Receptive fields of auditory cortical neurons in the cat

Receptive fields of auditory cortical neurons were studied by electrical stimulation of nerve fibers in different parts of the cochlea in cats anesthetized with pentobarbital. The dimensions of the receptive fields were shown to depend on the topographic arrangement of the neuron in the auditory cortex. The more caudad the neuron on the cortical projection of the cochlea in the primary auditory cortex, the more extensive its receptive field. The receptive fields were narrowest in the basal turn of the cochlea and were symmetrical with respect to their center. It is suggested that the region of finest discrimination of acoustic stimuli in cats is located in the basal region of the cochlea, i.e., in that part of its receptor system which has the narrowest receptive field and is represented by significantly more (than the middle and apical regions of the cochlea) nerve cells in the primary auditory cortex [1].A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 13, No. 5, pp. 467–473, September–October, 1981.     

Cochleotopic organization of the primary auditory cortex in cats

The cochleotopic organization of the primary auditory cortex was studied by the evoked potentials method in cats anesthetized with pentobarbital. Two foci of maximal activity (dorsal and ventral) were found in the primary auditory cortex of 85% of animals during local electrical stimulation of different areas of the cochlea. Analysis of projection maps of the primary auditory cortex of the cats showed that different areas of the cochlea are presented in this region disproportionately. The basal portion projects to a larger cortical surface than the middle and apical portions together, evidence of inequality of representation of different parts of the receptor apparatus of the cochlea in the primary auditory area. Considerable differences were observed in the arrangement of projections of the cochlea in the primary auditory cortex of different animals.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 11, No. 2, pp. 117–124, March–April, 1979.     

Functional properties distinguishing individual auditory cortical neurons

Responses of 246 auditory cortical neurons to paired and repetitive stimulation of geniculo-cortical fibers were studied in experiments on cats immobilized with tubocurarine. The refractory period (RP) varied from 1 to 200 msec in different neurons. For neurons excited antidromically it varied from 1 to 3 msec. Among neurons excited monosynaptically there were some with a short (1.3–6 msec), medium, (8–16 msec) or long (30–100 msec) refractory period. Most neurons excited polysynaptically had a RP of mean length. RPs 30–200 msec in length were due to inhibition arising in the neuron after conditioning stimulation. In some neurons, after a short (1.5–2.0 msec) initial period of refractoriness there was a temporary (for 2–3 msec) recovery of responsiveness, followed by another period of ineffectiveness of the testing stimulus lasting 30–100 msec. Barbiturates selectively inhibited long-latency unit responses in the auditory cortex and during their action the number of responding neurons with a mean RP decreased sharply. The results demonstrate functional heterogeneity of auditory cortical neurons responding to an incoming volley of afferent impulses.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 5, No. 3, pp. 236–245, May–June, 1973.     

Unit responses in the cat auditory cortex to medial geniculate body stimulation

Responses of 98 auditory cortical neurons to electrical stimulation of the medial geniculate body (MGB) were recorded (45 extracellulary, 53 intracellularly) in experiments on cats immobilized with tubocurarine. Responses of the same neurons to clicks were recorded for comparison. Of the total number of neurons, 75 (76%) responded both to MGB stimulation and to clicks, and 23 (24%) to MGB stimulation only. The latent period of extracellularly recorded action potentials of auditory cortical neurons in response to clicks varied from 7 to 28 msec (late responses were disregarded), and that to MGB stimulation varied from 1.5 to 12.5 msec. For EPSPs these values were 8–13 and 1–4 msec respectively. The latent period of IPSPs arising in response to MGB stimulation varied from 2.2 to 6.5 msec; for 34% of neurons it did not exceed 3 msec. The difference between the latent periods of responses to clicks and to MGB stimulation varied for different neurons from 6 to 21 msec. Responses of 11% of neurons to MGB stimulation, recorded intracellularly, consisted of sub-threshold EPSPs, while responses of 23% of neurons began with an EPSP which was either followed by an action potential and subsequent IPSP or was at once cut off by an IPSP; 66% of neurons responded with primary IPSPs. Neurons responding to MGB stimulation by primary IPSPs are distributed irregularly in the depth of the cortex: there are very few in layers III and IV and many more at a depth of 1.6–2 mm. Conversely, excited neurons are predominant in layer III and IV, and they are few in number at a depth of 1.6–2 mm. It is concluded that the afferent volley reaching the auditory cortex induces excitation of some neurons therein and, at the same time, by the principle of reciprocity, induces inhibition of others. This afferent inhibition takes place with the participation of inhibitory interneurons, and in some cells the inhibition is recurrent. The existence of reciprocal relationships between neurons in different layers of the auditory cortex is postulated.A. A. Bogomolets' Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 4, No. 1, pp. 23–31, January–February, 1972.     

Cochleotopic organization of the second auditory cortical area in the cat

The cochleotopic organization of the second auditory cortical area was investigated in cats anesthetized with pentobarbital by the evoked potentials method. Two independent representations of the cochlea were shown to exist in area AII: One in the dorsocaudal portion, the other in its ventrorostral portion. These projections of the cochlea differ in size and in the order of representation of its different parts. The dorsocaudal part of the auditory projection area of the cochlea, which extends over a distance of 2.6–2.8 mm from the center of the basal to the center of the apical focus, is arc-shaped. The order of arrangement of projections of different parts of the cochlea in this region of the auditory cortex coincides with that in the first auditory area, whereas the projection of the cochlea in the ventrorostral part of area AII, the length of which is 1.4–1.6 mm, has the opposite order of representation. The localization of projections of the cochlea in different cats shows considerable variability not only as regards anatomical topography of the auditory cortex, but also from one animal to another. The basal region of the cochlea was shown to project to a larger area of the cortex than the middle and apical portions taken together. It is suggested that the basal turn of the cochlea is functionally the most important for perception and primary analysis of auditory information.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 12, No. 1, pp. 18–27, January–February, 1980.     

Mechanisms of the effect of diazepam on paroxysmal electrical activity of an isolated cortical slab in cats

The effect of diazepam on paroxysmal global electrical activity of a neuronally isolated slab of auditory cortex and on inhibitory responses of its neurons due to intracortical electrical stimulation was investigated in cats. Diazepam (2 mg/kg, intravenously) caused inhibition of paroxysmal electrical activity and increased the number of inhibited neurons in both the acutely isolated slab and three weeks after isolation, compared with the intact cortex. However, the number of disynaptic responses was reduced under these circumstances, especially in the long-isolated slab. It is postulated that diazepam exerts its action through activation of GABA-ergic inhibitory neurons, by synchronizing inhibition and increasing the duration of the IPSPs. The action of diazepam is manifested first, probably, in the initial links of cortical neuron chains.I. I. Mechnikov Odessa State University. Translated from Neirofiziologiya, Vol. 17, No. 1, pp. 3–10, January–February, 1985.     

Neuronal and synaptic mechanisms of cortical inhibition

The latent periods, amplitude, and duration of IPSPs arising in neurons in different parts of the cat cortex in response to afferent stimuli, stimulation of thalamocortical fibers, and intracortical microstimulation are described. The duration of IPSPs evoked in cortical neurons in response to single afferent stimuli varied from 20 to 250 msec (most common frequency 30–60 msec). During intracortical microstimulation of the auditory cortex, IPSPs with a duration of 5–10 msec also appeared. Barbiturates and chloralose increased the duration of the IPSPs to 300–500 msec. The latent period of 73% of IPSPs arising in auditory cortical neurons in response to stimulation of thalamocortical fibers was 1.2 msec longer than the latent period of monosynaptic EPSPs evoked in the same way. It is concluded from these data that inhibition arising in most neurons of cortical projection areas as a result of the arrival of corresponding afferent impulsation is direct afferent inhibition involving the participation of cortical inhibitory interneurons. A mechanism of recurrent inhibition takes part in the development of inhibition in a certain proportion of neurons. IPSPs arise monosynaptically in 2% of cells. A study of responses of cortical neurons to intracortical microstimulation showed that synaptic delay of IPSPs in these cells is 0.3–0.4 msec. The length of axons of inhibitory neurons in layer IV of the auditory cortex reaches 1.5 mm. The velocity of spread of excitation along these axons is 1.6–2.8 msec (mean 2.2 msec).A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 16, No. 3, pp. 394–403, May–June, 1984.     

Neuronal responses of a chronically isolated slab of auditory cortex to intracortical stimulation during paroxysmal electrical activity in cats

Responses of 155 neurons 3 weeks after neuronal isolation of a slab of auditory cortex (area AI) to single intracortical stimulating pulses at the level of layer IV were studied in unanesthetized, curarized cats during paroxysmal electrical activity evoked by series of high-frequency (10–20 Hz) electrical stimulation by a current 2–5 times above threshold for the direct cortical response. In response to such stimulation a discharge of paroxysmal electrical activity, lasting from a few seconds to tens of seconds, appeared in the slab. As a rule it consisted of two phases — tonic and clonic. This indicates that cortical neurons can form both phases of paroxysmal cortical activity. Depending on behavior of the neurons during paroxysmal electrical activity and preservation of their ability to respond to intracortical stimulation at this time, all cells tested in the isolated slab were divided into four groups. Their distribution layer by layer and by duration of latent periods was studied. Two-thirds of the neurons tested were shown to generate spike activity during paroxysmal discharges whereas the rest exhibited no such activity. A special role of neurons in layer II in generation of paroxysmal activity in the isolated slab was noted. The view is expressed that at each moment functional neuronal circuits, independent of each other, exist in the slab and also, evidently in the intact cortex, which can interact with one another when conditions change.I. I. Mechnikov Odessa State University. Translated from Neirofiziologiya, Vol. 16, No. 1, pp. 3–11, January–February, 1984.     

Single unit activity related to acoustic stimulus meaning in the cat auditory cortex during instrumental food reflex

Responses of 93 neurons to isolated presentation of a single click and a series of 10 clicks with following frequency of 1000 Hz and responses of 66 neurons after the click had become a positive conditioned stimulus, and a series of 10 clicks had become a differential, negative stimulus, were investigated in chronic experiments on cats. Formation and realization of differential inhibition of an instrumental food reflex was shown not to lead to strengthening of inhibition in the auditory cortex, and the process of differential inhibition itself within the primary auditory cortex is not essentially an inhibitory process. Identical changes were found in responses of auditory cortical neurons to both positive and negative conditioned stimuli after training, evidence that neurons of the primary auditory cortex play a similar role in realization of the instrumental reflex and in its differential inhibition. It is suggested that the presence of groups of neurons responding by excitation or inhibition only to presentation of a stimulus with definite informative value is of great importance for differentiation.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukranian SSR, Kiev. Translated from Neirofiziologiya, Vol. 17, No. 2, pp. 212–221, March–April, 1985.     

Investigation of habituation reaction of neurons of the cat motor cortex

The neuronal and total surface activity of the cortical representation of the motor analyzer in the region of the posterior sigmoid gyrus of the cat brain in response to rhythmical light, sound, and electrical stimuli and their complexes was analyzed. Two groups of neurons were found, of which the first is characterized by a gradual decrease in the number of peaks in the response and by their subsequent disappearance and the second by the absence of a discharge in response to stimulation and by its development before the application of the next stimulus. The first group was comprised of neurons which do not have background activity and the second was made up of neurons with a background activity of 0.4–3.7 imp/sec. This reorganization of the activity of cortical neurons in response to rhythmical stimulation is considered to be a habituation phenomenon.A. A. Zhdanov Leningrad State University. Translated from Neirofiziologiya, Vol. 3, No. 3, pp. 245–251, May–June, 1971.     

Monosynaptic inhibitory postsynaptic potentials of cortical neurons

Monopolar intracortical stimulation of the auditory cortex was carried out in cats immobilized with D-tubocurarine. A macroelectrode (tip diameter 100 µ) or a microelectrode (tip diameter 10–15 µ) was used for stimulation. In both cases, besides excitatory responses, primary IPSPs with latent periods of 0.4–1.2 and 1.4–6.0 msec were recorded in cortical neurons close to the point of stimulation. The first group of IPSPs are considered to be generated in response to direct stimulation of bodies or axons of inhibitory cortical neurons, i.e., monosynaptically. The amplitude of these IPSPs varied in different neurons from 3 to 15 mV, and their duration from 4 to 150 msec. Additional later inhibitory responses were superposed on many of them. Of the IPSPs generated in auditory cortical neurons in response to stimulation of geniculocortical fibers 1.5% had a latency of 0.8–1.3 msec. They also are assumed to be monosynaptic. It is concluded that the duration of synaptic delay of IPSPs in cortical neurons and spinal motoneurons is the same, namely 0.3–0.4 msec. Axons of auditory cortical inhibitory neurons may be 1.5 mm long. The velocity of impulse conduction along these axons is 1.6–2.8 m/sec. The genesis of some special features of IPSPs of cortical neurons is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 7, No. 5, pp. 458–467, September–October, 1975.     

Unit responses in the auditory cortex to stimulation of geniculocortical fibers

Extracellular and intracellular single unit responses of neurons of the auditory cortex to electrical stimulation of geniculocortical fibers (GCF) were recorded in experiments on cats immobilized with tubocurarine. The latent period of responses of 15% of neurons to GCF stimulation was 0.3–1.5 msec. It is postulated that they were excited anti-dromically. The latent period of spikes generated by neurons responding to GCF stimulation orthodromically varied from 1.6 to 12 msec. In 28.6% of neurons the latent period was 1.6–2.5 msec. It is postulated that these neurons were excited monosynaptically. Intracellular recording revealed primary IPSPs in response to GCF stimulation in 63.3% of neurons, a brief EPSP followed by a prolonged IPSP in 17.7%, an EPSP-spike-IPSP complex in 12.3%, and subthreshold EPSPs in 7% of neurons. The latent period of the primary IPSPs varied from 1.8 to 11 msec, being 1.8–3.7 in 72%, 3.8–5.7 in 20.0%, and 5.8–11 msec in 8.0% of neurons. The latent period of responses beginning with an EPSP was 1–4 msec (mean 1.8 msec). Orthodromic responses arising 3–10 msec after the antidromic response, and consisting of 3–5 spikes, were recorded in some antidromically excited neurons. Hypotheses regarding the functional organization of the auditory cortex and mechanisms of inhibition in its neurons are put forward on the basis of the results obtained.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 4, No. 3, pp. 227–235, May–June, 1972.     

Responses of cat auditory cortical neurons to tones of different frequencies and electrical stimulation of corresponding regions of the cochlea

Characteristic frequencies of neurons in the cat auditory cortex (area AI) whose receptive fields are located in different parts of the basilar membrane of the cochlea were determined in cats anesthetized with pentobarbital. The higher the characteristic frequency of a neuron in area AI, the nearer its receptive field lies to the base of the cochlea. Receptive fields of neurons with a characteristic frequency higher than 4 kHz lie on the first 10 mm of the basilar membrane. Receptive fields of neurons with a characteristic frequency below 4 kHz lie on the remaining 11–12 mm of the membrane. The effect of electrical stimulation of the center of the receptive field of a neuron corresponds to its response to a tone of characteristic frequency. The more the frequency of the acting tone differs from the characteristic frequency, or the further the point of stimulation from the center of the receptive field of the neuron, the less likely is the neuron to respond with an action potential. Neurons with a low characteristic frequency have wider receptive fields than neurons with a high characteristic frequency. Receptive fields of neurons with close characteristic frequencies on the basilar membrane overlap considerably. It was shown by the method of paired stimulation that excitation evoked in neurons in area AI by the action of a tone of a particular frequency is followed by long-lasting inhibition. This inhibition lasts longest and is most effective if a tone of the characteristic frequency is used.     

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.     

Characteristics of inhibition in receptive fields of the cat visual cortex

Unit responses of neurons of zone 17 in the cat striate cortex to stripes of different widths were studied. Changes in the number of spikes during different time intervals (cuts) from the beginning of the response were analyzed in relation to stimulus area. Comparison of the results with those obtained by the study of receptive fields of the lateral geniculate body showed a significant difference in the dynamics of inhibition between cortical and geniculate receptive fields. Similar results were obtained when cortical unit responses to simultaneous and consecutive appearance of two stripes in the receptive field, one in the excitatory zone and the other at the inhibitory periphery, were studied. Evidence of the longer duration of cortical inhibition also was obtained by the same technique. When both stripes were placed in the excitatory center of the field another feature of cortical inhibition was revealed: its dependence on the order of stimulus application. If the order of stimulus application coincided with the optimal direction of movement of the stripe for the given field, the unit response to the next stimulus was strongly facilitated by the action of the stimulus applied previously. Application of stimuli in the opposite order invoked inhibition. The sensitivity of inhibition to the order of stimulus application was observed in the center of the field; it diminished toward the periphery, where application of the stimuli in any order evokes inhibition of the response.Medical Academy, Sofia, Bulgaria, I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 9, No. 4, pp. 339–346, July–August, 1977.     

Interaction between responses evoked by acoustic and somatosensory stimuli in neurons of the magnocellular part of the medial geniculate body

Interaction between responses to acoustic clicks and to electrodermal stimulation of the contralateral forelimb was investigated in 78 neurons in the magnocellular part of the medial geniculate body of curarized cats. Of this number, 33 neurons responded by discharges both to clicks and to electrodermal stimulation, 25 responded to clicks only, and 20 to electrodermal stimulation only, or to stimulation of the dorsal funiculus of the spinal cord. Conditioning stimulation evoked inhibition of the response to the testing stimulus in 32 of 33 neurons responding by spike discharges to both clicks and electrodermal stimulation. Electrodermal stimulation inhibited responses to clicks in all the neurons tested which responded only to clicks, whereas clicks evoked inhibition of responses to electrodermal stimulation (or to stimulation of the dorsal funiculus) in only four of the 20 neurons which responded to these types of stimulation only. It is suggested that inhibition of excitability arising in neurons of the magnocellular part of the medial geniculate body during interaction between auditory and somatosensory afferent volleys is based on postsynaptic inhibition.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 12, No. 4, pp. 368–374, July–August, 1980.     

电刺激大马蹄蝠听皮层对下丘神经元听觉敏感性的影响

实验在１２只大马蹄蝠上进行。用常规电生理学方法研究了电刺激听皮层对下丘２１２个神经元的听反应的影响,结果表明,有３２个神经元的听反应被抑制,１９个神经元的听反应褐易化。     

Responses of somatic cortical neurons during the conditioned placing reflex in cats

Unit activity was studied in areas 3 and 4 during the conditioned placing reflex in cats. Responses of somatic cortical neurons in this case were shown to develop comparatively late — 80–100 or, more often, 200–450 msec after the conditioned stimulus. In the motor cortex responses preceded movement by 50–550 msec, whereas in the somatosensory cortex they usually began simultaneously with or after the beginning of the movement. Judging from responses of somatic cortical neurons, the placing reflex is realized by the same neuronal mechanism as the corresponding voluntary movement. The differential stimulus and positive conditioned stimulus, after extinction of the conditioned placing reflex, evoked short-latency spike responses lasting 250–350 msec in the same neurons as took part in the reflex itself. In these types of internal inhibition, responses of the neurons were thus initially excitatory in character. Participation of the neurons in the conditioned placing reflex and its extinction, disinhibition, and differentiation, is the result of a change in the time course of excitatory processes and is evidently connected with differential changes in the efficiency of the various synaptic inputs of the neuron.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 14, No. 4, pp. 392–401, July–August, 1982.     

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.     

Electrophysiological investigation of medullary neurons connected with lateral line organs of the catfish (Ictalurus nebulosus)

Unit responses in the acoustic-lateral region of the medulla to electrical and mechanical stimulation of the lateral line organs were investigated in acute experiments on curarized catfish. Of the total number of neurons 70% possessed spontaneous activity. An electrical stimulus evoked a tonic response both in spontaneously active and in "silent" cells. Three main types of firing pattern of the neurons were distinguished: fast-adapting, slow-adapting, and grouped. As regards the relation of the neurons to polarity of the stimulus they were subdivided into two groups. The thresholds of unit responses to electrical stimulation varied considerably: from 2.5·10^–9 to 5·10^–12 A/mm². The effect of intensity of the electrical stimulation on unit responses in the medulla is analyzed. The precise dependence of on- and off-responses of each neuron on stimulus intensity of any polarity was determined. The neurons were shown to be sensitive to both electrical and mechanical stimuli. It is postulated that this phenomenon is due to convergence of impulses from electrical and mechanical receptors of the lateral line on the neurons. The properties of the central neurons are compared with those of the peripheral electroreceptor system in catfish.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 5, No. 2, pp. 156–163, March–April, 1973.