Comparative analysis of background activity of ventrolateral thalamic neurons in patients with parkinsonism and torsion muscle dystonia

In the course of neurosurgical interventions in 40 patients with parkinsonism and torsion muscle dystonia, the background activity (BA) was recorded from 124 neurons of the ventrolateral nucleus (VL) of the thalamus with the aid of microelectrodes during demarcation of boundaries between nuclear structures, and identification of zones within a nucleus. The following characteristic features of the BA in patients with parkinsonism were found: a relatively large proportion (71%) of cells with a burst pattern of activity; a great diversity of burst duration in the activity (short or prolonged bursts); a large proportion (67%) of cells with cyclic changes in the BA frequency; diverse patterns of cyclic modulation, where periods varied from fractions of a second (0.2–0.8 sec) to seconds (2–10 sec) or to tens of seconds (20–40 sec); cyclic successions of spike bursts with the rhythm typical of the peripheral tremor (3–7/sec) in a substantial proportion (40%) of the units.The background spike trains recorded in patients with torsion muscle dystonia consisted of separated spikes in the majority (69%) of units. Only short high-frequency discharges were found in the burst activity in this group; a cyclic BA pattern occurred much less frequently (in 23% of neurons); burst discharges at a 3–4/sec frequency were found only in 4% of the examined cells.The possible nature of motor disorders in patients with parkinsonism and torsion muscle dystonia and the mechanisms of the curing effects of cryodestruction of theVL of the thalamus in the treatment of the disease are discussed.Neirofiziologiya/Neurophysiology, Vol. 25, No. 4, pp. 246–253, July–August, 1993.     

Effects of systemic injection of neuroleptics on neuronal background activity in the cat ventrolateral thalamic nucleus

Background activity was recorded in 272 neurons of the ventrolateral thalamic nucleus before and after systemic haloperidol and droperidol injection at a cataleptic dose using intracellular techniques during chronic experiments on cats in a drowsy condition. Brief burster discharges lasting 5–50 msec and following on at a high intraburst spike rate (of 200–450 Hz) were characteristic of neuronal activity in intact animals. Regular discharges occurred at the rate of 2–2.5 Hz or occasionally 3–4 Hz in 15% of cells. Numbers of neurons with the latter activity pattern rose to 22 and 30%, respectively, following haloperidol and droperidol injection. Both irregular and prolonged (80–300 msec) regular discharges were recorded in one third of the total. A relatively low intraburst spike rate (of 60–170 Hz) was observed in 37% of cells following 10 days' haloperidol injection. These changes are thought to be produced by intensified inhibitory effects on neurons of the thalamic ventrolateral nucleus from the substantia nigra and reticular thalamic nucleus following blockade of dopaminergic and -adrenergic receptors.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 21, No. 5, pp. 675–685, September–October, 1989.     

Rhythmic background thalamic neuronal activity in patients with extrapyramidal motor disorders

The background activity of 123 thalamic neurons was recorded in 30 patients with motor extrapyramidal disorders applying microelectrode techniques to neurosurgical practice. Recordings were taken from the ventro-oral anterior and posterior thalamic nuclei and the adjacent reticular nucleus. A computer analysis was performed of neuronal activity in 44 units and plots produced of autocorrelation and spectral density functions. In patients with parkinsonism and double athetosis, rhythmic activity was found in 48% of cells. A wide variety of regular fluctuations in background neuronal discharges was noted: in the range of theta and delta rhythms (5–7 and 1–4 Hz respectively) with a periodicity of seconds (2–10 sec) and decaseconds (15–40 sec). It was thought possible that several types of regular waves may coexist: phenomena of 2 or 3 accelerated waves and reduced frequency of spike activity of differing periodicity were observed in eight neurons within the same train of spikes. The origin and significance of rhythmically occurring changes in thalamic neuronal spike activity are discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR; Institute of Neurosurgery, Ministry of Public Health of the Ukrainian SSR. Translated from Neirofiziologiya, Vol. 19, No. 2, pp. 192–201, March–April, 1987.     

Responses of cat cerebellar cortical neurons to stimulation of the caudate nucleus,globus pallidus,and substantia nigra

Stimulation of the head of the caudate nucleus in cats anesthetized with chloralose and pentobarbital evoked spike responses of the Purkinje cells and other cerebellar cortical neurons in the paramedian lobes, lobulus simplex, and the tuber of the vermis. Phasic responses in the form of simple discharges (on account of activation of the neurons through mossy fibers) appeared mainly after a latent period of 5–12 and 14–20 msec; the latent period of responses consisting of complex discharges (on account of activation of Purkinje cells through climbing fibers) was 5–6, 9–22 msec, or more. Depending on the latent period, the spike responses differed in their rhythm of generation. In response to stimulation of the caudate nucleus with a frequency of 4–6/sec recruiting responses were found. An inhibitory pause was an invariable component of the tonic responses. During stimulation of the globus pallidus responses of the same types (phasic and tonic) appeared as during stimulation of the caudate nucleus, but they differed in the distribution of the neurons by latent period of spike responses. The minimal latent period was 4 msec. Recruiting also was observed during repetitive stimulation of the globus pallidus. During stimulation of the substantia nigra Pukinje cells activated by climbing fibers responded. Evoked complex discharges appeared after a stable latent period of 8.5±0.3 msec. Arguments are put forward regarding the role of the substantia nigra, the globus pallidus, nuclei of the inferior olive, and also the thalamic nuclei in the mechanism of caudato-cerebellar oligosynaptic and polysynaptic connections.N. I. Pirogov Medical Institute, Vinnitsa. Translated from Neirofiziologiya, Vol. 10, No. 4, pp. 375–384, July–August, 1978.     

Pattern and correlation of fast and slow spontaneous unit activity in the visual cortex

Spontaneous unit activity in the visual cortex and its changes during stimulation by continuous light or flashes were investigated in waking rabbits. The study of distributions of adjacent intervals showed that the neurons differ in the ratio of burst (fast, with intervals of up to 15–40 msec) and nonburst (slow) activity and in the character of changes from one type of activity to the other. Of the total number of spikes 63% were outside bursts; the ratio of their number to the number of spikes within bursts consisting of two or of three or more spikes was 27:3:1. The relative stability of the burst structure of spontaneous activity and the limited number of spikes in them (on average 2.4) were demonstrated. Bursts of three or more spikes (mean 3.6) were irregular, and in 79% of them a longer interval (18.6±2.4 msec) was observed before the shortest interval (7.9±0.9 msec). Bursts of spikes of most neurons during photic stimulation contain more spikes with shorter intervals; they also began more frequently with the shortest interval, possibly signifying an increase in the steepness and amplitude of the EPSPs lying at their basis. However, in 20% of neurons spontaneous bursts included more spikes and with shorter intervals than bursts evoked by flash stimulation.Research Institute of Psychiatry, Ministry of Health of the RSFSR, Moscow. Translated from Neirofiziologiya, Vol. 11, No. 4, pp. 311–320, July–August, 1979.     

Analysis of spontaneous unit activity of the cat caudate nucleus

Spontaneous unit activity recorded extracellularly from the caudate nucleus in acute experiments on cats was analyzed. A graph of the sliding mean frequency, an interspike interval histogram, correlogram, intensity function, and histogram of correlation between adjacent intervals were plotted for the spontaneous activity of each neuron. The spontaneous activity of neurons of the caudate nucleus showed considerable variability in time and its mean frequency varied for different neurons from 0.5 to 20 spikes/sec. Depending on the temporal pattern of the spikes and also on the statistical indices, spontaneous unit activity in the caudate nucleus was conventionally divided into two types: single and grouped. A switch from one type of activity to the other was observed for the same neuron. On the basis of the data as a whole it is impossible to regard the spontaneous unit activity of the caudate nucleus as a simple random (Poissonian) spike train.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 9, No. 4, pp. 369–376, July–August, 1977.     

Unitary responses of the caudate nucleus to direct stimulation

Unitary responses of the caudate nucleus to stimulation of various parts of it were investigated by extracellular recording. Latent periods of response discharges varied from 3.5 to 40 msec. Most neurons were excited by stimulation of the most rostral part of the head of the caudate nucleus. Irrespective of the site of stimulation, in most cases responses consisted of initial excitation in the form of one or, less frequently, two discharges followed by a period of depression of spontaneous activity. Recovery of activity took place gradually, without postinhibitory facilitation. No afterdischarges or periodic repetitions of spikes were observed after the initial response. Repetitive stimulation of the caudate nucleus showed that the neurons of this nucleus reproduce frequencies of stimulation badly above 30/sec, and under these circumstances in many cases they continued to discharge on average at a frequency of 5–15/sec. The results are examined from the standpoint of participation of the caudate nucleus in the formation of spindle activity.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 8, No. 5, pp. 497–506, September–October, 1976.     

Neuronal firing activity in the lateral vestibular nucleus during locomotion in the guinea pig

The effects of spontaneous locomotor activity on neuronal background firing in the lateral vestibular nucleus was investigated during experiments on decerebrate guinea pigs. The onset of rhythmic muscular activity in the extramities was found to produce a rise in the rate of such discharges, which increased from 10–15 to 100 spikes/sec in most neurons. A higher rate occurred as activity began in the ipsilateral forelimb extensor muscles (the stage corresponding to the end of the swing phase and start of the stance phase in the locomotor cycle). The alterations noted in vestibular neuronal activity during locomotion are thought to ensure the background of high anti-graveity muscle tonus against which rhythmic limb movements take place.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 23, No. 5, pp. 536–541, September–October, 1991.     

Effects of experimentally induced deficiency of catecholaminergic transmission on neuronal activity in the ventrolateral thalamic nucleus

Background activity was investigated in 272 neurons of the ventrolateral thalamic nucleus (VLTN) before and after systemic administration of neuroleptics (haloperidol and droperidol) at cataleptic doses by means of extracellular techniques during chronic experiments on cats. Autocorrelation and spectral analysis revealed regularly-occurring changes in the background activity rate of VLTN neurons, the periodicity of which changed by fractions of seconds (0.2–0.8 sec), seconds (1.5–10 sec), or tens of seconds (12–30 sec). While numbers of neurons with individual types of periodic activity did not exceed 6–8% in intact animals, it did increase to 18–30% after administering neuroleptics. Raised numbers of neurons with two types of regularly occurring processes within a single spike train were also noted. Experimentally-produced data were compared with findings from clinical observations. Quantities of neurons with different variations in the periodicity of their firing activity reached 19–46% in patients with parkinsonism but did not exceed 4–8% in those with torsion dystonia. The genesis of raised rhythmic firing in thalamic neurons occurring with parkinsonism is thought to be associated with impaired catecholaminergic (both dopaminergic and -adrenergic) transmission.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 22, No. 3, pp. 359–368, May–June, 1990.     

Postsynaptic neuronal response of a cat sensorimotor cortex during evoked and self-sustained rhythmic "spike and wave" activity

Intracellular correlates of complex sets of rhythmic cortical "spike and wave" potentials evoked in sensorimotor cortex and of self-sustained rhythmic "spike and wave" activity were examined during acute experiments on cats immobilized by myorelaxants. Rhythmic spike-wave activity was produced by stimulating the thalamic relay (ventroposterolateral) nucleus (VPLN) at the rate of 3 Hz; self-sustained afterdischarges were recorded following 8–14 Hz stimulation of the same nucleus. Components of the spike and wave afterdischarge mainly correspond to the paroxysmal depolarizing shifts of the membrane potential of cortical neurons in length. After cessation of self-sustained spike and wave activity, prolonged hyperpolarization accompanied by inhibition of spike discharges and subsequent reinstatement of background activity was observed in cortical neurons. It is postulated that the negative slow wave of induced spike and wave activity as well as slow negative potentials of direct cortical and primary response reflect IPSP in more deep-lying areas of the cell bodies, while the wave of self-sustained rhythmic activity is due to paroxysmal depolarizing shifts in the membrane potential of cortical neurons.I. S. Beritashvili Institute of Physiology, Academy of Sciences of the Georgian SSR, Tbilisi. Translated from Neirofiziologiya, Vol. 18, No. 3, pp. 298–306, May–June, 1986.     

Responses of cat pulvinar neurons to moving visual stimuli

Responses of 114 pulvinar neurons to moving visual stimuli were studied. Most (79) neurons possessed spontaneous activity (10–25 spikes/sec). Of 59 neurons tested, 31 responded to stimulation of both retinas and 28 to stimulation only of the contralateral retina. Of 114 neurons, 41 responded only to movement of black objects, while the rest responded to movement of both black and light objects. According to the character of their responses to movement the neurons were divided into two main groups. The first group consisted of neurons sensitive to the direction of motion and responding with a spike discharge to movement in one direction and by inhibition to movement in the opposite direction. The second group included neurons insensitive to the direction of motion and responding by an equal number of discharges to movements in two opposite directions. Besides these two main groups, other neurons responding to movement in two opposite directions by discharges with different temporal distribution and also neurons which changed the character of their response from nondirectional to directional depending on the size of the moving stimulus, were found.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 10, No. 4, pp. 348–354, July–August, 1978.     

Tranformation of long reticulofugal spike trains by interneurons connected monosynaptically with the reticulospinal tract

Characteristics of transmission of activity evoked by stimulation of the reticular formation through interneurons located in the ventromedial zones of the gray matter of the lumbar division of the spinal cord and connected monosynaptically with reticulospinal fibers were investigated in cats. Responses of the neurons to relatively low-frequency (not exceeding 80–100/sec) stimulation consisted mainly of stationary discharges; with a further increase in frequency the response became nonstationary (the initial, relatively high-frequency discharge was followed by partial or complete suppression of the discharge). The maximal frequency of the initial phase of the response to high (over 400/sec) frequencies of stimulation was 180–230 spikes/sec. The "transmission factor" (ratio between the frequency of spikes in the response to the frequency of stimulation), calculated for stationary discharges, reached 0.7–0.8 at low frequencies of stimulation, and then decreased significantly. On the basis of the statistical characteristics of the stationary portions of the evoked activity and analysis of these data by the use of a mathematical model, indirect estimates were obtained of the parameters of processes lying at the basis of the transforming properties of this cell population.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 10, No. 3, pp. 278–286, May–June, 1978.     

Modification of evoked unit activity of the dorsomedial thalamic nucleus following destruction of the basolateral amygdala in rats

Neuronal responses of the dorsomedial thalamic nucleus of anesthetized rats to stimulation of the anterior periamygdalar cortex were compared before and after destruction of the basolateral nuclei of the ipsilateral amygdala. After destruction of the afferent inflow from the nuclei of the amygdala a new type of response appeared, in the form of prolonged generation of bursts of spikes, and also a significant decrease in the number of cells with a phasic activation response and an increase in the number of tonically activated neurons. Destruction of the amygdala was shown to lead to an increase in the number of cells of the dorsomedial nucleus with spontaneous activity. The duration of the tonic spike response in animals undergoing the operation was found to be significantly increased from 142.5±42.4 to 375.4±53.8 msec, whereas the parameters of other types of neuronal responses did not differ statistically significantly.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 16, No. 6, pp. 784–789, November–December, 1984.     

Electrophysiological Actions of Oxytocin on the Rabbit Myometrium

The electrical activities of myometrial cells of the pregnant rabbit uterus have been studied by means of sucrose-gap and intracellular micro-electrode recording techniques. The resting potential of the myometrial cell was about -50 mv, and it is unaffected by the duration of pregnancy or placental attachment. Action potentials of the myometrium, although dependent on external Na⁺, were not always of the regenerative type; preparations from nonparturient uteri often produce mainly small spikes. The mean spike amplitude was 35 mv, rising at a mean maximum rate of 3 v/sec. Oxytocin, in concentrations less than 500 µU/ml, increased the mean spike amplitude to 48 mv and the mean maximum rate of rise to 7 v/sec, without affecting the resting potential. The relation between membrane potential and dV/dt of the spike was steepened by oxytocin, suggesting that oxytocin increased the number of normally sparse sodium gates in the myometrial membrane. By this action, oxytocin is believed to increase the probability of successful regenerative spikes and thereby initiate electrical activity in quiescent preparations, increase the frequency of burst discharges, the number of spikes in each burst, and the amplitude of spikes in individual cells.     

Electrophysiological properties of neurons and neuronal organization of the crayfish somatogastric ganglion

Experiments with intracellular recording from neurons of the isolated crayfish somatogastric ganglion established that the membrane potential of the neurons is 53±3 mV. Single stimulation of the central branches of the ganglion evoked EPSP and a spike in the neurons. The spike amplitude was 7.5±0.6 mV. The small amplitude of the spike is explained by the fact that it arises at some distance from the body of the neuron and propagates electrotonically in it. Summation of several EPSP is necessary in most cases for initiation of the spike. When the orthodromic stimulus was strong enough, and IPSP occurred in some cells in addition to the EPSP and spike. Stimulation of the peripheral nerves of the ganglion induced in most neurons antidromic excitation and in some neurons orthodromic excitation. Some neurons spontaneously discharged rhythmically with an unstable frequency (11–27 impulses/sec). An investigation of the effect on neurons of chemical agents [acetylcholine, adrenalin, noradrenalin, gamma-aminobutyric acid (GABA), glutamic acid, and dopamine] showed that acetylcholine has the strongest and most stable depolarizing action and apparently is a synaptic transmitter in the ganglion. The other agents excited some neurons — depolarized them and evoked rhythmic discharges — and, coversely, hyperpolarized and suppressed the rhythmic activity of other neurons. A scheme of neuronal organization of the somatogastric ganglion of the crayfish is proposed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 2, No. 3, pp. 307–313, May–June, 1970.     

Various forms of potentials recorded from caudate nucleus neurons

Spontaneous and evoked activity of caudate nucleus neurons was recorded extracellularly in acute experiments on cats. Different forms of potentials were found by analysis of the results. The potentials recorded belong to three types: ordinary action potentials; prepotentials or incomplete spikes differing from ordinary action potentials in their lower amplitude and slower decline, and complex discharges in which a spike of somewhat reduced amplitude is followed by a slow positive-negative wave. In the spontaneous activity prepotentials were observed both in complete action potentials and in isolation. The frequency of the complex discharges was 0.5–1 per second. The slow wave of these discharges blocked prepotential and action potential formation. The origin of these forms of potentials in neurons of the caudate nucleus is discussed and they are compared with analogous forms of potentials described for the Purkinje cells of the cerebellum.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukranian SSR, Kiev. Translated from Neirofiziologiya, Vol. 9, No. 2, pp. 149–156, March–April, 1977.     

Responses of caudate neurons to acoustic stimulation in cats

Responses of 141 neurons of the caudate nucleus to acoustic stimuli — tones (500 and 2000 Hz) and clicks of different frequency (0.2 and 0.8/sec) and intensity (75, 80, 95 dB) — were recorded extracellularly in chronic experiments on cats. The responses recorded showed great variability with respect to character (phasic, tonic), structure (one or two phases of excitation), latent periods (from 7.5 to 300.0 msec), and burst discharge frequency (from 90 to 800 spikes/sec). Analysis of averaged poststimulus histograms and graphs of the dynamics of the responses showed that responses of 74% of neurons were much better expressed if less frequent stimuli were used: The regularity of the responses and the number of spikes in each response increased. Responses of neurons also increased and acquired a more distinct temporal structure if the intensity of the clicks increased. The character of responses to clicks and tones differed qualitatively in 17% of neurons studied: Phasic excitation arose in response to clicks, tonic changes in spike activity to tones. The particular features of responses of caudate neurons to acoustic stimulation with different parameters are discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 12, No. 6, pp. 588–595, November–December, 1980.     

Effect of diazepam on cyclic change of excitability in a cortical epileptic focus

On application of penicillin solution to the motor cortex of rats and electrical stimulation of the ventroposterolateral thalamic nucleus (VPL) with a frequency of 1/sec and with a strength three times greater than the threshold for the primary evoked potential, alternation of "active" and "passive" neurons was observed in the zone of the epileptic focus. "Active" neurons were characterized by the regular appearance of epileptiform discharges in response to each stimulation of VPL. In the course of the "passive" periods stimulation of VPL led only to the appearance of primary evoked potentials. Diazepam in a dose of 2 mg/kg completely disturbed spike generation in the epleptic focus, but in a dose of 0.1 mg/kg it caused disturbance of the established cyclic change of excitability in the focus, which was restored after stimulation with a frequency of 2/sec.Institute of General Pathology and Pathological Physiology, Academy of Medical Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 12, No. 6, pp. 563–570, November–December, 1980.     

Connections of neurons of the cat somatosensory cortex with the thalamic relay nuclei

Of 103 neurons in the rostral part of the posterior sigmoid gyrus of the cat cortex 30 responded to stimulation of the ventro-posterolateral and ventrolateral nuclei of the thalamus (VPL and VL), 42 responded to stimulation of VL only, and 31 to stimulation of VPL only. It was shown by intracellular recording that stimulation of VPL induces a spike response with or without subsequent IPSPs in some neurons and an initial IPSP in others. The spike frequency of single neurons reached 60/sec, but the IPSP frequency never exceeded 10–20/sec. Stimulation of VL was accompanied by: a) antidromic spike responses; b) short-latency monosynaptic EPSPs and spikes capable of following a stimulation frequency of 100/sec; c) long-latency polysynaptic EPSPs and spikes appearing in response to stimulation at 4–8/sec; d) short-latency IPSPs; e) long-latency IPSPs increasing in intensity on repetition of infrequent stimuli. It is concluded that the afferent inputs from the relay nuclei to neurons of the somatosensory cortex are heterogeneous. An important role is postulated for recurrent inhibition in the genesis of the long-latency IPSPs arising in response to stimulation of VL, and for direct afferent inhibition during IPSPs evoked by stimulation of VPL. It is shown that the rostral part of the posterior sigmoid gyrus performs the role of somatic projection and motor cortex simultaneously.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 4, No. 3, pp. 245–255, May–June, 1972.     

Characteristics of background unit activity in the nucleus reticularis of the human thalamus

Background firing activity was examined in 240 neurons belonging to the thalamic nucleus reticularis (Rt) in the unanesthetized human brain by extracellular microelectrode recording techniques during stereotaxic surgery for dyskinesia. The cellular organization of Rt was shown to be nonuniform, and distinguished by the presence of three types of neuron: one with arrhythmic single discharge (A-type, 40%), another with rhythmic (2–5 Hz) generation of short high-frequency (of up to 500/sec) burster discharges (B-type, 49%) and a third with aperiodic protracted high-frequency (of up to 500/sec) bursting discharges separated by "silent" intervals of a constant duration of 80–150 msec (i.e., C-type, 11%). Differences between the background activity pattern of these cell types during loss of consciousness under anesthesia are described. Tonic regulation of neuronal type was not pronounced but a tendency was noticed in the cells towards a consistent rise in firing rate, rhythmic frequency and variability, etc. in both A and B units, especially in the latter. Findings pointing to the absence of a direct relationship between rhythmic activity in the Rt and parkinsonian tremor were confirmed. Background activity in B-type cells was found to increase and then stabilize with a rise in the degree of tremor. The nature of regular bursting activity patterns in B and C neurons is discussed in the light of our findings.Institute of Chemical Physics, Academy of Sciences of the USSR, Moscow. Institute of Neurosurgery, Academy of Medical Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 19, No. 4, pp. 456–466, July–August, 1987.