Postsynaptic effects evoked by activation of long propriospinal pathways in cervical spinal neurons in cats

Effects induced in motoneurons and interneurons of the cervical enlargements of the cat spinal cord by stimulation of the lateral and ventral funiculi at the lower thoracic level were studied under conditions producing degeneration of fibers of descending brain systems. Stimulation of this sort evoked PSPs (mainly of mixed character) in 57 of 90 motoneurons tested. In nine motoneurons the primary response consisted of monosynaptic EPSPs evoked by activity of fibers of the lateral funiculus, and in the rest it consisted of polysyanptic (at least disynaptic) EPSPs and IPSPs. Polysynaptic effects arising in the neuron in response to stimulation of the lateral and ventral funiculi usually differed only quantitatively. The intensity of excitatory synaptic action on motoneurons of the proximal muscle (especially thoracid) was much greater than that on motoneurons of distal muscles. Nearly all motoneurons with no synaptic action belonged to the latter group. Stimulation of the lateral and ventral funculi facilitated synaptic action induced in motoneurons by stimulation of high-threshold segmental afferents and led to excitation of interneurons located in the vectral quadrant, and had no effect on interneurons in the dorsal regions of gray matter. These effects are regarded mainly as the result of excitation of long ascending propriospinal pathways in the cervical parts of the cord; it is also postulated that some of them are evoked by the arrival of activity along collaterals of descending propiospinal pathways to the neurons in this region.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 11, No. 4, pp. 339–347, July–August, 1979.     

Synaptic potentials of motoneurons of the masseter muscle

Postsynaptic potentials of 93 motoneurons of the masseter muscle evoked by stimulation of different branches of the trigeminal nerve were studied. Stimulation of the most excitable afferent fibers of the motor nerve of the masseter muscle evoked monosynaptic EPSPs with a latent period of 1.2–2.0 msec, changing into action potentials when the strength of stimulation was increased. A further increase in the strength of stimulation produced an antidromic action potential in the motoneurons with a latent period of 0.9 msec. In some motoneurons polysynaptic EPSPs and action potentials developed following stimulation of the motor nerve to the masseter muscle. The ascending phase of synaptic and antidromic action potentials was subdivided into IS and SD components, while the descending phase ended with definite depolarization and hyperpolarization after-potentials. Stimulation of cutaneous branches of the trigeminal nerve, and also of the motor nerve of the antagonist muscle (digastric) evoked IPSPs with a latent period of 2.7–3.5 msec in motoneurons of the masseter muscle. These results indicate the existence of functional connections between motoneurons of the masseter muscle and its proprioceptive afferent fibers, and also with proprioceptive afferent fibers of the antagonist muscle and cutaneous afferent fibers.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 1, No. 3, pp. 262–268, November–December, 1969.     

EPSPs of masseter motoneurons evoked by stimulation of low-threshold infraorbital nerve afferents in cat

Stimulation of the infraorbital nerve at strengths 1.4–2.5 times higer than the threshold of excitation of A fibers in cats anesthetized with chloralose and pentobarbital evoked EPSPs with an amplitude up to 3.0 mV and a duration of 9–15 msec in 69% of masseter motoneurons after 1.5–3.0 msec. These EPSPs were complex and formed by summation of simpler short-latency and long-latency EPSPs. The short-latency EPSPs appeared in response to infraorbital nerve stimulation at 1.1–1.5 thresholds and had a slow rate of rise (2.5–4.5 msec, mean 3.7±0.4 msec), low amplitude (under 2.0 mV), and short duration (5–6 msec). Their latent period varied from 1.5 to 3.0 msec (mean 2.1±0.2 msec). The shortness of the latent period and its constancy during stimulation of the nerve at increasing strength, and also the character of development of facilitation and inhibition of the EPSP during high-frequency stimulation suggests that these EPSPs are monosynaptic. The slow rate of rise suggested that these EPSPs arise on distal dendrites of the motoneurons. Long-latency EPSPs appeared 7–9 msec after stimulation of the infraorbital nerve at 1.1–1.5 thresholds. Their amplitude reached 1.5–2.0 mV and their duration 7–9 msec. The long duration of the latent period combined with low ability to reproduce high-frequency stimulation (up to 30/sec) points to the polysynaptic origin of these EPSPs.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 9, No. 6, pp. 583–591, November–December, 1977.     

Monosynaptic excitation of motoneurons of the accessory nerve nucleus by reticulofugal impulses

Acute experiments on cats anesthetized with chloralose and pentobarbital showed that excitation of fast-conducting (130 m/sec) reticulospinal fibers, arising during stimulation of the ipsilateral medullary reticular gigantocellular nucleus evoked monosynaptic EPSPs in motoneurons of the accessory nerve nucleus. The EPSPs had latent periods of between 0.6 and 1.0 msec (mean 0.7 msec), they reached their maximal amplitude (4.0 mV) after 2.0–2.5 msec, and lasted about 10 msec. The EPSPs underwent only weak potentiation through the different types of stimulation of the gigantocellular nucleus and were not transformed into action potentials.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 12, No. 1, pp. 62–66, January–February, 1980.     

Reticulospinal and vestibulospinal synaptic effects on the lumbar motor neurons of the monkey

It was established in experiments on rhesus monkeys by intracellular recording and computer averaging that fast-conducting reticulo- and vestibulospinal fibers form monosynaptic excitatory links with lumbar alpha-motor neurons. The monosynaptic bulbospinal effects are retained after sectioning of the pyramids or after chronic destruction of the motor cortex. The mean amplitude of the reticulomotor neuronal EPSP is less than that of the corticomotor neuronal EPSP, however, the mean amplitudes of the EPSP of individual motor neurons can reach similar values. In contrast to the corticomotor neuronal projections, the bulbomotor neuronal projections are directed primarily toward the motor neurons of the proximal muscles. In addition to monosynaptic EPSP, reticulo- and vestibulospinal impulses evoke disynaptic EPSP and IPSP in the lumbar motor neurons of monkeys which are very similar to the disynaptic effects found in subprimates.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 3, No. 4, pp. 408–417, July–August, 1971.     

Amygdaloid or cortical facilitation of antidromic activity of trigeminal motoneurons in the rat

1. Electrical stimulation of the rat's contralateral central amygdaloid (CAm) nucleus or the contralateral frontal cortex markedly augmented the antidromic field potential evoked by stimulation of mylohyoid (Myl) nerve. 2. This facilitation was shown to be due to EPSPs of the mylohyoid-anterior digastric (Myl-Dig) motoneurons. 3. In a few motoneurons, cortical EPSPs had fixed short latencies following high-frequency double stimuli and this is believed to be due to a monosynaptic pathway. 4. The amygdaloid or cortically evoked EPSPs relieved IS-SD blockade in a few motoneurons and also facilitated antidromic discharge in others which did not show any IS or M spike response to the same subthreshold antidromic stimulation. The underlying mechanisms are discussed.     

Excitation of accessory motoneurons by stimulation of Deiters' nucleus in the cat

Acute experiments on cats under chloralose-pentobarbital anesthesia showed that application of single stimuli to Deiters' nucleus evoked monosynaptic EPSPs in motoneurons of the accessory nucleus. Latent periods of EPSPs ranged from 1.3 to 2.3 msec (mean 1.8±0.3 msec), their rise time was 0.5–1.0 msec, and their duration 7–10 msec. During repetitive stimulation the EPSPs were weakly potentiated, but with an increase in the strength of stimulation applied to Deiters' nucleus they readily changed into action potentials. In some motoneurons polysynaptic EPSPs with latent periods of the order of 6.0 msec appeared on the descending phase of these EPSPs.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 13, No. 5, pp. 515–519, September–October, 1981.     

Synaptic processes evoked by intracortical microstimulation in lumbar motoneurons of rats

Experiments with intracortical microstimulation and intracellular recording from motoneurons of the rat hind limb showed that synaptic effects due to activation of pyramidal neurons of the motor cortex and transmitted along the pyramidal tract are exclusively polysynaptic in character. Mainly excitatory effects were found in motoneurons of flexors and extensors, and of distal and proximal muscles. The minimal intensity of intracortical stimulation required for synaptic excitation of -motoneurons is 5–10 µA. Low-threshold synaptic effects in lumbar motoneurons and movements of the hind limbs are evoked from the same zones.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 5, No. 2, pp. 174–180, March–April, 1973.     

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.     

Corticofugal influences on postsynaptic processes in rubrospinal neurons in the cat brain

Composite and unitary EPSPs of red nucleus neurons evoked by stimulation of the sensomotor and association parietal cortex and nucleus interpositus of the cerebellum were studied in acute experiments on cats anesthetized with pentobarbital. A monosynaptic connection was shown to exist between not only the sensomotor, but also the association cortex, and rubrospinal neurons, in which unitary EPSPs appeared during stimulation of the association cortex after a latent period of 1.5–2.7 msec, with a peak rise time of 1.1–3.1 msec and an amplitude of 0.22–0.65 mV. Analysis of the temporal characteristics of the unitary EPSP suggested that synapses formed by fibers from the association cortex occupy a position nearer the soma than synapses formed by axons of sensomotor cortical cells.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 16, No. 1, pp. 67–74, January–February, 1984.     

Changes in postsynaptic responses in spinal motoneurons during repetitive stimulation of the locus coeruleus

Experiments on cats anesthetized with chloralose showed that repetitive stimulation of the locus coeruleus is accompanied by a decrease in IPSPs evoked by stimulation of flexor reflex afferents in extensor motoneurons. The effect appeared 600 msec after the beginning of stimulation and reached its maximum after 1500–2000 msec. Repetitive stimulation of the locus coeruleus did not change the membrane potential and did not affect EPSPs or IPSPs evoked by stimulation of low-threshold muscle afferents; EPSPs due to activation of high-threshold cutaneous and muscle afferents likewise remained unchanged. Repetitive stimulation of more central regions of the brain stem was accompanied not only by a decrease in IPSPs evoked by stimulation of flexor reflex afferents in extensor motoneurons, but also by a decrease in amplitude of EPSPs arising in response to stimulation of these same afferents in flexor motoneurons. These effects were not connected with activation of monoaminergic structures, for unlike effects arising during stimulation of the locus coeruleus, they were also found in previously reserpinized animals.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 14, No. 1, pp. 51–59, January–February, 1982.     

Synaptic effects evoked in motoneurons by stimulation of single propriospinal neurons

Experiments on cats with simultaneous extracellular recording, stimulation of single propriospinal neurons, and intracellular recording of unitary postsynaptic potentials from motoneurons, followed by computer averaging showed that direct stimulation of individual propriospinal cells receiving mono- and disynaptic influences from the medial reticular formation can evoke monosynaptic EPSPs and IPSPs in lower lumbar motoneurons. The amplitude of these EPSPs was 49.6±6.0 and of the IPSPs 28.9±2.9 µV and their synaptic delay was 0.34±0.05 msec. The same propriospinal neuron of the ventral horn of the upper lumbar segments may be connected with several motoneurons of the hind limb muscles.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 9, No. 3, pp. 300–306, May–June, 1977.     

Role of different types of fibers of the infraorbital nerve in synaptic activation of masseter and digastric motoneurons

Postsynaptic potentials of motoneurons of the masseter and digastric muscles evoked by stimulation of the infraorbital nerve with a strength of between 1 and 10 thresholds were investigated in cats anesthetized with a mixture of chloralose and pentobarbital. Depending on their ability to be activated by low-threshold afferents of this nerve, motoneurons of the masseter were divided into two groups. Stimuli with a strength of 1.2–2.5 times above threshold for the most excitable fibers of the infraorbital nerve evoked short-latency EPSPs in the motoneurons of the first group; a further increase in stimulus strength (3–9 thresholds) led to the appearance of IPSPs with latent periods of 2.8–3.5 msec. Motoneurons of the second group responded to stimulation of the infraorbital nerve with a strength of 3–9 thresholds by IPSPs whose latent periods varied from 6 to 8 msec. Stimuli below 3 thresholds in strength evoked no responses in these motoneurons. Stimulation of the infraorbital nerve with pulses of between 1 and 2 thresholds in strength evoked EPSPs in digastric motoneurons, but an increase in the strength of stimulation led to action potential generation. The presence of many excitatory and inhibitory inputs formed by afferent fibers of different types evidently provides a basis for functional diversity of jaw-opening and jaw-closing reflexes.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 12, No. 6, pp. 596–603, November–December, 1980.     

Synaptic organization of supraspinal control over propriospinal ventral horn interneurons in cat and monkey spinal cord

Synaptic responses evoked in propriospinal neurons of the upper lumbar segments (L₃–L₄) by reticulo-, vestibulo-, and corticospinal impulses were studied in experiments on cats and monkeys. Propriospinal cells, identified by antidromic stimulation, were stained with Procion red, so that they could be localized in the different zones of the ventral horn. Monosynaptic reticular and vestibular excitatory influences were discovered in cats; convergence of these influences on the same neurons was demonstrated. In monkeys bulbospinal monosynaptic effects were supplemented by monosynaptic influences arriving from the motor cortex; convergence of monosynaptic excitatory influences from all supraspinal sources studied was found on some propriospinal neurons. The propriospinal neurons studied also had synaptic inputs from primary afferents.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 9, No. 2, pp. 177–184, March–April, 1977.     

The role of reflex mechanisms in postinhibitory rebound studied by analysis of human single motor unit activity

The mechanism of onset of rebound after inhibition induced by electrical stimulation of a nerve of maximal and submaximal strength for M-response was studied in single motor units of normal human soleus, rectus femoris, and hand muscles. Poststimulus histograms and changes in the duration of interspike intervals were compared with mechanical recordings of muscle contractions. In all muscles tested, during strong isotonic contraction, the increase in motor unit activity after a silent period was partly due to synchronization of their emergence from inhibition. However, it also contained a component of true facilitation of motoneurons, which was evidently a reflex response to lengthening of the muscle in the relaxation phase after evoked contraction. The latent period of this facilitation in the soleus and rectus femoris muscles coincided in value with the latent period of the monosynaptic spinal reflex, whereas in the hand muscles, in which a monosynaptic response to electrical nerve stimulation could not be evoked, the latent period of facilitation as a result of spindle activation during muscle relaxation was significantly longer than the latent period of the monosynaptic reflex. These findings support the hypothesis of presynaptic suppression of monosynaptic connections of Ia afferents with the motoneurons of some human muscles by descending tonic influences and of the use of information coming from spindles by supraspinal levels of the CNS.     

Connections of neurons of the cat somatosensory cortex with the thalamic relay nuclei

Of 103 neurons in the rostral part of the posterior sigmoid gyrus of the cat cortex 30 responded to stimulation of the ventro-posterolateral and ventrolateral nuclei of the thalamus (VPL and VL), 42 responded to stimulation of VL only, and 31 to stimulation of VPL only. It was shown by intracellular recording that stimulation of VPL induces a spike response with or without subsequent IPSPs in some neurons and an initial IPSP in others. The spike frequency of single neurons reached 60/sec, but the IPSP frequency never exceeded 10–20/sec. Stimulation of VL was accompanied by: a) antidromic spike responses; b) short-latency monosynaptic EPSPs and spikes capable of following a stimulation frequency of 100/sec; c) long-latency polysynaptic EPSPs and spikes appearing in response to stimulation at 4–8/sec; d) short-latency IPSPs; e) long-latency IPSPs increasing in intensity on repetition of infrequent stimuli. It is concluded that the afferent inputs from the relay nuclei to neurons of the somatosensory cortex are heterogeneous. An important role is postulated for recurrent inhibition in the genesis of the long-latency IPSPs arising in response to stimulation of VL, and for direct afferent inhibition during IPSPs evoked by stimulation of VPL. It is shown that the rostral part of the posterior sigmoid gyrus performs the role of somatic projection and motor cortex simultaneously.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 4, No. 3, pp. 245–255, May–June, 1972.     

Convergence of reticulo-, vestibulo-, and rubrospinal influences on cervical spinal motoneurons in cats

Intracellular recordings were made of synaptic responses of 93 motoneurons in the cervical region of the cat spinal cord to stimulation of the medial longitudinal bundle, the brain-stem reticular formation, the lateral vestibular nucleus of Deiters, and the red nucleus. In response to stimulation of the medial longitudinal bundle and the vestibular nucleus responses in the motoneurons of the distal groups of muscles of the forelimb were predominantly excitatory, whereas in motoneurons of the proximal extensor muscles they were predominantly inhibitory. During stimulation of the red nucleus, excitatory and inhibitory responses were recorded in almost equal numbers of cells regardless of their functional class. Monosynaptic EPSPs appeared in one-fifth of motoneurons in response to stimulation of the medial longitudinal bundle and, in a few cases, to stimulation of the vestibular and red nuclei. Otherwise, during stimulation of these structures polysynaptic responses were recorded in the motoneurons. In 62% of cases postsynaptic potentials arising in response to stimulation of the various suprasegmental structures tested were identical in direction in the same motoneurons. A mutually facilitatory effect was observed during stimulation of different suprasegmental inputs. The results are evidence that interaction between influences of the structures tested takes place largely at the level of spinal interneurons.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 10, No. 4, pp. 391–399, July–August, 1978.     

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.     

Synaptic processes in facial motoneurons evoked by stimulation of the caudal trigeminal nucleus

Field potentials and postsynaptic potentials of facial motoneurons evoked by stimulation of the caudal trigeminal nucleus were investigated in acute experiments on cats by extra- and intra-cellular recording. Pre- and postsynaptic components of field potentials were found. Four types of motoneuron response were distinguished: EPSP with generation of single action potentials; a gradual shift of depolarization inducing grouped action potentials; a rhythmic discharge of action potentials arising at a low level of depolarization; and EPSPs or EPSP-IPSP sequences. The monosynaptic and (chiefly) polysynaptic nature of these responses was demonstrated. The possible mechanism of afferent control over facial motoneurons are discussed.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 12, No. 3, pp. 272–282, May–June, 1980.     

Monosynaptic excitation of facial motoneurons during electrical stimulation of the red nucleus

Synaptic effects of the red nucleus on motoneurons of the facial nucleus were studied in cats. Impulses from the red nucleus activate motoneurons innervating the auricular, buccal, and orbicularis oculi muscles. Monosynaptic EPSPs appeared in all motoneurons which responded to stimulation. Their mean latent period was 1.5±0.04 msec, duration 12.3 ± 0.34 msec, and rise time between 1.5 and 3.2 msec. Repetitive stimulation of the red nucleus led to marked facilitation of the testing EPSP. Facilitation was maximal when the interval between stimuli was 3.5 msec; it was reduced by either a decrease or an increase in the interval. The functional role of the monosynaptic connections of neurons of the red nucleus and of the facial motoneurons is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 4, No. 3, pp. 272–279, May–June, 1972.