Investigation of electrical responses recorded from the fenestra cochleae in cats

Electrical responses of the fenestra cochleae to stimulation by clicks of different intensity, polarity, and frequency, were studied in anesthetized cats. The absolute values of amplitude and latent period of the neural component of the response reflect the physiological state of the auditory nerve. Besides ordinary potentials characterized by peaks N1 and N2, specific responses were observed when clicks with an intensity of 85 dB or "rarefaction" clicks were used. Dependence of the amplitude of these responses on the intensity of acoustic stimuli of different polarity was investigated during a change in the rhythm of the stimulation; the effect of different rhythms of stimulation on the gradient of the curve reflecting this relationship was examined. The possible mechanisms of the effect of stimulus frequency are discussed.Scientific-Research Institute of Otolaryngology, Ministry of Health of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 11, No. 2, pp. 151–157, March–April, 1979.     

Characteristics of unit responses of the cat cerebellar cortex to acoustic stimuli

Investigation of unit responses of the cerebellar cortex (lobules VI–VII of the vermis) to acoustic stimulation showed that the great majority of neurons responded by a discharge of one spike or a group of spikes with a latent period of 10–40 msec and with a low fluctuation value. Neurons identified as Purkinje cells responded to sound either by inhibition of spontaneous activity or by a "climbing fiber response" with a latent period of 40–60 msec and with a high fluctuation value. In 4 of 80 neurons a prolonged (lasting about 1 sec or more), variable response with a latent period of 225–580 msec was observed. The minimal thresholds of unit responses to acoustic stimuli were distributed within the range from –7 to 77 dB, with a mode from 20 to 50 dB. All the characteristics of the cerebellar unit responses studied were independent of the intensity, duration, and frequency of the sound, like neurons of short-latency type in the inferior colliculi. In certain properties — firing pattern, latent period, and threshold of response — the cerebellar neurons resemble neurons of higher levels of the auditory system: the medial geniculate body and auditory cortex.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 5, No. 1, pp. 3–12, January–February, 1973.     

Response of cat auditory cortex neurons to tonal stimulation during and after nembutal anesthesia

During acute experiments on 20 cats a comparative study was made of neuronal reaction to a tone, as recorded during the first few hours after administration of Nembutal and after an interval of 10–30 h. No spontaneous activity was seen in 89% of auditory cortex neurons of the anesthetized cats; these produced a sterotyped on- response to the optimal frequency tone. Late neuronal spike discharges at distinct intervals of 100–150 msec appeared in response to the setting up of acoustic stimulation after a brief latent reaction lasting 9–15 msec. It was shown that this stimulation did not produce an off-response in the cortical neurons. When the animals emerged from Nembutal anesthesia, the neurons reacted very differently to the optimal frequency tone. About 76% of the cells produced an on, on-off or off response, while about 21% responded with either tonic spike discharges or total inhibition of these throughout the acoustic stimulation. In unanesthetized cats the vast majority of AI cortical neurons were capable of reacting as long as the stimulus lasted. It is shown how this ability is lost under deep Nembutal anesthetic.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 17, No. 6, pp. 728–737, November–December, 1985.     

Tympanic and extratympanic sound transmission in the leopard frog

Summary The inner ear of the leopard frog,Rana pipiens, receives sound via two separate pathways: the tympanic-columellar pathway and an extratympanic route. The relative efficiency of the two pathways was investigated. Laser interferometry measurements of tympanic vibration induced by free-field acoustic stimulation reveal a broadly tuned response with maximal vibration at 800 and 1500 Hz. Vibrational amplitude falls off rapidly above and below these frequencies so that above 2 kHz and below 300 Hz tympanic vibration is severely reduced. Electrophysiological measurements of the thresholds of single eighth cranial nerve fibers from both the amphibian and basilar papillae in response to pure tones were made in such a way that the relative efficiency of tympanic and extratympanic transmission could be assessed for each fiber. Thresholds for the two routes are very similar up to 1.0 kHz, above which tympanic transmission eventually becomes more efficient by 15–20 dB. By varying the relative phase of the two modes of stimulation, a reduction of the eighth nerve response can be achieved. When considered together, the measurements of tympanic vibration and the measurements of tympanic and extratympanic transmission thresholds suggest that under normal conditions in this species (1) below 300 Hz extratympanic sound transmission is the main source of inner ear stimulation; (2) for most of the basilar papilla frequency range (i.e., above 1.2 kHz) tympanic transmission is more important; and (3) both routes contribute to the stimulation of amphibian papilla fibers tuned between those points. Thus acoustic excitation of the an uran's inner ear depends on a complex interac tion between tympanic and extratympanic sound transmission.Abbreviations dB SPL decibels sound pressure level re: 20 N/ m² - AP amphibian papilla - BP basilar papilla - BEF best excitatory frequency     

Responses of the neurons of the cortical secondary auditory area to acoustic and non-acoustic stimuli in cats

The responses of the cortical secondary auditory area (AII) to the non-acoustic stimuli (electrical stimulation of the skin in the vibrissae area and light flash) and their combination with acoustic stimulation (sound click or tone) were studied in experiments on cats anesthetized by kalipsol using extra- and intracellular recording. Of the total number of neurons, 69% of the units generating spike responses to the acoustic stimulation responded to the non-acoustic stimulation too. The responses to the modal-nonspecific stimulation, as a rule, were weak and variable; they were mostly represented by a tonic change in the neuronal discharge frequency. The nonspecific stimulation evoked primary excitatory and inhibitory postsynaptic potentials in 77% and 20% of the examined neurons, respectively. We found that synaptic effects of the nonspecific and specific stimulations interact with each other, ensuring considerable modulation of the latter (mostly a significant facilitation resulting from the EPSP summation and suppression of an inhibitory component of the response to acoustic stimulation). Possible participation of the midbrain reticular formation in the transmission of the modal-nonspecific influences to the cortical neurons is considered; stimulation of this structure evoked responses similar to those evoked by the modal-nonspecific sensory stimuli.Neirofiziologiya/Neurophhysiology, Vol. 26, No. 5, pp. 356–364, September–October, 1994.     

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.     

Multisensory convergence on neurons of the motor cortex in unanesthetized cats

Postsynaptic potentials (PSPs) of 83 neurons in the motor cortex of unanesthetized cats in response to electrodermal, photic, and acoustic stimulation were investigated by intra-and quasi-intracellular recording methods. Most cells responded to stimulation of at least one limb. About 60% of neurons of the posterior and over 75% of neurons of the anterior sigmoid gyrus responded to stimulation of two (or more) limbs. In 29 of 39 neurons of the anterior and 12 of 44 of the posterior sigmoid gyrus PSPs with a short (less than 50 msec) and stable latent period were evoked by flashes and clicks. On presentation of two somesthetic stimuli complete blocking (if the interval was less than 30–60 msec) or weakening (interval 30–200 msec) of responses to the second (testing) stimulus was observed. On presentation of paired photic (or acoustic) stimuli or paired stimuli of different modalities at various intervals from 0 to 100 msec, the testing response was often potentiated. The character of the responses and their interaction thus differed from those obtained under chloralose anesthesia [6, 7]. It is postulated that under the action of chloralose a system of neurons with strong excitatory feedback is formed in the motor cortex which may respond to stimuli of different modalities by something resembling the "all or nothing" principle.Brain Institute, Academy of Medical Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 3, No. 6, pp. 563–573, November–December, 1971.     

Neuronal responses in the limbic cortex of awake cats during defensive conditioning

Spike activity was investigated in limbic cortex neurons during defensive conditioning to acoustic stimulation in chronic experiments on cats. A relationship was found between the numbers of neurons responding, their contribution to formation of a temporal connection, and the duration of the acoustic stimulus. Phasic responses of 50–500 msec duration with latencies of 15–50 msec were observed for the most part. Intensive spike response with a minimum latency of 15 msec and a duration of between 200 msec and 2.5 sec evolved in most cells (95.1% in field 24 and 83% in field 32) in response to electrical stimulation. Response to acoustic stimulation rose during defensive conditioning in 33.3% cells and declined and finally disappeared in 13.3%, but response at the site where reinforcement was abolished was reproduced in all these cells. It was thus found that the numbers of limbic cortex neurons responding to sound not only fails to increase but actually decreases after training. The limbic cortex is thought to play its most active part in conditioning response to a recognized signal during the period preceding the awaited painful reinforcement.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 18, No. 5, pp. 660–669, September–October, 1986.     

Neuronal response in nucleus reticularis thalami and neighboring thalamic structures to natural stimulation during chronic experiments on cats

Spike response was investigated in 104 neurons of the nucleus reticularis thalami (R) and adjoining thalamic nuclei to acoustic, tactile, and visual stimuli during chronic experiments on cats. Of the test neurons, 29% responded to acoustic stimulation and 11% showed no preference in relation to different acoustic stimuli. Minimum latencies of response to sounds measured 12–37 msec in excitatory and 18–27 msec in inhibitory cells. Duration of excitation produced by acoustic stimuli reached 50–250 msec; inhibition lasted 27–190 msec. Most cells belonging to this nucleus were excited by different stimuli; the proportion of inhibitory neurons did not exceed 4–10%.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 21, No. 4, pp. 451–461, July–August, 1989.     

Dynamic characteristics of motor responses evoked by stimulation of the motor cortex in unanesthetized cats

Temporal characteristics of motor responses evoked in unanesthetized cats by stimulation of the motor cortex through bipolar needle electrodes were investigated in chronic experiments. Isometric and isotonic contractions of the flexor muscles of the hip and knee joints of the limb contralateral to the point of stimulation were recorded. The latent period of response varied from 100 msec or more in the case of low-frequency (100–150 Hz) and low-threshold (1.1–1.2 thresholds) stimulation of the motor cortex to 30–35 msec in the case of "optimal" parameters of stimulation (300–400 Hz, 1.5–1.6 thresholds). If the intensity of stimulation was high enough the rising time constant of evoked contraction was 50–80 msec; values of the falling time constant of muscular contraction after cessation of stimulation were much greater, namely 150–300 msec. The rising time constant of contraction decreased with an increase in both the frequency and strength of motor cortical stimulation. The results are examined and discussed from the standpoint of methods of automatic control theory.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 12, No. 5, pp. 451–458, September–October, 1980.     

Responses of neurons of reticular and ventral anterior nuclei of the cat thalamus to afferent stimuli of different modalities

Responses of 146 spontaneously active neurons of the reticular nucleus (R) and of 98 neurons of the ventral anterior (VA) nucleus of the thalamus to electrical stimulation of the skin of the footpads, to flashes, and to clicks were studied in experiments on cats immobilized with D-tubocurarine or myorelaxin. Stimulation of the contralateral forelimb was the most effective: 24.9% of R neurons and 31.3% of VA neurons responded to this stimulation. A response to clicks was observed in only 4.4% of R neurons and 2.4% of VA neurons. Nearly all responding neurons did so by phasic (one spike or a group of spikes) or tonic excitation. Depression of spontaneous activity was observed only in response to electrical stimulation of the skin. Depending on the site of stimulation, it was observed in 2.6–4.3% of R neurons and 1.7–2.1% of VA neurons tested. The latent period of the phasic responses of most neurons was 6–64 msec to electrical stimulation of the contralateral forelimb, 11–43 msec in response to stimulation of the hindlimb on the same side, 10–60 msec to photic and 8–60 msec to acoustic stimulation. Depending on the character of stimulation, 75.1–95.6% of R neurons and 68.7–97.6% of VA cells did not respond at all to the stimuli used. Of the total number of cells tested against the whole range of stimuli, 25% of R neurons and 47% of VA neurons responded to stimulation of different limbs, whereas 16% of R neurons and 22% of VA cells responded to stimuli of different sensory modalities. The functional role of the convergence revealed in these experiments is to inhibit (or, less frequently, to facilitate) the response of a neuron to a testing stimulus during the 40–70 msec after conditioning stimulation.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 7, No. 6, pp. 563–571, November–December, 1975.     

Two patterns of bulbar neuronal response to microstimulation of locomotor and inhibitory brain stem sites

Synaptic responses (postsynaptic potentials and action potentials) were evoked in mesencephalic decerebellated cats by stimulating pontine bulbar locomotor and inhibitory sites (LS and IS, respectively) with a current of not more than 20 µA in "medial" and "lateral" neurons of the medulla. Some neurons even produced a response to presentation of single (actually low — 2–5 Hz — frequency) stimuli. The remaining cells responded to stimulation at a steady rate of 30–60 Hz only. Both groups of medial neurons were more receptive to input from LS. Lateral neurons responding to even single stimuli reacted more commonly to input from LS and those responding to steady stimulation only to input from IS. Many neurons with background activity (whether lateral or medial) produced no stimulus-bound response, but rhythmic stimulation either intensified or inhibited such activity. This response occurs most commonly with LS stimulation. Partial redistribution of target neurons in step with increasing rate of presynaptic input may play a major part in control of motor activity.Institute for Research into Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 22, No. 2, pp. 257–266, March–April, 1990.     

Role of the midbrain periaqueductal gray matter in conditioned reflexes

Unit activity in the midbrain periaqueductal gray matter (PGM) during an instrumental placing reflex, its extinction, differentiation, and conditioned inhibition, was studied in chronic experiments on cats. Spike responses 1–2 sec in duration in 69 (36.7%) of 182 neurons preceded by 400–800 msec the beginning of conditioned-reflex and voluntary intertrial movements. These advanced responses appeared 200 msec before the corresponding advance responses of motor cortical neurons. Fifty-eight neurons (30.9%) responded directly to acoustic stimulation with a latent period of 10–50 msec for 2–6 sec, 19 neurons (10.1%) generated double responses, linked with both the acoustic stimulus and subsequent conditioned-reflex movement, and 42 neurons (22.3%) did not respond to acoustic stimulation, although individual neurons of this group changed the level of their spontaneous activity in response to repeated conditioned stimulation, and this change was maintained for some tens of minutes. Extinction, differentiation, and conditioned inhibition all abolished conditioned-reflex movements, but each type of internal inhibition was accompanied by its own characteristic changes in the firing pattern of PGM neurons. Functional independence of neurons of the first and second groups was demonstrated during extinction and recovery of the conditioned-reflex. The results indicate the important role of PGM not only in the mechanism of the conditioned reflex, but also in the development of its internal inhibition.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 16, No. 3, pp. 403–419, May–June, 1984.     

Postsynaptic responses of lamina II neurons of the spinal cord of the cat to activation of primary afferent input

Development of a complex response evokedin vivo in the neurons of lamina II of the spinal cord gray matter in cats by single electrical stimulation of primary afferents was simulated using mathematical models of these neurons, including the electrically excitable soma and axon and passive equivalent nonuniform dendrite. The intracellular response consisted of an excitatory postsynaptic potential (EPSP) with an action potential (AP) followed by a two-component hyperpolarization determined by the afterprocesses of hyperpolarization. The fast early hyperpolarization component appeared at the threshold stimulation of the most fast-conducting fibers; with an increase in the stimulation intensity it became superimposed on a slow later component. The direction of the early component changed after the hyperpolarizing shift of the membrane potential by 10 to 20 mV with respect to the resting level of –60÷–70 mV. The later component was abolished but not reversed even by the 50-mV shifts (to the –120-mV level). Simulation experiments showed that observedin vivo hyperpolarization-induced modification of the complex response is determined principally by a local interaction of electrotonus with synatic processes and does not depend on the behavior of the usual potential-activated sodium and potassium conductances in the soma. Inhibitory chloride synapses located on the soma and close to it represent the main source of fast early hyperpolarization, while distal dendritic potassium synapses are responsible for its late phase.Neirofiziologiya/Neurophysiology, Vol. 26, No. 5, pp. 382–390, September–October, 1994.     

Response in tonic type neurons of the cat auditory cortex to tones of different frequencies and intensities

In experiments on immobilized cats, intra- and extracellular response in tonic type neurons to tones of differing frequencies and intensities were investigated, as well as the organizational pattern of receptive fields in these units. Tonic type neurons were encountered at different cortical layers, but mostly (93% of the total) were located at a depth of 1.0–2.2 mm. Minimum thresholds required for response in these neurons were on average 7.7 dB below that found in neurons generating a phasic reaction in response to a tone. "Tonic" differed from "phasic" neurons in their inferior frequency-discriminative ability, with a Q₁₀ value averaging 4.1±0.4 as against 9.1±0.7 in phasic neurons. Size of receptive fields in tonic neurons (as revealed by occurrence of spike response in these units) was 3.5 times that observed in phasic cells. Length of action potentials in the majority (80%) of tonic neurons was about one and a half times to twice that found in phasic units. Tonic neurons also displayed a high degree of sensitivity to changes in the duration and intensity of acoustic stimulation.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 21, No. 4, July–August, pp. 498–506, 1969.     

Response of caudate nucleus neurons to acoustic stimulation during functional blockade of the thalamic centrum medianum in the cat

The response of caudate nucleus neurons to acoustic stimulation (a click at 0.5 Hz) was investigated during chronic experimentation in cats using intracellular techniques and reversible blockage of the thalamic centrum medianum produced by anode polarization. Having analyzed poststimulus histograms it was found that the response of phasic activation to an acoustic signal decreased, and disappeared in 52% of neurons. A reduction in the level of spontaneous activity was also observed in neurons of the caudate nucleus. The significance of a direct pathway from the thalamic centrum medianum to the caudate nucleus is discussed from the viewpoint of acoustic signal transmission to caudate nucleus neurons.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 18, No. 1, pp. 92–99, January–February, 1986.     

Dynamics of cortical unit responses during defensive conditioning to sound

Several phases were distinguished in single-unit responses in areas 3 and 4 during defensive conditioning to acoustic stimulation: an initial response, short inhibition of the spike discharge, early and late after-discharges, and changes arising after the end of acoustic stimulation. The initial spike response appeared or intensified (if present already) in the first period of defensive conditioning parallel with an increase in spontaneous unit activity. After-discharges appeared later. The conditioned-reflex movement usually began 100–400 msec after stimulation began. This latent period of the first movement was the same whether for a real conditioned reflex or an after-discharge. Comparison of the latent periods of conditioned movements with the phases of the unit responses showed that the conditioned responses of the cortical neuron were primarily modified after-discharges of neurons evoked by a conditioned stimulus. Differential unit responses to acoustic stimulation, also based on after-discharges, were formed just as actively as positive. The basic role of reinforcement during conditioning is not to increase the excitability of the neurons, which is important in connection with their acquisition of polysensory properties, but to modify the after-discharges of the neurons.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 10, No. 4, pp. 339–347, July–August, 1978.     

Efferent activity in the phrenic nerve during startle reflex in chloralose anesthetized cats

Efferent activity was investigated in the phrenic nerve during startle reflex manifesting as somatic nerve discharges (lower intercostal nerves and the nerve endings) in chloralose anesthetized cats. Inhibition (usually of short duration, lasting 23–36 msec) of inspiration activity was found to be the main component of response in the phrenic nerve in the shaping of "low threshold" startle reflex produced by acoustic and tactile stimuli and stimulation of low threshold peripheral afferents. Reflex discharge prevailed amongst the response patterns produced in the phrenic nerve by stimulating high threshold afferents, i.e., early (propriospinal) and late (suprasegmental, arising from stimulating intercostal nerve) or late only (when stimulating the hindlimb nerves). Two patterns of late response could be distinguished, one on inspiration (found in roughly 3 out of 4 experiments) and other on exhalation — the respiratory homologs of somatic startle reflex. Response pattern is described throughout the respiratory cycle. Structure and respiratory modulation of reflex responses produced in the phrenic nerve by stimulating bulbar respiratory structure are also examined. Possible neurophysiological mechanisms underlying phrenic response during the shaping of startle reflex are discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 19, No. 4, pp. 473–482, July–August, 1987.     

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.     

Dynamics of single unit responses of the rabbit visual cortex to repetitive photic stimulation

Single unit responses in the visual cortex of the waking rabbit to repetitive photic stimulation at a frequency of once every 2.5 sec were studied. Depending on the total number of spikes in the response, the dynamics of the responses could be divided into two types: "fast" and "slow." From 5 to 15 stimuli were required to establish a stable level of response with changes of the first type, but 50 to 100 stimuli were needed for the response with changes of the second type. About 50% of all neurons did not change the characteristics of response. In the group of neurons with changing responses, partial habituation was found in 55–59% of cells; there were 25% of neurons with sensitization of discharge and 17–20% with a humpbacked type of response dynamics. A "slow" dynamics of unit responses in most cases was accompanied by changes in the duration of inhibitory pauses in the response; negative correlation of these values was observed in 65% of neurons.M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 10, No. 5, pp. 451–459, September–October, 1978.