Formation of "caudate spindles" in the rabbit sensomotor cortex during ontogeny

Electrical activity of the sensomotor and visual areas of the neocortex during stimulation of the caudate nucleus was recorded in young rabbits aged 3–60 days and in adults. Single stimulation of the caudate nucleus was found to cause the appearance of characteristic bursts of spindle-like rhythmic activity ("caudate spindles"), described previously in cats and monkeys, in the adult rabbit cortex. The latent period of the caudate spindle was about 200 msec, its duration 1–3 sec, and the frequency of its oscillations of the order of 12 Hz. Caudate spindles were most marked in the sensomotor cortex of the ipsilateral hemisphere. In rabbits under 10 days old caudate spindles were not found even if the intensity of stimulation was increased many times. Starting from the age of 15 days bursts of rhythmic activity resembling caudate spindles, but with lower frequency (about 8 Hz), longer latent period (up to 350 msec), and also with a higher threshold, appeared in the sensomotor cortex. The definitive type of caudate spindles was established toward the end of the first month of postnatal life, corresponding to the time of formation and complication of conditioned-reflex activity in developing animals.Brain Institute, Academy of Medical Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 17, No. 1, pp. 11–15, March, 1985.     

Unit responses of the cat caudate nucleus to cortical stimulation before and after destruction of nonspecific thalamic nuclei

Responses of caudate neurons to stimulation of the anterior sigmoid and various parts of the suprasylvian gyrus were studied in acute experiments on cats. The experiments consisted of two series: on animals with an intact thalamus and on animals after preliminary destruction of the nonspecific thalamic nuclei. Stimulation of all cortical areas tested in intact animals evoked complex multicomponent responses in caudate neurons with (or without) initial excitation, followed by a phase of inhibition and late activation. The latent periods of the initial responses to stimulation of all parts of the cortex were long and averaged 14.5–25.5 msec. Quantitative and qualitative differences were established in responses of the caudate neurons to stimulation of different parts of the cortex. Considerable convergence of cortical influences on neurons of the caudate nucleus was found. After destruction of the nonspecific thalamic nuclei all components of the complex response of the caudate neurons to cortical stimulation were preserved, and only the time course of late activation was modified.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 12, No. 5, pp. 464–471, September–October, 1980.     

Single unit responses of the reticular and ventral anterior nuclei of the thalamus to electrical stimulation of the centrum medianum

In acute experiments on cats anesthetized with thiopental (30–40 mg/kg, intraperitoneally) and immobilized with D-tubocurarine (1 mg/kg) responses of 145 neurons of the reticular and 158 neurons of the ventral anterior nuclei of the thalamus to electrical stimulation of the centrum medianum were investigated. An antidromic action potential appeared after a latent period of 0.3–2.0 msec in 4.1% of cells of the reticular nucleus and 4.4% of neurons of the ventral anterior nucleus tested in response to stimulation. The conduction velocity of antidromic excitation along axons of these neurons was 1.7–7.6 m/sec. Neurons responding with an antidromic action potential to stimulation both of the centrum medianum and of other formations were discovered, electrophysiological evidence of the ramification of such an axon. Altogether 53.8% of neurons of the reticular nucleus and 46.9% of neurons of the ventral anterior nucleus responded to stimulation of the centrum medianum by orthodromic excitation. Among neurons excited orthodromically two groups of cells were distinguished: The first group generated a discharge consisting of 6–12 action potentials with a frequency of 130–640 Hz (the duration of discharge did not exceed 60 msec), whereas the second responded with a single action potential. Inhibitory responses were observed in only 0.7% of neurons of the reticular nucleus and 4.4% of the ventral anterior nucleus tested. Afferent influences from the relay nuclei of the thalamus, lateral posterior nucleus, and motor cortex were shown to converge on neurons responding to stimulation of the centrum medianum.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 12, No. 1, pp. 36–45, January–February, 1980.     

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.     

Cortical and subcortical spreading depression in rats produced by focused ultrasound

Steady potential shifts produced by focused ultrasond were recorded in the cerebral cortex, hippocampus, thalamus, and caudate nucleus. Impulses of 50–100 msec duration were presented at a frequency of 5 and 10 Hz. Negative steady potential shifts were produced in each of the structures investigated, which gradually increased during rhythmic electrical reaction to reach –3 to –7 mV within 10–30 sec, often succeeded by a wave of spreading depression (SD). In each structure analyzed amplitude of SD waves measured 20–30 mV, lasting 30–40 sec in the cortex, the caudate nucleus and the thalamus, and 80–120 sec in the hippocampus. In unanesthetized and lightly anesthetized animals SD waves were on occasions the precursors of convulsive discharges forming under the action of focused ultrasound. Ultrasound at threshold doses proved ineffective for 5–7 min after the occurrence of an SD wave, but again evoked repeated SD waves once the refractory period had ended. Accordingly, local effects produced by focused ultrasound can result in functional blockage of the brain structures due to cortical and subcortical spreading depression.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Institute of Brain Research, All-Union Research Center of Mental Health, Academy of Medical Sciences of the USSR, Moscow. N. N. Andreev Acoustic Institute, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 18, No. 1, pp. 55–61, January–February, 1986.     

The neuronal organization of the dentate nucleus of the cerebellum

The intracellular activity of the neurons of the dentate nucleus was studied in cats anesthetized with Nembutal by means of their antidromic and synaptic excitation through stimulation of the red nucleus (RN) and the ventrolateral nucleus of the thalamus (VL), as well as the sensomotor cerebral cortex (CC) and the peripheral nerves of the posterior and anterior extremities. Several functionally delimited groups of neurons were isolated and studied. Efferent neurons, antidromically activated from nuclei of the brain stem, which did not react to stimulation of the peripheral nerves were placed in group I. Group II neurons were synaptically activated from the nuclei of the brain stem, and in a majority of cases also reacted to stimulation of the peripheral nerves and CC. Cells with a rhythmic background activity, which did not react to any of the types of stimulation used, comprised group III. Group IV was made up of neurons having the properties of intermediate neurons with a selective reaction to stimulation of a specific peripheral nerve or which experience broad convergence of the effects of different afferent impulsations.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 3, No. 2, pp. 154–165, March–April, 1971.     

Role of the caudate nucleus in the development of evoked synchronized activity

Synchronized activity (spindles, augmentation response) evoked by stimulation of thalamic nonspecific, association, and specific nuclei was investigated in chronic experiments on 11 cats before and after successive destruction of the caudate nuclei. After destruction of the caudate nuclei the duration of spindle activity in the frontal cortex and subcortical formations (thalamic nuclei, globus pallidus, putamen) was reduced to only three or four oscillations. In the subcortical nuclei its amplitude fell significantly (by 50±10%); in the cortex the decrease in amplitude was smaller and in some cases was not significant. Different changes were observed in the amplitude of the augmentation response, depending on where it was recorded. In the subcortical formations it was considerably and persistently reduced (by 50±10%); in the cortex these changes were unstable in character. Unilateral destruction of the caudate nucleus inhibited synchronized activity evoked by stimulation of the thalamic nuclei on the side of the operation only. Destruction of the basal ganglia (caudate nucleus, globus pallidus, entopeduncular nucleus, and putamen) did not prevent the appearance of synchronized activity; just as after isolated destruction of the caudate nucleus, after this operation synchronized activity was simply reduced in duration and amplitude. It is suggested that the caudate nucleus exerts an ipsilateral facilitatory influence on the nonspecific system of the thalamus during the development of evoked synchronized activity.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 9, No. 3, pp. 239–248, May–June, 1977.     

电刺激大鼠尾核头部镇痛的中枢效应—[^3H]—2脱氧...

Sokoloff's 2-deoxyglucose (2-DG) autoradiographic technique was used to identify changes of glucose metabolic rate in the rat brain following unilateral stimulation of the head of the caudate nucleus. The results were as follows. The local glucose metabolic rate after noxious stimulation was increased in the somatosensory cortex, cingulate cortex, ventroposterior and parafascicular nucleus of the thalamus, septal area, habenular nucleus, head of caudate nucleus, periaqueductal gray (PAG) and dorsal raphe nucleus (P < 0.05). After stimulating the head of the caudate nucleus, the local glucose metabolic rate of nucleus raphe magnus (rm) and nucleus paragigantocellularis (pgcl) was increased significantly and that of the PAG and dorsal raphe nucleus had a tendency to increase, while stimulation of the head of caudate nucleus could partially abolish the increased glucose metabolic rate in the somatosensory cortex, cingulate cortex, ventroposterior and parafascicular nucleus of the thalamus, septal area and habenular nucleus as induced by noxious stimulation. These results suggest that caudate stimulation is able to depress the activation of some brain structures related to nociception and to activate those related to antinociception. The pgcl, rm, PAG and dorsal raphe nucleus might be the key structures participating in the caudate stimulation produced analgesia.     

Afferent connections of the cat caudate nucleus studied by the retrograde axonal transport method

Sources of direct and indirect afferent connections of the caudate nucleus were investigated in cats by the retrograde axonal transport of horseradish peroxidase method. Different parts of the neocortex were shown to form different types of projections to the caudate nucleus; the sources of these projections have a laminar organization. Connections of the globus pallidus with the caudate nucleus, not previously described, were found. Among the sources of the thalamo-caudate projections, besides nuclei of the intralaminar complex, an important place is occupied by the ventral anterior and mediodorsal nuclei. After injection of horseradish peroxidase into the caudate nucleus, retrograde axonal transport of the enzyme was observed in the caudal direction, as far as cells of the locus coeruleus. ON the basis of these results a general scheme of afferent projections to the caudate nucleus is drawn up, including its connections with the spinal cord mediated by the thalamic nuclei and mesencephalic reticular formation.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 12, No. 2, pp. 146–154, March–April, 1980.     

Electrophysiological study of connections between the entorhinal cortex and the neocortex

In acute experiments in rabbits immobilized by d-tubocurarine, stimulation of the entorhinal area with rectangular electric impulses led to the appearance of evoked potentials (EP) with a latent period of 6–12 msec in the occipital, temporal, parietal, and cingular areas of the neocortex. The amplitude of the positive response component was 500 µV, and its duration 25–50 msec. The negative component was not always discernible. When rhythmic stimulation was used, these EPs followed stimulation frequencies not exceeding 20 per sec. Stimulation of the medial parts of the entorhinal area with a frequency of one to three per sec was accompanied by recruitment of the EP in the occipital and temporal neocortex areas. Nembutal depressed the amplitude of the neocortex EP appearing in response to stimulation of the entorhinal cortex. With the aid of double stimulation it could be established that, after conditioning stimulation of the entorhinal area, the positive component of the primary response (PR) evoked by stimulation of the contralateral sciatic nerve in the projection zone of the somatosensory cortex is strengthened during the first 50 msec, and subsequently after 80–120 msec. In these cases, the negative component was depressed. These findings are discussed with a view to the influence of limbic structures on the neocortex.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 2, No. 1, pp. 73–78, January–February, 1970.     

Preferential ipsilateral influence of the posterior hypothalamus on the neocortex

On the 10th–20th day after precollicular transection of the brain stem weak low-frequency electrical stimulation of the posterior hypothalamus preferentially activates the ipsilateral neocortex. After unilateral injury to the posterior hypothalamus, synchronous activity predominates in the ipsilateral neocortex. In premesencephalic animals weak single electrical stimulation of the posterior hypothalamus evokes the appearance of spindles in ipsilateral zones of the frontal cortex, whereas stronger single stimulation causes the diffuse generation of spindles in various parts of the neocortex. Besides this preferential unilateral effect, the influence of the posterior hypothalamus is found to be more strongly expressed in the frontal than in the occipital cortex. It is postulated that the posterior hypothalamus exerts its influence on the neocortex through the thalamic nuclei.Institute of Physiology, Academy of Sciences of the Georgian SSR, Tbilisi. Translated from Neirofiziologiya, Vol. 8, No. 2, pp. 139–145, March–April, 1976.     

INCORPORATION OF 14C FROM [U-14C]GLUCOSE INTO FREE AMINO ACIDS IN MOUSE BRAIN REGIONS UNDER CYANIDE INTOXICATION

—During anoxia induced by the administration of potassium cyanide, [U-¹⁴C]glucose was injected intraperitoneally into adult mice and they were decapitated at 5, 15 and 30 min after the injection. After freeze-drying in vacuo, differences in the uptake of radioactive carbon from [U-¹⁴C]glucose into free amino acids (glutamate + glutamine, aspartate + asparagine, GABA, alanine and glycine) in mouse cerebral neocortex, cerebellar hemisphere, caudate nucleus, thalamus, hypothalamus and medulla oblongata were investigated (by macroautoradiography and GLC separation) and compared with those obtained under normal conditions. (1) During anoxia, autoradiographical densities in the thalamus and medulla oblongata were higher than that in the cerebral neocortex and caudate nucleus. (2) Among specific radioactivities (d.p.m./μmol) of free amino acids, alanine gave the highest value during anoxia, except in the cerebellar hemisphere and hypothalamus at 5 min and the medulla oblongata at 30 min. (3) During anoxia, the specific radioactivities of alanine and glycine in each brain region did not significantly decrease at 15 and 30 min compared with those under normal conditions. During anoxia, the specific radioactivity of glutamate + glutamine in the cerebellar hemisphere and hypothalamus did not significantly decrease compared with the normal conditions, while that of GABA, aspartate + asparagine and glutamate + glutamine in the cerebral neocortex, caudate nucleus, thalamus and medulla oblongata showed an increase. (4) The percentage decrease of glutamate + glutamine and aspartate + asparagine at 5 and 15 min was highly significant in the cerebral neocortex and caudate nucleus.     

电刺激大鼠尾核头部镇痛的中枢效应——〔~3H〕-2脱氧葡萄糖放射自显影研究

本文利用[~3H]-2脱氧葡萄糖定量放射自显影方法,研究了电刺激大鼠尾核头部镇痛时中枢神经系统有关结构的葡萄糖代谢率变化。结果表明,痛刺激后,皮层躯体感觉Ⅰ,Ⅱ区、扣带回皮质、丘脑束旁核、丘脑中央中核、丘脑腹后核、尾核、外侧缰核、外侧隔核、中缝背核及中脑导水管周围灰质等结等的葡萄糖代谢率均明显升高(P<0.05)。电刺激大鼠尾核头部后,中缝大核及延髓旁巨细胞网状外侧核的葡萄糖代谢率显著升高,中脑导水管周围灰质和中缝背核的葡萄糖代谢率亦有升高趋势。电刺激大鼠尾核头部可部份降低痛刺激引起的有关结构葡萄糖代谢率升高(如皮层躯体感觉Ⅰ、Ⅱ区、扣带回皮质、丘脑束旁核、丘脑中央中核、丘脑腹后核、外侧隔核及外侧缰核等)。上述结果提示,电刺激大鼠尾核头部镇痛时抑制了与痛感觉有关的结构,同时激活了与镇痛有关的结构。中缝大核、中缝背核、中脑导水管周围灰质及延髓旁巨细胞网状外侧核等结构是实现尾核镇痛的重要环节。     

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.     

Effect of posterior hypothalamic stimulation on neocortical electrical activity in the chronic premesencephalic cat

Chronic experiments were carried out on 19 adult cats after preliminary division of the brain stem at the level of the anterior border of the superior colliculus. High-frequency electrical stimulation of the posterior hypothalamus of these animals caused desynchronization of electrical activity in different parts of the neocortex. The influence of the posterior hypothalamus was predominantly on activity in the frontal zones of the neocortex. It is postulated on the basis of these results that the posterior hypothalamus has an activating effect on the neocortex through the thalamic nuclei.Institute of Physiology, Academy of Sciences of the Georgian SSR, Tbilisi. Translated from Neirofiziologiya, Vol. 8, No. 1, pp. 47–53, January–February, 1976.     

Afferent connections of the ventral magnocellular section of the cat red nucleus

The location within the brain of labeled neurons giving rise to projections to the ventral magnocellular section of the red nucleus were investigated by means of microiontophoretically injected horseradish peroxidase. Projections were identified from many cortical, thalamic, and hypothalamic structures and from the head of the caudate nucleus, septum, globus pallidus, anterior commissure nucleus, central amygdalar nucleus, field of Forel, Zona incerta, and a number of brainstem structures. Findings in accordance with those found in the literature were obtained on projections to the red nucleus from the coronary and cruciate cortical sulci, the midbrain and dentate (lateral) cerebellar nuclei, subststantia nigra, nucleus gracilis, and the cuneate nucleus. Trajectories of retrogradely labeled fiber systems of the red nucleus are described.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 20, No. 5, pp. 680–687, September–October, 1988.     

Neurophysiological analysis of postnatal development of corticostriate connections in rabbits

Evoked potentials were recorded in the caudate nucleus of adult rabbits and young rabbits aged 2–30 days in response to stimulation of the ipsilateral motor cortex. The response of the caudate nucleus in the adult rabbit consisted of a positive-negative complex with latent period of 3–5 msec. Maximal amplitude of the response was observed in the dorsorostral region of the nucleus. As the recording electrode was inserted deeper, the amplitude of the response gradually decreased but without reversal of its polarity. Responses of the caudate nucleus to stimulation of the motor cortex were recorded as early as on the 3rd day after birth. These responses were indistinguishable in configuration from responses of the nucleus of adult rabbits. Their latent period was about 10 msec. Between the 16th and 20th day after birth the latent period of the response decreased considerably — from 9 to 5 msec, and by the 30th day of life it had reached its definitive value. With age the amplitude of the response increased but the threshold of stimulation decreased, The results indicate early functional maturation of connections of the motor cortex with the caudate nucleus and they agree with the results of morphological investigations of the structural development of the afferent systems of this nucleus.Brain Institute, Academy of Medical Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 14, No. 3, pp. 284–289, May–June, 1982.     

Efferent connections of the cat caudate nucleus studied by retrograde axonal transport of horseradish peroxidase

A well-developed descending efferent system of the caudate nucleus has been revealed by retrograde axonal transport of horseradish peroxidase. It consists of numerous projections into the thalamus. A topical differentiation of the connections between the caudate nucleus and the paleostriatum and substantia nigra was found. It was established that the main source of efferent connections of the caudate nucleus were small and medium-sized neurons. It was demonstrated that the subthalamic nucleus has a special role in the descending efferent system of the caudate nucleus. In addition to the direct connections into the caudate nucleus itself the subthalamic nucleus has direct connections with the main output structures of the caudate nucleus, the paleostriatum, and the substantia nigra. The concept that the descending and ascending connections are interlinked in the mammalian central nervous system is supported by the results of this investigation into the caudate nucleus.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 17, No. 4, pp. 509–517, July–August, 1985.     

Analysis of activity of neuron pairs in the cat caudate nucleus

Activity of neuron pairs in the caudate nucleus, derived simultaneously by a single microelectrode, was investigated in experiments on lightly anesthetized, immobilized cats. Strong temporal correlation was shown to be characteristic of the spontaneous activity of a neuron pair if grouped discharges were present in that activity. If, however, spontaneous activity was characterized by impulses randomly distributed in time, temporal correlation was observed in only 50% of cases, it was weaker, and it reflected excitatory and inhibitory interactions equally. In many cases negative correlation was observed in discharges of neurons within the time interval of 0–4 msec. Electrical stimulation of the various afferent inputs of the caudate nucleus not only did not cause correlation to appear in the discharges of the neurons but, on the contrary, it abolished correlation which existed for that same pair of neurons discharging spontaneously. Comparison of the results with data in the literature indicates that, by the character of interaction of its neurons, the caudate nucleus is one of the group of associative nuclei of the brain.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 10, No. 5, pp. 486–493, September–October, 1978.     

Electrophysiological analysis of the organization of somatosensory thalamic inputs in the frog

Potentials produced in the frog thalamus by electrical stimulation of the peripheral nerves were investigated by sink and current source-density analysis. Sinks, which are viewed as potential generation sites, were located in three regions: the cell-free zone of the ventral thalamus adjoining the ventrolateral nucleus, the ventromedial and ventrolateral nuclei, and the caudal section of the dorsal thalamus. Evoked activity was recorded in individual neurons in the area of the second and third of these sinks. The first sink failed to form after section of the dorsal tracks of the spinal cord, while the remaining two only appeared after a considerably extended latency. It is suggested that nuclei of the ventral and caudal sections of the dorsal thalamus receive somatic impulses through the systems connected with the dorsal as well as the ventrolateral columns of the spinal cord. The direct projections of the primordial nuclei of dorsal columns may be involved in afferentation the ventral thalamus.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 18, No. 5, pp. 687–695, September–October, 1986.