The effects of wrist muscle vibration on human voluntary elbow flexion-extension movements

Summary The effect of forearm muscle tendon vibration during alternating step flexion-extension movements about the elbow was studied in normal humans. In one experiment, a vibrator was mounted over either the forearm flexor or the extensor muscle. In a second experiment, a vibrator was mounted over either the forearm muscle or the biceps muscle. In both experiments, vibration was applied either to a single muscle or simultaneously to both muscles during elbow flexion-extension movements. After a period of practice, subjects learned the required movements and were able to make them with their eyes closed. Application of vibration to the forearm and the biceps muscle during extension movements produced an undershoot of the required end movement position. Moreover, application of high frequency vibration (100 Hz) to the forearm extensor and flexor muscle produced an overshoot of the required end-movement position. The observed results are consistent with vibration induced activation of muscle spindle receptors not only in the lengthening muscle during movement but also in the forearm muscles. It is suggested that the pattern of distribution of muscle spindle afferent from the forearm muscle onto -motoneurons of muscles acting at the elbow has played an important role of alternating step flexion-extension movements.     

The effect of agonist/antagonist muscle vibration on human position sense

Summary During voluntary movement, muscle spindles of both the agonist and antagonist muscles potentially can supply information about position of the limb. Muscle vibration is known to increase muscle spindle discharge and cause systematic distortions of limb position sense in humans. The following two experiments attempted to examine these contributions by separately vibrating over the triceps and biceps muscles during forearm positioning. In the first experiment, subjects performed a horizontal flexion or extension of the right arm to a mechanical stop randomly positioned at 20, 40 or 60°. Vision was occluded and vibration was applied to the right arm. The perceived position of the right limb was assessed by instructing subjects to simultaneously match the right arm position with the left limb. Vibration of the shortening, agonist muscle had no effect on limb matching accuracy. However, antagonist muscle vibration resulted in a significant overestimation of the vibrated limb position by 6–13°. The procedures for the second experiment were similar to the first, except that movements of the right limb were self-terminated and only flexion movements were performed. A screen was mounted over the arms and subjects were instructed to move the right arm until it was positioned beneath a marker on the screen. Vibration of the shortening agonist muscle had no effect on either the positioning accuracy of the right limb or matching accuracy of the left limb. However, antagonist muscle vibration resulted in significantly shorter movements (6–10°) by the right limb and an overestimation of right limb position by the left, matching limb. These findings support the hypothesis that muscle spindle afferent information from the lengthening antagonist muscle contributes to limb position sense during voluntary movement.     

The effects of muscle vibration on the attainment of intended final position during voluntary human arm movements

Summary Muscle tendon vibration was applied during voluntary step-tracking arm target-movements performed by normal human subjects. Vibration (freq. = 120 Hz) was applied over either the biceps or triceps tendons. During non-visually guided (eyes closed) trials, vibration of the muscle antagonistic to the movement being performed resulted in an undershoot of the required target. Thus, biceps vibration produced an undershoot of the extension target and triceps vibration an undershoot of the flexion target. The same effect occurred if the vibration was applied continuously over several movements or only during the course of individual movements. In contrast, vibration of the muscle acting as the prime mover had no effect on the correct attainment of the required target. It is suggested that the central nervous system may monitor muscle afferent activity of the lengthening (antagonist) muscle during simple, step movements.Supported by the Medical Research Council of Canada, Grant MA-6699     

The effect of muscle vibration on human position sense during movements controlled by lengthening muscle contraction

Summary Muscle vibration studies suggest that during voluntary movement limb position is coded by muscle spindle information derived from the lengthening, antagonist muscle. However, these investigations have been limited to movements controlled by shortening contractions. This study further examined this property of kinesthesia during movements controlled by lengthening contraction. Subjects performed a horizontal flexion of the right forearm to a mechanical stop randomly positioned at 30, 50 and 70° from the starting position. The movement was performed against a flexor load (1 kg) requiring contraction of the triceps muscle. Vision was occluded and movements were performed under three conditions: no vibration, vibration of the right biceps and vibration of the right triceps. The perceived position of the right forearm was assessed by instructing subjects to simultaneously match the right limb position with the left limb. Vibration of the shortening biceps muscle had no effect on limb matching accuracy. However, triceps vibration resulted in significant overestimation of the vibrated limb position (10–13°). The variability in movement distance was uninfluenced by muscle vibration. During movements controlled by lengthening contraction, there is a concurrent gamma dynamic fusimotor input that would enhance primary afferent discharge. Despite this additional regulating input to the muscle spindle, it appears that muscle spindle information from the lengthening muscle is important for the accurate perception of limb movement and/or position.     

Electromyographic responses to constant position errors imposed during voluntary elbow joint movement in human

The role of reflexes in the control of stiffness during human elbow joint movement was investigated for a wide range of movement speeds (1.5–6 rad/s). The electromyographic (EMG) responses of the elbow joint muscles to step position errors (step amplitude 0.15 rad; rise time 100 ms) imposed at the onset of targeted flexion movements (1.0 rad amplitude) were recorded. For all speeds of movement, the step position disturbance produced large modulations of the usual triphasic EMG activity, both excitatory and inhibitory, with an onset latency of 25 ms. In the muscles stretched by the perturbation, the early EMG response (25–60 ms latency) magnitude was greater than 50% of the activity during the unperturbed movements (background activity). In all muscles the EMG responses integrated over the entire movement were greater than 25% of the background activity. The responses were relatively greater for slower movements. Perturbations assisting the movement caused a short-latency (25–60 ms) reflex response (in the antagonist muscle) that increased with movement speed and was constant as a percentage of the background EMG activity. In contrast, perturbations resisting the movement caused a reflex response (in the agonist muscle) that was of the same absolute magnitude at all movement speeds, and thus decreased with movement speed as a percentage of the background EMG activity. There was a directional asymmetry in the reflex response, which produced an asymmetry in the mechanical response during slow movements. When the step perturbation occurred in a direction assisting the flexion movement, the antagonist muscle activity increased, but the main component of this response was delayed until the normal time of onset of the antagonist burst. When the step perturbation resisted the movement the agonist muscles responded briskly at short latency (25 ms). A reflex reversal occurred in two of six subjects. A fixed reflex response occurred in the antagonist muscle, regardless of the perturbation direction. For the extension direction perturbations (resisting movement), this response represented a reflex reversal (50 ms onset latency) and it caused the torque resisting the imposed step (stiffness) to drop markedly (below zero for one subject). Reflex responses were larger when the subject was prevented from reaching the target. That is, when the perturbation remained on until after the normal time of reaching the target, the EMG activity increased, with a parallel increase in stiffness. Similarly, when the perturbations prevented the subject from reaching the target during a 1-rad voluntary cyclic movement, the EMG and stiffness increased markedly. Coactivation of the antagonist muscle with the agonist muscle was not prominent (<30% of antagonist activity) during unperturbed movements. The perturbations were resisted with reciprocal activity, and thus reflex action did not increase the coactivation. However, as a result of the low-pass properties of muscle, substantial cocontraction of the agonist and antagonists muscle forces may have occurred during rapid movements, thus leading to increased stiffness. As the relative changes in normal EMG activity produced by the perturbation were often comparable with the changes in mean muscle torque (stiffness) reported in the first paper of this series, we conclude that the action of reflexes produced a significant portion of the resistance to perturbations. This reflexive portion was greater for slower movements, it was greater when the subject neared the target, and it was variable according to the perturbation direction and the particular subject. Given that the perturbations were of similar frequency content to the movement itself (though of smaller amplitude) and that the reflexes contributed substantially to the resistance to these perturbations, we suggest that in normal unperturbed movements the observed EMG is likewise substantially determined by the reflex activity.     

Effects of gravitational forces on single joint arm movements in humans

We have examined the kinematics and muscle activation patterns of single joint elbow movements made in the vertical plane. Movements of different amplitudes were performed during a visual, step-tracking task. By adjusting shoulder position, both elbow flexion and extension movements were made under three conditions: (a) in the horizontal plane, (b) in the vertical plane against gravity, and (c) in the vertical plane with gravity. Regardless of the gravitational load, all movements were characterized by time symmetric velocity profiles. In addition, no differences were found in the relationships between movement duration, peak velocity, and movement amplitude in movements with or against gravity. The pattern of muscle activation was influenced however, by the gravitational load. Both flexion and extension movements made with gravity were characterized by a reciprocally organized pattern of muscle activity in which phasic agonist activity was followed by phasic antagonist activity. Flexion and extension movements made against gravity were characterized by early phasic antagonist activity occurring at about the same time as the initial agonist burst. These findings suggest that EMG patterns are modified in order to preserve a common temporal structure in the face of different gravitational loads.     

Kinaesthetic role of muscle afferents in man,studied by tendon vibration and microneurography

Summary The characteristics of vibration-induced illusory joint movements were studied in healthy human subjects. Unseen by the subject, constant frequency vibration trains applied to the distal tendon of the Triceps or Biceps induced an almost constant velocity illusory movement of the elbow whose direction corresponded to that of a joint rotation stretching the vibrated muscle. Vibration trains of the same duration and frequency applied alternatively to the Biceps and Triceps evoked alternating flexion-extension illusory movements.During successive application of vibration trains at frequencies from 10 to 120 Hz, the perceived velocity of the illusory movements increased progressively from 10 to 70–80 Hz, then decreased from 80 to 120 Hz. The maximal perceived velocity was three times higher during alternating vibration of the Biceps and Triceps than during single muscle stimulation.Unit activity from 15 muscle spindle primary endings and five secondary endings located in Tibialis anterior and Extensor digitorum longus muscles were recorded using microneurography in order to study their responses to tendon vibration and passive and active movements of the ankle.Primary endings were all activated by low amplitude tendon vibration (0.2–0.5 mm) previously used to induce illusory movements of the elbow. The discharge of some was phase-locked with the vibration cycle up to 120 Hz, while others responded one-to-one to the vibration cycle up to 30–50 Hz, then fired in a sub-harmonic manner at higher frequencies. Secondary endings were much less sensitive to low amplitude tendon vibration.Primary and secondary ending responses to ramp and sinusoïdal movements of the ankle joint were compared. During the movement, the primary ending discharge frequency was almost constant, while the secondary ending activity progressively increased. During ankle movements the primary ending discharge appeared mainly related to velocity, while some secondary activities seemed related to both movement velocity and joint angle position.Muscle spindle sensory ending responses to active and passive ankle movements stretching the receptor-bearing muscle (plantar flexion) were qualitatively and quantitatively similar. During passive reverse movements (dorsiflexion) most of the sensory endings stopped firing when their muscle shortened. Active muscle shortening (isotonic contraction) modulated differently the muscle spindle sensory ending discharge, which could stop completely, decrease or some times increase during active ankle dorsiflexion. During isometric contraction most of the muscle spindle sensory endings were activated.The characteristics of the vibration-induced illusory movements and the muscle spindle responses to tendon vibration and to active and passive joint movements strengthened the possibility of the contribution of primary endings to kinaesthesia, as suggested by several previous works. Moreover, the present results led us to attribute to proprioception in the muscle stretched during joint movement a predominant, but not exclusive, role in this kind of perception.     

Control of simple arm movements in elderly humans

Eight elderly subjects (aged 68-95 years) and 6 young adults (aged 21-24 years) performed elbow flexion and extension movements in a visual step-tracking paradigm. Movement amplitudes ranging from 10 degrees-80 degrees were made under two instructions: "move at own speed" and "move fast and accurate." In a second experiment, 5 elderly subjects practiced 30 degrees movements for a total of 180 flexion and 180 extension movements under the instruction to increase movement speed, while maintaining accuracy, during practice. Movement trajectories became more variable as both movement amplitude and speed increased. Trajectory variability was greater in the elderly subjects for both the acceleratory and deceleratory phases of movements. This was due primarily to a greater rate of increase in trajectory variability during the acceleration phase in the elderly. With practice, elderly subjects could substantially reduce trajectory variability with little change in movement speed. The agonist burst initiating movements was qualitatively normal in the elderly subjects. However, there was considerable tonic cocontraction of agonist and antagonist muscles prior to and during movement. Phasic antagonist EMG activity was obviously abnormal in many elderly subjects. There was often no clear antagonist burst associated with deceleration of the movements or, if present, it was timed inappropriately early. With practice, combined agonist-antagonist EMG variability decreased. A clear antagonist burst also developed during practice in most elderly subjects, but its inappropriate timing remained in all but one subject. The results show that movement trajectories are less accurately controlled in the elderly.(ABSTRACT TRUNCATED AT 250 WORDS)     

The Ia afferent feedback of a given movement evokes the illusion of the same movement when returned to the subject via muscle tendon vibration

The aim of the present study was to further investigate the contribution of primary muscle spindle feedback to proprioception and higher brain functions, such as movement trajectory recognition. For this purpose, complex illusory movements were evoked in subjects by applying patterns of muscle tendon vibration mimicking the natural Ia afferent pattern. Ia afferent messages were previously recorded using microneurographic method from the six main muscle groups acting on the ankle joint during imposed “writing like” movements. The mean Ia afferent pattern was calculated for each muscle group and used as a template to pilot each vibrator. Eleven different vibratory patterns were applied to ten volunteers. Subjects were asked both to copy the perceived illusory movements by hand on a digitizing tablet and to recognize and name the corresponding graphic symbol. The results show that the Ia afferent feedback of a given movement evokes the illusion of the same movement when it is applied to the subject via the appropriate pattern of muscle tendon vibration. The geometry and the kinematic parameters of the imposed and illusory movements are very similar and the so-called “two-thirds power law” is present in the reproduction of the vibration-induced illusory movements. Vibrations within the “natural” frequency range of Ia fibres firing (around 30 Hz) produce clear illusions of movements in all the tested subjects. In addition, increasing the mean frequency of the vibration patterns resulted in a linear increase in the size of the illusory movements. Lastly, the subjects were able to recognize and name the symbols evoked by the vibration-induced primary muscle spindle afferent patterns in 83% of the trials. These findings suggest that the “proprioceptive signature” of a given movement is associated with the corresponding “perceptual signature”. The neural mechanisms possibly underlying the sensory to perceptual transformation are discussed in the general framework of “the neuronal population vector model”.     

Effects of unexpected perturbations on trajectories and EMG patterns of rapid wrist flexion movements in humans

To investigate how motor programs can be modified by sensory inputs we recorded kinematic and EMG patterns from normal human subjects performing well-practised wrist flexion movements in response to an auditory tone. On random trials unexpected wrist perturbations were introduced at varying times after the signal to move had been given. Extension perturbations delivered before agonist EMG onset resulted in an increased maximum velocity (MV) during the wrist flexion movement and in an increased target overshoot even though the wrist was further from the target than expected by the subject at the onset of the movement. The first agonist EMG burst and the antagonist burst were both increased in magnitude in these perturbed trials. Flexion perturbations delivered before the agonist EMG onset moved the hand nearer to the target just prior to movement onset. These resulted in a reduced MV, but the expected increased target overshoot did not occur. The first agonist burst was reduced in magnitude, and the antagonist burst was increased in magnitude. Perturbations delivered after agonist EMG onset produced less change in the first agonist and antagonist EMG burst, and less compensation for the perturbation was evident in wrist position and velocity recordings. These results indicate that, at least in some situations, motor programs for rapid voluntary movements can be modified by afferent inputs. This interaction between central motor commands and sensory feedback might occur at the cortical or spinal level, depending on when perturbations occur relative to onset of EMG and movement. The timing of the EMG changes suggest that both reflex mechanisms and longer latency 'voluntary' adjustments contribute to the compensatory changes in movement trajectory.     

Role of proprioceptive information in the temporal coordination between joints

Ten subjects made rapid, simultaneous movements of jaw (elevation or lowering) and right foot (ankle flexion or extension) in two experimental situations: (1) in response to an external signal (reaction-time situation), and (2) in a self-paced situation. We calculated the mean time intervals between the onsets of electromyographic (EMG) activity of agonist muscles (tibialis anterior or gastrocnemius lateralis compared with masseter or digastricus pars anterior) and those between the onsets of movement acceleration at each joint. Despite the fact that subjects reported simultaneous jaw-foot movements, there was always a short time interval between the two movements as between the agonist EMG activities. When the subjects were asked to perform a jaw elevation movement simultaneously with an ankle movement (flexion or extension), the sign of the time interval was dependent on the situation of movement initiation. In the reaction-time situation, the jaw motor activity preceded that of the ankle, whereas the reverse temporal order was observed in the self-paced situation. This is consistent with a previous hypothesis suggesting that the simultaneity of two motor actions is centrally established through two separate central processes: reactive or predictive. When subjects tried to perform simultaneous jaw lowering and foot flexion or extension movements, the strict temporal order observed when considering jaw elevation and ankle movements disappeared. The jaw motor activity generally preceded that of ankle in the reactive situation, but, depending on the subjects, it preceded or followed the ankle motor activity during self-paced movements. It is likely that the specific spindle supply of jaw muscles accounts for these results. Indeed, the jaw depressor muscles, in contrast to the elevators, lack muscle spindles. Our results suggest that the kinesthetic inputs used by the upper central nervous system to synchronize two rapid voluntary movements are mainly those from spindles located in the muscles that accelerate the movement, suggesting a strong α-γ linkage. Received: 31 October 1996 / Accepted: 1 September 1997     

Arm movement performance during reversible basal ganglia lesions in the monkey

Summary Arm motor performance of eight Cebus monkeys was examined during reversible cooling in the ventral lateral region of the putamen and globus pallidus (primarily the external segment), where neurons discharging during arm movements have been found (DeLong 1972).When attempting to hold a handle stationary during basal ganglia cooling, all monkeys developed flexion at the wrist and some developed a slow flexion drift of the arm at the elbow. The prominence of wrist flexion emphasizes that the basal ganglia may normally influence distal musculature.During basal ganglia cooling an increase in segmental stretch reflexes (15–30 ms) was sometimes observed following arm perturbations, but no consistent increase occurred in the later EMG responses (30–95 ms) in contrast to results obtained in Parkinsonian patients (Tatton and Lee 1975).No major changes were observed in the time of onset of the earliest EMG activity in the agonist muscle in a simple reaction time elbow movement task during basal ganglia cooling.Basal ganglia lesions produced major disorders in both flexion and extension movements including slowing of movements and rebound of the arm towards its initial position after onset of movement. These disorders were accompanied by an increase in tonic activity of both flexors and extensors while holding and by increased levels of cocontraction of agonists and antagonists during attempted movements.It is suggested that this basal ganglia disorder is due to a failure to achieve the correct balance of activity between agonists and antagonists that is appropriate for a particular motor act.This study was supported by the Canadian MRC PG-1     

Trained slow tracking. I. Muscular production of wrist movement

Electromyographic (EMG) activity was recorded from those forearm muscles that act across the wrist as highly trained monkeys tracked slow hold-ramp-hold target trajectories with angular wrist position. During performance of this task, the forearm flexors and extensors had a common "basic pattern" of EMG activity. Flexor digitorum sublimis (FDS) and extensor digitorum communis (EDC), though commonly classified as prime movers of the fingers, were the most active flexor and extensor muscles during these movements at the wrist. The basic pattern of EMG activity was analyzed by varying independently 1) the movement direction, 2) the initial and final held wrist positions, 3) the ramp-movement velocity, and 4) the direction and magnitude of maintained external torque load. Most of the modulation of the basic pattern was related to wrist position: EMG amplitude was greatest at the extreme of muscle shortening. There was a slight difference in EMG activity between flexion and extension ramps that was related purely to the direction of movements, independent of wrist position, velocity, and external load; EMG amplitude was greater when a muscle was shortening and less when it was lengthening. During ramp movement, there was little or no observed EMG activity related to velocity (8-28 degrees/s). The magnitude of EMG activity varied in proportion to the external torque load, but this load-related component was additive, and the basic pattern of activity (related to direction and position) did not change with load. From these results we infer that a muscle's EMG activity was determined by 1) passive elastic properties of the wrist and the active length-tension characteristics of the muscle itself (position), 2) asymmetries in the muscle's contractile force depending on whether it was lengthening or shortening (direction), and 3) magnitude of the external torque load (force). By contrast, since no EMG activity was related to velocity in these slow movements, passive viscous properties and velocity-related cross-bridge kinetics were apparently so slight as to make undetectable the small additional EMG activity and contractile force presumably required to overcome them. A model of the muscle forces acting at the wrist incorporates these experimental observations.     

Cutaneous receptors contribute to kinesthesia at the index finger, elbow, and knee

The neural mechanisms underlying the sense of joint position and movement remain controversial. While cutaneous receptors are known to contribute to kinesthesia for the fingers, the present experiments test the hypothesis that they contribute at other major joints. Illusory movements were evoked at the interphalangeal (IP) joints of the index finger, the elbow, and the knee by stimulation of populations of cutaneous and muscle spindle receptors, both separately and together. Subjects matched perceived movements with voluntary movements of homologous joints on the contralateral side. Cutaneous receptors were activated by stretch of the skin (using 2 intensities of stretch) and vibration activated muscle spindle receptors. Stimuli were designed to activate receptors that discharge during joint flexion. For the index finger, vibration was applied over the extensor tendons on the dorsum of the hand, to evoke illusory metacarpophalangeal (MCP) joint flexion, and skin stretch was delivered around the IP joints. The strong skin stretch evoked the illusion of flexion of the proximal IP joint in 6/8 subjects (12 +/- 5 degrees, mean +/- SE). For the group, strong skin stretch delivered during vibration increased the perceived flexion of the proximal IP joint by eight times with a concomitant decrease in perceived flexion of the MCP joint compared with vibration alone (P < 0.05). For the elbow, vibration was applied over the distal tendon of triceps brachii and skin stretch over the dorsal forearm. When delivered alone, strong skin stretch evoked illusory elbow flexion in 5/10 subjects (9 +/- 4 degrees). Simultaneous strong skin stretch and vibration increased the illusory elbow flexion for the group by 1.5 times compared with vibration (P < 0.05). For the knee, vibration was applied over the patellar tendon and skin stretch over the thigh. Skin stretch alone evoked illusory knee flexion in 3/10 subjects (8 +/- 4 degrees) and when delivered during vibration, perceived knee flexion increased for the group by 1.4 times compared with vibration (P < 0.05). Hence inputs from cutaneous receptors, muscle receptors, and combined inputs from both receptors likely subserve kinesthesia at joints throughout the body.     

Changes in movement final position associated with agonist and antagonist muscle fatigue

We have tested the hypothesis that agonist and antagonist muscle fatigue could affect the final position of rapid, discrete movements. Six subjects performed consecutive elbow flexion and extension movements between two targets, with their eyes closed prior to, and after fatiguing the elbow extensor muscles. The results demonstrate that elbow extension movements performed in the post-test period systematically undershot the final position as compared to pre-test movements. However, attainment of the aimed final position in elbow flexion movements was unaffected by fatiguing of the extensor muscles. Undershoot of the final position obtained in extension movements was associated with agonist muscle fatigue, a result that was expected from the point of view of current motor control theories, and that could be explained by a reduced ability of the shortening muscle to exert force. On the other hand, the absence of the expected overshoot of the final position when the antagonist is fatigued, indicates the involvement of various reflex and/or central mechanisms operating around the stretched muscle that could contribute to returning the limb to the standard final position after a brief prominent overshoot.     

Jaw-opening accuracy is not affected by masseter muscle vibration in healthy men

There is a functional integration between the jaw and neck regions with head extension–flexion movements during jaw-opening/closing tasks. We recently reported that trigeminal nociceptive input by injection of hypertonic saline into the masseter muscle altered this integrated jaw–neck function during jaw-opening/closing tasks. Thus, in jaw-opening to a predefined position, the head–neck component increased during pain. Previous studies have indicated that muscle spindle stimulation by vibration of the masseter muscle may influence jaw movement amplitudes, but the possible effect on the integrated jaw–neck function is unknown. The aim of this study was to investigate the effect of masseter muscle vibration on jaw–head movements during a continuous jaw-opening/closing task to a target position. Sixteen healthy men performed two trials without vibration (Control) and two trials with bilateral masseter muscle vibration (Vibration). Movements of the mandible and the head were registered with a wireless three-dimensional optoelectronic recording system. Differences in jaw-opening and head movement amplitudes between Control and Vibration, as well as achievement of the predefined jaw-opening target position, were analysed with Wilcoxon’s matched pairs test. No significant group effects from vibration were found for jaw or head movement amplitudes, or in the achievement of the target jaw-opening position. A covariation between the jaw and head movement amplitudes was observed. The results imply a high stability for the jaw motor system in a target jaw-opening task and that this task was achieved with the head–neck and jaw working as an integrated system.     

Perceptual and motor effects of agonist-antagonist muscle vibration in man

Summary Perceptual and motor effects of vibration applied simultaneously to the distal tendons of the Biceps and Triceps muscles, in isometric conditions and without sight of the stimulated arm, have been studied in human volunteers. Motor effects, measured by surface EMG, are inexistent when the flexor and extensor muscles are simultaneously vibrated at the same frequency. However, EMG activity appears in the muscle being vibrated at the lower frequency when simultaneous vibration is applied at different frequencies. The sensations felt by the subjects were reproduced by the nonvibrated arm and recorded by a goniometer. The studies show that the velocity and the amplitude of the ilusory movement is related to the difference in vibration frequency applied to the two muscles. The direction of movement felt (flexion or extension) is that produced by shortening of the muscle being vibrated at the lower frequency. When the two vibration frequencies are the same, there is either no sensation of movement, or a sensation of very slow movement. These results support the notion that the sensation of movement at a joint may be derived from a central processing of the proprioceptive inflow data obtained from flexor and extensor muscles. This interpretation may also be valid for the results obtained earlier by vibration of a single muscle. Furthermore, it is coherent with data on spindle afferent fibres obtained by microneurography in man during passive or active movements.This work was supported by grants from the Ministère de l'Industrie et de la Recherche     

Relationship between EMG patterns and kinematic properties for flexion movements at the human wrist

Summary EMG patterns associated with voluntary wrist flexion movements were studied in normal human subjects. Initially, subjects were trained to produce movements within five specified velocity ranges while the amplitude of the movement and the opposing load remained constant. In a second set of experiments, subjects were required to produce movements at four different amplitudes, moving as rapidly as possible against a constant load. Finally, with movement velocity and amplitude kept constant, the external load was varied so that different forces were required to generate the movements. The slowest movements were associated with a prolonged burst of EMG activity from the agonist muscle with little or no antagonist activity. With increasing movement velocity, there was a gradual evolution to the characteristic triphasic pattern associated with rapid voluntary movements. As velocity of movement increased further, the amplitude and area of the EMG bursts increased while burst duration and interburst intervals decreased. Increases in movement amplitude were accomplished mainly by changing the timing of the EMG bursts; with larger amplitude movements the antagonist burst occurred later. With movements against larger loads there was an increase in the size of the agonist burst and a decrease in the antagonist burst, but no change in the relative timing of the EMG bursts. These systematic changes in EMG patterns associated with different types of movement provide an indirect method of obtaining information concerning the motor programs which generate the movements.     

Trunk muscle fatigue and associated EMG changes during a dynamic iso-inertial test

This study was designed to investigate the relationship between trunk muscle fatigue and associated changes in the electromyographic (EMG) signals during a dynamic iso-inertial test. Eleven subjects performed dynamic trunk flexion/extension movements against 40% maximum voluntary contraction (MVC) torque until exhaustion in a tri-axial trunk dynamometer. EMG parameters in the time and frequency domain were studied by analysing changes of the signal amplitudes and the spectral density (using the zero-crossing-rate and the median frequency). The kinematics of the movement were analysed according to the movement velocities and the deviations from the required movement plane. The flexion and extension velocities decreased from the beginning to the end of the test. Movement deviations from the sagittal plane into the frontal and transverse plane increased with increasing test duration, as did the EMG amplitude. The median frequency during periods with maximum muscle activity decreased, as did the zero-crossing-rate. The increase in amplitude and decrease in median frequency were more pronounced in the trunk flexors than in the trunk extensors. The parameters of median frequency, zero-crossing-rate and amplitude seem to be sensitive identifiers of muscle fatigue during well-controlled dynamic contractions. While the kinematic data did not yield any information on the mechanisms of the fatigue, changes in the EMG parameters demonstrated that the duration of the test was limited by the fatigue of the trunk flexors.     

Supraspinal control of spinal centers for antagonist muscles in man

Summary Monosynaptic testing with the H-reflex was used to determine reflex excitability of motoneurons of the gastrocnemius muscle during single voluntary movements: extension or flexion of the ankle. For the last 60 msec of the latent period before the onset of voluntary extension of the foot, reflex excitability of motoneurons of the gastrocnemius (the agonist in extension) gradually increases. With voluntary flexion of the ankle reflex excitability of motoneurons to the gastrocnemius (the antagonist in flexion) is unchanged throughout the latent period until the onset of movement. Simultaneously (accuracy to 10 msec) with the beginning of the myogram of voltary foot flexion, reflex excitability of motoneurons of the gastrocnemius (antagonist) drops sharply. These results provide a basis for discussing an hypothesis concerning supraspinal control of spinal centers for antagonist muscles in man.