla neurons indicates that (1) they are fed by local elementary motion-detectors consistent with the correlation model and (2) there is a non-linear spatial integration mechanism in operation.     

Inhibitory components in the neuronal response of the lateral suprasylvian cortical area in the cat

Inhibitory components in the response evoked by presentation of mobile visual stimuli in neurons belonging to the lateral suprasylvian area of the cerebral cortex were investigated in cats. It was demonstrated by comparing poststimulus histograms of neuronal response to movement in two opposite directions that the location of discharge centers within the receptive fields changed in relation to movement direction. No spatial area giving rise to the inhibitory component of response could be found in any of the neurons with monotone stationary structure of their receptive fields. Findings from experiments involving techniques of stimulating a test area of the receptive field separately indicated that inhibitory components of response in neurons of the lateral suprasylvian area with monotone organization of the receptive field could represent inhibitory after-response following the neuronal excitation produced by the visual stimulus traveling across this field.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 19, No. 3, pp. 299–308, May–June, 1987.     

Orientation-selective neurons in the squirrel visual cortex

The organization of receptive fields of neurons sensitive to orientation of visual stimuli was investigated in the squirrel visual cortex. Neurons with mutually inhibitory on- and off-areas of the receptive field, with partially and completely overlapping excitatory and inhibitory mechanisms, were distinguished. Neurons of the second group are most typical. They exhibit orientation selectivity within the excitatory area of the receptive field because, if the stimulus widens in the zero direction, perpendicular to the preferred direction, lateral inhibition is much stronger than if it widens in the preferred direction. Additional inhibitory areas (outside the excitatory area) potentiate this inhibition and increase selectivity. It is suggested that there is no strict separation of simple (with separate excitatory and inhibitory mechanisms in the receptive field) and complex (with overlapping of these mechanisms) neurons in the squirrel visual cortex.A. N. Severtsov Institute of Evolutionary Morphology and Ecology of Animals, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 11, No. 6, pp. 540–549, November–December, 1979.     

Spontaneously emerging direction selectivity maps in visual cortex through STDP

It is still an open question as to whether, and how, direction-selective neuronal responses in primary visual cortex are generated by feedforward thalamocortical or recurrent intracortical connections, or a combination of both. Here we present an investigation that concentrates on and, only for the sake of simplicity, restricts itself to intracortical circuits, in particular, with respect to the developmental aspects of direction selectivity through spike-timing-dependent synaptic plasticity. We show that directional responses can emerge in a recurrent network model of visual cortex with spiking neurons that integrate inputs mainly from a particular direction, thus giving rise to an asymmetrically shaped receptive field. A moving stimulus that enters the receptive field from this (preferred) direction will activate a neuron most strongly because of the increased number and/or strength of inputs from this direction and since delayed isotropic inhibition will neither overlap with, nor cancel excitation, as would be the case for other stimulus directions. It is demonstrated how direction-selective responses result from spatial asymmetries in the distribution of synaptic contacts or weights of inputs delivered to a neuron by slowly conducting intracortical axonal delay lines. By means of spike-timing-dependent synaptic plasticity with an asymmetric learning window this kind of coupling asymmetry develops naturally in a recurrent network of stochastically spiking neurons in a scenario where the neurons are activated by unidirectionally moving bar stimuli and even when only intrinsic spontaneous activity drives the learning process. We also present simulation results to show the ability of this model to produce direction preference maps similar to experimental findings     

Dependence of functional organization of receptive fields of the cat lateral geniculate body on visual stimulus contrast

The spatial organization of receptive fields of the lateral geniculate body in response to visual stimuli with different degrees of contrast was studied in cats. During variation of contrast changes in organization of the central zone were found to take place in some receptive fields. Inside the central zone of the receptive field as revealed by the use of low stimulus contrasts, an additional inhibitory ring appears in response to a stimulus of high contrast. The weighting function of the central zone of the receptive field becomes variable in sign. The role of this phenomenon in transmission of information on high spatial frequencies (increase in visus) at high contrasts is discussed.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 16, No. 6, pp. 789–796, November–December, 1984.     

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.     

Peculiarities of summation processes in visual-sensitive neurons in the pretectal area of cat

Summation processes occurring in single neurons of the pretectal area in response to either moving or stationary light stimuli were studied in acute experiments on cats. In most neurons studied (85%), gradual increase of the angular size of stimulus resulted in clearly defined summation. In neurons lacking directional sensitivity (nondirectional neurons) the stimulus movement in two opposite directions caused, as a rule, similar and symmetrical changes in the number of spikes, whereas under the same conditions direction-sensitive neurons, in addition to symmetrical development of summation processes, could exhibit substantial differences in the summation curves. The responses to a preferred movement direction could be significantly inhibited or facilitated, while the responses to a non-preferred direction remained stable or changed reciprocally. Neuronal responses to different directions of the movement of stimulus might change independently of each other. This was also the case whenon andoff responses of theon—off neurons to stationary stimuli were compared. It is concluded that neurons of the pretectal area have a complex infrastructure of receptive fields that significantly influences the integration of incoming information.Neirofiziologiya/Neurophysiology, Vol. 25, No. 5, pp. 376–382, September–October, 1993.     

The spatial substructure of visual receptive fields in the cat's superior colliculus

Although the direction selective properties of the superficial layer cells of the cat's superior colliculus have been extensively studied, the mechanisms underlying this property remain controversial. With the aim to understand the mechanism(s) underlying directional selectivity of collicular neurons we examined the substructure of their visual receptive fields. 1. The strength of cell responses and the direction selectivity indices varied in relation to the location of the tested region within the receptive field and the amplitude of stimulus movement. 2. Decrease of the amplitude of motion resulted in a decrease of direction selectivity index both in the group of direction-selective cells and in the group of cells classified as direction nonselective but with a directional bias. 3. The decrease of direction selectivity for small amplitude movement resulted mainly from increase in the magnitude of response in the nonpreferred direction of movement. 4. These results suggest that the receptive fields of most collicular cells are composed of subregions with different response profiles and indicate that inhibitory mechanisms dictate direction selectivity of collicular cells.     

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.     

Heterogeneity of neuron populations with complex receptive fields in the cat visual cortex

The selectivity of striate neurons with complex receptive fields to the orientation, direction, and velocity of movement of various stimuli was investigated in unanesthetized and uncurarized cats. On the basis of all characteristics obtained by the study of single-unit responses to a stationary flickering slit, a moving spot of light, and a moving oriented stimulus, four groups of complex neurons were distinguished. The characteristics of group I neurons indicate a mechanism of orientation selectivity in the organization of their receptive fields, group IV neurons have a mechanism of directional selectivity, and neurons of groups II and III possess both mechanisms. The existence of separate neuronal systems coding the orientation and direction of stimulus movement is suggested.V. Kapsukas State University, Vilnius. Translated from Neirofiziologiya, Vol. 11, No. 2, pp. 109–116, March–April, 1979.     

“Regular” visual receptive fields of neurons of the cat lateral geniculate body

The spatial organization of receptive fields (RF) of neurons was studied in the lateral geniculate body (LGB) of cats with pretrigeminal transection of the brainstem (without general anesthesia). Using systematic point testing of the entire RF area and adjacent regions, the RF configuration and distribution of the response types for a stable flickering stimulus throughout the RF area were determined. Only 40% (64 units of 160 studied) LGB neurons had simple RF configuration. Such RF of ellipsoid or round shape were called regular receptive fields, RRF. Most RRF (51, or about 80%) demonstrated spatially homogeneous organization with similar-type (on, off, oron-off) responses to stimulation of the entire RF area. The RRF of 13 neurons, i.e., about 20%, included subfields with qualitatively different responses to application of a stable flickering light spot. The position of subfields was asymmetrical in 8 neurons (13%), while a nearly concentric RF arrangement, with the center surrounded by an antagonistic area, was found only in 5 units (7%) with RRF. Nearly all neurons with heterogeneous RRF demonstrated directional selectivity to moving stimuli.Neirofiziologiya/Neurophysiology, Vol. 27, No. 5/6, pp. 413–424, September–December, 1995.     

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.     

Selectivity of visual cortical neurons of rabbits to magnitude of moving stimuli

Unit responses of the rabbit visual cortex were investigated in relation to size of visual stimuli moving in their receptive field. With an increase in size of the stimulus in a direction perpendicular to the direction of movement ("width" of the stimulus) an initial increase in the intensity of the unit response through spatial summation of excitory effects is followed by a decrease through lateral inhibition. This inhibition is observed between zones of the receptive field which behave as activating when tested by a stimulus of small size. Each neuron has its own "preferred" size of stimuli evoking its maximal activation. No direct correlation is found between the "preferred" stimulus size and the size of the receptive field. With a change in stimulus size in the direction of movement ("length" of the stimulus) the responses to stimuli of optimal size may be potentiated through mutual facilitation of the effects evoked by the leading and trailing edges of the stimulus and weakened in response to stimuli of large size. The selective behavior of the neurons with respect to stimulus size is intensified in the case of coordinated changes in their length and width. It is postulated that the series of neurons responding to stimuli of different "preferred" dimensions may constitute a system classifying stimuli by their size.A. N. Severtsov Institute of Evolutionary Morphology and Ecology of Animals, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 4, No. 6, pp. 636–644, November–December, 1972.     

Mechanism of directional selectivity of neurons with complex receptive fields in the cat visual cortex

Spatio-temporal interactions within complex receptive fields in the cat visual cortex were investigated by sequential presentation of two stationary stimuli. When two stimuli were presented in phase (on-on or off-off) in the order corresponding to preferred direction of movement, facilitation or weak inhibition of the response to the second stimulus was observed, whereas if it corresponded to zero direction of movement, the response was strongly inhibited. In the case of stimulation out of phase (on-off or off-on), in the order corresponding to the preferred direction of movement, considerable inhibition of the response to the second stimulus was observed, whereas in the opposite order, facilitation or weak inhibition was observed. The strength of interaction between different parts of the field depended on the distance between them and the duration of the interval between stimuli. Directional selectivity of "complex" neurons is thus ensured by asymmetry of spatio-temporal interactions between receptive field inputs of the same type. Interactions between inputs of different types, arising when a multiedge stimulus (bar, grating) can be used by the visual system to distinguish an object from the background and to assess changes in size of objects and the relative velocity of their movement.V. Kapsukas State University, Vilnius. Translated from Neirofiziologiya, Vol. 16, No. 4, pp. 505–512, July–August, 1984.     

Structure of receptive fields of hyppocampal visually-sensitive neurons in the cat

We studied the structure of receptive fields (RF) in the visually sensitive CA1 and CA3 fields of the dorsal hippocampus in alert cats subjected to pretrigeminal section of the brain stem. Scanning with moving stimuli and mapping the whole RF area with point by point application of a stationary stimulus, we studied 76 RF neurons. We found no significant differences in the response characteristics of neurons of fields CA1 and CA3. Our data are a basis for classifying the neurons in the following groups: homogeneous RF structure of the RF (54%) and nonhomogeneous structure (28%) and weak reaction to the standard stimulus, but a strong reaction to movement (18%). The results showed that responses of some of the hippocampal neurons exhibit high specificity. Thus, 9% of the neurons with nonhomogeneous RF structure disclosed with static stimuli reacted variously to a change in contrast and contours of moving stimuli. The data presented indicate that visually sensitive neurons of the hippocamus have a relatively well-developed mechanism for processing visual sensory information and this obviously ensures participation of the limbic system in visually controlled behavior of the animal.Institute of Experimental Biology, Academy of Sciences of the Armenian SSR, Yerevan. Translated from Neirofiziologiya, Vol. 23, No. 2, pp. 160–167, March–April, 1991.     

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.     

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 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.