Phase-dependent modulation of primary afferent depolarization in single cutaneous primary afferents evoked by peripheral stimulation during fictive locomotion in the cat

Previous results from our laboratory have shown with intra-axonal recordings that hindfoot cutaneous primary afferents are subjected to rhythmic depolarizations during fictive locomotion (L-PAD) suggesting that cutaneous presynaptic mechanisms are activated by the central locomotor program. In this study, we examined the transmission in pathways responsible for primary afferent depolarizations (PAD) of cutaneous fibres during spontaneous fictive locomotion in decorticate cats and in spinal cats injected with nialamide and L-DOPA. PADs were evoked (E-PADs) by electrical stimulation of peripheral nerves and recorded intra-axonally with micropipettes in identified superficialis peroneal (SP; n = 7) and tibialis posterior (TP; n = 17) cutaneous primary afferents. Results showed that the amplitude of E-PADs, which were superimposed on the L-PAD, was deeply modulated throughout the locomotor cycle; decreasing to reach a minimum during the flexor phase and increasing to a maximum during the extensor phase. The results were not statistically different in fibres of the two nerves and in both types of preparation. The amplitude of E-PADs was always maximum during the extensor phase whether there was a large L-PAD or not during that phase. This suggests that the presynaptic mechanisms activated by central locomotor networks (L-PAD) and those activated by peripheral inputs (E-PAD) may in part be controlled differently. The results thus show that the transmission in PAD pathways activated by cutaneous inputs is phasically modulated by the central pattern generator for locomotion. This strongly suggests that the presynaptic inhibition in cutaneous fibres evoked by the movement-related feedback during real locomotion could be similarly modulated.     

Stimulus time-locked responses of motoneurons during forelimb fictive locomotion evoked by repetitive stimulation of the lateral funiculus

In cat forelimb fictive locomotion evoked by repetitive stimulation of the upper cervical lateral funiculus, locomotor discharges consisted of activities time-locked to each stimulus, which were rhythmically modulated. The stimulus time-locked activities were investigated by intracellular recording from motoneurons. In both elbow flexor and extensor motoneurons, there observed stimulus time-locked disynaptic EPSPs, trisynaptic IPSPs and polysynaptic EPSPs, all of which were rhythmically modulated with specific patterns. The disynaptic EPSPs of flexor motoneurons were facilitated in the flexor phase of locomotion, whereas those of extensor motoneurons were facilitated from the flexor phase to the flexor-to-extensor transition phase. Modulation depth was larger in flexor motoneurons. Trisynaptic IPSPs changed in amplitude in parallel with the disynaptic EPSPs of the antagonistic motoneurons. Late, polysynaptic EPSPs of both flexor and extensor motoneurons increased in amplitude along with corresponding nerve discharges. After lesions of the lateral funiculus at C6/C7, both the disynaptic EPSPs and trisynaptic IPSPs were abolished in the motoneurons located caudally to the lesions. However, only trisynaptic IPSPs were lost in the rostrally located motoneurons. Furthermore, the lesions disclosed that extensor motoneurons received another kind of stimulus time-locked EPSPs, trisynaptic EPSPs, which were transmitted through the ventral part of the spinal cord, and rhythmically facilitated in the extensor phase. Stimulus time-locked PSPs observed in this study may at least in part be evoked by last-order interneurons of the central pattern generator, which may be reciprocally organized.     

Cutaneous reflex activity of the cat forelimb during fictive locomotion

To investigate mechanisms of rhythmic modulation of cutaneous reflex by the central pattern generator for locomotion, we made intracellular recordings from elbow flexor motoneurons, biceps brachii (Bi) and brachialis (Br), and quantitatively analyzed excitatory postsynaptic potentials (EPSPs) evoked by stimulation of superficial radial (SR) nerves during forelimb fictive locomotion, which was evoked in immobilized, decerebrate cats with the low thoracic spinal cord transected. We found that (1) in both Br and Bi, SR stimulation evoked trisynaptic EPSPs (segmental latencies were 2.10±0.39 ms for Br and 2.07±0.93 ms for Bi), (2) the SR-EPSPs of Br and Bi were rhythmically modulated with a similar pattern; the maximal and minimal EPSPs appeared in the flexor and extensor phase, respectively, (3) however, there were differences as follows. In the control state where fictive locomotion was absent, the mean amplitude of SR-EPSPs of Br (4.65±2.76 mV, n=14) was much larger than that of Bi (1.25±1.22 mV, n=10). During fictive locomotion, the maximum amplitude of SR-EPSPs of Br was 5.63±1.79 mV (n=18), and the minimum was 3.82±1.69 mV (n=18); that is, the maximum during fictive locomotion was larger or smaller than that of the control, while the minimal were always smaller. In contrast, SR-EPSPs of Bi were modulated over the control level; the maximum was 3.97±1.71 mV (n=16) and the minimum was 2.24±1.11 mV (n=16), both the maximum and the minimum during locomotion being larger than that of the control. These results may suggest that two mechanisms are involved in the rhythmic modulation, first, cyclic facilitation in the flexor phase, and secondly, cyclic inhibition in the extensor phase. The former was involved in both cases of Bi and Br motoneurons, while the latter would be involved only in the case of Br motoneurons. © 1997 Elsevier Science B.V. All rights reserved.     

Phasic modulation of transmission from vestibular inputs to reticulospinal neurons during fictive locomotion in lampreys

The aim of this study was to determine whether the transmission from sensory inputs to reticulospinal neurons is modulated during fictive locomotion in lampreys. Reticulospinal neurons play a key role in the control of locomotion; modulation of sensory transmission to these neurons might be of importance for the adaptation of the control they exert during locomotion. In this series of experiments, intracellular synaptic responses of reticulospinal neurons of the posterior rhombencephalic reticular nucleus elicited by electrical stimulation of vestibular nerves on each side were studied during fictive locomotion induced by 50 microM N-methyl-D-aspartate (NMDA). Interestingly, shortly after NMDA had reached the bath and much before locomotor discharges were apparent in the recorded ventral roots, there was a significant depression of the synaptic transmission from vestibular nerves. The effect was reversed by washing out the NMDA and persisted in the isolated brainstem after spinal transection at the first segmental level. As locomotor discharges appeared in the ventral roots, synaptic responses elicited by vestibular nerve stimulation showed a clear phasic modulation of their amplitude during the locomotor cycle. Responses to stimulation of the ipsilateral vestibular nerve were smaller during the ipsilateral burst discharge than during the contralateral activity, whilst responses to stimulation of the contralateral vestibular nerve were minimal during contralateral activity and maximal during ipsilateral activity. This opposite pattern of modulation observed in the same reticulospinal neuron suggests that the phasic modulation of vestibular transmission is not due to changes in the membrane properties of the reticulospinal cell but is produced at a pre-reticular level.(ABSTRACT TRUNCATED AT 250 WORDS)     

Effect of chronic undernourishment on the cord dorsum potentials and the primary afferent depolarization evoked by cutaneous nerves in the rat spinal cord

The effect of chronic undernourishment on the cord dorsum potentials (CDPs) and the dorsal root potential (DRP), closely related to primary afferent depolarization (PAD) and presynaptic inhibition in the spinal cord of the rat, was analyzed in this study. Single electrical pulses applied to the sural nerve (SU) of control (n=14) and chronically undernourished (n=16) Wistar rats produced CDPs, which are composed of four components: afferent volley (AV), two negative components (N(1) and N(2)), and one positive component (P wave) and negative DRPs recorded in a small rootlet of the L6 segment of the rat. The CDPs of the control and undernourished rats with AV components of comparable amplitude (U(AV)/C(AV)=0.96), showed N(1) components of similar amplitude (U(N1)/C(N1)=0.94), but smaller P wave (U(PW)/C(PW)=0.23). A comparable reduction in the amplitude of the DRPs was obtained in the undernourished rats (U(DRP)/C(DRP)=0.36). When normalized as a function of the body mass of the animals, the CDPs and DRPs produced in undernourished rats were of significantly smaller normalized amplitude than those evoked in the control. According to these results, it is suggested that chronic undernourishment induce a depressive effect on the mechanisms generating the P wave component in the CDP and the DRPs either by decreasing the sensory input and/or the excitability of the dorsal horn neurones involved in the generation of PAD and presynaptic inhibition in the spinal cord of the rat.     

The role of spinal cord inputs in modulating the activity of reticulospinal neurons during fictive locomotion in the lamprey

Lamprey reticulospinal neurons are rhythmically modulated during fictive swimming. The present study examines the possibility that this modulation may originate from the spinal cord locomotor networks rather than from the brainstem. To test this, the in vitro preparation of the lamprey brainstem-spinal cord was separated into two compartments which could be exposed to different chemical environments. Locomotor activity was induced pharmacologically in the caudal spinal cord compartment and reticulospinal (RS) neurons from the posterior rhombencephalic reticular nucleus (PRRN) were recorded intracellularly in the rostral compartment containing normal lamprey Ringer. Under these conditions, the membrane potential of RS neurons showed clear rhythmic oscillations which are correlated with the ongoing locomotor activity in the caudal spinal cord bath, although no locomotor discharges were present in the ventral roots of the rostral bath. Such oscillations were not present in the absence of locomotion. These results indicate that the spinal cord locomotor networks can contribute to the rhythmic oscillations which occur in RS neurons during fictive locomotion. Moreover, the latter oscillations of membrane potential are due to both phasic excitation and Cl- -dependent inhibition in the opposite phase.     

Comparison between ventral spinocerebellar and rubrospinal activities during locomotion in the cat

During fictive locomotion of the thalamic cat, rhythmic activity related to the efferent discharges in hindlimb nerves was found in rubrospinal neurons (Arshavsky et al., this issue). Since the movements were abolished by curarization, this modulation could not result from rhythmic peripheral inputs and had therefore a central origin. Taking into account the existence of spinal generators, it was suggested that ascending pathways transmit rhythmic activity from these spinal centers to the supraspinal ones. Preliminary results have been obtained for neurons of the ventral spinocerebellar tract (VSCT) recorded during fictive locomotion: (1) their discharge is rhythmically modulated at the periodicity of the locomotor rhythm; (2) their discharge pattern can be complex and variable in relation with the complexity and variability of the pattern of efferent activity in various muscle nerves of the ipsilateral hindlimb; (3) their responses to phasic afferent stimulation of the ipsilateral hindlimb are modulated in parallel with their locomotor-related activity. These results show that VSCT neurons convey information on the central spinal activity during locomotion, and suggest that these neurons contribute to the activity of lumbar-projecting rubrospinal neurons which have similar characteristics.     

Modulation of reflex responses during fictive locomotion in the forelimb of the cat

During cat forelimb fictive locomotion, short-latency reflex pathways were examined by recording nerve discharges and intracellularly from motoneurones. Stimulation of cutaneous afferents, superficial radial nerves, evoked trisynaptic excitation of the elbow flexors, biceps brachii and brachialis, and stimulation of muscle afferents, deep radial nerves, evoked oligosynaptic, i.e. monosynaptic and disynaptic excitation of the flexors. The short-latency excitatory postsynaptic potentials (EPSPs) evoked from both nerves were rhythmically modulated; they were facilitated during the flexion phase and suppressed during the extension phase. Stimulation of high threshold muscle afferents evoked EPSPs with a central delay of ca. 4.2 ms, which were depressed throughout episodes of fictive locomotion. Since the short-latency EPSPs and longer-latency EPSPs in the same motoneurone were differently influenced during fictive locomotion, the effects observed could not be explained by changes occurring at only the motoneuronal level but they probably occurred at the premotoneuronal level. In addition, short-latency cutaneous excitation of the distal muscles, innervated by the median and ulnar nerves, was little modulated during fictive locomotion.     

Descending pathways eliciting forelimb stepping in the lateral funiculus: Experimental studies with stimulation and lesion of the cervical cord in decerebrate cats

The funicular pathways that elicit forelimb stepping were investigated with stimulation and lesion of the cervical white matter in decerebrate cats with the lower thoracic cord transected. We localized cross-sectional areas where stimulation evoked rhythmic motor-nerve discharges imitating those of stepping (fictive locomotion) in the immobilized animal, and further examined whether or not lesions made in the corresponding areas abolished actual locomotor movements. Stimulation of the C3 lateral funiculus (LF) produced fictive locomotion in the ipsilateral forelimb. The effective areas of stimulation were located separately in the dorsolateral funiculus (DLF) and in the ventrolateral funiculus (VLF), while the VLF was more effective than the DLF. Effective stimuli were pulse trains with a frequency of about 30 Hz, with a rather wide pulse duration of about 0.5 ms. Blocking axonal conduction through the lower thoracic cord by cooling reproducibly facilitated fictive locomotion in both amplitude and frequency. In the lesion experiments, forelimb locomotor movements were elicited spontaneously or by stimulation of the mesencephalic locomotor region (MLR). The locomotor movements were abolished by complete lesions of the C2-C3 LFs on both sides, but these remained when either the DLF or the VLF was intact on one side. These findings together suggested that the descending pathways for the activation of the spinal locomotor network of the single forelimb are located ipsilaterally in the DLF as well as in the VLF. Both the DLF and the VLF pathways can initiate locomotion, while the VLF pathways have a higher potential for its initiation. Lesion experiments further showed that cats can walk with both forelimbs, even though the spinal locomotor network of the single forelimb was deprived of its main descending input by unilateral lesions of the LF. However, when the unilateral lesion extended to the medial part of the LF, the bilateral walking was abolished; the limb on the lesioned side showed only rhythmic extension movements without active flexion movements, which was out of phase with the stepping movements on the intact side. This finding suggested that the medial part of the LF is important for producing flexion movements during the swing phase of stepping.     

Activity of fin muscles and fin motoneurons during swimming motor pattern in the lamprey

Coordination of motoneuron activity is a fundamental prerequisite for the generation of functional locomotor patterns. We investigate the neural mechanisms that coordinate activity of motoneuron pools in the vertebrate spinal cord with differing phases of activity in the locomotor cycle in a simple motor system, the lamprey swimming network. In the region of dorsal fins the lamprey spinal cord contains two groups of motoneurons: the myotomal motoneurons that innervate the trunk muscles; and the fin motoneurons controlling muscle fibres of the dorsal fins. We investigated the activity of fin muscles during swimming in vivo and that of fin motoneurons during fictive swimming in vitro. During swimming in vivo with cycle periods of 4-8 Hz, fin muscle activity covered a broad portion of the cycle, with the peak of activity out-of-phase to the ipsilateral myotomal muscles. During fictive swimming evoked by N-methyl-d-aspartate in the isolated spinal cord, fin motoneurons expressed similar out-of-phase activity. The phase relationship of the synaptic drive to fin motoneurons was examined by recording their activity intracellular during fictive swimming. Three different forms of membrane potential oscillation with different time courses in the locomotor cycle could be distinguished. Sagittal lesions of the spinal cord in the segment where fin motoneurons are recorded and up to one segment rostral and caudal from it did not influence the out-of-phase activity pattern of the motoneurons. Our results indicate that coordination of fin motoneuron activity with the locomotor activity of myotomal motoneurons does not depend on intrasegmental contralateral premotor elements.     

Phase-linked modulation of excitability of presynaptic terminals of low-threshold afferent fibers in the inferior alveolar nerve during cortically induced fictive mastication in the guinea pig

Excitability of presynaptic terminals of low-threshold primary afferent fibers in the inferior alveolar nerve was tested in the trigeminal spinal nucleus of the ketamine-anesthetized, paralyzed guinea pig, by Wall's method. Fictive mastication was induced by repetitive stimulation of the cortical masticatory area, and was monitored by rhythmical burst activity in the jaw-opening anterior digastric motoneuron pool. The excitability was rhythmically modulated in a phase-linked manner during the masticatory cycle: it was decreased coincidentally with the digastric burst activity (jaw-opening phase) and increased during the middle and late periods of the interburst phase (jaw-closing phase) of the masticatory cycle. The results imply that presynaptic modulation of synaptic transmission of peripheral inputs from primary afferents to interneurons in the jaw-opening reflex pathway may contribute to the rhythmical modulation of the jaw-opening reflex evoked by innocuous stimulation of the intraoral structures during mastication; presynaptic inhibition contributing to the depression of the jaw-opening reflex during the jaw-closing phase and presynaptic facilitation to its enhancement during the jaw-opening phase.     

Depolarization of primary afferents evoked by cyclically modulated natural activity of proprioceptors of the hindlimb of the cat

The primary afferent depolarization (PAD) evoked during passive sinusoidal movements of a hindlimb in the ankle joint was investigated in decerebrated cats. The frequency of movements varied within 0.14-5.0 Hz, the amplitude of the joint angle with respect to the axis of the tibia changed from 90 degrees to 130 degrees. The dorsal root potential (DRP) negativity increased both during flexion and during extension of the joint. The amplitude of the evoked DRPs was about 50-100 mV. A strong negative correlation was observed between the latency and rise time of the DRP and the frequency of the joint angle changes. During flexion the latency changed from 650 ms at 0.14-0.16 Hz frequency to 100-110 ms at 2.0 Hz and higher frequencies; during extension at the same frequencies the latency changed from 300 ms to 80-85 ms. The latency and rise time became minimal at 2.0 Hz frequency and practically did not change during the further increase of the oscillation frequency. The cord dorsum potential (CDP) evoked by the cutaneous nerve stimulation was recorded in parallel with the DRP. Periodical changes of the N-component of the CDP were in the opposite phase to changes of the DRP. Mechanisms of the observed changes of the PAD and functional significance of these changes during rhythmical motor acts are discussed.     

Sensory interactions with a central motor program in anuran larvae

Swimming in anuran larvae is directed by a central motor program that is modulated by spinal sensory input. Dorsal root stimulation activates the locomotor program in vitro and can produce either an increase or a decrease in the rate of ongoing fictive swimming. Records obtained from the distal stumps of cut dorsal roots during passive tail bending show that receptors respond to tail movement but not to tail position. During episodes of fictive swimming, primary afferent terminals are depolarized and their sensitivity to antidromic stimulation increased, indicating that the motor program exerts presynaptic inhibitory control over spinal sensory transmission. These results suggest that the central program is very sensitive to dorsal root inputs and modulates these inputs during swimming.     

Effects of stimulating the reticular formation during fictive locomotion in lampreys

The effects of stimulating the reticular formation were studied during fictive locomotion in lampreys (Ichthyomyzon unicuspis). The in vitro isolated preparation of the brainstem and spinal cord was used and fictive locomotion was induced by bath application of N-methyl-

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-aspartate (NMDA; 50–100 μM). During different phases of the locomotor cycle, short trains of stimuli (10 pulses at 80–100 Hz; 10 μA) were delivered through glass-coated tungsten microelectrodes positioned within the middle rhombencephalic reticular nucleus (MRRN) and their effects were studied on ipsi- and contralateral ventral root locomotor discharges. Irrespective of the locomotor phase during which the stimulation train was delivered, a resetting effect occurred. It was characterized by a re-synchronization of the locomotor discharges with a constant latency for each ventral root on the ipsilateral side. The latency increased as the recorded root was located further caudally. This increase in latency was in the range of the phase lag observed between roots during control bouts of locomotion. These results suggest that reticulospinal neurones exert strong resetting effects on spinal locomotor networks. These effects may play a significant role with respect to changes of direction during swimming.     

The spinal 5-HT system contributes to the generation of fictive locomotion in lamprey

Activation of NMDA receptors evokes sustained fictive locomotion in the isolated spinal cord of the sea lamprey Petromyzon marinus (P. marinus), but in the river lamprey Lampetra fluviatilis (L. fluviatilis) the ventral root activity is often irregular. A previous study showed that the number of 5-HT immunoreactive fibres, neurones and varicosities are much lower in the spinal cord of L. fluviatilis than in P. marinus. To further analyse the underlying mechanisms, the present study investigated the role of the 5-HT system in stabilising fictive locomotion. In P. marinus a blockade of 5-HT1A receptors by spiperone reversibly increased the frequency and the coefficient of variation. This implies that there is an endogenous release of 5-HT during fictive locomotion that is important for the generation of locomotor activity. In L. fluviatilis bath applied NMDA or -glutamate evoked in most cases irregular activity. An addition of 5-HT (0.5–2 μM) rapidly stabilised the burst generation and led to a sustained fictive locomotion. In a split-bath configuration, NMDA administered to the rostral part of the spinal cord in P. marinus evoked fictive locomotion in both the rostral part and the first few segments of the caudal part. When spiperone was added to the caudal part, the burst activity changed into tonic activity within 10 min. Taken together, these results indicate that activity in the intrinsic 5-HT system in the lamprey spinal locomotor network contributes significantly to the rhythm generation. The quantitative differences with regard to the 5-HT plexus between P. marinus and L. fluviatilis may account for the observed discrepancy between the two species.     

Effects of bicuculline and strychnine on synaptic inputs to edge cells during fictive locomotion

Edge cells are mechanoreceptive neurones located in the lateral tracts of the lamprey spinal cord. Phasic activation of these cells by lateral bending can entrain the activity of the locomotor central pattern generator. During fictive locomotion induced by bath-applied NMDLA (N-methyl-D,L-aspartate) or sensory stimulation, edge cells receive synaptic input. In this paper we provide evidence that the tonic inhibition received during sensory-induced fictive locomotion is glycinergic. We also show that during fictive locomotion induced by application of NMDLA to the spinal cord, bicuculline unveils phasic synaptic activity in edge cells. During sensory-evoked fictive locomotion, by contrast, only tonic synaptic activity is apparent in the presence of bicuculline.     

Organization of mammalian locomotor rhythm and pattern generation

Central pattern generators (CPGs) located in the spinal cord produce the coordinated activation of flexor and extensor motoneurons during locomotion. Previously proposed architectures for the spinal locomotor CPG have included the classical half-center oscillator and the unit burst generator (UBG) comprised of multiple coupled oscillators. We have recently proposed another organization in which a two-level CPG has a common rhythm generator (RG) that controls the operation of the pattern formation (PF) circuitry responsible for motoneuron activation. These architectures are discussed in relation to recent data obtained during fictive locomotion in the decerebrate cat. The data show that the CPG can maintain the period and phase of locomotor oscillations both during spontaneous deletions of motoneuron activity and during sensory stimulation affecting motoneuron activity throughout the limb. The proposed two-level CPG organization has been investigated with a computational model which incorporates interactions between the CPG, spinal circuits and afferent inputs. The model includes interacting populations of spinal interneurons and motoneurons modeled in the Hodgkin-Huxley style. Our simulations demonstrate that a relatively simple CPG with separate RG and PF networks can realistically reproduce many experimental phenomena including spontaneous deletions of motoneuron activity and a variety of effects of afferent stimulation. The model suggests plausible explanations for a number of features of real CPG operation that would be difficult to explain in the framework of the classical single-level CPG organization. Some modeling predictions and directions for further studies of locomotor CPG organization are discussed.     

Presynaptic GABAA and GABAB Receptor-mediated Phasic Modulation in Axons of Spinal Motor Interneurons

The lamprey spinal cord has been utilized to investigate the role of presynaptic inhibition in the control of the spinal motor system. Axons of the lamprey spinal cord are comparatively large because of their lack of myelination. Axons impaled with microelectrodes demonstrate depolarizing responses to the application of GABAA and GABAB receptor agonists, muscimol and baclofen. These depolarizing effects are counteracted by the specific GABAA and GABAB receptor antagonists, bicuculline and phaclofen. GABAA receptor activation leads to a gating of Cl- channels on the axons. However, the ionic mechanism leading to axonal depolarization following GABAB receptor activation is unknown. After initiation of fictive locomotion, these axons demonstrate oscillations in axonal membrane potential related to the locomotor cycle. During ficitive locomotion they depolarize in phase with the bursting of the ipsilateral ventral root of the same segment. These axonal membrane potential oscillations are due to a phasic GABAA and GABAB receptor-mediated gating of ion channels on the axonal membrane. Fictive locomotion in the lamprey spinal cord is largely unaffected by antagonism of one or other GABA receptor subtype alone, but is severely disrupted by simultaneous antagonism of both subtypes. In conclusions, we demonstrate, for the first time, an agonist-gated depolarization of a vertebrate presynaptic element measured by direct impalement of the axon under study. We also demonstrate that GABA-mediated presynaptic inhibition occurs in axons of spinal interneurons. It is not limited to the primary afferents as has previously been believed.     

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

Rearrangement of the locomotor generator activity evoked by phasic electrical stimulation of different descending systems has been investigated on decerebrated immobilized cats. The rearrangement heavily depends on the stimulation phase. A maximal increase of the locomotor cycle duration by electrical stimulation of the Deiters nucleus takes place in the end of an "extensor" phase. Stimulation of the Deiters nucleus during the flexor phase decreases the locomotor cycle duration. A maximal increase of the locomotor cycle duration by electrical stimulation of the red nucleus, pyramidal tract and reticular gigantocellular nucleus takes place in the first half of a "flexor" phase. Electrical activation of these descending pathways during the flexor phase promotes an increase in intensity of this phase and a decrease in intensity of the "extensor" phase. Electrical activation of the Deiters nucleus increases intensity of the extensor phase and decreases that of the "flexor" phase. Possible principles of the suprasegmental correction of the locomotor generator activity are discussed.     

Spinal cord reflexes induced by epidural spinal cord stimulation in normal awake rats

Motor responses in hindlimb muscles to epidural spinal cord stimulation in normal awake rats during bipedal standing were studied. Stimulation at L2 or S1 induced simultaneous and bilateral responses in the vastus lateralis, semitendinosus, tibialis anterior, and medial gastrocnemius muscles. Stimulation at S1 evoked an early (ER), middle (MR) and late (LR) response: stimulation at L2 elicited only a MR and LR. Vibration and double epidural stimulation testing suggests that the ER is a direct motor response, whereas the MR and LR are mediated synaptically. MR has properties of a monosynaptic reflex, i.e., inhibited during vibration and depressed during the second pulse of a double stimulation. Some components of the LR seem to be mediated by afferents associated with the flexor reflex and probably involve group II afferents. During bipedal treadmill stepping, the MR was modulated in extensors, whereas the LR was modulated in flexors. These results show differential modulation of monosynaptic and polysynaptic reflexes in flexor and extensor motor pools during locomotion. Monosynaptic responses to stimulation at either L2 or S1 generally were amplified in extensors during the stance phase and in flexors during the swing phase of the step cycle. No correlation was found between the ER and the EMG background during stepping, whereas both the MR and LR were closely correlated with the changes in the EMG activity level of the corresponding muscle. These data demonstrate the feasibility of using epidural stimulation for examining monosynaptic and polysynaptic pathways to motor pools associated with multiple muscles during movement and over a prolonged period.