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.     

Inhibitory effects of reticulospinal fibers of the lateral funiculus on neurons of thoracic spinal reflex pathways

Experiments on anesthetized cats with partial transection of the spinal cord showed that reticulo-spinal fibers in the ventral part of the lateral funiculus participate in the inhibition of polysynaptic reflexes evoked by stimulation of the ipsi- and contralateral reticular formation. The reticulo-fugal wave in the ventrolateral funiculus evoked comparatively short (up to 70 msec) IPSPs in some motoneurons of the internal intercostal nerve investigated and at the same time evoked prolonged (up to 500 msec) inhibition of IPSPs caused by activation of high-threshold segmental afferents. This wave also led to the appearance of IPSPs in 14 of 91 (15.5 %) thoracic spinal interneurons studied. The duration of these IPSPs did not exceed 100 msec; meanwhile, segment excitatory responses of 21 of 43 interneurons remained partly suppressed for 120–500 msec. It is concluded that the inhibitory action of the lateral reticulo-spinal system on segmental reflexes is due to several synaptic mechanisms, some of them unconnected with hyperpolarization of spinal neurons. The possible types of mechanisms of this inhibition are discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 10, No. 2, pp. 162–172, March–April, 1978.     

Peculiarities of the synaptic adaptation of the intermediate neurons of the thoracic segments of the spinal cord by cutaneous,muscle, and visceral afferents

The responses of the interneurons of the thoracic segments of the spinal cord to stimulation of the intercostal and splanchnic nerves were studied on decerebrated and narcotized cats. It was established that the neurons of different layers of the gray matter (according to Rexed) differ substantially in type of afferent inputs. Cells in laminae I–III and IV are activated chiefly by somatic afferents: primarily high-threshold in laminae I–III and low-threshold in lamina IV. The neurons of lamina V and most of the neurons of laminae VII and VIII respond to stimulation of high-threshold somatic afferents (cutaneous fibers of the A group and muscle afferents of groups II and III), as well as visceral afferents of group A, conducting impulses at a rate of 9–35 m/sec. Cells of laminae VII and VIII, monosynaptically activated by muscle afferents of group I, do not respond to stimulation of the visceral afferents. The peculiarities of the "functional" laminar organization of the thoracic segments of the spinal cord are discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 2, No. 6, pp. 563–572, November–December, 1970.     

Effects of vasopressin and oxytocin on evoked dorsal horn cell activity in an isolated segment of spinal cord

During experiments on an isolated segment of the spinal cord of 2- to 3-week-old rats, a study was made of the effects of vasopressin and oxytocin on the activity of dorsal horn cells produced by stimulating the afferent root. Both field and action potentials were recorded in single cells. It was established that vasopressin and oxytocin produced reversible inhibition of the postsynaptic component of field potentials. The amplitude of potentials was reduced by 33–39% by vasopressin and by 12–34% using oxytocin. The effect of the test substances depended on the concentration used and the duration of their action on the brain. Both vasopressin and oxytocin reversibly depressed discharges of single dorsal horn cells evoked by stimulating the dorsal root. These two neuropeptides prolonged latency, and reduced the number of evoked potentials or completely suppressed response. A facilitatory effect was recorded in a small number of cells. We deduced from our findings that their hypothalamospinal neurohormonal system inhibits transmission of afferent impulses at the level of interneurons of the dorsal horn.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 17, No. 5, pp. 634–640, September–October, 1985.     

Depolarization of primary afferents during fictitious scratching in thalamic cats

The dorsal cord and dorsal root potentials were recorded in immobilized thalamic cats during fictitious scratching evoked by mechanical stimulation of the ear. Depolarization of primary afferents was shown to be simulated by the central scratching generator. Antidromic spike discharges appeared at the peak of the primary afferent depolarization waves in certain afferent fibers. Similar discharges arise in the resting state in response to stimulation of limb mechanoreceptors. It is suggested that during real scratching primary afferent depolarization and antidromic spikes evoked by it may effectively modulate the level of the afferent flow to spinal neurons.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 10, No. 2, pp. 173–176, March–April, 1978.     

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.     

Scratch reflex evoked by strychnine applied to the spinal cord

Effective facilitation of the scratch reflex was observed in experiments on recently decerebrated and spinal cats in response to application of strychnine solution to the dorsal surface of the spinal cord. The receptive fields of the scratch reflex in this case depended on the segments to which the strychnine was applied. The receptive fields of a scratch reflex evoked by D-tubocurarine coincided with those of the intact adult animals, but the receptive fields for strychnine application were wider and corresponded to those of the scratch reflex in kittens [1, 10]. It is suggested that strychnine application abolishes the inhibition of the afferent inputs of the scratch reflex in the lower cervical segments of the spinal cord which has developed during ontogeny.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 10, No. 6, pp. 622–625, November–December, 1978.     

Synaptic processes in neurons of the cervical enlargement of the cat spinal cord during stimulation of propriospinal pathways of the dorsolateral tract

Responses of motoneurons and interneurons of the cervical enlargement of the cat spinal cord were studied by a microelectrode technique during selective stimulation of propriospinal fibers of the dorsolateral tract of the lateral white column. The long descending and ascending pathways were blocked by preliminary (10–16 days earlier) hemisection of the spinal cord cranially and caudally to the segments studied. Stimulation of the dorsolateral tract at a distance of 15–25 mm from the site of recording evoked complex postsynaptic potentials consisting of several successive waves in the motoneurons. The character of the PSPs was not clearly linked with the function of the motoneurons. By their latent periods the components of the PSPs could be placed in three groups. The "primary" components were reproduced in response to stimulation at 50–100/sec whereas the "secondary" and "tertiary" components were weakened or blocked. It is postulated that the "primary" components are evoked through monosynaptic connections between propriospinal fibers of the dorsolateral tract and motoneurons of the forelimb muscles, while the late components are evoked through polysynaptic pathways, including segmental interneurons. Many of these interneurons, located in the ventral horn and intermediate zone, were strongly excited during stimulation of the dorsolateral tract.A. A. Bogomolets' Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 5, No. 1, pp. 61–69, January–February, 1973.     

Neurochemical mechanisms in inhibitory effects of stimulation of the periaqueductal gray matter on high-threshold reflex activity of the reticular formation

A study was made of microinjections of antagonists of various neuromodulators on the dynamics of inhibition of the spino-bulbo-spinal reflexes which were evoked by stimulation of the central gray matter (PAG) in rats anesthetized with chloralose. Injections were made into the reticular gigantocellular nucleus (GN), which is the basic supraspinal center of this reflex. Administering methysergide (a blocker of serotonin receptors) was accompanied by significant (two to four times) diminution of inhibition evoked by PAG stimulation with a short, high-frequency series of stimuli. Long inhibition caused by long, rhythmic stimulation of the PAG was diminished less significantly: from 6–10 to 2.5–4 min. When the opiate receptors of the GN neurons were blocked with naloxone, duration of inhibition was reduced by two to five times. The most clearly expressed diminution of both types of inhibitions was noted with injections of haloperidol, an antagonist of catecholamines. Our data indicate that evidently all of these neuromediator (neuromodulator) systems participate in inhibition of high-threshold, reflex activity of the reticular formation evoked by stimulation of the PAG, but their participation in this process is unequal.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 23, No. 4, pp. 455–463, July–August, 1991.     

Analysis of the organization of afferent inputs in the projection cortex

Fast fluctuations in the evoked potentials (EP) at a local point of the projection cortex following stimulation of different pathways may reflect the activity of pyramidal neurons of different cortical layers. Analysis shows that the afferent and interarea projections to the somatic sensory cortex terminate on different neurons which can be regarded as relay neurons for a given pathway. Each group of neurons has its own system of inhibition for selective control of impulses coming along this pathway at the cortical level.Institute of Normal and Pathological Physiology, Academy of Medical Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 2, No. 4, pp. 368–372, July–August, 1970.     

Primary afferent depolarization in the cervical enlargement of cats evoked by activation of cervical sensory nerves

Depolarization of primary afferent terminals in the cervical enlargement of the spinal cord evoked by activation of sensory nerves of the upper cervical segments (C₂) was studied in cats anesthetized with pentobarbital. It was shown that low-threshold muscular and high-threshold cutaneous afferents of nerves of the forelimb were depolarized most strongly. Parallel with this depolarization, prolonged (over 0.5 sec) inhibition of the monosynaptic and polysynaptic flexor reflex developed. It is suggested that these influences are transmitted via pathways running in the posterior and lateral white columns. The results are discussed in connection with regulation of postural motor activity in vertebrates.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 14, No. 2, pp. 190–197, March–April, 1982.     

Projections of afferent ventral root fibers on dorsal horn neurons in the cat spinal cord

Mechanisms of the effect of stimulation of afferent fibers in ventral roots on dorsal horn interneurons were investigated in experiments on anesthetized cats. Dorsal horn interneurons on which such fibers project were shown to exist. In particular, some dorsal horn interneurons can exert an inhibitory influence on effects of dorsal root fiber activation.Institute of Physiology, Academy of Sciences of the Kazakh SSR, Alma-Ata. Translated from Neirofiziologiya, Vol. 17, No. 3, pp. 300–305, May–June, 1985.     

Interaction between excitation and inhibition in presumptive hippocampal inhibitory interneurons

Electrical responses of 25 presumptive hippocampal inhibitory interneurons to stimulation of two afferent systems of fibers, originating in the contralateral hippocampus, were investigated in acute experiments on unanesthetized, immobilized rabbits. Inhibitory neurons were found to have a relatively ineffective inhibitory input as well as a very effective excitatory input. On interaction between synaptic processes during spontaneous and evoked activity the excitatory input clearly predominates over the inhibitory and plays a definite role in behavior of the neurons.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 12, No. 6, pp. 580–587, November–December, 1980.     

Influence of afferent impulses from the nerves of the anterior limbs on the insertion neurons of the lumbar section of the spinal cord

In anesthetized cats in conditions of muscular relaxation we have studied the participation of the interneurons of the lumbar section of the spinal cord in the interaction of the FRA systems of the fore and hind limbs. Using microelectrodes we have made extra- and intracellular recordings of the potentials. It has been shown that from the flexor afferents of the fore limbs both facilitating and inhibitory influences are transmitted. The former are expressed in increased frequency of the background impulse activity of the neurons, in the appearance of evoked responses of the "silent" cells and intensification of the test responses for short time intervals with paired heteronymous stimulation. The inhibitory influences prevail over the facilitating and are manifest in depression of the background activity and evoked segmental responses of the neurons. The maximum inhibition of the segmental responses was noted for intervals of 40–140 msec. The duration of inhibition varied from 100–500 msec and more. Depending on the intensity and duration of the inhibitory influences two groups of interneurons have been isolated. The role of the pre- and postsynaptic mechanisms in the transmission of inhibitory influences from the afferents of the fore limbs on the afferents of the hind limbs is discussed.Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 1, No. 3, pp. 235–242, November–December, 1969.     

Visceral afferent pathways relaying synaptically in the caudal mesenteric sympathetic ganglion

The caudal mesenteric sympathetic ganglion of cats was isolated and perfused, and responses of the preganglionic trunks of the ganglion to electrical stimulation of the central end of the hypogastric nerve were studied. Stimulation of the nerve with single square pulses gives rise to early and late responses. Early responses appear after various latent periods and are the result of excitation of transit fibers of groups A, B, and C, whereas the appearance of late responses is associated with the synaptic transmission of excitation in the sympathetic ganglion from afferent sympathetic neurons at the first level (from the pelvic organs to the caudal ganglion) to afferent sympathetic neurons of the second level (from the caudal ganglion and above). Early responses are not blocked, but late responses are blocked by perfusion of the ganglion with azamethonium bromide and magnesium salts, and also by tetanization of the hypogastric nerve at 10–50 Hz. Other facts indicating the synaptic relaying of visceral sympatho-sympathetic afferent pathways in the ganglion are also described.Institute of Physiology, Academy of Sciences of the Belorussian SSR, Minsk. Translated from Neirofiziologiya, Vol. 2, No. 5, pp. 507–514, September–October, 1970.     

Investigation of spinal neuronal mechanisms of movement control systems

Morphological and electrophysiological investigations of the means whereby the principal descending motor systems (the cortico-, rubro-, reticulo-, and vestibulo-spinal tracts) are connected with the segmental interneuronal apparatus and motoneurons show that these connections can be based on two different principles. Descending systems either activate motoneurons directly (monosynaptically) or are connected primarily with various interneuron systems, exerting their influence in that case by regulating the activity of simpler or more complex spinal mechanisms. The older descending system (reticulo- and vestibulo-spinal) possess a monosynaptic excitatory action of motoneurons; the evolutionarily newer descending systems, which transmit the most complex motor signals from the cerebral and cerebellar cortex to the spinal cord (cortico- and rubro-spinal), terminate synaptically in every case on interneurons. It is only in primates that a few cortico-spinal fibers form monosynaptic connections with motoneurons. The chief ways of action of the descending systems on interneurons are: control of the afferent inflow into the interneuron system by presynaptic inhibition of the corresponding synapses; control of the interneuron system by postsynaptic interaction with afferent influences; control of motoneurons through the specialized interneuron apparatus. The investigation shows that the last of these mechanisms functions in the cortico- and rubro-spinal, and possibly also in the reticulo- and vestibulo-spinal systems. The functional role of the various means of connection of the descending systems with the spinal neurons in the system of movement control is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 2, No. 2, pp. 189–202, March–April, 1970.     

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

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

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.     

Evoked activity in the ventral horn of the lumbar segments of the spinal cord during cortical inhibition of cortical motor responses

Electrical activity in the flexor nerve and focal potentials (FP) in the medial and lateral zones of the ventral horn (VH) of segments L₆ and L₇ of the spinal cord, evoked by excitation of the contralateral motor cortex, were recorded in delicate experiments on cats. These focal potentials were studied during inhibition of the flexor response that developed as a result of prior excitation of the ipsilateral cortex ("cortical inhibition"). During the inhibition the FP's of the medial zone (layer VIII, according to Rexed) were greatly increased, mainly in their negative components, their time-characteristics being altered. When the interval between excitations was 50 msec (in that case the inhibition was most pronounced) the medial FP's arose against a negative background, which was a late component of the previous activity evoked by conditioning excitation. The appearance of this late component was correlated with the development of inhibition of the cortical flexor response. At the same time a positive condition developed in the lateral zone, in the region of the nucleus biceps-semitendinosus, which indicated orientation in a lateral direction of the interneurons discharging in the medial zone at late periods after the conditioning excitation. Inhibition of the flexor response was accompanied by depression of the lateral FP's without change in their sign or in the time-structure of their components. It is suggested that cortical inhibition of the cortical flexor response arises at the interneuron level. The functional structure of that inhibitory pathway is discussed.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 3, No. 2, pp. 185–193, March–April, 1971.     

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.