Convergence of corticospinal and rubrospinal influences on cervical motoneurons

Synaptic responses of 121 identified cervical motoneurons to stimulation of the pyramidal tract and red nucleus were investigated by intracellular recording in cats. Responses of EPSP or EPSP-IPSP type were predominant in motoneurons of distal groups of muscles and proximal flexors, while responses of IPSP type were predominant in motoneurons of the proximal extensors. The minimal effective number of stimuli for most motoneurons was 2 or 3. The mean latent period, counted from the first stimulus in the series, was 7.86 msec for EPSPs for stimulation of the pyramidal tract and 7.91 msec for stimulation of the red nucleus, while the corresponding periods for IPSPs were 8.68 and 8.75 msec. The segmental delay of 1.3–2 msec for EPSPs and IPSPs generated in certain motoneurons in response to stimulation of both structures indicates that the shortest pathway for transmission of activity from the fibers of these tracts to the motoneurons may be disynaptic. At the same time, the possible presence of an additional neuron for most inhibitory pathways cannot be ruled out. Analysis of the results also suggests the presence of a common interneuronal apparatus for both systems.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol.3, No.6, pp. 599–608, November–December, 1971.     

Effects of repetitive stimulation of the locus coeruleus on spinal inhibitory responses to suprasegmental stimulation in cats

Effects of repetitive stimulation of the locus coeruleus on spinal responses to activation of cortico-, reticulo-, and vestibulospinal tracts were studied in decerebellate cats anesthetized with chloralose. Descending influences of these structures were assessed from changes in amplitude of extensor and flexor monosynaptic discharges or from the magnitude of postsynaptic potentials recorded from the corresponding motoneurons. Stimulation of the motor cortex or modullary reticular formation as a rule evoked two-component inhibitory responses in extensor motoneurons and excitatory-inhibitory responses in flexor motoneurons. Stimulation of locus coeruleus effectively depressed the amplitude of the late component and, to a lesser degree, that of the early component of inhibition arising after stimulation of the cerebral cortex or reticular formation. During stimulation of the locus coeruleus no marked changes were found in inhibitory responses evoked by vestibulospinal influences in flexor motoneurons, and also in excitatory responses arising after stimulation of the above-mentioned descending pathways in both groups of motoneurons.     

Reticulospinal and propriospinal monosynaptic influences on motoneurons in frogs

Monosynaptic effects evoked by electrical stimulation of suprasegmental structures and the ventral and lateral columns were recorded intracellularly from motoneurons of the lumbar and cervical enlargements after isolation of the spinal cord and medulla in frogs. Reticulospinal fibers arising from cells of the medial reticular formation of the medulla and running in the ventro-lateral columns evoke monosynaptic excitation of cervical and lumbar motoneurons. The reticulo-motoneuronal E PSPs do not exceed 2–3 mV in amplitude and do not reach the threshold for action potential generation. Division of the spinal cord and interaction between all synaptic inputs tested in chronic experiments showed that monosynaptic E PSPs evoked by direct stimulation of the ventral and lateral columns are due to activation of the descending system of propriospinal fibers. By transmembrane polarization experiments the equilibrium potentials of the reticulo-motoneuronal and propriospinal monosynaptic E PSPs could be determined.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Leningrad. Translated from Neirofiziologiya, Vol. 5, No. 2, pp. 164–173, March–April, 1973.     

Synaptic processes in thoracic motoneurons evoked by visceral impulses

Synaptic processes in various functional groups of thoracic motoneurons (segments T9–T11) were investigated in anesthetized (chloralose and Nembutal), decerebrate, and spinal cats. Visceral stimulation in animals with an intact CNS during artificial respiration evokes the development of primary (latent period under 12 msec) and secondary (latent period over 30 msec) PSPs in the motoneurons. The primary PSPs consist of early and principal components. The early component is due to excitation of group A₂ and A visceral afferents, the principal PSP to excitation of the A group. The principal component in motoneurons of the internal and external intercostal muscles and abdominal muscles is excitatory, while in motoneurons innervating the spinal muscles it is excitatory—inhibitory or inhibitory. The secondary PSPs as a rule are excitatory and are due to activation of fibers of the A group. During natural respiration the primary PSPs of motoneurons of the intercostal muscles and abdominal muscles are predominantly inhibitory. In spinal animals no secondary responses are present and the primary becomes entirely excitatory regardless of the functional class of the motoneurons. The mechanisms of reciprocal activation of thoracic motoneurons by visceral impulses in animals during artificial and natural respiration are discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 4, No. 3, pp. 286–295, May–June, 1972.     

Changes in the excitability of and synaptic effects on motoneurons during formation of the scratch motion generator in the cat

Studies on immobilized decerebrate (at intracollicular level) cats in which the scratch generator had been set up following bicuculline application to the upper cervical segments of the spinal cord, showed that the state of the segmental apparatus of the lumbosacral section of the spinal cord differs substantially from that seen in the spinal animal. Direct excitability of motoneurons of the "aiming" and "scratching" muscles rises, while recurrent and reciprocal Ia inhibition of motoneurons intensifies and the influence of Ib afferents on motoneurons declines. Afferents of the flexor reflex exert a primarily inhibitory influence on motoneurons of the "aiming" muscles. This influence becomes predominantly excitatory following spinalization, while the inhibitory effects of these afferents on motoneurons of the "scratch" muscles declines. The functional significance of the changes discovered in generation of scratch routine is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 19, No. 2, pp. 244–250, March–April, 1987.     

An intracellular study of pudendal afferent inputs onto tail motoneurons in the spinalized cat

Postsynaptic potentials, elicited by stimulation of the sensory pudendal (SPud) and superficial perineal nerves (SPeri) on both sides, were recorded from motoneurons innervating tail muscles in the non-anaesthetized and spinalized cat. The stimulation of SPud and SPeri on both sides predominantly produced excitatory postsynaptic potentials (EPSPs) in all kinds of tail motoneurons (70-95%). The inhibitory postsynaptic potentials (IPSPs) were often observed in motoneurons innervating ventral tail muscles (30-33%). The means of averaged central latencies of EPSPs and IPSPs ranged from 4.3 to 7.3 ms, and from 4.6 to 8.4 ms, respectively. The findings suggests that polysynaptic neuronal pathways from pudendal nerve to tail motoneurons produce tonic activities of all tail muscles to raise the tail in micturation, defecation and sexual movements which are induced by stimulation of pudendal nerves.     

Corticofugal influences on the activity of vestibular nuclei units in the cat

Neuronal responses to stimulation of vestibular motor and orbital cerebral cortex were recorded by extracellular techniques in the lateral and medial vestibular nuclei of the bulbar complex during experiments on unanesthetized, immobilized cats. Both phasic and (mostly) tonic response of predominantly inhibitory type were observed. Horseradish peroxidase was injected into the aforementioned nuclei of the vestibular complex during the course of morphological experiments. Labeled neurons were found in the anterior supra- and ectosylvian cerebral gyri, the region of the cruciform sulcus, and that of the orbital cerebral cortex. Findings are discussed from the aspect of corticovestibular interaction.Ivano-Frankovsk Medical Institute, Soviet Ministry of Health. Translated from Neirofiziologiya, Vol. 19, No. 6, pp. 802–809, November–December, 1987.     

Different Sensitivity of Motoneuron Synaptic Inputs in the Frog, Rana ridibunda, to Antagonists of Excitatory Amino Acids

The effects of antagonists of excitatory amino acids (AP-5, kynurenate, and CNQX) on PSP recorded intracellularly in lumbar motoneurons of a preparation of the isolated spinal cord in the frog, Rana ridibunda, in response to activation of three different synaptic inputs (stimulation of DR, RF, and VC or LC) were analyzed. It is shown that the effects of the antagonists were non-uniform in different motoneurons. Inputs of suprasegmental and sensomotor projections substantially differed from each other. A considerable amount of DC-PSP resistant to kynurenate and CNQX was found, whereas the latter regularly inhibited DR-PSP in the same cell. The disynaptic, as judged by its latency, plane-shaped component was always relatively more stable to kynurenate as compared with other components. Unlike kynurenate that inhibited the early and late components, CNQF selectively depressed the early components of DR-PSP.     

Rubrospinal synaptic influences on the lumbar motoneurons of the rat

Intracellular recording was employed in experiments on rats with the nervous system intact and after acute pyramidotomy to study the postsynaptic effects produced in the lumbar motoneurons on stimulation of the nucleus ruber. Stimulation of this nucleus with single stimuli and with a short series of stimuli caused excitatory and inhibitory postsynaptic potentials (EPSP and IPSP) to develop in the motoneurons. Most of the EPSP recorded were disynaptic, but response development involved a monosynaptic segmental delay in five of the 124 cells that exhibited EPSP. A capacity for high-frequency potentiation was a characteristic feature of the disynaptic excitatory and inhibitory effects. Transmembrane polarization of the motoneurons had a marked influence on the amplitude of the disynaptic EPSP and IPSP. The properties of the disynaptic rubrospinal influences were similar to those described for the cat.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 3, No. 3, pp. 266–273, May–June, 1971.     

Responses of units in the magnocellular part of the medial geniculate body to acoustic and somatosensory stimulation

Responses of 150 neurons in the magnocellular part of the medial geniculate body to clicks and to electrodermal stimulation of the contralateral forelimb were investigated in cats immobilized with myorelaxin. Of the total number of neurons 65% were bimodal, 16.6% responded only to clicks, and 15.4% only to electrodermal stimulation. The unitary responses were excitatory (spike potentials) and inhibitory (inhibition of spontaneous activity). Responses beginning with excitation occurred more frequently to stimulation by clicks than to electrodermal stimulation, whereas initial inhibition occurred more often to electrodermal stimulation. The latent period of the initial spike potentials in response to clicks and to electrodermal stimulation was 5–27 and 6–33 (mean 11.6 and 16.2) msec respectively. Positive correlation was found between the latent periods of spike potentials recorded in the same neurons in response to clicks and to electrodermal stimulation, and also to electrodermal stimulation and to stimulation of the dorsal funiculus of the spinal cord. It is concluded that the magnocellular division of the medial genicculate body is a transitional structure between the posterior ventral nucleus and the parvocellular division of the medial geniculate body, and that in addition, it is connected more closely with the auditory than with the somatosensory system. It is suggested that the somatosensory input into the magnocellular division of the medial geniculate body is formed mainly by fibers of the medial lemniscus.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 10, No. 2, pp. 133–141, March–April, 1978.     

The effect of electrical stimulation of vestibular-complex formations on systemic arterial pressure and regional blood flow

Electrical stimulation of the descending and medial vestibular nuclei produced increased arterial blood pressure and also changes in regional blood flow. On stimulation of the descending vestibular nucleus there was a decrease in blood flow in the hind-limb muscles, which might be a response to stimulation of the ventrolateral parts of the nucleus; when the central parts of the nucleus were stimulated the blood flow decreased both in the hind-limb muscles and in some of the viscera. Stimulation of the medial vestibular nucleus, particularly its ventral parts, caused a decrease in blood flow in the skin of the hind limbs, often accompanied by a decrease in visceral blood flow. On the basis of analysis of the afferent and efferent connections of the descending and medial vestibular nuclei, it is suggested that the cerebellum takes part in the transmission of vasomotor influences.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 2, No. 1, pp. 35–42, January–February, 1970.     

Function of alpha-motoneurons of the human foot muscles in normal conditions and with descending and segmental deprivation

The characteristics of the central response of the abductor hallucis muscle to stimulation of the posterior tibial nerve were studied in healthy subjects and neurological patients. In normal subjects over 22–23 years old the response has the usual properties of a late potential recorded from the small muscles of the foot (F-wave) whereas in children and adolescents it has the features of the central response of a leg muscle. A unilateral lesion of the posterior sacral roots leads to an increase in amplitude and regularity of the F-wave of the foot muscle on the side of the lesion. This is evidence of the antidromic origin of the principle component of the F-wave and points to increased antidromic excitability of the spinal motoneurons in man in the presence of partial segmental deafferentation. A disturbance of descending control, resulting in the loss predominantly of pyramidal influences, is accompanied by regeneration of the characteristic features of the late potential found in responses recorded in adolescent subjects. The same phenomenon can also be observed in pathology of the posterior horns of the cervical portion of the spinal cord when no pyramidal disorders can be detected clinically. The central response is much less frequently changed in lesions of the ventral structures of higher segments. The significance of descending control for establishment of the level of excitability in the population of motoneurons innervating the muscles of the human foot is discussed.O. V. Kuusinen Petrozavodsk State University. Translated from Neirofiziologiya, Vol. 5, No. 6, pp. 563–570, November–December, 1973.     

Electrophysiological analysis of corticofugal influences on preoptic neurons

Evoked potential (EPs) and responses of the medial (MPO) and lateral (LPO) preoptic region (RPO) and adjacent structures of the hypothalamus to stimulation of the prefrontal (area 8) and cingulate (area 24) cortex, piriform lobe (periamygdaloid cortex), and hippocampus (area CA3) were investigated in acute experiments on cats under ketamine anesthesia. The most pronounced EPs were observed in the RPO after stimulating the piriform and cingulate cortex. A close relation was found between neuronal responses and EP components. The majority of neurons responding to stimulation of various cortical structures were localized in the LPO, where primarily excitatory responses dominate. The MPO contained somewhat fewer neurons responding to cortical stimulation, and the dominant response here was primarily inhibitory. The ratio of inhibitory and excitatory responses in the LPO was 0.6:1 and in the MPO 5.8:1. Primarily in-inhibitory responses dominated also in the LPO zone adjacent to the bed nucleus stria terminalis (BST) and primarily excitatory in the region surrounding the supraoptic nucleus (SO) (respective ratios 4.9:1 and 0.7:1). The RPO is a broad convergence zone, where 3/4 of the neurons responded to stimuli of two and more cortical regions.A. M. Gorky Medical Institute, Ukrainian Minstry of Health, Donetsk. Translated from Neirofiziologiya, Vol. 23, No. 6, pp. 709–719, November–December, 1991.     

Synaptic processes in spinal cord interneurons under rubrospinal effects

Synaptic processes of the spinal cord interneurons under rubrospinal effects have been investigated. A recording was made of 156 interneurons from the different parts of the gray matter, 111 of the interneurons were activated by descending effects from the red nucleus and 47 were not activated. Sixty nine interneurons of the first group responded only to rubrospinal impulsation and 42 neurons were also activated by afferent volleys. Interneurons activated only by the rubrospinal tract were located in the most lateral part of the VII Rexed's gray matter layer; the majority of interneurons activated by both rubrospinal and peripheral afferent volleys were located in the nucleus propius of the dorsal horn and the Cajal intermediate nucleus. The mean latencies of EPSP's and action potentials in interneurons activated only by a rubrospinal tract were 64±0.2 and 9.5±0.62 msec, respectively. The mean latency of EPSP's in motoneurons of flexor muscles was 10.3±0.62 msec and of IPSP's in motoneurons of extensor muscles, it was 11.5±1.28 msec. It is assumed that rubrospinal impulsation evokes excitatory PSP's in the motoneurons via the disynaptic pathway with the participation of special interneurons located in the lateral part of the VII layer. Inhibitory and late excitatory responses are, apparently, evoked via additional interneurons.A. A. Bogomolets Institute of Physiology of the Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 1, No. 2, pp. 158–166, September–October, 1969.     

Synaptic potentials of digastric-muscle motoneurons

We studied the antidromic and synaptic potentials evoked from 32 digastric-muscle motoneurons by stimulation of the motor nerve to this muscle, different branches of the trigeminal nerve, and the mesencephalic trigeminal nucleus. Antidromic potentials appeared after 1.1 msec and lasted about 2.0 msec. Stimulation of the infraorbital, lingual, and inferior alveolar nerves led to development of excitatory postsynaptic potentials (EPSP) and action potentials in the motoneurons. The antidromically and synaptically evoked action potentials of the digastric-nerve motoneurons were characterized by weak after-effects. We were able to record EPSP and action potentials in two of the motoneurons investigated in response to stimulation of the mesencephalic trigeminal nucleus, the latent period being 1.3 msec. This indicates the existence of a polysynaptic connection between the mesencephalic-nucleus neurons and the digastric-muscle motoneurons. Eight digastric-muscle motoneurons exhibited inhibitory postsynaptic potentials (IPSP), which were evoked by activation of the afferent fibers of the antagonistic muscle (m. masseter). The data obtained indicate the presence of reciprocal relationships between the motoneurons of the antagonistic muscles that participate in the act of mastication.A. A. Bogomol'ts Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 3, No. 1, pp. 52–57, January–February, 1971.     

Neuromuscular transmission of pectoral fin muscles of the goldfish Carassius auratus

The electrical properties and neuromuscular transmission of white and red fibers of pectoral fin muscles of the goldfish Carassius auratus were studied using an intracellular recording technique. The pectoral fin muscles consist mainly of white and red fibers. Almost all of white fibers elicited action potentials with overshoot by direct stimulation, but graded responses appeared in the red fibers. However, overshooting action potentials were often recorded from the red fibers in saline containing 20 microM tetraethylammonium (TEA) chloride. In response to single nerve stimulations, excitatory (EJPs) and inhibitory junction potentials (IJPs) were obtained from both white and red fibers in common. Both EJPs and IJPs were blocked completely or partially by d-tubocurarine, a nicotinic acetylcholine (ACh) receptor antagonist. Nicotine, a nicotinic ACh receptor agonist, and oxotremorine, a muscarinic ACh receptor agonist, depolarized both fiber types. The results suggest that white and red fibers receive double innervation from excitatory and inhibitory nerves, and have nicotinic and muscarinic ACh receptors. In the resting muscle, miniature excitatory junction potentials were generated spontaneously in both white and red fibers. Occasionally, miniature inhibitory junction potentials were recorded from the red fibers. The results indicate that the release of both excitatory and inhibitory transmitters is quantal in nature.     

Effect of epidural spinal cord stimulation on the activity of lateral vestibular nucleus neurons in the cat

The activity of neurons in Deiters' lateral vestibular nucleus was recorded in decerebrate cats before, during and after spinal cord stimulation. An almost equal number of units were inhibited and excited early during stimulation. Later during stimulation the majority of units was inhibited. Early after cessation of stimulation an ever larger number of units were inhibited to an even larger extent (for about 2 imp/s on the average). Later after stimulus cessation the predominant inhibitory effect could still be noted, as well as excitation in some units. The results could support the hypothesis that the inhibition of Deiters' neurons during and for some time after epidural cord stimulation may play a part in the decrease of limb spasticity. The mechanism of inhibitory and excitatory unitary responses, side effects during stimulation and differences between the experimental model and human state are discussed.     

Identification of common excitatory motoneurons in Drosophila melanogaster larvae

In insects, four types of motoneurons have long been known, including fast motoneurons, slow motoneurons, common inhibitory motoneurons, and DUM neurons. They innervate the same muscle and control its contraction together. Recent studies in Drosophila have suggested the existence of another type of motoneuron, the common excitatory motoneuron. Here, we found that shakB-GAL4 produced by labels this type of motoneuron in Drosophila larvae. We found that Drosophila larvae have two common excitatory motoneurons in each abdominal segment, RP2 for dorsal muscles and MNSNb/d-Is for ventral muscles. They innervate most of the internal longitudinal or oblique muscles on the dorsal or ventral body wall with type-Is terminals and use glutamate as a transmitter. Electrophysiological recording indicated that stimulation of the RP2 axon evoked excitatory junctional potential in a dorsal muscle.     

Corticofugal influences on the neurons of different regions of the hypothalamus

Responses of the neurons of the lateral and ventromedial hypothalamic regions (HL andHvm, respectively), as well as of the area of the dorsal hypothalamus (aHd) and the projection region of the medial forelimb bundle (MFB), evoked by stimulation of the proreal cortex (field 8), cingular cortex (field 24), pyriform lobula (periamigdalar cortex), and hippocampus (CA3) were studied in acute experiments on cats under ketamine anesthesia. Distributions of the latent periods of the responses recorded from hypothalamic neurons at stimulation of the above cortical structures were analyzed. The responses were classified into primary excitatory and primary inhibitory. Stimulation of the proreal gyrus evoked four times more excitatory responses than inhibitory responses. With stimulation of the cingular gyrus, the ratio of excitatory/inhibitory responses was 1.5∶1. Stimulation of the pyriform cortex evoked activatory and inhibitory responses with a similar probability. With hippocampal stimulation, inhibitory responses appeared two times more frequently than excitatory reactions. The hypothalamus was found to be a zone of wide convergence: one-half of all responding neurons in theHL andHvm responded to stimulations of two or more tested cortical zones. In 26% of the cells, only excitatory convergence was observed, while in 10% only inhibitory convergence was found; 21% of the cells revealed mixed convergence.     

Synaptic processes in thoracic α-motoneurons evoked by segmental afferent stimulation

Synaptic processes in various functional groups of thoracic motoneurons (Th₉-Th₁₁) evoked by stimulation of segmental nerves were investigated in anesthetized and decerebrate cats. No reciprocal relations were found between these groups of motoneurons. Only excitatory mono- and polysynaptic responses were recorded in the motoneurons of the principal intercostal nerve following stimulation of the homonymous nerve. Activation of the afferents of the external intercostal muscle and dorsal branches does not cause noticeable synaptic processes in these motoneurons; much more rarely it is accompanied by the development of low-amplitude polysynaptic EPSP's. In motoneurons of the dorsal branches, stimulation of homonymous nerves leads to the appearance of simple, short-latent EPSP's. Late responses of the IPSP or EPSP - IPSP type with a predominance of the inhibitory component were observed in most motoneurons of this type following activation of the afferent fibers of the principal intercostal nerve. In other motoneurons of the dorsal muscles, stimulation of the main intercostal nerve (and nerve of the external intercostal muscle) did not evoke apparent synpatic processes. EPSP's (mono- and polysynaptic) appeared in the motoneurons of the external intercostal muscle following stimulation of the homonymous and main intercostal nerves. Activation of the afferents of the dorsal branches was ineffective. The character of the synaptic responses of the respiratory motoneurons to segmental afferent stimulation, investigated under conditions of spontaneous respiration, was different. The characteristics of synaptic activation of thoracic motoneurons by segmental afferents under conditions of hyperventilation apnea and during spontaneous breathing of the animals are discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 2, No. 3, pp. 279–288, May–June, 1970.