Human group I excitatory projections from ankle dorsiflexors to quadriceps muscle

A short latency projection of group I afferent fibers from ankle dorsiflexors to knee extensor muscles has been categorized as species specific to humans. However, the effects of the pathway have only been inferred from conditioning homonymous reflexes in relaxed muscle. This study focused directly on the responses evoked in the electromyogram of the heteronymous muscles when active, in two experiments. In the first, preferential activation of group I afferents of ankle dorsiflexors, by electrical stimulation of the common peroneal nerve, excited both vastus medialis (mean latency, 26.3 ms) and rectus femoris (mean latency, 33.5 ms). No excitation or inhibition in either muscle was associated with stimulation of the tibial nerve. The second experiment compared vastus medialis responses with common peroneal nerve stimulation during three different movement conditions in which the muscle was equally contracted: rhythmic isotonic (pedalling); episodic isotonic; isometric contraction. Responses were identified in all three active states, with no significant differences in amplitude or latency. No responses were seen in the relaxed muscle.     

Synaptic processes in thoracic α-motoneurons evoked by segmental afferent stimulation

Synaptic processes in various functional groups of thoracic motoneurons (Th₉-Th₁₁) evoked by stimulation of segmental nerves were investigated in anesthetized and decerebrate cats. No reciprocal relations were found between these groups of motoneurons. Only excitatory mono- and polysynaptic responses were recorded in the motoneurons of the principal intercostal nerve following stimulation of the homonymous nerve. Activation of the afferents of the external intercostal muscle and dorsal branches does not cause noticeable synaptic processes in these motoneurons; much more rarely it is accompanied by the development of low-amplitude polysynaptic EPSP's. In motoneurons of the dorsal branches, stimulation of homonymous nerves leads to the appearance of simple, short-latent EPSP's. Late responses of the IPSP or EPSP - IPSP type with a predominance of the inhibitory component were observed in most motoneurons of this type following activation of the afferent fibers of the principal intercostal nerve. In other motoneurons of the dorsal muscles, stimulation of the main intercostal nerve (and nerve of the external intercostal muscle) did not evoke apparent synpatic processes. EPSP's (mono- and polysynaptic) appeared in the motoneurons of the external intercostal muscle following stimulation of the homonymous and main intercostal nerves. Activation of the afferents of the dorsal branches was ineffective. The character of the synaptic responses of the respiratory motoneurons to segmental afferent stimulation, investigated under conditions of spontaneous respiration, was different. The characteristics of synaptic activation of thoracic motoneurons by segmental afferents under conditions of hyperventilation apnea and during spontaneous breathing of the animals are discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 2, No. 3, pp. 279–288, May–June, 1970.     

Modulation of the presynaptic inhibition of spinal α-motoneurons during mechanical stimulation of different types

Using the method of assessing the presynaptic inhibition of heteronymous Ia afferents and α motoneurons of the m. soleus during a homonymous vibration effect on the tendo calcaneus in ten subjects, changes in the inhibition of spinal α motoneurons during 15 min of the aftereffect of mechanical stimulation of different types were studied. Intense mechanical stimuli and weak tactile, vibratory stimuli applied in combination intensify inhibitory processes in the afferent fibers of group Ia, their effects differing in the value of the increase in the presynaptic inhibition of spinal α motoneurons.     

Transmitter potentiality of homonymous and heteronymous monosynaptic reflex connections of individual motoneurons

An assemblage of 110 individual tricipital motoneurons has been examined with the aim of determining those factors that predispose certain motoneurons to heteronymous response in post-tetanic potentiation. Motoneurons that respond most readily to homonymous volleys are not those that respond most readily to post-tetanically potentiated heteronymous volleys. Hence differences in presynaptic organization rather than differences in mean postsynaptic threshold determine differences in readiness for response. Every motoneuron exhibits a distinct asymmetry in transmitter potentiality of homonymous and heteronymous monosynaptic reflex connections. The range of transmitter potentialities is wide and that of heteronymous connections to some motoneurons is greater than that of homonymous connections to some other motoneurons.     

The investigation of control mechanisms of stepping rhythm in human in the air-stepping conditions during passive and voluntary leg movements

In unloading condition the degree of activation of the central stepping program was investigated during passive leg movements in healthy subjects, as well as the excitability of spinal motoneurons during passive and voluntary stepping movement. Passive stepping movements with characteristics maximally approximated to those during voluntary stepping were accomplished by experimenter. The comparison of the muscle activity bursts during voluntary and imposed movements was made. In addition to that the influence of artificially created loading onto the foot to the leg movement characteristics was analyzed. Spinal motoneuron excitability was estimated by means of evaluation of amplitude modulation of the soleus H-reflex. The changes of H-reflexes under the fixation of knee or hip joints were also studied. In majority of subjects the passive movements were accompanied by bursts of EMG activity of hip muscles (and sometimes of knee muscles), which timing during step cycle was coincided with burst timing of voluntary step cycle. In many cases the bursts of EMG activity during passive movements exceeded activity in homonymous muscles during voluntary stepping. The foot loading imitation exerted essential influence on distal parts of moving extremity during voluntary as well passive movements, that was expressed in the appearance of movements in the ankle joint and accompanied by emergence and increasing of phasic EMG activity of shank muscles. The excitability of motoneurons during passive movements was greater then during voluntary ones. The changes and modulation of H-reflex throughout the step cycle without restriction of joint mobility and during exclusion of hip joint mobility were similar. The knee joint fixation exerted the greater influence. It is supposed that imposed movements activate the same mechanisms of rhythm generation as a supraspinal commands during voluntary movements. In the conditions of passive movements the presynaptic inhibition depend on afferent influences from moving leg in the most degree then on central commands. It seems that afferent inputs from pressure receptors of foot in the condition of "air-stepping" actively interact with central program of stepping and, irrespective of type of the performing movements (voluntary or passive), form the final pattern activity.     

The role of anticipatory postural adjustments in compensatory control of posture: 1. Electromyographic analysis

Anticipatory (APAs) and compensatory (CPAs) postural adjustments are the two principal mechanisms that the central nervous system uses to maintain equilibrium while standing. We studied the role of APAs in compensatory postural adjustments. Eight subjects were exposed to external predictable and unpredictable perturbations induced at the shoulder level, while standing with eyes open and closed. Electrical activity of leg and trunk muscles was recorded and analyzed during four epochs representing the time duration typical for anticipatory and compensatory postural control. No anticipatory activity of the trunk and leg muscles was seen in the case of unpredictable perturbations; instead, significant compensatory activation of muscles was observed. When the perturbations were predictable, strong anticipatory activation was seen in all the muscles: such APAs were associated with significantly smaller compensatory activity of muscles and COP displacements after the perturbations.The outcome of the study highlights the importance of APAs in control of posture and points out the existence of a relationship between the anticipatory and the compensatory components of postural control. It also suggests a possibility to enhance balance control by improving the APAs responses during external perturbations.     

The investigation of locomotor activity control system in humans under the air-stepping conditions during passive and voluntary leg movements

The degree of activation of the central stepping program during passive leg movement was studied in healthy subjects under unloading conditions; the excitability of spinal motoneurons was studied during passive and voluntary stepping movements. Passive stepping movements with characteristics maximally close to those during voluntary stepping were accomplished by the experimenter. The bursts of muscular activity during voluntary and imposed stepping movements were compared. In addition, the influence on the leg movement of artificially created loading onto the foot was studied. The excitability of spinal motoneurons was estimated by the amplitude of modulation of the m. soleus H reflex. Changes in the H reflex (Hoffmann’s reflex) after fixation of the knee and hip joints were also studied. In most subjects, passive movements were accompanied by bursts of electromyographic (EMG) activity in the hip muscles (sometimes in shank muscles); the timing of the EMG burst during the step cycle coincided with the burst’s timing during voluntary stepping. In many cases, the bursts in EMG activity exceeded the activity of homonymous muscles during voluntary stepping. Simulation of foot loading influenced significantly the distal part of the moving extremity during both voluntary and passive movements, which was expressed in the appearance of movements in the ankle joint and an increase in the phasic EMG activity of the shank muscles. The excitability of motoneurons during passive movements was higher than during voluntary movements. Changes and modulation of the H reflex throughout the step cycle were similar without restriction of joint mobility and without hip joint mobility. Fixation of the knee joint was of great importance. It is supposed that imposed movements activate the same mechanisms of rhythm generation as supraspinal commands during voluntary movements. During passive movements, presynaptic inhibition depends mostly on the afferent influences from the moving leg rather than on the central commands. Under the conditions of “air-stepping,” the afferent influences from the foot pressure receptors are likely to interact actively with the central program of stepping and to determine the final activity pattern irrespective of the movement type (voluntary or passive).     

Activation of walking by electrical stimulation in humans under the conditions of muscle unloading and its variations under the effect of afferent influences

The possibility of initiating an involuntary walking rhythm in a suspended human leg by electrical stimulation was studied. The subjects lay on the side with one leg suspended in an exoskeleton allowing horizontal rotation in three joints: the hip, knee, and ankle ones. To evoke involuntary walking of the suspended leg, two methods were used: continuous vibration of the quadriceps muscle of the hip and electrical stimulation of the cutaneous nerves innervating the foot of the immobile leg. The hip and ankle were involved in the involuntary movements, with reciprocal bursts of electromyographic activity being also observed in the antagonistic muscles of the hip. The application of an external load (4 N or 8 N) to the foot caused a perceptible intensification of its movements. An additional weight (0.5 kg) or a rubber band wrapped around the foot caused no substantial change in the pattern of stimulated walking. Electrical stimulation is an effective means of activating walking movements, and their characteristics confirm the assumption that the walking rhythm is of central origin. Additional afferentation from the sole’s receptors plays an important role in the modulation of the induced movements and the modification of the general walking pattern under the conditions of muscle unloading.     

The effect of short-term changes in body mass distribution on feed-forward postural control

It was recently shown that short-term changes in the whole body mass and associated changes in the vertical position of the center of mass (COM) modify anticipatory postural adjustments (APAs) [Li X, Aruin AS. The effect of short-term changes in the body mass on anticipatory postural adjustments. Exp Brain Res 2007;181:333–46]. In this study, we investigated whether changes in the body mass distribution and related changes in the anterior–posterior COM position affect APA generation. Fourteen subjects were instructed to catch a 2.2 kg load with their arms extended while standing with no additional weight or while carrying a 9.08 kg weight. Adding weight to a backpack, front pack or belly pocket was associated with an increase of the whole body mass, but it also involved changes in the anterior–posterior (A/P) and vertical positions of the COM. Electromyographic activity of leg and trunk muscles, body kinematics, and ground reaction forces were recorded and quantified within the typical time intervals of APAs. APAs were modified in conditions with changed body mass distribution: increased magnitude of anticipatory EMG activity in leg and trunk muscles, as well as co-activation of leg muscles and decreased anticipatory displacement of the COM in the vertical direction, were seen in conditions with increased body mass. Changes in the COM position induced in both A/P and vertical directions were associated with increased anticipatory EMG activity. In addition, they were linked to a co-activation of muscles at the ankle joints and significant changes in the center of pressure (COP) position. Modifications of the COM position induced in the A/P direction were related to increased anticipatory EMG activity in the leg and trunk muscles. At the same time, no significant differences in anticipatory EMG activity or displacement of COP were observed when changes of COM position were induced in the vertical direction. The study outcome suggests that the CNS uses different strategies while generating APAs in conditions with changes in the COM position induced in the anterior–posterior and vertical directions.     

The role of afferent feedback in the control of hamstrings activity during human gait

In vertebrates, possibly also in man, the pattern of activation of muscles during locomotion can be generated by the spinal cord (locomotor CPG, central pattern generator). However, sensory feedback is crucial to adapt the functioning of the CPG to the external requirements during gait. It is postulated that afferent input from skin and muscles can contribute to the EMG activation patterns as observed in various limb muscles during gait. The activity of the hamstrings at end swing may be partially due to stretch reflexes of these muscles. At end stance the hamstring activity may be assisted by reflexes from natural skin activation from the dorsum of the foot. In addition, more specific actions are also incorporated. For example, sural nerve stimulation induces an activation of biceps femoris (BF) whereas a suppression is usually obtained for semitendinosus (ST), indicating that the induced activation is aimed at exorotation of the lower leg. Similarly, the preferential activation of medial versus lateral gastrocnemius (GM versus GL) in sural nerve induced reflexes could favor such exorotation. It is concluded that the present evidence points towards a possible contribution of various reflexes to the motor output seen during gait for movements both inside and outside the sagittal plane.     

Larva-adult relationships in an ancestral dipteran: A re-examination of sensillar pathways across the antenna and leg anlagen of Chaoborus crystallinus (DeGeer, 1776; Chaoboridae)

 In one of his classical studies on insect metamorphosis, Weismann compared the imaginal anlagen of the ancestral phantom midge, Chaoborus, with those of advanced brachycerans. We have expanded his findings on the relationships between larval and imaginal organs using electron microscopy and cobalt backfilling of the antenna and leg anlagen and the axonal trajectories of corresponding larval sensilla. We show that both primordia are confluent with the larval antennae and ”leg” sensilla (an ancestral Keilin organ), respectively. These fully developed larval organs represent the distal tips of the imaginal anlagen rather than separate cell clusters. The axons of the larval antenna and leg sensilla project across the corresponding anlagen to their target neuromeres within the central nervous system (CNS). Within the discs, nerves composed of these larval axons, developing afferent fibres and efferences ascending from the CNS are found. Both the structure of the primordia and the axonal trajectories thus relate the situation found in advanced brachycerans with that seen in more ancestral insects. In addition, the larval antennae, legs, wings and even the eyes possess very similar afferent pioneer trajectories supporting the idea that the described pattern is generally used in the ontogeny of sensory systems. Received: 30 June 1998 / Accepted: 27 September 1998     

Renorenal reflexes: interaction between efferent and afferent renal nerve activity.

In rats, stimulation of renal mechanoreceptors by increasing ureteral pressure results in a contralateral inhibitory renorenal reflex response consisting of increases in ipsilateral afferent renal nerve activity, decreases in contralateral efferent renal nerve activity, and increases in contralateral urine flow rate and urinary sodium excretion. Mean arterial pressure is unchanged. To study possible functional central interaction among the afferent renal nerves and the aortic and carotid sinus nerves, the responses to renal mechanoreceptor stimulation were compared in sinoaortic denervated rats and sham-denervated rats before and after vagotomy. In contrast to sham-denervated rats, there was an increase in mean arterial pressure in response to renal mechanoreceptor stimulation in sinoaortic-denervated rats. However, there were no differences in the renorenal reflex responses among the groups. Thus, our data failed to support a functional central interaction among the renal, carotid sinus, and aortic afferent nerves in the renorenal reflex response to renal mechanoreceptor stimulation. Studies to examine peripheral interaction between efferent and afferent renal nerves showed that marked reduction in efferent renal nerve activity produced by spinal cord section at T6, ganglionic blockade, volume expansion, or stretch of the junction of superior vena cava and right atrium abolished the responses in afferent renal nerve activity and contralateral renal function to renal mechanoreceptor stimulation. Conversely, increases in efferent renal nerve activity caused by thermal cutaneous stimulation increased basal afferent renal nerve activity and its responses to renal mechanoreceptor stimulation. These data suggest a facilitatory role of efferent renal nerves on renal sensory receptors.     

家兔主动脉神经低阈压力感受反射快速重调中枢过程的特征

40只家兔,乌拉坦静脉麻醉。切断双侧主动脉神经(AN)、窦神经及迷走神经。以选择兴奋 AN 有髓传人纤维的条件刺激(0.02ms,50Hz,4—6V,5min)给予切断的 AN 中枢段,模拟导致低阈压力感受反射快速重调的保持压背景,借以诱导快速重调的中枢过程。实验表明:该中枢过程使 AN 有髓纤维传入所激发的压力感受反射降压效应衰减41.82%(P<0.01),肾交感神经活动抑制效应衰减19.31%(P相似文献   

The response of alpha-motoneurons of different size to stretch and vibration of extensor muscles after injection of 5-hydroxytryptophan in spinal cats.

1. Vibration of the GS muscle at 100/sec, and the peak-to-peak amplitude of about 100 mu, produced slight reflex activity of homonymous and heteronymous alpha-motoneurons in the spinal, unanesthetized cat. 2. Intravenous injection of the 5-HT precursor 5-HTP (25 mg/kg, L-form) performed in the spinal preparation facilitated the responses of single extensor alpha-motoneurons to both vibration and stretch of the corresponding muscle. The 5-HT antagonist methysergide (0-5 mg/kg i. v.) blocked almost completely the facilitation of the vibration and stretch induced motoneuronal discharges produced by 5-HTP in the spinal cat, indicating that the facilitation was 5-HT specific. 4. A comparison of the types of alpha-motoneurons participating in the reflex responses elicited by the different stimuli in the spinal cat following injection of 5-HTP showed that both small-tonic and large-phasic alpha-motoneurons take part in both stretch and vibration reflexes. 5. It is concluded that the alpha-motoneuron pool of the GS muscle represents an homogenous population, no matter what kind of synaptic drive is used for its activation, and that the susceptibility of a motoneuron to discharge depends on the amount of the afferent excitatory input rather than on the modalities of sensory stimulation.     

Interaction between the Postactivation Depression of H Reflex and Long-Lasting Inhibition,Evoked by Stimulation of a Nerve to the Antagonist Muscles,in Humans

In a study on healthy humans, we examined interaction of the inhibitory influences on the H reflex recorded from the m. soleus (the respective EMG discharge, evoked by stimulation of the n. tibialis comm.). Postactivation depression of the reflex was evoked by a preceding conditioning stimulation of the same nerve, while conditioning stimulation of a nerve to the antagonist muscles (n. peroneus comm.) evoked long-lasting inhibition of the reflex, which included two consecutive waves of depression, D ₁ and D ₂. When the intensity of conditioning stimulations slightly exceeded the threshold for the development of inhibitory effects, interaction between postactivation depression and both the D ₁ and D ₂ waves demonstrated mutual facilitation of these effects. When the intensity of conditioning stimuli was increased, facilitation was changed by occlusion. We conclude that afferent impulsation, evoked by homo- and heteronymous conditioning stimulations of the peripheral nerves, converges on common interneuronal populations providing long-lasting suppression of the H reflex, which develops due to depolarization of primary afferent Ia terminals.     

Peripheral specification of Ia synaptic input to motoneurons innervating foreign target muscles

Muscle sensory neurons, called Ia afferents, make monosynaptic connections with functionally related sets of motoneurons in the spinal cord. Previous work has suggested that peripheral target muscles play a major role in determining the central connections of Ia afferents with motoneurons. Here, we ask whether motoneurons can also be influenced by their target muscles in terms of the monosynaptic input they receive from Ia afferents, by transplanting thoracic motoneurons into the lumbosacral spinal cord so that they innervate foreign muscles. Three or four segments of thoracic neural tube from stage 14-15 chicken embryos were transplanted to the lumbosacral region of stage 16-17 embryos, and electrophysiological recordings were made from transplanted motoneurons after the embryos had reached stage 38-40. Transplanted thoracic motoneurons innervated limb muscles and received monosynaptic inputs from Ia afferents. These connections were not random: Most of the connections were formed between Ia afferents and motoneurons projecting to the same muscle (homonymous connections). Few aberrant connections were found although the anatomical distribution of afferents in the transplant indicated that they had ample opportunity to contact inappropriate motoneurons. We conclude that although peripheral target cues are not sufficient to respecify an already committed motoneuron (turn a thoracic motoneuron into a lumbosacral motoneuron), they do provide sufficient information for Ia afferent input to be functionally correct.     

Afferents from a single muscle of the forelimbs and fastigial neurons of the cerebellum

Responses of neurons in the medial nucleus of cerebellum (CBM) were studied on stimulation of ipsilateral and contralateral homonymous muscles, in decerebrated cats. The aim was to find out to what extent information from homonymous muscles of the forelimbs converge on the same CBM neurons and whether the probability of such a convergence depends on location (axial, proximal, distal) or function (flexor, extensor) of the tested muscles. The analysis was limited to the neurons belonging to the rostral part of the nucleus which is known to control the ipsilateral muscle periphery. Neuronal activity was recorded extracellularly using tungsten microelectrodes (5-12 M omega) and muscle stimulation was performed by bipolar coated steel electrodes, with the exception of the tip. At least 6 pairs of homonymous muscles were generally stimulated: two axial, two proximal and two distal in both forelimbs. Care was taken that, when a muscle was stimulated, the others were not activated either directly or in a reflex way. Out of the 65 neurons studied, 60 (92%) were responsive to muscle stimulation. It was specifically observed that a high percentage of cells reacted to stimulation of distal muscles (74% to ipsilateral and 71% to contralateral ones). More than half (55%) of the neurons were responsive to activation of a pair of homonymous distal muscles and about one third of them (31%) to both the pairs of distal muscles. On the contrary the percentage of responses to proximal muscles was reduced foremost in the ipsilateral ones (23%) and only an exiquous percentage of cells (15%) received information from the homonymous proximal muscles.(ABSTRACT TRUNCATED AT 250 WORDS)     

Facilitation between extensor carpi radialis and pronator teres in humans: A study using a post-stimulus time histogram method

Group I muscle afferents modulate the excitability of motor neurons through excitatory and inhibitory spinal reflexes. Spinal reflex relationships between various muscle pairs are well described in experimental animals but not in the human upper limb, which exhibits a fine control of movement. In the present study, spinal reflexes between the extensor carpi radialis (ECR) and pronator teres (PT) muscles were examined in healthy human subjects using a post-stimulus time histogram method. Electrical stimulation of low-threshold afferents of ECR nerves increased the motor neuron excitability in 31 of 76 PT motor units (MUs) in all eight subjects tested, while stimulation of low-threshold afferents of PT nerves increased the motor neuron excitability in 36 of 102 ECR MUs in all 10 subjects. The estimated central synaptic delay was almost equivalent to that of homonymous facilitation. Mechanical stimulation (MS) of ECR facilitated 16 of 30 PT MUs in all five subjects tested, while MS of PT facilitated 17 of 30 ECR MUs in all six subjects. These results suggest excitatory reflex (facilitation) between PT and ECR. Group I afferents should mediate the facilitation through a monosynaptic path.     

Biocybernetic studies on reflectory heart modification in the rabbit]

In rabbits the depressor nerves and cardiac vagal branches were stimulated. Their actions on heart rate, atrio-ventricular conduction time, myocardial action potential and mean central blood pressure were recorded. The frequency-effect characteristics of the chronotropic, dromotropic and electrotropic actions on the heart, resulting from afferent and efferent nerve stimulation, are compared. The participation of each of the depressor nerves in their total effects on heart rate and blood pressure is studied. Time courses of heart rate and blood pressure decrease by afferent and efferent nerve stimulation with sinusoidally modulated pulse rates are presented. The results are discussed with respect to the different dynamics of blood pressure and heart rate control. It is concluded that at least two mechanisms are involved in blood pressure control by the depressor nerves: 1. Decrease of vascular resistance by lowering the sympathetic tone. 2. Decrease of heart rate by enhancing the cardiac vagal activity. It is suggested that the parasympathetic control unit compensates rapid disturbances, whereas the slow-acting sympathetic vascular mechanism exerts a long-time pressure control of high efficiency.     

Effects of peripheral inputs from hindlimb on the monosynaptic reflex of motoneurons innervating tail muscles.

The effects of group II muscle (PBSt, GS) and cutaneous afferent (Sur, SPc, Tib) inputs from the hindlimb on the monosynaptic reflexes of motoneurons innervating tail muscles were studied in lower spinalized cats. Stimulation of the cutaneous nerves at the conditioning-test stimulus interval of about 10-20 ms facilitated and inhibited the monosynaptic reflexes of ipsilateral and contralateral tail muscles, respectively. The effects of the muscle nerve stimulation were not so prominent as those elicited by cutaneous nerve stimulation. The monosynaptic reflex was also inhibited by muscle nerve stimulation at 10-50 ms intervals. The effects of conditioning stimulation of the hindlimb peripheral nerves at short intervals were depressed or blocked by section of the ipsilateral lateral funiculus at S1 spinal segment. These findings show that the neuronal pathway from hindlimb afferents to tail muscle motoneurons passed the lateral funiculus of the spinal cord and modulates the motoneuronal activity of tail muscles.