Quantitative changes in efferent activity of the scratch generator produced by eliminating influences from the descending systems

The effects of signals proceeding along descending systems of the spinal cord on the parameters of efferent activity in the scratch generator (i.e., discharges in ventral roots L₅ and L₇) were investigated in decerebrate immobilized cats. It was found that eliminating cerebellar modulatory influences on nuclei of the descending system (decerebellation) leads to an altered ratio between the activity of generator hemicenters, increased instability of generator operation, a reduced statistical relationship between alteration in parameters of hemicenter activity, and raised sensitivity to afferent signals in the generator. Complete elimination of descending system influences (after severing the spinal cord at the anterior boundary of the first cervical segment) produced a reduction in the intensity of efferent activity and in the afferent sensitivity of the scratch generator. The functional significance of descending signals for operation of the scratch generator is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 22, No. 1, pp. 123–131, January–February, 1990.     

Analysis of the mechanisms of interaction between generators of cyclic motor reactions and inputs from suprasegmental systems

The simulation mathematical model of neuronal generator systems was used for analyzing the interaction between inputs from descending (afferent) systems and generators of scratching (locomotion). The data obtained indicate that in cases when generators of cyclic motor reactions influence the effectiveness of synaptic transmission from the fibers of descending (or afferent) systems, the differences in the main characteristics of signals that are produced by these generators themselves and those that come to them are emphasized. These data allow us to conclude that the system supplying interaction between inputs from suprasegmental (afferent) fiber systems and generators of locomotion of scratching can be interpreted as an adaptive filter which processes spatial and temporal information coming to the spinal cord via different suprasegmental or primary afferent inputs and allows generators of cyclic motor reactions to correct their functioning in accordance with changing external conditions.Neirofiziologiya/Neurophysiology, Vol. 25, No. 3, pp. 211–215, May–June, 1993.     

Possible organization pattern of interaction between the generators of cyclic motor activity and inputs from supraspinal and afferent systems

Possible organization patterns of scratching and locomotor generators that allow interpretation of experimentally demonstrated reorganizations in temporal parameters of these generator activities after electrical stimulation of descending and peripheral afferent systems were analyzed with application of mathematical simulation of neuronal generator systems. The results obtained led to the conclusion that patterns of such reorganizations influenced by signals from suprasegmental and/or peripheral systems may be determined by only two factors: 1) the structure of synaptic connections between interneuronal functional groups underlying these generator associations, and 2) the structure of connections between these groups of interneurons and fibers from suprasegmental and peripheral afferent sources. The existence of inhibitory-excitatory actions from descending and afferent systems upon the neurons of locomotor or scratching generator half-centers is a sufficient condition to ensure phasic changes in the sensitivity of these generators to supraspinal and afferent signals. The locomotor generator, unlike the scratching generator, is apparently characterized by a more complex organization of connections between functional neuronal groupings and descending fibers.Translated from Neirofiziologiya, Vol. 25, No. 1, pp. 45–50, January–February, 1993.     

Alterations produced in the quantitative characteristics of efferent activity parameters of the locomotor generator by decerebellation

Changes in the parameters of activity in hindlimb locomotor generators following decerebellation were quantified during experiments on decerebrate immobilized cats. Eliminating modulating cerebellar influences on nuclei of descending systems was found to lead to a slight increase in the length of activity in the flexor generator half-center and less intensive activity, as well as shortening of the period and more intensive activity in the extensor half-center, together with increased instability in generator operation, reduced statistical dependence between alterations in parameters of activity at the hindlimb half-center generators, and finally intensified effects of afferent inputs on generator activity. A comparison is drawn between the functional role of the spino-cerebellar loop in the operation of locomotor and scratch generators.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 23, No. 2, pp. 142–150, March–April, 1991.     

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.     

Rearrangement of scratch generator efferent activity during electrical activation of descending systems

Effects of signals proceeding along descending pathways on parameters of scratch generator activity were investigated in decerebrate immobilized cats. Certain phase-linked alterations in these parameters were shown to occur under the effects of electrical activation of the main descending systems. The biggest increase in scratch cycle duration under electrical stimulation of Deiter's nucleus, the red nucleus, and pyramidal tract is produced when stimuli are applied during the first half of the aiming stage. Stimulation during the second half of the aiming phase and at the start of the scratching movement hardly affect the scratching cycle. The main increase in length of scratch cycle during electrical stimulation of the reticular gigantocellularis nucleus is noted when stimuli are presented during the second half of the aiming stage. Electrical activation of descending pathways during the latter induces a rise in intensity in this phase and reduced intensity of the actual scratching stage. Activation of the pathways during this scratching motion causes heightened intensity of the motion while hardly affecting intensity of the aiming phase. The principles of suprasegmental rectification of scratch generator operation are discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 22, No. 3, pp. 300–309, May–June, 1990.     

Rearrangement of efferent activity of the locomotor generator under electrical activation of descending systems in immobilized cats

The effects of signals travelling through vestibulo-, rubro-, reticulo-, and corticospinal systems on the parameters of locomotor generator activity were investigated in decerebrate immobilized cats. Certain phase-linked alterations in these parameters were found to occur under the effects of electrical stimulation applied to these systems (brief trains of stimuli). The biggest increase in locomotor cycle length was produced by electrical stimulation of Deiter's nucleus — stimulus presentation at the end of the extension phase; stimulation at the flexor stage leads to a shortening of this cycle. Maximum increase in locomotor cycle length produced by electrical stimulation of the red nucleus and nucleus gigantocellaris reticularis together with the pyramidal tract takes place during the first half of the flexion phase. Electrical activation of these descending pathways during the flexion phase induces intensification of this phase and reduced intensity of the extension phase. Activation of the vestibulospinal tract produces increased and reduced intensification of the extension phase respectively. Principles of suprasegmental correction of locomotor and scratch generators are compared.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 23, No. 2, pp. 151–160, March–April, 1991.     

Activity of lumbar interneurons during fictitious locomotion in thalamic cats

Unit activity of the lumbar interneurons was recorded in thalamic cats during fictitious locomotion. Neurons whose activity was modulated in the rhythm of fictitious locomotion were found in the lateral parts of the intermediate zone of gray matter and ventral horn. Of these neurons, 41.2% were activated mainly in the phase of "flexion," 48.5% in the phase of extension, and 10.3% in both phases. Neurons with tonically increasing or decreasing activity during rhythmic discharges and neurons whose activity was unchanged during fictitious locomotion also were observed. During later discharges all these neurons were similarly activated, although a depth of modulation of unit activity was lower than during fictitious locomotion. Afferent inputs to the recorded interneurons also were studied. The neuronal organization of the spinal locomotor generator is discussed on the basis of these results.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 11, No. 4, pp. 329–338, July–August, 1979.     

Limb movement,intensity of integral afferent impulses from limb receptors,and level of primary afferent terminal polarization during locomotion in cats decerebrated at high level

The relationship between parameters of electrical muscle activity, changes at hindlimb joint angles, intensity of integral afferent flow, and dorsal root potential during real-life locomotion was investigated in cats decerebrated at high level. Characteristics of rear limb movements before and after deafferentation were described. It was found that afferent activity during locomotion motion consists, of two components — a tonic and a periodic phasic stage. Three main waves may be distinguished in the latter, each of which gives rise to associated changes in the level of primary afferent terminal polarization. These changes in turn are summated with the effects produced by the central generator. Correlations, between the parameters of these processes were investigated and the mechanisms underlying afferent control of locomotion generator function discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 20, No. 1, pp. 119–127, January–February, 1988.     

Neuronal activity of the lateral vestibular nucleus of the guinea pig evoked by tilting the animal about the longitudinal axis during locomotion

In experiments on decerebrate guinea pigs, the impulse activity of neurons of the lateral vestibular nucleus evoked by tilting the animal about the longitudinal axis was investigated under conditions of spontaneous and mesencephalon stimulation-evoked locomotor activity. In most investigated neurons, locomotor activity led to changes in their responses to adequate vestibular stimulation. The dominant reaction was intensification of such responses, which was observed in almost all vestibulospinal neurons and in 2/3 of cells not having descending projections. Responses were suppressed only in 1/4 of the neurons not projecting to the spinal cord. The changes in the evoked responses had an amplitude character; the lag of the changes in the discharge frequency relative to the acceleration that caused them was constant. It is suggested that intensification of dynamic reactions of vestibular neurons during locomotion provides maintenance of the animal's equilibrium during movements in space by various gaits and along different trajectories.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 23, No. 5, pp. 541–549, September–October, 1991.     

Polarization of primary afferent terminals in the lumbar spinal cord during fictitious locomotion

Changes in electrical polarization of primary afferent terminals in the lumbosacral portion of the spinal cord were investigated during fictitious locomotion in immobilized decorticated and spinal cats. Fictitious locomotion was accompanied by stable hyperpolarization of the afferent terminals, against the background of which they were periodically depolarized in rhythm with efferent activity. These tonic and phasic changes were observed in terminals of all groups of afferent fibers tested: cutaneous and muscular (Ia and Ib). Periodic in-phase depolarization was carried out in different ipsilateral segments of the lumbosacral enlargement. During ficitious galloping changes in depolarization of the primary efferents were in phase on different sides; during fictitious walking, these periodic changes were out of phase. On the basis of these results the physiological importance of changes in electrical polarization of primary afferent terminals of the spinal locomotor generator is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 12, No. 5, pp. 481–489, September–October, 1980.     

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.     

Brainstem pathways of the initiation of locomotion

Conclusion This overview of the brainstem pathways of the initiation of locomotion can be summarized as follows. There are locomotor regions (hypothalamic and mesencephalic) whose stimulation leads to the appearance of rhythmic stepping movements. These regions are nonuniform in composition: transient fibers as well as cells are found. The locomotor effects of electrical stimulation of these regions can largely be explained on the basis of the presence of efferent projections of the neurons to the medial reticular formation, as well as the activation of the transient fibers of other brain systems. The ventromedial parts of the reticular formation of the medulla oblongata, including areas of the macrocellular and (to a lesser extent) gigantocellular nuclei, form the final element in the suprasegmental system of locomotion initiation. The inconclusive data of different scientists who have used chemical microinjections into the locomotor regions make it impossible at present to specify precisely the neurochemical mechanisms underlying the initiation of locomotion. NMDA has been found to play an important role. The activation of the reticular formation during the triggering of stepping movements can take place either from the locomotor regions or by means of signals coming from the collaterals of the ascending sensory tracts. The wide spectrum of possible pathways of the initiation of locomotion apparently affords the organism a choice of ways by which to realize this process and is an important factor in its adaptation to its environment.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 23, No. 4, pp. 488–505, July–August, 1991.     

Mathematical model of activity of spinal generators producing rhythmic movement

Different organizational arrangements of scratching and locomotor rhythm generators were simulated by a computer-aided mathematical model. A functional group of neurons (a hemicenter constructed on the basis of a stochastically arranged neuronal network) served as the basis for the generator. Several organizational arrangements of scratching and locomotor rhythm generators are considered: two hemicenters with reciprocal inhibitory connections and tonic excitatory influences on both; two hemicenters with inhibitory-excitatory connections and tonic excitatory influences on only one of these; circular structures consisting of more than two functional groups of neurons with excitatory and inhibitory connections between them. All these arrangements would allow for generation of rhythmic activity with a similar time course to that of scratching and locomotor rhythm. It was found that the transition from locomotor to scratching rhythm could be based on fairly simply organized effects on generator neurons. Principles possibly guiding the construction of spinal generators of scratching and locomotor movements are discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 20, No. 5, pp. 586–597, September–October, 1988.     

Transmission of descending activity evoked by prolonged stimulation of the pyramids and red nucleus by groups of spinal interneurons

Responses of two groups of spinal interneurons to prolonged stimulation of the medullary pyramids and the red nucleus by repetitive and random sequences of stimuli were investigated in cats anesthetized with pentobarbital. Interneurons specialized for transmission of descending, but not peripheral, effects were excited by impulses from both higher structures; the evoked activity was stable and stationary in character in response to stimulation within the range from 20–30 to 75–100/sec. The response of interneurons activated by flexor reflex afferents was more complex and, as a rule, it was not stationary. Prolonged pyramidal stimulation led to an increase in the mean spontaneous firing rate, while stimulation of the red nucleus inhibited spontaneous activity. Statistical analysis of the records of unit activity on the "Dnepr'-1" computer was used to study some possible mechanisms of the transformation of activity in the lateral descending systems by these groups of interneurons.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 5, No. 6, pp. 644–653, November–December, 1973.     

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.     

Comparative analysis of the kinematics of hind limb movements in rats during different kinds of locomotion

A comparative analysis of phases of the locomotor cycle and the dynamics of changes in hind limb joint angles during swimming and stepping movements (on a treadmill), involving the fore- and hind limbs to different degrees, were undertaken in rats. Differences in the sequence and degree of changes in joint angles during locomotion of the types investigated were participation of the forelimbs in locomotion was found to be accompanied by more marked forward carrying of the hind limb. Dependence of the swing phase on duration of the cycle was observed and differences were found in the period of protraction of the limb (F period) during swimming and stepping. The role of central spinal processes and influences of peripheral afferents in the formation of different types of locomotion is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 17, No. 2, pp. 189–198, March–April, 1985.     

Neuronal mechanisms of interaction between the red nucleus and other brain stem structures

Axon collaterals of rubrospinal neurons running to many brain stem structures were identified in acute experiments on cats by a technique of intracellular recording of antidromic action potentials in conjunction with collision testing. A systemic principle of organization of rubrospinal influences and also a tendency toward synchronous arrival of rubrospinal impulses at various brain stem centers were demonstrated. Most of these centers are relay nuclei, sending direct afferent projections to regions of the cerebellum which, in turn, control activity of the red nucleus. Besides such a loop, effecting dynamic cerebellar control over motor function, transmission of somatosensory information from nuclei of the dorsal columns of the spinal cord directly to the red nucleus was demonstrated. Special features of mono- and polysynaptic EPSPs evoked by stimulation of nuclei of the dorsal columns indicate that such PSPs arise in different regions of the soma-dendritic membrane of red nucleus neurons. The mechanisms of integration of descending motor volleys by the red nucleus are discussed.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 16, No. 5, pp. 665–678, September–October, 1984.     

Travelling wave patterns in a model of the spinal pattern generator using spiking neurons

The aim of this study is to produce travelling waves in a planar net of artificial spiking neurons. Provided that the parameters of the waves – frequency, wavelength and orientation – can be sufficiently controlled, such a network can serve as a model of the spinal pattern generator for swimming and terrestrial quadruped locomotion. A previous implementation using non-spiking, sigmoid neurons lacked the physiological plausibility that can only be attained using more realistic spiking neurons. Simulations were conducted using three types of spiking neuronal models. First, leaky integrate-and-fire neurons were used. Second, we introduced a phenomenological bursting neuron. And third, a canonical model neuron was implemented which could reproduce the full dynamics of the Hodgkin–Huxley neuron. The conditions necessary to produce appropriate travelling waves corresponded largely to the known anatomy and physiology of the spinal cord. Especially important features for the generation of travelling waves were the topology of the local connections – so-called off-centre connectivity – the availability of dynamic synapses and, to some extent, the availability of bursting cell types. The latter were necessary to produce stable waves at the low frequencies observed in quadruped locomotion. In general, the phenomenon of travelling waves was very robust and largely independent of the network parameters and emulated cell types.     

Rearrangement of parameters of efferent activity in generators of cyclic motor response produced by electrical stimulation of cerebellar inputs in cats

Rearrangement of the parameters of scratch and locomotor generators produced by electrical stimulation of the inferior olive and nucleus reticularis lateralis as well as the cerebellar fastigial nucleus and nucleus interpositus was investigated in decerebrate immobilized cats. Results showed that a comparable rearrangement of the time course of activity in both locomotor and scratch generators was produced by altering the nature of signals proceeding along mossy and climbing fibers alike. Maximum rearrangement of scratch and locomotor generator activity, as induced by electrical activation of the inferior olive and lateral reticular nucleus, is observed during the first half of flexor half-center operation in these generators. The scratch (unlike the locomotor) generator typically shows considerably rearranged efferent activity following electrical activation of nuclei of the cerebellum and cerebellar afferents. The article discusses mechanisms of cerebellar origin which may be responsible for exerting a corrective action on scratch and locomotor generators during change in the phase and amplitude parameters of cerebellar input signals.A. A. Bogomolets Institute of Physiology, Ukrainian Academy of Sciences, Kiev. Translated from Neirofiziologiya, Vol. 24, No. 2, pp. 131–140, March–April, 1992.