Coordination and inhomogeneous activation of human arm muscles during isometric torques

1. In this study we have recorded the activity of motor units of the important muscles acting across the elbow joint during combinations of voluntary isometric torques in flexion/extension direction and supination/pronation direction at different angles of the elbow joint. 2. Most muscles are not activated homogeneously; instead the population of motor units of muscles can be subdivided into several subpopulations. Inhomogeneous activation of the population of motor units in a muscle is a general finding and is not restricted to some multifunctional muscles. 3. Muscles can be activated even if their mechanical action does not contribute directly to the external torque. For example, m. triceps is activated during supination torques and thus compensates for the flexion component of the m. biceps. On the other hand, motor units in muscles are not necessarily activated if their mechanical action contributes to a prescribed torque. For example, there are motor units in the m. biceps that are activated during flexion torques, but not during supination torques. 4. The relative activation of the muscles depends on the elbow angle. Changing the elbow angle affects the mechanical advantage of different muscles differently. In general, muscles with the larger mechanical advantage receive the larger input. 5. We have calculated the relative contributions of some muscles to isometric torques. These contributions depend on the combination of the torques exerted. 6. Existing theoretical models on muscle coordination do not incorporate subpopulations of motor units and therefore need to be amended.     

Short- and long-latency reflex responses during different motor tasks in elbow flexor muscles

 Stretch reflex responses in three elbow flexor muscles – the brachioradialis and the short and long heads of the biceps brachii – were studied during different motor tasks. The motor tasks were iso-velocity (8 deg/s) elbow flexion movements in which the muscles performed shortening or lengthening contractions, or were isometric contractions. Care was taken to maintain constant background electromyographic (EMG) activity in the brachoradialis muscle at a 50-deg elbow angle across the tasks by changing the magnitude of the initial load. During each task, mechanical perturbations (duration 170 ms) were applied at pseudorandom intervals when the elbow angle was 50 deg. The magnitude of the perturbation was varied across tasks in order to induce an elbow extension velocity of 80 deg/s over the first 50 ms after the onset of perturbation. The stretch reflex EMG responses in all muscles varied across the three tasks, despite a constant EMG level and similar perturbation-induced angular velocity in the direction of elbow extension. In particular, both the short- and long-latency reflex EMG components were reduced during the lengthening contractions. Further, the task-dependent variations in the early (M2) and the late (M3) components of the long-latency reflex were different, i.e., the magnitude of M3 was considerably enhanced during the shortening task as compared with that of M2. These findings suggest that central modification was responsible for the task-dependent modulation of late EMG responses. Received: 24 April 1996 / Accepted: 24 January 1997     

Differences in activation patterns in elbow flexor muscles during isometric,concentric and eccentric contractions

Summary To investigate the relative activation of the synergistic muscles during three different types of muscle contraction, the electromyograms (EMG) of two elbow flexor muscles, the biceps brachii (BB) and the brachioradialis (BR), have been compared. To accomplish this eight healthy human subjects performed the following elbow flexions against the same load — concentric, eccentric and isometric contractions. The isometric contractions were performed at three elbow angles: 10, 45 and 90° (0° equal to full expension). The EMG were recorded by bipolar surface electrodes, and the relative activation between the two muscles was evaluated as the quotient of mean EMG activities (BR/BB). For the isotonic elbow flexions, BR/BB were calculated at three angle divisions: 0–30°, 30–60° and 60–90°. Results indicated that the relative activation of the BR during the concentric contractions was higher than that of the eccentric contraction, particularly at the extended elbow angles, i.e. the BR/BB of the concentric contractions for the elbow joint angles ranging from 0–30° and 30–60° were significantly greater (P<0.05) than those of the eccentric contractions. During the isometric and eccentric contractions, the BR/BB at the flexed joint angles tended to be greater than those at the extended angles. In contrast, there were no angle-dependent BR/BB variations during the concentric elbow flexions. Further, changing patterns in the EMG power spectra due to the type of contraction were different between BB and BR. These results indicated that the activation pattern in the two elbow flexor muscles varied with the muscle contraction pattern.     

EMG patterns in antagonist muscles during isometric contraction in man: Relations to response dynamics

Summary We studied the EMG activity of biceps and triceps in human subjects during isometric force adjustments at the elbow. Rapid targeted force pulses exhibited stereotyped trajectories in which peak force was a linear function of the derivatives of force and the time to peak force was largely independent of its amplitude. These responses were associated with an alternating triphasic pattern of EMG bursts in agonist and antagonist muscles similar to that previously described for rapid limb movements. When the instructions demanded rapid force pulses, initial agonist bursts were of constant duration, and their magnitude was strongly related to peak force achieved. The timing of EMG bursts in antagonist pairs was closely coupled to the dynamics of the force trajectory, and the rising phase of the force was determined by both agonist and antagonist bursts. When peak force was kept constant and rise time systematically varied, the presence and magnitude of antagonist and late agonist bursts were dependent on the rate of rise of force, appearing at a threshold value and then increasing in proportion to this parameter. It is proposed that antagonist activity compensates for nonlinearity in muscle properties to enable the linear scaling of targeted forces which characterizes performance in this task.Supported by the Dystonia Medical Research Foundation and NS 19205     

Intramuscular EMG from the hip flexor muscles during human locomotion

The purpose was to investigate the activation pattern of five major hip flexor muscles and its adaptation to changing speed and mode of progression. A total of 11 healthy subjects performed walking and running on a motor-driven treadmill at speeds ranging from 1.0 to 6.0 m s^?1. Intramuscular fine-wire electrodes were used to record myoelectric signals from the iliacus, psoas, sartorius, rectus femoris and tensor fascia latae muscles. The basic pattern, with respect to number of activation periods, remained the same irrespective of speed and mode of progression. However, differences in the relative duration and timing of onset of activation occurred between individual muscles. Over the speed range in walking, a progressively earlier onset was generally seen for the activation period related to hip flexion. Changes in EMG amplitude were measured in the iliacus and psoas muscles and showed a marked increase and difference between walking and running at speeds above 2.0 m s^?1. Thus, the alternating flexion–extension movements at the hip during locomotion appear to be governed by a rather fixed `neural program' which normally only needs minor modulations to accomplish the adjustments accompanying an increase in speed of progression as well as a change from walking to running.     

Matching different levels of isometric torque in elbow flexor muscles after eccentric exercise

Human subjects generated a specified level of isometric torque with elbow flexor muscles of one arm, the reference arm, under visual feedback. They were then asked to generate what they perceived to be the same level, with the other arm, the indicator, but with no visual feedback. A number of torque levels, between 2% and 30% of maximum were used in the matching trials. Elbow flexors of one arm were then exercised eccentrically on a dynamometer. Immediately after the exercise, there was a large (40%) drop in maximum voluntary torque, as well as some soreness and swelling 24 h later, indicative of muscle damage. When the torque-matching experiment was repeated after the indicator arm had been exercised, the indicator signalled torque levels significantly below the reference level (P<0.05). When the reference arm was exercised, errors were in the opposite direction. Over the 4 days of testing post-exercise, errors became less as torque levels returned to normal. When errors were expressed in terms of maximum torque post-exercise, they were significantly reduced. This suggested that subjects were using as a matching cue the perceived effort required to generate a given level of torque rather than the level of torque itself. Persisting matching errors, from 24 h onwards after the eccentric contractions, were proposed to include a component attributable to the muscle soreness. Changes in electromyogram recorded after eccentric exercise were consistent with the effort-matching hypothesis. The muscle's torque–angle relationship was used to estimate matching ability in the absence of fatigue. One forearm was placed at various angles and its reference torque was matched by the other, the indicator, always at 90°. Again, matching errors were consistent with an interpretation based on a match of effort rather than torque. Electronic Publication     

Electrical and mechanical response of finger flexor muscles during voluntary isometric contractions in elite rock-climbers

To determine the differences between rock-climbers and controls in finger flexor (FF) motor units (MUs) features and activation strategy, eleven climbers and ten controls volunteered for the study. After maximal voluntary contraction (MVC) assessment, five levels of isometric contractions at 20, 40, 60, 80 and 100% MVC were performed. During contractions, electromyogram (EMG) and mechanomyogram (MMG) were recorded, from which the root mean square (RMS) and mean frequency (MF) were calculated. Climbers showed significantly higher MVC. EMG RMS was statistically higher in climbers than in controls from 60 to 100% MVC. In climbers MMG RMS increased up to 80% MVC, whereas in controls it increased only up to 60% MVC. MMG MF was higher in climbers than in controls from 60 to 100% MVC (P < 0.05). EMG–MMG combined analysis revealed significant differences in MU activation strategy between the two groups. The results are compatible with a shift of climbers’ muscles toward faster MUs.     

Motor unit firing behavior in human arm flexor muscles during sinusoidal isometric contractions and movements

Simultaneous recordings of action potentials (APs) of multiple single motor units (MUs) were obtained in brachialis and biceps (caput breve) muscles during sinusoidally modulated isometric contractions of elbow flexor muscles and during sinusoidal flexion/extension movements in the elbow against a preload in the extension direction. The results show that MUs typically fire in one short burst for each sinusoidal cycle. The mean phase lead of the bursts of APs relative to a sinusoidally modulated isometric torque in the elbow joint or relative to sinusoidal movements in the elbow increases gradually with frequency. The increase of the mean phase lead during isometric contractions was very similar for all MUs and could be explained well by modeling the force production of MUs with a second-order linear low-pass system. For sinusoidal flexion/extension movements each MU reveals a specific, reproducible phase lead as a function of frequency. However, there is a large variability in phase behavior between MUs. Also, the modulation of the firing rate for sinusoidal isometric contractions versus sinusoidal movements appeared to be different for various MUs. In simultaneous recordings some MUs clearly revealed a larger firing rate in each burst for movements relative to isometric contractions, whereas other MUs revealed a smaller firing rate. This suggests that some MUs are preferentially activated during movements whereas others are preferably activated during isometric contractions. The results demonstrate task-dependent changes in the relative activation of MUs within a single muscle for sinusoidal isometric contractions and movements. Received: 18 November 1996 / Accepted: 3 April 1997     

The relationship between the electromyogram-amplitude and isometric extension torques of neck muscles at different positions of the cervical spine

A group of 12 healthy men volunteered for the experiment. Electromyograms (EMG) were obtained from semispinalis capitis, splenius capitis, levator scapulae, and trapezius muscles. The flexion angle of the cervical spine was precisely adjusted to 0°, 10°, 20°, and 30° relative to the horizontal, with a constant angle of the atlanto-occipital joint. The subjects made eight short (about 2 s) vertical extension forces (6%, 12%,18%, 24%, 30%, 36%, 42%, and 48% of maximal voluntary peak contraction force). For each position, the centre of pressure under the head was determine as the basis for the calculation of the external lever arm. The presence of motor endplate regions was ascertained by multiple surface electrodes. The slopes of individual linear regression lines for the root mean square (rms)-values were dependent on the existence of endplates in the area of the electrodes — endplates caused smaller rms values per Newton metres of external torque. Significant intersubject differences between regression equations could not be eliminated by the normalization of EMG-parameters and/or torques. The elimination of gravity, the continuous monitoring of positions, and the consideration of localization of motor endplate regions were essential prerequisites for the acquisition of reliable relationships between EMG of different neck muscles and external torques. Two important conclusions were derived for the prediction of torques from EMG measurements: firstly, individual regression equations which take into account the position of the head and neck should be used; secondly, normalization procedures do not justify the application of average regressions to a group of subjects.     

Differences in stretch reflex responses of elbow flexor muscles during shortening, lengthening and isometric contractions

Stretch reflexes were evoked in elbow flexor muscles undergoing three different muscle contractions, i.e. isotonic shortening (SHO) and lengthening (LEN), and isometric (ISO) contractions. The intermuscle relationships for the magnitude of the stretch reflex component in the eletromyographic (EMG) activities of two main elbow flexor muscles, i.e. the biceps brachii (BB) and the brachioradialis (BRD), were compared among the three types of contractions. The subjects were requested to move their forearms sinusoidally (0.1?Hz) against a constant pre-load between elbow joint angles of 10° (0°?=?full extension) and 80° during SHO and LEN, and to keep an angle of 45° during the ISO. The perturbations were applied at the elbow angle of 45° in pseudo-random order. The EMG signals were rectified and averaged over a period of 100?ms before and 400?ms after the onset of the perturbation 40–50 times. From the ensemble averaged EMG waveform, the background activity (BGA), short (20–50?ms) and long latency (M2, 50–80, M3, 80–100?ms) reflex and voluntary activity (100–150?ms) components were measured. The results showed that both BGA and reflex EMG activity of the two elbow flexor muscles were markedly decreased during the lengthening contraction compared to the SHO and ISO contractions. Furthermore, the changes of reflex EMG components in the BRD muscle were more pronounced than those in the BB muscle, i.e. the ratios of M2 and M3 magnitudes between BRD and BB (BRD:BB) were significantly reduced during the LEN contractions. These results would suggest that the gain of long latency stretch reflex EMG activities in synergistic muscles might be modulated independently according to the model of muscle contraction.     

Differences in coordination of elbow flexor muscles in force tasks and in movement tasks

Summary Motor-unit activity in m. biceps brachii, m. brachialis and m. brachioradialis during isometric contractions has been compared with motor-unit activity during slow voluntary (extension and flexion) movements made against external loads. During these slow movements the recruitment threshold of m. biceps motor units is considerably lower than it is during isometric contractions but the recruitment threshold of both m. brachialis and m. brachioradialis motor units is considerably higher. For all three elbow flexor muscles the motor-unit firing frequency seems to depend on the direction of movement: the firing frequency is higher during flexion movements (3 deg/s) and lower during extension movements (−3 deg/s) than during isometric contractions. The relative contribution of the biceps to the total exerted flexion torque during slow voluntary movements is estimated to increase from 36% to about 48% and that of the brachialis/brachioradialis is estimated to decrease from 57% to about 45% compared to the relative contribution of these muscles during isometric contractions. This difference in the relative contribution of the three major elbow flexor muscles is shown to be caused by differences in the central activation in force tasks and movement tasks.     

EMG power spectra of trunk muscles during graded maximal voluntary isometric contraction in flexion-rotation and extension-rotation

The purpose of this study was to determine the electromyographic (EMG) power spectral characteristics of seven trunk muscles bilaterally during two complex isometric activities extension-rotation and flexion-rotation, in both genders to describe the frequency-domain parameters. Eighteen normal young subjects volunteered for the study. The subjects performed steadily increasing isometric extension-rotation and flexion-rotation contractions in a standard trunk posture (40° flexed and 40° rotated to the right). A surface EMG was recorded from the external and internal oblique, rectus abdominis, pectoralis, latissimus dorsi, and erector spinae muscles at the 10th thoracic and the 3rd lumbar vertebral levels, at 1?kHz and 25%, 50%, 75% and 100% of maximal voluntary contraction (MVC). The median frequency (MF), mean power frequency (MPF), frequency spread and peak power were obtained from fast Fourier transform analysis. The MF and MPF for both extension-rotation and flexion-rotation increased with the grade of contraction for both males and females. The EMG spectra in flexion-rotation were different from those of extension-rotation (P?P?相似文献   

Energy metabolism in different human skeletal muscles during voluntary isometric contractions

Summary The energy turnover in contracting skeletal muscle was studied by measuring the rate of temperature rise during voluntary, isometric contractions and circulatory arrest in M. soleus, M. sacrospinalis and M. biceps brachii in 14 males, by thermoelements inserted in the muscles. A linear relationship between rate of temperature rise and force intensity given as per cent of maximal voluntary contraction (MVC) was demonstrated in biceps (r=0.95), but not so clearly confirmed in soleus (r=0.73). Muscle biopsies were taken from the same muscles and fibre type distribution was determined histochemically by staining for ATPase. The rate of heat production at MVC showed positive correlation to the percentage of fast twitch (FT) fibres in the muscles (r=0.90). Linear extrapolation indicates that the maximal energy turnover in human FT fibres is approximately six times that of slow twitch (ST) fibres during voluntary isometric contractions.This work was submitted by G. Bolstad as a thesis to the University of Bergen in June 1975, in partial fulfilment of the requirements for the degree of Candidatus realium     

Influence of neural adjustments and muscle oxygenation on task failure during sustained isometric contractions with elbow flexor muscles

This study investigated the adjustments in muscle activation and oxygenation in biceps and triceps brachii during two tasks sustained to failure at 20 and 60% of the maximal voluntary contraction (MVC) force. The tasks required participants either to push against a rigid restraint (force task) or to support an inertial load (position task) with the elbow flexor muscles. The surface EMG was recorded for biceps brachii, brachioradialis, triceps brachii and trapezius superior muscles. Muscle oxygenation of biceps and triceps brachii was measured by near-infrared spectroscopy. The position task was briefer (404 ± 159 s) than the force task (533 ± 194 s) when performed at 20% MVC (P = 0.011), but endurance time did not differ at 60% MVC (54 ± 19 versus 64 ± 16 s, respectively; P = 0.13). Biceps brachii oxygenation decreased slightly (by ~7%) during tasks performed at 20% MVC, whereas it dropped (-40%) for tasks sustained at 60% MVC. However, the decrease in muscle oxygenation was not a significant predictor of time to failure at the two target forces, although its contribution to muscle fatigue cannot be completely ruled out at 60% MVC. In contrast, time to failure was predicted by the increase in EMG of biceps brachii for both tasks at 20% MVC, and EMG of brachioradialis and trapezius for both tasks at 60% MVC. These results suggest that neural adjustments rather than muscle oxygenation limited the time to failure for the force and position tasks at low and high target forces.     

Activation of agonist and antagonist muscles at different joint angles during maximal isometric efforts

The purpose of this study was to investigate the influence of different angles of the knee joint on the activation level of an agonist (quadriceps femoris muscle) and antagonist (biceps femoris muscle) from electromyographic activities and activation levels (twitch interpolation). Isometric torque measurements were performed on 23 healthy subjects at 10° intervals between 40° and 110° of knee joint flexion. Superimposed twitches at maximal voluntary contraction were applied and the voluntary activation estimated. To quantify the antagonist muscle activity, we normalized its integrated EMG (iEMG) value at each joint angle with respect to its iEMG value at the same angle when acting as an agonist at maximal effort. The activation levels at the knee-flexed position (80–110°) were higher than that at the knee-extended position (40–70°). The co-activation levels at 90, 100, and 110° were significantly higher than that the other knee angle. These results suggest that the activation level of an agonist (quadriceps femoris) muscle and the co-activation level of an antagonist (biceps femoris) muscle were higher in longer muscles than in shorter muscles. It was also concluded that the risk of knee injuries could be reduced by applying these mutual relationships between activation levels of agonist and antagonist muscles.     

Computer analysis of the surface EMG during isometric exercise

A computer program is described which can provide either on-line or off-line analysis of the surface electromyogram (EMG) in man during submaximal isometric contractions. The intention of the program was to quantify the tension developed by, and the degree of fatigue induced in muscle during this form of exertion from the surface EMG. Six male and female volunteers served as subjects to test the program by exerting brief isometric contractions of their handgrip muscles at tensions ranging from 3 to 100% of their maximum strength and fatiguing isometric contractions at tensions ranging between 20 and 95% of their maximum strength. The results of these experiments showed that the program was able to estimate the tension developed by muscle within an average of 2.4% of the subject's maximum strength and estimate the duration of fatigue contractions within an average of 3.5% of the actual recorded endurance. The applications of this form of analysis to work physiology are discussed.     

Tendon vibration during submaximal isometric strength and postural tasks

To establish the effect of tendon vibration (TV) on steady submaximal strength and static balance control, 29 women performed isometric plantarflexions at 10, 20, 30 and 50% of MVC and postural tasks of increasing difficulty (Normal Quiet Stance, NQS, Sharpened Tandem Stance and One Leg Stance) with vibrators (80?Hz) applied to the Achilles tendon. Both tasks were performed under four conditions (10?s each): eyes open, eyes closed, eyes closed with TV, eyes open with TV. During the isometric tasks, the application of TV increased the plantarflexion torque at 20 and 30% of MVC, accompanied by a greater (p?<?0.001) activation of agonists (Medial Gastrocnemius, MGAS and Soleus, SOL), as well as a greater (p?<?0.05) coactivation of the antagonist (tibialis anterior, TA). Non-significant differences were observed at 10 and 50% of MVC. During posture, TV resulted in a backward displacement of the Centre of Pressure only for NQS and an increase (p?<?0.01) in EMG activity of the ankle muscles in all balance tasks. The MGAS contraction during TV application was confirmed (by ultrasonography) at ten participants by an increase of the pennation angle and decrease of its fascicle length (p?<?0.05). The vibration-induced muscle excitation and accompanying increase in motor output, probably due to excitatory Ia afferent input, was confirmed during strength and postural tasks. However, motor output attenuates when the magnitude of central drive to the ankle muscles increases.     

Physiological aspects of MMG and EMG spectra during load-varying isometric dorsiflexion

The aim of the study was to examine spectral features of the mechanomyogram (MMG) and electromyogram (EMG) during static and load-varying isometric dorsiflexion to characterize force control strategies of the tibialis anterior. Twelve healthy subjects performed two motor tasks including 1) four exertion levels of static isometric dorsiflexion and 2) load-varying isometric dorsiflexion while tracking a target quasi-sinusoidal curve of three different amplitudes. Generally speaking, for both static and load-varying isometric contractions, the mean frequency of MMG-EMG cross spectra (MMG-EMG MF) progressively increased with effort level, whereas the median frequency of EMG auto spectra among higher effort levels remained unchanged. The MMG-EMG MF versus EMG root mean square regression slope was significantly larger for load-varying isometric contraction than for static contraction control measurements. These findings highlight effort-dependent and task-specific rate coding for force regulation of the tibialis anterior.     

EMG power spectra of elbow extensors during ramp and step isometric contractions

Summary The goal of the present study was to compare electromyogram (EMG) power spectra obtained from step (constant force level) and ramp (progressive increase in the force level) isometric contractions. Data windows of different durations were also analysed for the step contractions, in order to evaluate the stability of EMG power spectrum statistics. Fourteen normal subjects performed (1) five ramp elbow extensions ranging from 0 to 100% of the maximum voluntary contraction (MVC) and (2) three stepwise elbow extensions maintained at five different levels of MVC. Spectral analysis of surface EMG signals obtained from triceps brachii and anconeus was performed. The mean power frequency (MPF) and the median frequency (MF) of each power spectrum were obtained from 256-ms windows taken at 10, 20, 40, 60 and 80% MVC for each type of contraction and in addition on 512-, 1024-and 2048-ms windows for the step contractions. No significant differences (P>0.05) were found in the values of both spectral statistics between the different window lengths. Even though no significant differences (P>0.05) were found between the ramp and the step contractions, significant interactions (P<0.05) between these two types of contraction and the force level were found for both the MPF and the MF data. These interactions point out the existence of different behaviours for both the MPF and the MF across force levels between the two types of contraction.