Influence of afferent impulses from the nerves of the anterior limbs on the insertion neurons of the lumbar section of the spinal cord

In anesthetized cats in conditions of muscular relaxation we have studied the participation of the interneurons of the lumbar section of the spinal cord in the interaction of the FRA systems of the fore and hind limbs. Using microelectrodes we have made extra- and intracellular recordings of the potentials. It has been shown that from the flexor afferents of the fore limbs both facilitating and inhibitory influences are transmitted. The former are expressed in increased frequency of the background impulse activity of the neurons, in the appearance of evoked responses of the "silent" cells and intensification of the test responses for short time intervals with paired heteronymous stimulation. The inhibitory influences prevail over the facilitating and are manifest in depression of the background activity and evoked segmental responses of the neurons. The maximum inhibition of the segmental responses was noted for intervals of 40–140 msec. The duration of inhibition varied from 100–500 msec and more. Depending on the intensity and duration of the inhibitory influences two groups of interneurons have been isolated. The role of the pre- and postsynaptic mechanisms in the transmission of inhibitory influences from the afferents of the fore limbs on the afferents of the hind limbs is discussed.Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 1, No. 3, pp. 235–242, November–December, 1969.     

Interaction between segmental and descending intersegmental reflexes in lumbar motoneurons

The effects of segmental reflexes on descending intersegmental reflexes to stimulation of forelimb afferents were studied in anesthetized cats by recording postsynaptic responses from single motoneurons. Interaction between these influences was found to be reciprocal in character for groups of neurons with primary connections with afferents of the superficial and deep branches of the peroneal nerve and afferents of the nerve to the gastrocnemius muscle. Excitatory postsynaptic responses arising in groups of motoneurons of the peroneal nerve to stimulation of forelimb afferents underwent profound and prolonged inhibition during conditioning stimulation of afferents in the deep and superficial peroneal nerves. Activation of segmental afferents during conditioning stimulation of the gastrocnemius nerve was accompanied by inhibition of excitatory intersegmental responses and deinhibition of inhibitory responses in motoneurons of the gastrocnemius muscle. Segmental inhibition of intersegmental descending impulse activity appeared in the interneuron system of the segmental reflex centers connecting the descending propriospinal tracts with the motoneurons of these centers.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 4, No. 2, pp. 16872-175, March–April, 1972.     

Convergence of synaptic influences of contralateral somatic afferents on interneurons in segmental inhibitory pathways to motoneurons

Convergence of contralateral somatic afferent synaptic influences on segmental inhibitory neurons was investigated by intracellular recording of postsynaptic potentials of -motoneurons in experiments on cats. Excitatory synaptic influences of afferents of the contralateral flexor reflex were shown to converge on interneurons of both segmental inhibitory systems studied: afferents of flexor reflex and group Ia muscle afferents. Interneurons of inhibitory systems are exposed not only to excitatory but also to inhibitory contralateral influences. Contralateral inhibitory PSPs of montoneurons are produced through ipsilateral inhibitory systems; a leading role is played by inhibitory neurons of the flexor reflex system of afferents. Inhibitory neurons of the Ia system as a rule do not make an important contribution to generation of contralateral IPSPs.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 5, No. 5, pp. 476–484, September–October, 1973.     

Mechanism of inhibition following reflex discharge in spinal cord interneurons

Using the method of microelectrode (intracellular and extracellular) recording, the mechanism of inhibition following reflex discharge in interneurons of the lumbosacral section of the spinal cord of cats on activation of cutaneous and high-threshold muscle afferents was studied. It was shown that the postdischarge depression of the reflex responses 10–20 msec after the moment of activation of the neuron is due to afterprocesses in the same neuron and presynaptic pathways. The depression of spike potentials from the 20th to the 100th msec is produced by inhibitory postsynaptic potentials (IPSP). During the development of IPSP the inhibition of spike potentials can be due to both a decrease of the depolarization of the postsynaptic membrane below the critical threshold and a decrease of sensitivity of the cell membrane to the depolarizing action of the excitatory postsynaptic potential (EPSP). At intervals between the stimuli of 30–100 msec the duration of EPSP after the first stimulus does not differ from that after the second stimulus. Hence, it is suggested that the presynaptic mechanisms do not play an essential part in this type of inhibition of interneurons. The inhibition following the excitation favors the formation of a discrete message to the neurons of higher orders.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 2, No. 1, pp. 3–9, January–February, 1970.     

Investigation of spinal neuronal mechanisms of movement control systems

Morphological and electrophysiological investigations of the means whereby the principal descending motor systems (the cortico-, rubro-, reticulo-, and vestibulo-spinal tracts) are connected with the segmental interneuronal apparatus and motoneurons show that these connections can be based on two different principles. Descending systems either activate motoneurons directly (monosynaptically) or are connected primarily with various interneuron systems, exerting their influence in that case by regulating the activity of simpler or more complex spinal mechanisms. The older descending system (reticulo- and vestibulo-spinal) possess a monosynaptic excitatory action of motoneurons; the evolutionarily newer descending systems, which transmit the most complex motor signals from the cerebral and cerebellar cortex to the spinal cord (cortico- and rubro-spinal), terminate synaptically in every case on interneurons. It is only in primates that a few cortico-spinal fibers form monosynaptic connections with motoneurons. The chief ways of action of the descending systems on interneurons are: control of the afferent inflow into the interneuron system by presynaptic inhibition of the corresponding synapses; control of the interneuron system by postsynaptic interaction with afferent influences; control of motoneurons through the specialized interneuron apparatus. The investigation shows that the last of these mechanisms functions in the cortico- and rubro-spinal, and possibly also in the reticulo- and vestibulo-spinal systems. The functional role of the various means of connection of the descending systems with the spinal neurons in the system of movement control is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 2, No. 2, pp. 189–202, March–April, 1970.     

Inhibitory effects of reticulospinal fibers of the lateral funiculus on neurons of thoracic spinal reflex pathways

Experiments on anesthetized cats with partial transection of the spinal cord showed that reticulo-spinal fibers in the ventral part of the lateral funiculus participate in the inhibition of polysynaptic reflexes evoked by stimulation of the ipsi- and contralateral reticular formation. The reticulo-fugal wave in the ventrolateral funiculus evoked comparatively short (up to 70 msec) IPSPs in some motoneurons of the internal intercostal nerve investigated and at the same time evoked prolonged (up to 500 msec) inhibition of IPSPs caused by activation of high-threshold segmental afferents. This wave also led to the appearance of IPSPs in 14 of 91 (15.5 %) thoracic spinal interneurons studied. The duration of these IPSPs did not exceed 100 msec; meanwhile, segment excitatory responses of 21 of 43 interneurons remained partly suppressed for 120–500 msec. It is concluded that the inhibitory action of the lateral reticulo-spinal system on segmental reflexes is due to several synaptic mechanisms, some of them unconnected with hyperpolarization of spinal neurons. The possible types of mechanisms of this inhibition are discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 10, No. 2, pp. 162–172, March–April, 1978.     

Synaptic processes in neurons of the cervical enlargement of the cat spinal cord during stimulation of propriospinal pathways of the dorsolateral tract

Responses of motoneurons and interneurons of the cervical enlargement of the cat spinal cord were studied by a microelectrode technique during selective stimulation of propriospinal fibers of the dorsolateral tract of the lateral white column. The long descending and ascending pathways were blocked by preliminary (10–16 days earlier) hemisection of the spinal cord cranially and caudally to the segments studied. Stimulation of the dorsolateral tract at a distance of 15–25 mm from the site of recording evoked complex postsynaptic potentials consisting of several successive waves in the motoneurons. The character of the PSPs was not clearly linked with the function of the motoneurons. By their latent periods the components of the PSPs could be placed in three groups. The "primary" components were reproduced in response to stimulation at 50–100/sec whereas the "secondary" and "tertiary" components were weakened or blocked. It is postulated that the "primary" components are evoked through monosynaptic connections between propriospinal fibers of the dorsolateral tract and motoneurons of the forelimb muscles, while the late components are evoked through polysynaptic pathways, including segmental interneurons. Many of these interneurons, located in the ventral horn and intermediate zone, were strongly excited during stimulation of the dorsolateral tract.A. A. Bogomolets' Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 5, No. 1, pp. 61–69, January–February, 1973.     

Corticofugal influences on segmental responses of lumbar spinal ventral horn interneurons in cats

Experiments on cats using extra- and intracellular recording methods showed that stimulation of the motor cortex of both hemispheres leads to considerable modulation of responses to stimulation of cutaneous and muscular lower limb afferents in spinal ventral horn interneurons in segments L6, 7. Three types of conditioning corticofugal effect were observed: facilitation, inhibition, and facilitation followed by inhibition (biphasic effect), and inhibitory effects predominated. The duration of facilitation of responses did not exceed 30–40 msec. The characteristics of the time course of inhibition varied: in some cases it began with relatively short intervals (8–15 msec), in other cases with an interval of 30–40 msec; its duration was 125–500 msec, or sometimes more. The effect of cortical stimulation on responses to stimulation of various afferent inputs of the same interneuron was shown to differ. The character of the conditioning corticofugal effect correlated with the latent period of segmental responses: facilitation was observed only in responses with a relatively short latent period (under 5 msec); responses with a longer latent period were mainly inhibited. The type of cortical effect also depended on the function performed by the activated afferent input. It is suggested that differential descending control of segmental polysynaptic responses recorded in ventral horn interneurons with wide convergence of afferent influences takes place in the initial stages of the reflex are. The mechanism of this control is discussed.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neiorofizologiya, Vol. 14, No. 6, pp. 563–571, November–December, 1982.     

Synaptic effects of contralateral somatic afferents on spinal motoneurons

Electrical activity of flexor and extensor alpha-motoneurons of the lumbar segments of cat's spinal cord as recorded intracellularly during electric stimulation of afferents of the contralateral posterior limb. Contralateral postsynaptic potentials (PSP) were shown to be evoked by activation of cutaneous and high-threshold muscle afferents. The high-threshold afferents of various muscle nerves participate to varying degrees in the generation of contralateral PSP. Contralateral inhibitory postsynaptic potentials (IPSP) were recorded in both flexor and extensor motoneurons along with contralateral excitatory postsynaptic potentials (EPSP). There are no fundamental differences in their distribution between flexor and extensor neurons. Inhibitory influences as a rule are predominant in both during the first 20 msec, and EPSP are predominant in the interval between 20 and 100 msec. The balance of excitatory and inhibitory pathway activity was found to be not as stable as that of the homolateral pathways.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 3, No. 4, pp. 418–425, July–August, 1971.     

Evoked activity of spinal neurons in the early period after sciatic nerve division

The character of dorsal horn motoneurons and interneurons evoked by stimulation of the dorsal root, and activity of Renshaw cells in response to stimulation of the ventral root were studied in albino rats in the lower lumbar segments of the spinal cord 5 days after sciatic nerve division. A significant increase in the mean amplitude of excitatory postsynaptic potentials of motoneurons was observed on the side of division of the nerve. No significant change in membrane potential and in the threshold of appearance of the action potential of these motoneurons took place. The mean number of action potentials and the duration of discharge of the Renshaw cells and dorsal horn interneurons likewise were not significantly changed.Dnepropetrovsk Medical Institute, Ukrainian Ministry of Health. Translated from Neirofiziologiya, Vol. 24, No. 3, pp. 306–314, May–June, 1992.     

Postsynaptic effects evoked by activation of long propriospinal pathways in cervical spinal neurons in cats

Effects induced in motoneurons and interneurons of the cervical enlargements of the cat spinal cord by stimulation of the lateral and ventral funiculi at the lower thoracic level were studied under conditions producing degeneration of fibers of descending brain systems. Stimulation of this sort evoked PSPs (mainly of mixed character) in 57 of 90 motoneurons tested. In nine motoneurons the primary response consisted of monosynaptic EPSPs evoked by activity of fibers of the lateral funiculus, and in the rest it consisted of polysyanptic (at least disynaptic) EPSPs and IPSPs. Polysynaptic effects arising in the neuron in response to stimulation of the lateral and ventral funiculi usually differed only quantitatively. The intensity of excitatory synaptic action on motoneurons of the proximal muscle (especially thoracid) was much greater than that on motoneurons of distal muscles. Nearly all motoneurons with no synaptic action belonged to the latter group. Stimulation of the lateral and ventral funculi facilitated synaptic action induced in motoneurons by stimulation of high-threshold segmental afferents and led to excitation of interneurons located in the vectral quadrant, and had no effect on interneurons in the dorsal regions of gray matter. These effects are regarded mainly as the result of excitation of long ascending propriospinal pathways in the cervical parts of the cord; it is also postulated that some of them are evoked by the arrival of activity along collaterals of descending propiospinal pathways to the neurons in this region.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 11, No. 4, pp. 339–347, July–August, 1979.     

Neuronal pathways from low-threshold muscle and cutaneous afferents innervating tail to trunk muscle motoneurons in the cat

We studied neuronal pathways from low-threshold muscle (group I, II) and cutaneous afferents (group A(alpha)beta) innervating the tail to motoneurons innervating trunk muscles (m. iliocostalis lumborum and m. obliquus externus abdominus) in 18 spinalized cats. Stimulation of group I muscle afferents produced excitatory postsynaptic potentials or excitatory postsynaptic potentials followed by inhibitory postsynaptic potentials in all motoneurons innervating the m. iliocostalis lumborum which showed effects (32%), and predominantly inhibitory postsynaptic potentials in motoneurons innervating the m. obliquus externus abdominus (47%). Stimulation of group I+II afferents produced significant increases of the incidence of motoneurons showing postsynaptic potentials (the notoneurons innervating the m. iliocostalis lumborum, 87%; the motoneurons innervating the m. obliquus externus abdominus, 82%). The effects of low threshold cutaneous afferents were bilateral, predominantly producing inhibitory postsynaptic potentials in motoneurons innervating both muscles. These results suggest that neuronal pathways from muscle afferents to back muscle motoneurons mainly increase the stiffness of the trunk to maintain its stability, while those to abdominal muscles help to extend the dorsal column by decreasing their activities. The results also indicate that neuronal pathways from cutaneous afferents to trunk motoneurons functionallY disconnect the tail from the trunk.     

Synaptic processes in facial motoneurons evoked by stimulation of the caudal trigeminal nucleus

Field potentials and postsynaptic potentials of facial motoneurons evoked by stimulation of the caudal trigeminal nucleus were investigated in acute experiments on cats by extra- and intra-cellular recording. Pre- and postsynaptic components of field potentials were found. Four types of motoneuron response were distinguished: EPSP with generation of single action potentials; a gradual shift of depolarization inducing grouped action potentials; a rhythmic discharge of action potentials arising at a low level of depolarization; and EPSPs or EPSP-IPSP sequences. The monosynaptic and (chiefly) polysynaptic nature of these responses was demonstrated. The possible mechanism of afferent control over facial motoneurons are discussed.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 12, No. 3, pp. 272–282, May–June, 1980.     

Reciprocal inhibition studied in discharging human motor units

The effect of excitation of group Ia afferents, evoked by stimulation of a mixed nerve, on the firing pattern of voluntarily activated single motor units of an antagonist muscle (biceps femoris, triceps surae, and tibialis anterior muscles) was studied. Poststimulus histograms were constructed for rhythmic sequences of motor unit potentials recorded by needle electrodes and the duration of interspike intervals was analyzed. Reciprocal inhibition and other effects accompanying nerve stimulation were discovered in the motoneurons of all three muscles. Distinguishing features of the manifestation of reciprocal inhibition in a discharging motoneuron were investigated; the effect was shown to depend on the time of occurrence of the inhibitory action in the interspike interval.Institute for Problems in Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 10, No. 6, pp. 626–636, November–December, 1978.     

Changes in the excitability of and synaptic effects on motoneurons during formation of the scratch motion generator in the cat

Studies on immobilized decerebrate (at intracollicular level) cats in which the scratch generator had been set up following bicuculline application to the upper cervical segments of the spinal cord, showed that the state of the segmental apparatus of the lumbosacral section of the spinal cord differs substantially from that seen in the spinal animal. Direct excitability of motoneurons of the "aiming" and "scratching" muscles rises, while recurrent and reciprocal Ia inhibition of motoneurons intensifies and the influence of Ib afferents on motoneurons declines. Afferents of the flexor reflex exert a primarily inhibitory influence on motoneurons of the "aiming" muscles. This influence becomes predominantly excitatory following spinalization, while the inhibitory effects of these afferents on motoneurons of the "scratch" muscles declines. The functional significance of the changes discovered in generation of scratch routine is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 19, No. 2, pp. 244–250, March–April, 1987.     

Opiate-Induced Suppression of Rat Hypoglossal Motoneuron Activity and Its Reversal by Ampakine Therapy

Background

Hypoglossal (XII) motoneurons innervate tongue muscles and are vital for maintaining upper-airway patency during inspiration. Depression of XII nerve activity by opioid analgesics is a significant clinical problem, but underlying mechanisms are poorly understood. Currently there are no suitable pharmacological approaches to counter opiate-induced suppression of XII nerve activity while maintaining analgesia. Ampakines accentuate α-amino-3-hydroxyl-5-methyl-4-isoxazole-propionate (AMPA) receptor responses. The AMPA family of glutamate receptors mediate excitatory transmission to XII motoneurons. Therefore the objectives were to determine whether the depressant actions of μ-opioid receptor activation on inspiratory activity includes a direct inhibitory action at the inspiratory premotoneuron to XII motoneuron synapse, and to identify underlying mechanism(s). We then examined whether ampakines counteract opioid-induced depression of XII motoneuron activity.

Methodology/Principal Findings

A medullary slice preparation from neonatal rat that produces inspiratory-related output in vitro was used. Measurements of inspiratory burst amplitude and frequency were made from XII nerve roots. Whole-cell patch recordings from XII motoneurons were used to measure membrane currents and synaptic events. Application of the μ-opioid receptor agonist, DAMGO, to the XII nucleus depressed the output of inspiratory XII motoneurons via presynaptic inhibition of excitatory glutamatergic transmission. Ampakines (CX614 and CX717) alleviated DAMGO-induced depression of XII MN activity through postsynaptic actions on XII motoneurons.

Conclusions/Significance

The inspiratory-depressant actions of opioid analgesics include presynaptic inhibition of XII motoneuron output. Ampakines counteract μ-opioid receptor-mediated depression of XII motoneuron inspiratory activity. These results suggest that ampakines may be beneficial in countering opiate-induced suppression of XII motoneuron activity and resultant impairment of airway patency.     

The amplitude and phase responses of the firing rates of some motoneuron models

A vertebrate motoneuron receives an enormous amount of synaptic activity from descending pathways, from spinal cord interneurons and directly from mechanoreceptor afferents. The intrinsic characteristics of the motoneuron will determine how its output spike train will encode the activities of all its inputs. Therefore, the essence of the intrinsic motoneuron characteristics should be well studied and modelled if the roles of the motoneuron as a processing or encoding element are to be well understood. Mathematical models of motoneurons have been described in the literature and tested mostly under static conditions. To increase the reality of the validation of such models, the objective of the present work is to test a few selected models described in the literature using sinusoidal injected current of different frequencies. The resulting frequency responses are compared with data available in the literature from cat type F motoneurons. Discrepancies between some of the models' responses and real motoneuron data suggest that improvements are needed in the modelling of the afterhyperpolarization mechanism.     

Neuronal organization of coeruleo-spinal connections in cats

Experiments on cats showed that electrical stimulation of the locus coeruleus leads to diminution of the inhibitory action of flexor reflex afferents (FRA) on the extensor monosynaptic reflex and to a decrease in amplitude of the IPSP evoked by FRA in extensor motoneurons. Injection of microdoses of aspartic acid and chlorpromazine into the locus coeruleus depresses the inhibitory effects of FRA, whereas injections of procaine and noradrenalin potentiate the inhibitory action of FRA. Data are given on the character of the descending influence of the locus coeruleus on different groups of spinal interneurons. It is concluded that depression of the inhibitory action of FRA is effected at the level of the final inhibitory interneurons of the "FRA system."A. A. Bogolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 16, No. 3, pp. 362–374, May–June, 1984.     

Corticofugal influences on the motoneurons of the trigeminal nucleus

We studied the postsynaptic potentials evoked from 76 trigeminal motoneurons by stimulation of the motor (MI) and somatosensory (SI) cortex in the ipsilateral and contralateral hemispheres of the cat. Stimulation of these cortical regions evoked primarily inhibitory postsynaptic potentials (PSP) in the motoneuron of the masseter muscle, but we also observed excitatory PSP and mixed reactions of the EPSP/IPSP type. The average IPSP latent period for the motoneurons of the masseter on stimulation of the ipsilateral cortex was 6.1±0.3 msec, while that on stimulation of the contralateral cortex was 5.2±0.4 msec; the corresponding figures for the EPSP were 7.6±0.5 and 4.5±0.3 msec respectively. Corticofugal impulses evoked only EPSP and action potentials in the motoneurons of the digastric muscle (m. digastricus). The latent period of the EPSP was 7.6 msec when evoked by afferent impulses from the ipsilateral cortex and 5.4 msec when evoked by pulses from the contralateral cortex. The duration of the PSP ranged from 25 to 30 msec. Postsynaptic potentials developed in the motoneurons studied when the cortex was stimulated with a single stimulus. An increase in the number of stimuli in the series led to a rise in the PSP amplitude and a reduction in the latent periods. When the cortex was stimulated with a series of pulses (lasting 1.0 msec), the IPSP were prolonged by appearance of a late slow component. We have hypothesized that activation of the trigeminal motoneurons by corticofugal impulsation is effected through a polysynaptic pathway; each functional group of motoneurons is activated in the same manner by the ipsilateral and contralateral cortex. The excitation of the digastric motoneurons and inhibition of the masseter motoneurons indicates reciprocal cortical control of their activity.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 3, No. 5, pp. 512–519, September–October, 1971.     

Changes in segmental reflex responses during stimulation of locus coeruleus and potentiation of the action of catecholamines

Repetitive stimulation of the locus coeruleus (up to 150 µA in strength) was accompanied by marked weakening of the inhibitory action of flexor reflex afferents and of the reciprocal inhibitory action on extensor motoneurons. Meanwhile stimulation of this sort had no significant effect on direct inhibition of flexor and extensor motoneurons, on the facilitatory action of flexor reflex afferents and the reciprocal inhibitory action on flexor motoneurons and also on dorsal root potentials. Intravenously injected pyrogallol had a similar action, but its effect was much weaker after spinalization of the animals or blocking of spinal cord conduction by cold. Enhancement of the monosynaptic reflex, which also was observed after injection of pyrogallol, was characterized by different temporal parameters; the intensity of this effect was unaffected both by spinalization and by cold block. These data, and also the results of experiments with partial divisions of the spinal cord, suggest that the effects of stimulation of the locus coeruleus are the result of activity of a descending coerulo-spinal tract, running in the ventral quadrant of the spinal cord.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 13, No. 1, pp. 39–47, January–February, 1981.