Neuronal inhibitory responses to intracortical stimulation of an isolated slab of cat auditory cortex

Neuronal responses of an isolated slab of cortex to intracortical stimulation were studied intracellularly. The predominant responses were primary IPSPs. Their latent periods did not exceed 10 msec. Within the volume of cortex studied, neurons inhibited in response to stimulation were most numerous in the upper layers (II, III). Predominance of disynaptic IPSPs is evidence of the important role of cortical interneurons in their genesis. It is concluded from the results that primary IPSPs limit the spread of excitation primarily in the activated area of cortex. Since involvement of neurons of the isolated slab in the inhibition process takes place for only 10 msec after stimulation, neurons giving spike responses to intracortical stimulation with a longer latent period can transmit information into other brain zones. The role of duration of IPSP in the dynamics of interneuronal interaction in the cerebral cortex is discussed.I. I. Mechnikov Odessa State University. Translated from Neirofiziologiya, Vol. 16, No. 1. pp. 42–49, January–February, 1984.     

Neuronal responses of an isolated slab of cat auditory cortex to intracortical stimulation

Neuronal responses of an isolated slab of auditory cortex (area AI) to intracortical stimulation at the level IV were studied in curarized cats by extracellular recording 3 weeks after isolation. Dispersion of response latencies in the isolated slab was reduced (compared with that observed soon after isolation); the predominant responses were mono- and disynaptic, and the number of discharges consisting of bursts of spikes increased. However, despite simplification of the structural and functional organization of the chronically isolated slab of auditory cortex, the conditions for complex polysynaptic interaction between neurons of all layers were preserved in it, and in each layer the character of such interaction depended on the distance of the neuron from the focus of origin of the excitation. [In the chronically isolated slab of auditory cortex, just as in the acutely isolated slab, late reponses of over 40 msec were absent.]I. I. Mechnikov Odessa State University. Translated from Neirofiziologiya, Vol. 14, No. 5, pp. 462–469, September–October, 1982.     

Primary neuronal inhibitory responses of a chronically isolated slab of auditory cortex to intracortical stimulation in cats

Responses of 579 neurons in a slab of cerebral cortex (3 weeks after its isolation) to intracortical stimulation, with a distance of 0.5, 1.0, and 2.0 mm between recording and stimulating electrodes, were tested intercellularly and histological changes in a similar slab were studied in experiments on cats. Primary IPSPs were shown to develop in the chronically isolated slab in a much larger number of neurons than in the acutely isolated slab. Latent periods of IPSPs in all series of experiments did not exceed 10 msec, and most IPSPs were mono- and disynaptic. The amplitude and duration of the IPSPs were similar to those observed in the acutely isolated slab and intact auditory cortex in cats. It is concluded that local intracortical neuronal chains along which impulses evoking an inhibitory effect in the terminal neuron of the chain are transmitted are of relatively short length. Such chains may participate in local processing of incoming information. Analysis of the distribution of neurons responding by primary IPSPs by duration of their latent periods and depth in the slab in each series of experiments revealed a spatial and temporal mosaic of inhibitory responses in the chronically isolated slab of auditory cortex and showed that this mosaic is due to intracortical mechanisms.I. I. Mechnikov Odessa State University. Translated from Neirofiziologiya, Vol. 16, No. 2, pp. 152–161, March–April 1984.     

Effect of pentobarbital on neuronal inhibitory response evoked in an isolated slab of cat auditory cortex by intracortical stimulation

Neuronal responses of an acutely isolated slab of auditory cortex (area AI) to intracortical electrical stimulation were studied intracellularly in cats anesthetized with pentobarbital. It was found that 77% of responses were primary IPSPs, and allowing for secondary inhibitory responses, an inhibitory response was observed in 92% of neurons. All types of neuronal responses in the slab were short-latency. The maximal response latency did not exceed 5 msec. Neurons responding to stimulation by IPSPs were found at all depths in the slab, with a maximum in layers II–III. Nearly all primary IPSPswere mono- and disynaptic. Pentobarbital increased the duration of individual neuronal inhibitory responses in the isolated slab of auditory cortex without affecting maximal duration of the IPSP. The mechanisms of the effect of pentobarbital on the amplitude and duration of IPSPs are discussed.I. I. Mechnikov Odessa State University. Translated from Neirofiziologiya, Vol. 16, No. 2, pp. 147–152, March–April, 1984.     

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.     

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.     

Somatosensory cortical unit responses in the waking cat to afferent stimuli

Extracellular and intracellular responses of 183 neurons in the primary projection area of the somatosensory cortex to electrical and tactile stimulation of the skin on the contralateral fore limb and to stimulation of the ventro-posterolateral thalamic nucleus of the ipsilateral hemisphere were studied in chronic experiments on cats. Spike responses to afferent stimuli are subdivided into three types: initial with a latent period of under 60 msec; initial followed by late responses with a latent period of over 60 msec; late with a latent period of over 60 msec. In addition another group of neurons responding to peripheral stimuli in the interval between the initial and the late response was identified. In nearly all cases the initial responses to peripheral stimulation had the form of a series of spikes, unlike responses to thalamic stimulation. It is concluded from the durations of the latent periods of these responses that about 70% of neurons in the primary projection area are activated mono- and disynaptically in response to peripheral stimulation; consequently, the intracortical spread of excitation in this zone is restricted.     

Unit responses of the first and second somatosensory cortical areas to peripheral,thalamic, and cortical stimulation

Unit responses of the first (SI) somatosensory area of the cortex to stimulation of the second somatosensory area (SII), the ventral posterior thalamic nucleus, and the contralateral forelimb, and also unit responses in SII evoked by stimulation of SI, the ventral posterior thalamic nucleus, and the contralateral forelimb were investigated in experiments on cats immobilized with D-tubocurarine or Myo-Relaxin (succinylcholine). The results showed a substantially higher percentage of neurons in SII than in SI which responded to an afferent stimulus by excitation brought about through two or more synaptic relays in the cortex. In response to cortical stimulation antidromic and orthodromic responses appeared in SI and SII neurons, confirming the presence of two-way cortico-cortical connections. In both SI and SII intracellular recording revealed in most cases PSPs of similar character and intensity, evoked by stimulation of the cortex and nucleus in the same neuron. Latent periods of orthodromic spike responses to stimulation of nucleus and cortex in 50.5% of SI neurons and 37.1% of SII neurons differed by less than 1.0 msec. In 19.6% of SI and 41.4% of SII neurons the latent period of response to cortical stimulation was 1.6–4.7 msec shorter than the latent period of the response evoked in the same neuron by stimulation of the nucleus. It is concluded from these results that impulses from SI play an important role in the afferent activation of SII neurons.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 8, No. 4, pp. 351–357, July–August, 1976.     

Analysis of evoked potentials of the cerebellar cortex

Evoked potentials (EP) of the cerebellar cortex in response to stimulation of peripheral nerves are characterized by a two-phase positive-negative oscillation of the potential having a latent period of 10–25 msec. The electropositive phase can contain up to three components. The latent period of component I comprises 3–9 msec. The latent period and amplitude of this component are distinguished by considerable stability, which indicates the predominant significance of presynaptic processes in its formation. The sign of component II changes at a depth of 500 µ (and more), which corresponds to the position of the granular cell layer. At this level there arises in the neurons a response with a latent period of 4–10 msec in the form of a group (3–10) of impulses with a frequency of up to 200 per sec. It is concluded that the granular cells participate in the formation of component II and partially participate in the formation of components I and III of the EP. Responses to stimulation of the nerves appear synchronously with the EP in 24% of responding Purkinje cells; they fall on the maximum electropositive deviation or component III of the EP. Microinjections of 1% strychnine into the cerebellar cortex cause an increase of EP amplitude; impulse activity of the neurons is intensified. This indicates participation of postsynaptic processes in the formation of EP. No shifts in the EP of the cerebellar cortex were observed after intracortical injection of 0.1% atropine.N. I. Pirogov Vinnitsa Medical Institute. Translated from Neirofiziologiya, Vol. 2, No. 4, pp. 429–433, July–August, 1970.     

Transformation of the afferent tactile signal into a motor command in the cat motor cortex

Activity of 112 neurons of the precruciate motor cortex in cats was studied during a forelimb placing reaction to tactile stimulation of its distal parts. The latent period of response of the limb to tactile stimulation was: for flexors of the elbow (biceps brachii) 30–40 msec, for the earliest reponses of cortical motor neurons about 20 msec. The biceps response was observed 5–10 msec after the end of stimulation of the cortex with a series of pulses lasting 25 msec. Two types of excitatory responses of the neurons were identified: responses of sensory type observed to each tactile stimulation of the limb and independent of the presence or absence of motion, and responses of motor type, which developed parallel with the motor response of the limb and were not observed in the absence of motion. The minimal latent period of the responses of motor type was equal to the latent period of the sensory responses to tactile stimulation (20±10 msec). Stimulation of the cortex through the recording microelectrode at the site of derivation of unit activity, which increased during active flexion of the forelimb at the elbow (11 stimuli at intervals of 2.5 msec, current not exceeding 25 µA), in 70% of cases evoked an electrical response in the flexor muscle of the elbow.M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 9, No. 2, pp. 115–123, March–April, 1977.     

Unit responses of the pericruciate cortex to stimulation of the medial geniculate body

Responses of 239 neurons of the pericruciate cortex to stimulation of the medial geniculate body and pyramidal tract were investigated (189 extracellularly, 50 intracellularly) in cats anesthetized with thiopental and immobilized with D-tubocurarine. In response to stimulation of the medial geniculate body, the mean spontaneous firing rate of 63.6% of neurons in the pericruciate cortex increased by 10–25%, in 23.6% of neurons it decreased within the same limits, and mixed effects were observed in 5.5% of neurons. Phasic responses to single stimulation of the medial geniculate body were observed in 20% of neurons of the pericruciate cortex. Responses with a latent period of 0.3–1.0 msec (16%) were classed as antidromic, those with a latent period of 1.5–2.0 msec (20%) as orthodromic, monosynaptic, and those with a latent period of 2.5–4.0 msec or more (64%) as polysynaptic. With intracellular recording, excitatory responses of the EPSP, EPSP-AP, and AP type with latent periods of between 1.3 and 19.5 msec developed in 78.2% of cells. IPSPs, which were recorded in 21.8% of neurons, were usually found as components of mixed responses; primary IPSPs were found in only two cases. Monosynaptic connection of the medial geniculate body was shown to take place with neurons of the pericruciate cortex that did not belong to the pyramidal tract.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 11, No. 1, pp. 18–24, January–February, 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.     

Unit responses in area 5b of the suprasylvian gyrus to stimulation of the posterior lateral thalmamic nucleus

In response to stimulation of the posterior lateral nucleus in unanesthetized cats immobilized with D-tubocurarine an evoked potential consisting of three components with a latent period of 3–5 msec appeared in area 5b of the suprasylvian gyrus. All three components were reversed at about the same depth in the cortex (1500–1600 µ). Reversal of the potential shows that it is generated in that area by neurons evidently located in deeper layers of the cortex and is not conducted to it physically from other regions. Responses of 53 spontaneously active neurons in the same area of the cortex to stimulation of the posterior lateral nucleus were investigated. A characteristic feature of these reponses was that inhibition occurred nearly all of them. In 22 neurons the responses began with inhibition, which lasted from 30 to 400 msec. In 30 neurons inhibition appeared immediately after excitation while one neuron responded by excitation alone. The latent periods of the excitatory responses varied from 3 to 28 msec. The short latent period of the evoked potentials and of some single units responses (3–6 msec) confirms morphological evidence of direct connections between the posterior lateral nucleus and area 5b of the suprasylvian gyrus. Repetitive stimulation of that nucleus led to strengthening of both excitation and inhibition. Influences of the posterior lateral nucleus were opposite to those of the specific nuclei: the posterior ventrolateral nucleus and the lateral and medial geniculate bodies. Stimulation of the nonspecific reticular nucleus, however, evoked discharges from neurons like those produced by stimulation of the posterior lateral nucleus.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 5, No. 5, pp. 502–509, September–October, 1973.     

Spread of excitation in upper layers of the cat auditory cortex with participation of intracortical interneuronal connections

In chronically isolated slabs of the cat auditory cortex with additional transection of lower layers and preservation of the structural integrity of one, two, or three upper layers of cortex just under the pial membrane, impulse responses of slab neurons to stimulation applied at the additionally undercut section were studied. High effectiveness of axodendritic and axospinal excitatory contacts formed by nerve elements of intracortical origin in upper cortical layers was demonstrated. The participation of geniculocortical fibers in spread of excitation in the cortex through synaptic contacts in layer I with dendrites of underlying-layer pyramidal neurons is discussed. The capacity for generation of polysynaptic excitation responses by the neurons indicates preservation of complex interneuronal interactions in the isolated cortex slab preparations with their undercut lower layers.I. I. Mechnikov State University of Odessa, Odessa. Translated from Neirofiziologiya, Vol. 23, No. 1, pp. 80–87, January–February, 1991.     

Conduction of excitation in the cat visual system

Unit responses in area 17 of the visual cortex to stimulation of the lateral geniculate body and optic tract were studied in experiments on unanesthetized cats immobilized with D-tubocurarine. Of the neurons tested, 53.6% responded to stimulation of the lateral geniculate body. In 92% of these cells the responses were orthodromic with latent periods of between 2 and 12.5 msec. Most cells responded with latent periods of 2.0–2.5, 3.0–3.5, and 4.0–4.5 msec, corresponding to latent periods of the components of the electropositive wave of the primary response. Antidromic responses to stimulation of the lateral geniculate body were given by 8% of neurons. The difference between the latent periods of responses of the same visual cortical neurons to stimulation of the optic tract and lateral geniculate body was 0.1–1.8 msec, but for most neurons (55.8%) it was 0.5–1 msec. The histograms of response latencies of visual cortical neurons to stimulation of the above-mentioned formations were found to be similar. It is concluded that the optic radiation contains three principal groups of fibers with conduction velocities of 28.5–16.6, 11.7–8.9, and 7.4–6.0 m/sec, respectively.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 7, No. 6, pp. 589–596, November–December, 1975.     

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.     

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.     

Responses of limbic cortical neurons during instrumental conditioned reflexes in cats

Unit activity in cortical areas 24 and 32 was studied during conditioned placing reflex formation in cats. Neuronal responses in the limbic cortex of trained animals correlated with acoustic stimulation, the motor response, and also with the presentation of food reinforcement. In untrained animals 16% of neurons responded to acoustic stimulation. After training the number of neurons responding to sound in area 32 increased to 51.3%. Of the total number of neurons, 34.6% responded by initial excitation and 26.7% by inhibition of spike activity. The latent period of these responses was about 50 msec and their duration up to 200 msec. Similar but weaker responses were observed in area 24. Short-latency activation responses to conditioned and differential stimulation were similar in character. It is suggested that after training processes taking place in the limbic cortex may contribute to better perception of both conditioned and differential acoustic stimuli, irrespective of their functional significance.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 16, No. 2, pp. 201–208, March–April, 1984.     

Unit responses of the ventral posterior and ventral lateral thalamic nuclei to electrical stimulation of the thalamic reticular nucleus

A microelectrode investigation was made of responses of 72 physiologically identified neurons of the ventral posterior (VP) and 116 neurons of the ventral lateral (VL) thalamic nuclei to electrical stimulation of the reticular (R) thalamic nucleus. Mainly those neurons of VP and VL (73.7 and 86.2% respectively) which responded to stimulation of the first motor area and nucleus interpositus of the cerebellum responded to stimulation of R; 19.8% of VL neurons tested responded to stimulation of R by an antidromic action potential with latent period of 0.5–2.0 msec and 46.6% of neurons responded by orthodromic excitation; 23% of orthodromic responses had a latent period of 0.9–3.5 msec and 77% a latent period of 4.0–21.0 msec; 19.8% of VL neurons tested were inhibited. Among IPSPs recorded only one was monosynaptic (1.0 msec) and the rest polysynaptic. It is postulated that both R neurons are excitatory and that the inhibition which develops in VL neurons during stimulation of R are connected mainly with activation of inhibitory interneurons outside the reticular nucleus.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 9, No. 5, pp. 477–485, September–October, 1977.