Corticofugal influences on reticulospinal neurons of the gigantocellular nucleus in cats

It was shown by intracellular recording that stimulation of the motor cortex evokes E PS Ps and I PS Ps in reticulospinal neurons of the gigantocellular nucleus of the cat medulla. The E PS Ps appeared in 94.3% and the I PS Ps in 5.7% of neurons tested. Analysis of the presynaptic pathway showed that 77.4% of E PS Ps studied arose through monosynaptic, and 22.6% through polysynaptic corticoreticular connections. By their latent period, duration, and rise time up to a maximum the monosynaptic E PS Ps were divided into two groups: "fast" and "slow." It is postulated that "fast" E PS Ps are generated in reticulospinal neurons which are activated by fast-conducting fibers and "slow" E PS Ps by slowly conducting corticobulbar fibers. I PS Ps were recorded from reticulospinal neurons that also were inhibited by stimulation of the ventral columns of the spinal cord. The hypothesis is put forward that cortical motor signals in cats can be transmitted to the spinal cord via monosynaptic and polysynaptic connections of "fast" and "slow" pyramidal neurons with reticulospinal neurons.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 8, No. 3, pp. 250–257, May–June, 1976.     

Reticular structures and reticulospinal pathways participating in the initiation and inhibition of spino-bulbo-spinal activity

Reflex discharges in intercostal nerves and activity of reticulospinal fibers of the ventral and lateral funiculi, evoked by stimulation of the reticular formation and of the splanchnic and intercostal nerves were investigated in cats anesthetized with chloralose (50 mg/kg). Brain-stem neuronal structures participating in the "relaying" of spino-bulbo-spinal activity were shown to lie both in the medial zones of the medullary and pontine reticular formation and in its more lateral regions; they include reticulospinal neurons and also neurons with no projection into the spinal cord. Structures whose stimulation led to prolonged (300–800 msec) inhibition of reflex spino-bulbo-spinal activity were widely represented in the brain stem, especially in the pons. Analogous inhibition of this activity was observed during conditioning stimulation of the nerves. Reticulospinal fibers of the ventral (conduction velocity 16–120 m/sec) and lateral (17–100 m/sec) funiculi were shown to be able to participate in the conduction of spino-bulbo-spinal activity to spinal neurons. In the first case fibers with conduction velocities of 40–120 m/sec were evidently most effective. Evidence was obtained that prolonged inhibition of this activity can take place at the supraspinal level.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Institute of Normal and Pathological Physiology, Slovak Academy of Sciences, Bratislava, Czechoslovakia. Translated from Neirofiziologiya, Vol. 8, No. 4, pp. 373–383, July–August, 1976.     

Functional properties of propriospinal pathways of the dorsolateral tract of the cat spinal cord

The characteristics of conduction of the excitation wave along propriospinal fibers of the dorsolateral tract of the spinal cord were studied in cats anesthetized with pentobarbital. At a preliminary operation, 10–18 days beforehand, lateral hemisection of the spinal cord was performed, cranially in the lumbar division and caudally and cranially in the cervical division to the segments to be studied, leading to degeneration of the long descending and ascending fibers. During stimulation, the dorsolateral tract developed a composite response consisting of a positive-negative wave recorded up to 60–65 mm (4 or 5 segments) from the point of stimulation. The mean conduction velocity of this wave in the lumbar division was 37.9 m/sec compared with 44.5 m/sec in the cervical division. From its properties as a whole this wave can be regarded as the result of excitation of relatively fast-conducting propriospinal fibers of the dorsolateral tract. If the strength of stimulation was increased, late components began to appear in the response. These were evidently connected with excitation of thinner propriospinal fibers and synaptic activation of other other groups of spinal neurons.A. A. Bogomolets' Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 5, No. 1, pp. 54–60, January–February, 1973.     

Effectiveness of ascending synaptic influences from various spinal funiculi on reticulospinal neurons in cats

Activity of reticulospinal neurons evoked by stimulation of the ventral, ventrolateral, dorsolateral, and dorsal funiculi of the spinal cord was recorded extracellularly in cats anesthetized with chloralose. Responses of 57 reticulospinal neurons, of which 22 projected into the ventral funiculus, 20 into the ventrolateral, and 15 into the dorsolateral, were studied. The functional properties (conduction velocity and refractory period) and the location of the neurons of the above-mentioned groups in the medulla did not differ appreciably. The most effective synaptic activation of all neurons was observed during stimulation of the dorsal and dorsolateral funiculi. Responses to stimulation of the dorsal funiculus had the lowest threshold. These responses arose in reticulospinal neurons of the ventral and ventrolateral funiculi after the shortest latent period. The effectiveness of synaptic influences from the dorsal and dorsolateral funiculi was identical in the group of neurons of the dorsolateral funiculus. Correlation between activity evoked by stimulation of the dorsal funiculus in reticulospinal neurons and peripheral nerves indicated that the responses appeared in these cells to stimulation of muscular (groups I and II) and cutaneous (group II) afferent fibers. The results indicate that impulses from low-threshold muscular and cutaneous afferents, which effectively activate reticulospinal neurons, are transmitted along ascending pathways of the dorsal funiculi.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 11, No. 3, pp. 254–263, May–June, 1979.     

Reticulofugal influences of interneurons in the lateral region of gray matter of the cat spinal cord

Responses of lumbar interneurons located in the most lateral regions of Rexed's laminae IV–VII to stimulation of the medial longitudinal bundle and gigantocellular reticular nucleus of medullary pyramids, red nucleus, and peripheral nerves were investigated in cats anesthetized with pentobarbital. Stimulation of the reticulospinal fibers evoked monosynaptic excitation of many interneurons specialized for transmitting activity of the lateral descending systems, but not of peripheral afferents. Convergence of excitatory influences of all three descending systems (cortico-, rubro-, and reticulospinal) was observed on some cells of this group. In addition, monosynaptic "reticular" E PSPs appeared in interneurons transmitting activity of group Ia muscle fibers and in some interneurons of the flexor reflex afferent system. Stimulation of reticulospinal fibers evoked IPSPs in some neurons of this last group. Neurons not exposed to reticulofugal influences (both specialized neurons and interneurons of segmental reflex arcs) were located chiefly in the dorsal zones of the region studied. Recordings were also obtained from single fibers of the lateral reticulospinal tracts (conduction velocity from 26 to 81 m/sec).A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 5, No. 5, pp. 525–536, September–October, 1973.     

Synaptic activation of thoracic spinal interneurons by reticulospinal pathways

Synaptic processes of 119 thoracic spinal interneurons (T10–11) were investigated in anesthetized cats in response to stimulation of the medial and central zones of the gigantocellular nucleus in the medulla and the ventral columns of the spinal cord. Fast (90–130 m/sec) reticulospinal fibers running in the ventral column were found to produce monosynaptic or disynaptic excitation of interneurons of Rexed's layers VII–VIII, which are connected monosynaptically with group I muscle afferents, and interneurons excited both by group I muscle afferents and low-threshold cutaneous afferents. In most neurons of layer IV, connected monosynaptically with low-threshold cutaneous afferents, and in neurons of layers VII–VIII excited by afferents of the flexor reflex no marked postsynaptic processes were observed during stimulation of the reticular formation. Excitatory, inhibitory, and mixed PS Ps during activation of reticulospinal fibers were found in 14 neurons, high-threshold afferents in which evoked predominantly polysynaptic IPSPs. Seventeen neurons activated monosynaptically by reticulospinal fibers and not responding to stimulation of segmental afferents were found in the medial part of the ventral horn (layers VII–VIII).A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 4, No. 6, pp. 566–578, November–December, 1972.     

Morphological study of the origin of spinal locomotor strip fibers

The origin of spinal locomotor strip fibers was investigated in cats by applying electrical stimulation and the retrograde axonal horseradish peroxidase transport technique. It was found to be mainly composed of corticospinal tract fibers. Moderate numbers of reticulospinal tract and trigeminal spinal tract fibers were also observed. Descending projections from brain stem catecholaminergic neuronal groups do not pass through the test sites of the dorsolateral funiculus, nor, apparently, do they go to make up the spinal locomotor strip. Specificity of the brain stem and spinal locomotor region is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 21, No. 3, pp. 327–335, May–June, 1989.     

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.     

Normal anatomy and physiology of the spinal cord dorsal horn

The dorsal horn of the spinal cord receives afferent input from innocuous primary afferent neurons via collaterals from the dorsal columns. This input is integrated and relayed primarily by neurons in laminae III-VI. Dorsal horn neurons which encode innocuous inputs project to the medulla and the cervical spinal cord via the dorsal columns and the dorsolateral funiculus. Nociceptive primary afferent neurons enter the spinal dorsal horn via collaterals from Lissauer's tract. Nociceptive input is integrated and relayed by neurons in laminae I, II and V which project to the reticular formation and thalamus via the anterolateral tract.     

Effect of nociceptive stimulation of signal transmission from low-threshold cutaneous afferents in the somatosensory system

In cats anesthetized with chloralose nociceptive heating of the skin of the foot to 44–60°C led to a two- to fourfold increase in amplitude of primary cortical responses to direct stimulation of neurons of the spinocervical tract receiving information from the heated area of skin, but did not affect primary responses evoked by stimulation of axons of these neurons in the dorsolateral funiculus, and actually inhibited the response to stimulation of the nerve innervating the heated area of skin. Inhibition was accompanied by depolarization of central terminal of low-threshold fibers of this nerve: During heating the amplitude of the antidromic discharges evoked in the nerve by stimulation of its presynaptic endings in the spinal cord was increased two- to threefold. After abolition of presynaptic depolarization with picrotoxin (0.2–0.7 mg/kg, intravenously) or as a result of asphyxia, nociceptive heating acquired the ability to facilitate primary responses arising as a result of stimulation of the nerve also. The amplitude of the responses was increased under these circumstances by 3–20 times. It is concluded that acute nociceptive stimulation causes such powerful presynaptic inhibition of impulse transmission from low-threshold fibers of the cutaneous nerve that it virtually abolishes the facilitating effect of nociceptive impulses on sensory neurons of the spinal cord. It is suggested that it is this inhibitory mechanism which prevents the development of hyperalgesia during acute nociceptive stimulation.Institute of General Pathology and Pathological Physiology, Academy of Medical Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 13, No. 6, pp. 621–627, November–December, 1981.     

Connections of the corticospinal and rubrospinal tracts with neurons of the cervical spinal cord in cats

The distribution of focal potentials over the cross section of the 7th cervical segment of the spinal cord was studied during stimulation of the pyramids, the red nucleus, and a peripheral nerve (ulnar) in adult cats anesthetized with chloralose and Nembutal. The earliest focal potentials in the fasciculus dorsolateralis were recorded 1.4–1.5 msec after stimulation of the pyramids and 0.8–0.9 msec after stimulation of the red nucleus. These times correspond to maximal condution velocities of 56–68 and 105–124 m/sec respectively. The earliest post-synaptic activity in response to pyramidal stimulation was found in the lateral areas of laminae V and VI, and in response to stimulation of the red nucleus in laminae VI and VII in Rexed's classification. The pyramidal wave also evoked considerable postsynaptic activity in medial areas of the dorsal horn. In response to stimulation of peripheral afferents activity was evoked in neurons in the central and medial parts of laminae V and VI. It is postulated on the basis of these results that corticospinal and rubrospinal fibers may be connected monosynaptically with specialized interneurons, free from peripheral influences, in the lateral areas of laminae V and VII respectively; in the lateral part of lamina VI convergence of both types of influences on the same cells is possible. Interaction between descending and afferent influences possibly takes place on more medially located neurons.A.A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 4, No. 2, pp. 158–167, March–April, 1972.     

Responses of pyramidal tract and corticorubral neurons to stimulation of lateral hypothalamic structures in cats

Responses of pyramidal tract neurons and corticorubral and unidentified neurons in the pericruciate area of the cortex to electrical stimulation in the posterior, tuberal, and anterior zones of the lateral hypothalamus and also to electrodermal stimulation of all four limbs were studied in cats anesthetized with chloralose. The proportion of pyramidal tract, corticorubral, and unidentified neurons which responded to hypothalamic stimulation was 73.3, 55.7, and 79.1% respectively. Data on the possibility of monosynaptic activation of some pyramidal tract neurons and unidentified cells were obtained. The presence of less stable and longer-latency responses of corticorubral neurons indicated the absence of a monosynaptic pathway for realization of ascending hypothalamic influences on neurons of the cortical extrapyramidal system. Some cortical neurons responded to stimulation of more than one zone of the hypothalamus. Pyramidal tract neurons and corticorubral neurons with axon collaterals running into the region of the hypothalamus were discovered for the first time. It was shown that most neurons of the pericruciate area of the cortex to which the ascending influence of the hypothalamus is directed have a large bilateral receptive field and respond to electrodermal stimulation of several limbs.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 14, No. 3, pp. 298–306, May–June, 1982.     

Synaptic activation of thoracic interneurons by reticulospinal fibers of the lateral funiculus

Synaptic responses of different functional groups of interneurons in segments T10 and T11 to stimulation of the ipsilateral and contralateral medullary reticular formation were investigated in anesthetized cats with only the ipsilateral lateral funiculus remaining intact. Activation of reticulospinal fibers of the lateral funiculus with conduction velocities of 30–100 m/sec was shown to induce short-latency and, in particular, monosynptic EPSPs in all types of cells tested: in interneurons excited by group Ia muscle afferents, in cells activated only by high-threshold cutaneous and muscle afferents (afferents of the flexor reflex), in cells activated mainly by descending systems, and, to a lesser degree, in neurons connected with low-threshold cutaneous afferents. These cell populations are located mainly in the central and lateral parts of Rexed's lamina VII. Most neurons in laminae I–V of the dorsal horn, except six cells located in the superficial layers of the dorsal horn, received no reticulofugal influences. The functional organization of connections of the lateral reticulospinal tract with spinal neurons is discussed and compared with the analogous organization of the medial reticulospinal tract, and also of the "lateral" (cortico- and rubrospinal) descending systems.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 10, No. 2, pp. 150–161, March–April, 1978.     

Interaction of direct afferent and recurrent signals in cat motor cortex neurons

Intracellular activity was recorded from the functionally identified motor cortex neurons (MI, area 4) in acute experiments on myorelaxin-immobilized cats under calypsol anesthesia. Changes in neuronal responses to testing stimulation of the ventrolateral thalamic nucleus or pyramidal tract fibers were studied; the same or another input was used for a conditioning stimulation. Excitatory and inhibitory components of test responses of variousMI neurons were found to be either facilitated or depressed. The facilitation of orthodromic excitation was more frequent in the case of thalamic testing stimulation. The depression of both excitatory and inhibitory components of the response was more pronounced with paired stimulation of the pyramidal tract fibers. The peculiarities of interaction between direct afferent and recurrent signals in theMI neurons are thought to be determined by different distribution of thalamocortical fiber terminals and recurrent collaterals of corticofugal axons in the cortex and nonuniform localization of their synapses on dendrites and somata of the studied cells. It seems possible that these peculiarities also are connected with different chemical mechanisms of synaptic transmission in the above synapses and different properties of postsynaptic membrane receptors.Neirofiziologiya/Neurophysiology, Vol. 26, No. 3, pp. 203–210, May–June, 1994.     

Morphology of lumbar-projecting lateral vestibulospinal neurons in the brainstem and cervical spinal cord in the squirrel monkey

The lateral vestibulospinal tract (LVST) is one of the major descending pathways controlling the extensor musculature of the body. To determine whether individual LVST neurons terminating in the lumbosacral spinal segments issue collaterals more rostrally to exert an influence of the cervical ventral horn intracellular recording and biocytin labeling techniques were used in the squirrel monkey. Only neurons monosynaptically related to the 8th nerve and antidromically identified to project below T12 were selected for study. The axon course through the brainstem and cervical spinal cord was examined in 37 LVST neurons. The average distance of recovered axon was 17.3 mm (4.5-31.7 mm). None could be antidromically activated from shocks applied to the rostral medial longitudinal fasciculus near the 3rd nuclei; and no collaterals were observed in the brainstem. Of the 37 neurons, only 1 axon issued a collateral to innervate the ventral horn, primarily in the region of the spinal accessory motoneurons; this single collateral provided a relatively minor input compared to that of LVST neurons terminating in the cervical cord. Thus, secondary, caudal-projecting LVST neurons represent a private, and mostly rapid, communication pathway between dorsal Deiters' nucleus and the motor circuits controlling the lower limbs and tail.     

Medullary reticular neurons in the Japanese toad: morphologies and excitatory inputs from the optic tectum

1. To elucidate the neural mechanisms that mediate visual responses of optic tectum (OT) to medullary and spinal motor systems, we analyzed medullary reticular neurons in paralyzed Japanese toads (Bufo japonicus). We examined their responses to electrical stimulation of OT, and stained some neurons intracellularly. Responses to stimulation of the glossopharyngeal nerve (IX) were also analyzed. 2. Extracellular single unit recording revealed excitatory responses of medullary neurons to OT and IX stimulation. Among 92 units encountered, 79 responded to OT stimuli, 10 to IX stimuli, and 3 to both. Some units responded to successive stimuli of short intervals with relatively stable lags. 3. Intracellular recording and staining experiments revealed morphologies of reticular neurons that received excitatory inputs from OT. Thirteen units were identified after complete reconstruction of somata and dendrites. Neurons in the nucleus reticularis medius received excitatory inputs from bilateral OT. They had wide dendrites in ventral, ventrolateral and lateral funiculi, and single axons descending in the ipsilateral ventral funiculus as far caudally as the cervical spinal cord. Some collaterals of these axons projected directly to the hypoglossal and spinal motor nuclei. Some neurons in other medullary nuclei (nuc. reticularis superior, pretrigeminal nucleus, nuc. reticularis inferior, and nuc. tractus spinalis nervi trigemini) also responded to the OT stimulation. 4. Activities in bilateral OT converge onto medullary reticular neurons, which may directly control medullary and spinal motor systems.     

Reticular pathways transmitting corticofugal impulses

Details of the organization of reticular pathways transmitting motor cortical influences were studied in cats anesthetized with chloralose. Of all reticular neurons studied 33.3% were reticulospinal cells, 12.2% were neurons with descending-ascending projection of their axons, 15.4% were purely ascending neurons, and 39.1% were unidentified cells. Analysis of responses evoked by cortical stimulation showed that influences of both fast- and slowly-conducting corticofugal fibers are transmitted through reticulospinal cells, neurons with descending-ascending projections, and cells projecting into the hypothalamus. It was shown that 37% of reticulospinal and 66.7% of neurons with two-way projection that were studied form a group of cells transmitting rapidly into the spinal cord impulses from slowly-conducting fibers only. Reticular neurons with projections to the thalamus also transmit influences of slowly-conducting fibers only. The organization and role of reticular pathways transmitting corticogugal impulses are discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 13, No. 5, pp. 491–499, September–October, 1981.     

Study of pathways transmitting sympathetic-activating influences from hypothalamus to spinal cord

The properties of hypothalamic pathways activating sympathetic preganglionic neurons in the lateral horns of the spinal cord were studied in cats. This activation was shown to take place through reticulospinal sympathetic-activating neurons in the medulla. The possible zone of location of hypothalamic sympathetic-activating output neurons was shown by the scanning stimulation method to be in the posterolateral hypothalamus. Reticulospinal sympathetic-activating neurons in the medulla are probably excited monosynaptically by hypothalamo-sympathetic activating fibers projected on them.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 14, No. 3, pp. 307–314, May–June, 1982.     

Localization and properties of reticulospinal neurons with axons descending in the dorsolateral parts of the lateral funiculi

The localization of reticulospinal neurons responding antidromically to stimulation of fibers in the dorsolateral parts of the lateral funiculi (shown previously to be the principal collector of fibers conveying bulbar pressor influences) was determined in experiments on anesthetized and curarized cats. Most of these neurons were found to occupy the medioventral portions of the medulla, but they were concentrated in the rostral portions of the gigantocellular and ventral nuclei of the reticular formation. The velocity of conduction of excitation along axons of most reticulospinal neurons was 10–50 m/sec. Reflex responses to stimulation of the sciatic nerve with a latent period of 10–40 msec were found in 35 of 125 such cells. Stimulation of the sinus nerve did not activate them. Spontaneous activity occurred in 29 reticulospinal neurons; the mean firing rate of the various cells varied from 5 to 20/sec.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 6, No. 3, pp. 266–272, May–June, 1974.     

Projection linkage from spinal neurons to both lateral cervical nucleus and solitary tract nucleus in the cat

Intracellular recordings from the lumbosacral dorsal horn were made to identify the axonal projection and the afferent innervation of the lateral cervical nucleus (LCN) and solitary tract nucleus (STN) on the spinal neurons of chloralose-anesthetized cats. A total of 49 neurons from laminae III-V in the spinal dorsal horn responded to stimulation of both the LCN and STN. Of these, 28 and 21 neurons responded antidromically and orthodromically to stimulation of the LCN and STN, respectively. Seven of the 28 antidromically activated neurons were followed by one or more responses synaptically driven from the LCN and/or STN. The diameter of these ascending or descending fibers was in the range of A delta fibers. The results indicate that (1) some spinal neurons, namely spinocervical tract-spinosolitary tract (SCT-SST) neurons, issue branched axons of A delta-fibers and dually project to both LCN and STN; (2) some SCT-SST neurons receive innervation from both the LCN and STN; (3) some spinal neurons and interneurons are dually innervated by descending fibers originating from both the LCN and STN, and (4) the convergence and integration between somatic and visceral sensory inputs might occur in the SCT-SST neurons.