Spatio-temporal organization of receptive fields of the cat striate cortex

The responses of cortical cells to gratings and bars were compared. The excitatory and inhibitory on-and off-zones of a simple cell are composed of on- and off-subfields of CGL. Any zone is formed by an opponent pair of subfields one of which gives an excitatory effect, the other — inhibitory. Such organization assumes the linear properties of a simple field. The deviations from linearity are due to spatial dis-placements of the subfields, heterogeneity of subfields, or the absence of one subfield in the opponent pair. Subfields may be both phasic and tonic, even in the same RF. Analysis of the most common type of a complex cell with modulated responses against unmodulated background shows that a mask eliminating stimulation of any half of the RF causes (according to the theory of filtres) increasing the bandwidth due to the increase or the appearance of responses to side low and high frequencies. The modulated components of the responses from both halves of the RF are out of phase. Analysis of this fact and the responses to thin bars suggests that a complex field is formed by linear and nonlinear subsystems converging onto output neuron. Other types of complex fields are organized by different combinations of subsystems. Limited in area by masking the RF responds to much higher spatial frequencies than the whole RF. The optimal frequency in two-dimensional spatial frequency characteristics of the RF does not change with orientation. Simple RFs and a part of complex RF calculate the amplitude and the phase of the stimulus, the other part of complex RFs (with unmodulated response) calculate only amplitude. Given all this, the RFs are grating filters of spatial frequency.     

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.     

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.     

The organization of exteroceptive information from the uropod to ascending interneurones of the crayfish

The organization of exteroceptive inputs to identified ascending interneurones of the crayfish, Procambarus clarkii (Girard), has been analyzed by stimulation of hairs on the uropod and simultaneous intracellular recordings from ascending interneurones. The spikes of single afferent neurones which innervated hairs on the distal ventral surface of the exopodite were consistently followed by a depolarizing synaptic potential in many identified ascending interneurones with a constant and short central delay of 0.7–1.5 ms. The amplitude of the potentials depended on the membrane potential of the ascending interneurones. Each afferent neurone made divergent outputs onto several ascending interneurones and each ascending interneurone received convergent inputs from several afferent neurones. Certain ascending interneurones made inhibitory or excitatory connections with other ascending interneurones. These central interactions were always one-way, and the spikes from one ascending interneurone consistently evoked excitatory or inhibitory post-synaptic potentials in other interneurones which followed with a constant and short latency of 0.7–1.0 ms. The inhibitory postsynaptic potential was reversed by injection of steady hyperpolarizing current.Abbreviations EPSP excitatory post-synaptic potential - IPSP inhibitory post-synaptic potential     

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.     

Stimulus size selectivity and receptive field organization of ectostriatal neurons in the pigeon

The avian ectostriatum is the telencephalic recipient zone of the tectofugal pathway, which may be homologous to the colliculo-pulvinar-cortical pathway in mammals. The present paper studies the visual response properties and receptive field organization of ectostriatal cells in pigeons with extracellular recording and computer mapping techniques. The results indicate that 90% of ectostriatal cells prefer forward, downward and upward motion of visual stimuli at velocities in the range of 16-128 degrees s(-1). They respond optimally to small stimuli (1-4 degrees visual angle), mostly preferring a target of 2 degrees. Most cells (78.8%) have one excitatory receptive field that usually possesses one or two hotspots, some cells (13.5%) have two excitatory receptive fields each having one or two hotspots, and a few cells (7.7%) have one excitatory receptive field with an unresponsive region in the center. An inhibitory receptive field is not found surrounding the excitatory receptive field in the ectostriatal cells examined. These response properties and receptive field organization may reflect the possible roles of the ectostriatum in stimulus discrimination and visual cognition.     

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.     

Spatiotemporal organization of unit activity in the frog tectum and its modification by visual stimuli

Unit activity was recorded in the tectum of curarized frogs during presentation of various visual stimuli (on- and off-responses to diffuse illumination and movement of an object of recognizable shape). It was shown that different types of stimulation lead to the organization of a different distribution of unit activity in the tectum, in the form of excitatory-inhibitory neuronal mosaics; inhibitory responses, limiting and exacerbating the excitatory responses of other neurons, predominate. The differences observed in the spatiotemporal characteristics of the neuronal mosaics under the influence of different stimuli may be evidence of specificity of coding of visual impulses carrying different information from the retina in the tectum.     

Structure of a complex receptive field in the cat visual cortex

The most common type of complex receptive field, whose response to the passage of sinusoidal gratings across it consisted of modulated and unmodulated components, was analyzed. The use of a mask to cover half the field, according to the filter theory, led to widening of the transmission band of the field as a spatial frequency filter, due to the appearance or enhancement of the response at lateral low and high frequencies. Modulated components of responses from the left and right halves of the field were out of phase. Analysis of this fact, and also of responses of the field to thin light and dark bars enabled the field structure to be described. It consists of linear and nonlinear subsystems, converging on the output neuron of the complex field. The former is composed of several pairs of on- and off-subfields of the lateral geniculate body. The on- and off-subfields in the pair overlap spatially and converge on the output neuron of the linear subsystem with opposite signs. The nonlinear subsystem is composed of either on- or off-subfields. Other types of complex fields may include different combinations of subsystems. The results indicate that complex fields are spatiotemporal grating filters.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 14, No. 1, pp. 19–25, January–February, 1982.     

A dynamic model of the vertebrate retina

In this paper a mathematical model of the retina was proposed to clarify the spatio-temporal information processing mechanism in the retina of vertebrates. In order to explain spatio-temporal characteristics of an on-center receptive field of a ganglion cell, excitatory and inhibitory cell layers were introduced of which time lags increased with the lateral distance from a point of stimulation. The characteristics of this model were found to agree well with the physiological data: e.g., this model shows on-response to the input stimulus given on the center, off-response to the input on the surround, and on-off response to the input on the border between on- and off-response regions of the on-center field.     

Characteristics of electrical activity of olfactory bulb neurons of fish

It was shown by intracellular recording of the activity of olfactory bulb neurons of the carp that their dendrites are excited both by synaptic activation and by direct stimulation with an electric current. The dendrites generate an action potential and probably conduct it for some distance toward the soma. The neurons can be divided into two groups: one responds to ortho- and antidromic stimuli with one, rarely with two peaks, the other responds with a rhythmical discharge. The presence of early and late IPSP is characteristic of neurons of both groups. Rhythmical variations in potential with a frequency of 26–33/sec, so-called oscillations, are recorded; they may be excitatory (in secondary neurons they correspond to EPSP) or inhibitory (they correspond to IPSP). Possible mechanisms of the excitatory oscillations and the rhythmical discharge in olfactory bulb neurons of the carp are discussed.M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol.3, No.5, pp. 505–511, September–October, 1971.     

A functional model of the wiring of the simple cells of visual cortex

A computer model of the simple cells in the mammalian visual cortex was constructed. The model cells received inputs from a great number of isopolar centre/surround cells assumed to be located in the lateral geniculate nucleus (LGN). The distribution of input to the model simple cells was either inhibitory/excitatory or inhibitory/excitatory/inhibitory. Such arrangements produced receptive fields containing four or five consecutively antagonistic subfields. Responses produced by the model cells to different types of stimuli (periodical as well as nonperiodical) were obtained and compared to responses of living cells reported from various laboratories under comparable stimulus conditions. In all the situations tested, the responses of the model cells corresponded qualitatively very well to those of living cells. It was seen that the same wiring mechanism was able to account for orientation selectivity, spatial frequency filtering, various phase relationships between stimulus and response, subfield orientational selectivity, and slight end-inhibition. Furthermore, the receptive fields of the model simple cells closely resemble Gabor functions.     

Differences in synaptic output between excitatory and inhibitory motoneurons in a crayfish muscle

A pair of antagonistic motoneurons, one excitatory and one inhibitory, innervates the distal accessory flexor muscle in the walking limb of the crayfish Procambarus clarkii. The number and size of synapses formed by these two axons on the muscle fibers (neuromuscular synapses) and on each other (axo-axonal synapses) were estimated using thin-section electron microscopy. Although profiles of nerve terminals of the two axons occur in roughly equal proportions, the frequency of occurrence of neuromuscular synapses differed markedly: 73% were excitatory and 27% were inhibitory. However, inhibitory synapses were 4–5 times larger than excitatory ones, and consequently, the total contact areas devoted to neuromuscular synapses were similar for both axons. Axo-axonal synapses were predominantly from the inhibitory axon to the excitatory axon (86%), and a few were from the excitatory axon to the inhibitory axon (14%). The role of the inhibitory axo-axonal synapse is presynaptic inhibition, but that of the excitatory axo-axonal synapse is not known. The differences in size of neuromuscular synapses between the two axons may reflect intrinsic determinants of the neuron, while the similarity in total synaptic area may reflect retrograde influences from the muscle for regulating synapse number.     

Interaction between neurons displaying tonic response to sound and other cells of the feline auditory cortex

Interaction between neurons with a tonic response pattern and either nearby or further removed (by about 400–500 µm) cortical neurons was investigated during acute experiments on 15 immobilized cats using cross-correlation analysis techniques. Synchronizing excitatory input common to the nerve cells was found in cross-correlation histograms (CCH) in 26 out of 36 test pairs of neurons (72%). Both positive and negative cross-correlation (five pairs in each case) were discovered, pointing to mono- or polysynaptic excitatory and inhibitory effects of the tonic neuron on spike activity in the other cell from CCH of 10 pairs of neurons. The functional diversity of neurons f distinguished by a tonic pattern of response to sound was deduced on the basis of findings from this research. The theory that some tonic type cells act as excitatory neurons and others fulfil the function of inhibitory interneurons is examined.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 21, No. 5, pp. 613–620, September–October, 1989.     

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.     

The Effect of STDP Temporal Kernel Structure on the Learning Dynamics of Single Excitatory and Inhibitory Synapses

Spike-Timing Dependent Plasticity (STDP) is characterized by a wide range of temporal kernels. However, much of the theoretical work has focused on a specific kernel – the “temporally asymmetric Hebbian” learning rules. Previous studies linked excitatory STDP to positive feedback that can account for the emergence of response selectivity. Inhibitory plasticity was associated with negative feedback that can balance the excitatory and inhibitory inputs. Here we study the possible computational role of the temporal structure of the STDP. We represent the STDP as a superposition of two processes: potentiation and depression. This allows us to model a wide range of experimentally observed STDP kernels, from Hebbian to anti-Hebbian, by varying a single parameter. We investigate STDP dynamics of a single excitatory or inhibitory synapse in purely feed-forward architecture. We derive a mean-field-Fokker-Planck dynamics for the synaptic weight and analyze the effect of STDP structure on the fixed points of the mean field dynamics. We find a phase transition along the Hebbian to anti-Hebbian parameter from a phase that is characterized by a unimodal distribution of the synaptic weight, in which the STDP dynamics is governed by negative feedback, to a phase with positive feedback characterized by a bimodal distribution. The critical point of this transition depends on general properties of the STDP dynamics and not on the fine details. Namely, the dynamics is affected by the pre-post correlations only via a single number that quantifies its overlap with the STDP kernel. We find that by manipulating the STDP temporal kernel, negative feedback can be induced in excitatory synapses and positive feedback in inhibitory. Moreover, there is an exact symmetry between inhibitory and excitatory plasticity, i.e., for every STDP rule of inhibitory synapse there exists an STDP rule for excitatory synapse, such that their dynamics is identical.     

“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.     

Rubrospinal synaptic influences on the lumbar motoneurons of the rat

Intracellular recording was employed in experiments on rats with the nervous system intact and after acute pyramidotomy to study the postsynaptic effects produced in the lumbar motoneurons on stimulation of the nucleus ruber. Stimulation of this nucleus with single stimuli and with a short series of stimuli caused excitatory and inhibitory postsynaptic potentials (EPSP and IPSP) to develop in the motoneurons. Most of the EPSP recorded were disynaptic, but response development involved a monosynaptic segmental delay in five of the 124 cells that exhibited EPSP. A capacity for high-frequency potentiation was a characteristic feature of the disynaptic excitatory and inhibitory effects. Transmembrane polarization of the motoneurons had a marked influence on the amplitude of the disynaptic EPSP and IPSP. The properties of the disynaptic rubrospinal influences were similar to those described for the cat.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 3, No. 3, pp. 266–273, May–June, 1971.     

Brainstem lateral line responses to sinusoidal wave stimuli in the goldfish, Carassius auratus

Extracellular recordings were made from single lateral line units in the medial octavolateralis nucleus in the brainstem of goldfish, Carassius auratus. Units were defined as receiving lateral line input if they responded to the water motions generated by a stationary, sinusoidally oscillating sphere and/or a moving sphere but not to airborne sound and vibrations. Units which responded to airborne sound or vibrations were assumed to receive input from the inner ear and were not further investigated. Responses of lateral line units were quantified in terms of the number of evoked spikes and the degree of phase-locking to a 50 Hz vibrating sphere presented at various stationary locations along the side of the fish. Receptive fields were characterized based on spike rate, degree of phase-locking and average phase angle as a function of sphere location. Four groups of units were distinguished: 1, units with receptive fields comparable to those of primary afferents; 2, units with receptive fields which consisted of one excitatory and one inhibitory area; 3, units with receptive fields which consisted of more than two excitatory and/or inhibitory areas; 4, units with receptive fields which consisted of a single excitatory or a single inhibitory area. The receptive fields of most units were characterized by adjacent excitatory and inhibitory areas. This organization is reminiscent of excitatory-inhibitory receptive field organizations in other vertebrate sensory systems.