Reflex and intrinsic changes induced by fatigue of human elbow extensor muscles

Fatigue-induced changes in intrinsic and reflex properties of human elbow extensor muscles and the underlying mechanisms for fatigue compensation were investigated. The elbow joint was perturbed using small-amplitude and pseudorandom movement patterns while subjects maintained steady levels of mean joint extension torque. Intrinsic and reflex properties were identified simultaneously using a nonlinear delay differential equation model. Intrinsic joint properties were characterized by measures of joint stiffness, viscous damping, and limb inertia and reflex properties characterized by measures of dynamic and static reflex gains. Fatigue was induced using 15 min of intermittent voluntary isometric (submaximal) exercise, and a rest period of 10 min was taken to allow the fatigued muscles to recover from acute fatigue effects. Identical experimental and data analysis procedures were used before and after fatigue. Our findings were that after fatigue, joint stiffness was significantly reduced at higher torque levels, presumably reflecting the reduced force-generating capacity of fatigued muscles. Conversely, joint viscosity was increased after fatigue potentially because of the reduced crossbridge detachment rate and prolonged relaxation associated with intracellular acidosis accompanying fatigue. Static stretch reflex gain decreased significantly at higher torque levels after fatigue, indicating that the isometric fatiguing exercise might be associated with a preferential change in properties of spindle chain fibers and bag(2) fibers. For matched pre- and postfatigue torque levels, dynamic reflexes contributed relatively more torque after fatigue, displaying higher dynamic reflex gains and larger dynamic electromyographic responses elicited by the controlled small-amplitude position perturbations. These changes appear to counteract the fatigue-induced reductions in joint stiffness and static reflex gain. The compensatory responses could be partly due to the effects of increasing the number of active motoneurons innervating the fatiguing muscles. This shift in operating point gave rise to significant compensation for the loss of contractile force. The compensation could also be due to fusimotor adjustment, which could make the dynamic reflex gain much less sensitive to fatigue than intrinsic stiffness. In short, the reduced contribution from intrinsic stiffness to joint torque was compensated by increased contribution from dynamic stretch reflexes after fatigue.     

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.     

Fatigue induced changes in phasic muscle activation patterns for fast elbow flexion movements

The present study investigated how muscle fatigue influences single degree-of-freedom elbow flexion movements and their associated patterns of phasic muscle activation. Maximal unfatigued voluntary isometric elbow flexor and extensor joint torque was measured at the beginning of the experiment. Subjects then performed elbow flexion movements over two distances as fast as possible, and movements over the longer distance at an intentionally slower speed. The slower speed was close to what would become the maximal speed in the fatigued state. Subjects then performed a fatiguing protocol of 20 sustained isometric flexion contractions of 25 s duration with 5 s rest at 50% maximal unfatigued voluntary force. After a recovery period they repeated the movements. The fatigue protocol was successful in inducing muscle fatigue, the evidence being decreased isometric maximal joint torque of over 20%. Fatigued movements had lower peak muscle torque and speed. Our principal finding was of changes in the timing of the phasic patterns of fatigued muscle activation. There was an increase in the duration of the agonist burst and a delay in the timing of the antagonist muscle as measured by the centroid of the EMG signals. We conclude that these changes serve as partial but incomplete, centrally driven compensation for fatigue induced changes in muscle function. An additional, unexpected finding was how small an effect fatigue had on movement performance when using a recovery time of 10 min that is long enough to allow muscle membrane conduction velocity to return to normal. This raises questions concerning the behavioral significance of classical laboratory studies of human fatigue mechanisms.     

The effects of muscle fatigue and movement height on movement stability and variability

Performing repetitive manual tasks can lead to muscle fatigue, which may induce changes in motor coordination, movement stability, and kinematic variability. In particular, movements performed at or above shoulder height have been associated with increased shoulder injury risk. The purpose of this study was to determine the effects of repetitive motion-induced muscle fatigue on posture and on the variability and stability of upper extremity movements. Ten healthy subjects performed a repetitive task similar to sawing continuously until volitional exhaustion. This task was synchronized with a metronome to control movement timing. Subjects performed the sawing task at shoulder (“High”) and sternum height (“Low”) on two different days. Joint angles and muscle activity were recorded continuously. Local and orbital stability of joint angles, kinematic variability (within subject standard deviations), and peak joint angles were calculated for five bins of data spaced evenly across each trial. Subjects fatigued more quickly when movements were performed at the High height. They also altered their kinematic patterns significantly in response to muscle fatigue. These changes were more pronounced when the task was performed at the High height. Subjects also exhibited increased kinematic variability of their movements post-fatigue. Increases in variability and altered coordination did not lead to greater instability, however. Shoulder movements were more locally stable when the task was performed at the High height. Conversely, shoulder and elbow movements were more orbitally unstable for the High condition. Thus, people adapt their movement strategies in multi-joint redundant tasks and maintain stability in doing so.     

Patterns of hypermetria and terminal cocontraction during point-to-point movements demonstrate independent action of trajectory and postural controllers

We examined elbow muscle activities and movement kinematics to determine how subjects combine elementary control actions in performing movements with one and two trajectory segments. In reaching, subjects made a rapid elbow flexion to a visual target before stabilizing the limb with either a low or a higher level of elbow flexor/extensor coactivity (CoA), which was cued by target diameter. Cursor diameter provided real-time biofeedback of actual muscle CoA. In reversing, the limb was to reverse direction within the target and return to the origin with minimal CoA. We previously reported that subjects overshoot the goal when attempting a reversal after first having learned to reach accurately to the same target. Here we test the hypothesis that this hypermetria results because reversals co-opt the initial feedforward control action from the preceding trained reach, thereby failing to account for task-dependent changes in limb impedance induced by differences in flexor/extensor coactivity as the target is acquired (higher in reaching than reversing). Instructed increases in elbow CoA began mid-reach, thus increasing elbow impedance and reducing transient oscillations present in low CoA movments. Flexor EMG alone increased at movement onset. Test reversals incorporated the initial agonist activity of previous reaches but not the increased coactivity at the target, thus leading to overshoot. Moreover, we observed elevated coactivity in reversals upon returning to the origin even though coactivity in reaching was centered at the goal target. These findings refute the idea that the brain necessarily invokes distinct unitary control actions for reaches and reversals made to the same target. Instead, reaches and reversals share a common control action that initiates trajectories toward their target and another later control action that terminates movement and stabilizes the limb about its final resting posture, which differs in the two tasks.     

The role of cocontraction in the impairment of movement accuracy with fatigue

The present experiment was designed to test the hypothesis that fatigue-induced impairment in movement accuracy is caused by a decrease in muscle cocontraction rather than a reduced ability to produce muscular force. Seven participants performed fast and accurate elbow extensions aimed at a target, before and after a fatigue protocol. The inertia of the manipulandum was decreased after the fatigue protocol so that the ratio of required to available force during movements was identical pre- and post-fatigue. After the fatigue protocol, movement endpoint accuracy decreased and movement endpoint variability increased. These alterations were associated with a decrease in cocontraction. We concluded that the impairment of movement accuracy during fatigue could not be explained by the lack of available force, but was likely to be due to a fatigue-induced decrease in muscular cocontraction. We then speculate that fatigue influences the relative weights of accuracy and energy economy in the optimisation of sensorimotor control.     

Optimal impedance control for task achievement in the presence of signal-dependent noise

There is an infinity of impedance parameter values, and thus different co-contraction levels, that can produce similar movement kinematics from which the CNS must select one. Although signal-dependent noise (SDN) predicts larger motor-command variability during higher co-contraction, the relationship between impedance and task performance is not theoretically obvious and thus was examined here. Subjects made goal-directed, single-joint elbow movements to either move naturally to different target sizes or voluntarily co-contract at different levels. Stiffness was estimated as the weighted summation of rectified EMG signals through the index of muscle co-contraction around the joint (IMCJ) proposed previously. When subjects made movements to targets of different sizes, IMCJ increased with the accuracy requirements, leading to reduced endpoint deviations. Therefore without the need for great accuracy, subjects accepted worse performance with lower co-contraction. When subjects were asked to increase co-contraction, the variability of EMG and torque both increased, suggesting that noise in the neuromotor command increased with muscle activation. In contrast, the final positional error was smallest for the highest IMCJ level. Although co-contraction increases the motor-command noise, the effect of this noise on the task performance is reduced. Subjects were able to regulate their impedance and control endpoint variance as the task requirements changed, and they did not voluntarily select the high impedance that generated the minimum endpoint error. These data contradict predictions of the SDN-based theory, which postulates minimization of only endpoint variance and thus require its revision.     

Effects of visual gain on force control at the elbow and ankle

Visual feedback is essential when minimizing force fluctuations. Varying degrees of somatotopic organization have been shown in different regions of the brain for the upper and lower extremities, and visual feedback may be processed differently based on the body effector where feedback-based corrections are used. This study compared the effect of changes in visual gain on the control of steady-state force at the elbow and ankle. Ten subjects produced steady-state isometric force to targets at 5 and 40% of their maximum voluntary contraction at seven visual gain levels. Visual gain was used effectively at both joints to reduce variability of the force signal and to improve accuracy, with a greater effect of visual gain at the elbow than the ankle. Visual gain significantly decreased the regularity of force output, and this effect was more pronounced at the elbow than the ankle. There were accompanying changes in the proportion of power in the 0–4, 4–8, and 8–12 Hz frequency bins of the force signal across visual gain at the elbow. Changes in visual gain were accompanied by changes in both agonist and antagonist electromyographic (EMG) activation at the elbow. At the ankle joint, there were only changes in agonist EMG. The results suggest better use of visual information in the control of elbow force than ankle force and this improved control may be related to the changes in the pattern of agonist and antagonist activation.     

The senses of force and heaviness at the human elbow joint

The present-day view of the neural basis for the senses of muscle force and heaviness is that they are generated centrally, within the brain, from copies of motor commands. A corollary of the motor discharge generates a sense of effort which underlies these sensations. In recent experiments on force and heaviness sensations using thumb flexor muscles, a rather different explanation has been invoked: Subjects were proposed to rely predominantly on inputs of a peripheral origin, in particular, the signals of muscle spindles. The present experiments have been carried out at the elbow joint to determine whether these new ideas apply more widely. The effects of fatigue of elbow flexor muscles have been studied in force and heaviness matching tasks using three exercise regimes, a sustained maximum voluntary contraction (MVC), a maintained contraction of 35 % MVC, and a maintained contraction of 35 % MVC combined with muscle vibration at 80 Hz. In force-matching experiments, subjects were required to contract both arms and while the reference arm generated the target force under visual control, it was matched by the indicator arm without visual feedback. During the 100 % MVC exercise, force in the exercising reference arm fell rapidly to almost a half of its original value over 90 s while force in the indicator did not fall, leading to a significant overestimation of the reference force. During the 35 % MVC exercise, subjects also overestimated the reference force and this persisted at 5 and 10 min after the exercise. When 35 % MVC was combined with vibration, the amount by which the indicator arm overestimated the reference force was significantly reduced. In heaviness matching experiments, subjects could move their arms through a small range. The reference arm was loaded with a weight, and weights were added or removed from the indicator until heaviness felt the same in the two arms. There was a small, but significant fall in the matching weight used after 100 % MVC exercise, that is, the weight held by the fatigued arm felt lighter. The 35 % exercise did not alter heaviness sensation while 35 % MVC exercise with vibration led to a significant reduction in perceived heaviness. To conclude, while the results of these experiments on elbow flexors are not as clear cut as for thumb flexors, the central effort hypothesis falls short, in a number of respects in explaining the data which are able to be interpreted in terms of a peripheral afferent contribution to the senses of force and heaviness.     

Multijoint movement control in Parkinson's disease

Impairments in the performance of complex actions in Parkinson's disease (PD) patients are well documented. The aim of the present study was to investigate potential mechanisms that may be contributing to impaired movement performance in PD patients. PD patients and age-matched control subjects performed rapid pointing movements to a series of four tabletop targets. The height of the table was adjusted until the targets could be achieved with arm movements in the horizontal plane. The targets were arranged such that target 1 required elbow extension only and targets 2–4 required increasing amounts of horizontal shoulder flexion in addition to the elbow extension. While the control subjects accelerated and decelerated the elbow and shoulder joints simultaneously regardless of the target location, the PD patients decomposed motion during the acceleration phase by accelerating first the shoulder and then the elbow joint. For PD patients this decomposition of arm segments was associated with greater coactivation of the muscles about the elbow when elbow extension and shoulder flexion were simultaneously required (targets 2–4), in contrast to the single joint action. The control subjects decreased elbow joint coactivation while the patients increased it across the four targets. The resulting peak interaction torques at both the elbow and shoulder joints occurred relatively later for the PD patients. The coactivation patterns observed in PD patients may reduce the ability to take advantage of interaction torques and may also contribute to joint motion decomposition. Electronic Publication     

A comparison of the effects of agonist and antagonist muscle fatigue on performance of rapid movements

The aim of this study was to investigate the effects of agonist and antagonist muscle fatigue on the performance of rapid, self-terminating movements. Six subjects performed rapid, consecutive elbow flexion and extension movements between two targets prior to and after fatiguing either the elbow flexor or elbow extensor muscles. The experiments demonstrated consistent results. Agonist muscle fatigue was associated with a decrease in peak velocity and peak deceleration, while a decrease in peak acceleration was particularly prominent. Antagonist muscle fatigue, however, was associated with a decrease in peak deceleration, while a decrease in both the peak velocity and peak acceleration was modest and, in some tests, non-significant. The relative acceleration time (i.e. acceleration time as a proportion of the total movement time) increased when agonists were fatigued, but decreased when antagonists were fatigued. Taken together, these results emphasize the mechanical roles of the agonist and antagonist muscles; namely, the fatigue of each muscle group particularly affected the movement phase in which that group accelerated a limb, while changes of the movement kinematics pattern provided more time for action of the fatigued muscles. In addition, the results presented suggest that agonist muscle fatigue affects movement velocity more than antagonist muscle fatigue, even in movements that demonstrate prominently both mechanical and myoelectric activity of the antagonist muscles, such as rapid, self-terminating movements.     

Repetitive arm motion-induced fatigue affects shoulder but not endpoint position sense

Neck/shoulder pain has previously been linked to repetitive work and muscle fatigue. We have shown that asymptomatic people performing repetitive upper limb tasks display signs of shoulder fatigue and of whole-body compensatory strategies. However, the role played by the proprioceptive system in the production of these compensatory strategies has not been studied. A group of asymptomatic adults (n = 18) performed a repetitive pointing task at shoulder height to fatigue. Before and after fatigue, they performed two position sense tasks, eyes closed: a single-joint task where they abducted their shoulder to the perceived horizontal and a multi-joint task, where they stood and placed their finger at the perceived location of a target in front of them at shoulder height. After fatigue, subjects made larger shoulder errors by raising their elbow higher above the horizontal (~+1.3 cm) than before fatigue; however, their finger position accuracy was not changed, despite all subjects performing the movement in less time (~−0.18 s) while fatigued. There were no gender differences in shoulder or finger position accuracy before or after fatigue; however, there were gender differences in the perceived finger-target location and in the temporal characteristics of the finger movement toward the target. Results suggest that healthy individuals are able to develop strategies to compensate for fatigue-induced deficits at one joint to maintain the endpoint accuracy of a multi-joint task constant. Gender differences in movement strategies and perception of endpoint location may play parts in the previously reported gender differences in work-related neck/shoulder symptoms.     

Adaptation of the precentral cortical command to elbow muscle fatigue

The control exerted by individual motor cortical cells on their fatigued target muscles was assessed by analyzing the discharge patterns and electromyographic (EMG) postspike effects of cortical cells in monkeys making repeated forceful, but submaximal, isometric flexions of the elbow to produce fatigue. Two monkeys were trained to perform self-paced isometric contractions (for longer than 2 s) at forces greater than 35% maximal contraction, with three sets of 20 consecutive contractions; the first and last sets were at the same force level. Pairs of EMG electrodes were implanted in the biceps brachii, brachioradialis, and triceps brachii. The cortical cell discharges were modulated with the active and passive movements of the elbow and produced consistent EMG postspike effects during isometric contraction. Muscle fatigue was assessed as a statistically significant (P<0.05) drop in the mean power frequency of the EMG power spectrum in one or both flexors in the last set of contractions. Clear signs of muscular fatigue occurred in 20 different experimental sessions. Before fatigue, cortical cells were classified as phasic-tonic (18), phasicramp (three), or tonic (five). Twenty cells briskly fired to passive elbow extension, and 9 also responded to passive flexion. Only 6 cells showed a decreased discharge to passive extension. A 22–30% increase in the contraction force produced a higher discharge frequency in 13 cells, and a lower frequency in 5 cells. All cells exerted EMG postspike effects in their target muscles: 20 cells facilitated the flexors, and some of these also inhibited (3 cells) or cofacilitated (5 cells) the extensor; the other 6 cells had mixed effects: 5 of them inhibited at least one flexor, and 1 cell only facilitated the extensor. Most cells (24/26) still produced EMG postspike effects in their target muscles during fatigue, and the number of facilitated muscles increased: 21 cells facilitated the flexors, and 12 of them cofacilitated the extensor. Only 3 cells still inhibited the flexors and were tonic cells. The cortical cell firing frequency increased during fatigue in 13 cells and decreased in 8 cells. Increases involved 10 cells excited by passive elbow extension. Fourteen cells showed parallel changes in firing frequency with fatigue and force, and 9 of these cells facilitated both extensors and flexors in fatigue. Increases were found in 8 cells, decreases in 5 cells and no change in 1 cell. As muscle afferents provide substantial information to cortical cells, which in turn establish functional linkages with their target muscles before and during fatigue, the changes in cell firing frequencies during fatigue demonstrate the active participation of the motor cortex in the control of compensation for the peripheral adjustments concomitant with muscle fatigue.     

Movement reorganization to compensate for fatigue during sawing

Peripheral (muscle) aspects of fatigue are well documented. However, little is known about the central aspects of fatigue that could influence, in particular, multijoint coordination. To investigate the central aspects of fatigue, we compared the multijoint kinematics of non-fatigued and fatigued individuals while sawing. Muscle fatigue was associated with decreases in sawing force and movement amplitude at the elbow whereas the basic characteristics of the saw trajectory, including the movement direction, extent and duration, remained invariant. This invariance was maintained by increasing the movement amplitude at the wrist, shoulder and trunk. The system thus takes advantage of the redundancy of the motor apparatus to maintain the endpoint trajectory despite fatigue.     

Human forearm position sense after fatigue of elbow flexor muscles

After a period of eccentric exercise of elbow flexor muscles of one arm in young, adult human subjects, muscles became fatigued and damaged. Damage indicators were a fall in force, change in resting elbow angle and delayed onset of soreness. After the exercise, subjects were asked to match the forearm angle of one arm, whose position was set by the experimenter, with their other arm. Subjects matched the position of the unsupported reference arm, when this was unexercised, with a significantly more flexed position in their exercised indicator arm. Errors were in the opposite direction when the reference arm was exercised. The size of the errors correlated with the drop in force. Less consistent errors were observed when the reference arm was supported. A similar pattern of errors was seen after concentric exercise, which does not produce muscle damage. The data suggested that subjects were using as a position cue the perceived effort required to maintain a given forearm angle against the force of gravity. The fall in force from fatigue after exercise meant more effort was required to maintain a given position. That led to matching errors between the exercised and unexercised arms. It was concluded that while a role for muscle spindles in kinaesthesia cannot be excluded, detailed information about static limb position can be derived from the effort required to support the limb against the force of gravity.     

Adaptation of motor behavior to preserve task success in the presence of muscle fatigue

To achieve task goals in the various contexts of everyday life, the CNS has to adapt to short time scale changes in the properties of the neuromuscular system, such as those induced by fatigue. Here we investigated how humans preserve task success despite fatigue-induced changes within the neuromuscular system, when they have to aim at a target as fast and as accurately as possible. In such a task, subjects generally choose a compromise between speed and accuracy that has been formalized as Fitts's law. We first characterized the effect of fatigue on Fitts's law in an experiment where participants had to perform fast but accurate elbow movements aimed at targets of different sizes, before and after a fatiguing exercise that reduced maximal voluntary force by ∼30%. We found that movements were slower to guarantee task success in the presence of fatigue. We then used an optimal control model to determine how fatigue-induced changes in variables such as noise in motor commands, muscle contraction and relaxation times, and the gain between neural activation and muscle force may contribute to changes in Fitts's law with fatigue. We concluded that the observed behavior was not due to the lack of available force, but very likely reflected the fact that the CNS uses the same optimal strategy with a fatigued neuromuscular plant that notably exhibits increased signal-dependent noise in motor commands. This strategy appears necessary to preserve task success in the presence of acute changes in the neuromuscular system.     

Optimality versus variability: effect of fatigue in multi-finger redundant tasks

We used two methods to address two aspects of multi-finger synergies and their changes after fatigue of the index finger. Analytical inverse optimization (ANIO) was used to identify cost functions and corresponding spaces of optimal solutions over a broad range of task parameters. Analysis within the uncontrolled manifold (UCM) hypothesis was used to quantify co-variation of finger forces across repetitive trials that helped reduce variability of (stabilized) performance variables produced by all the fingers together. Subjects produced steady-state levels of total force and moment of force simultaneously as accurately as possible by pressing with the four fingers of the right hand. Both before and during fatigue, the subjects performed single trials for many force–moment combinations covering a broad range; the data were used for the ANIO analysis. Multiple trials were performed at two force–moment combinations; these data were used for analysis within the UCM hypothesis. Fatigue was induced by 1-min maximal voluntary contraction exercise by the index finger. Principal component (PC) analysis showed that the first two PCs explained over 90% of the total variance both before and during fatigue. Hence, experimental observations formed a plane in the four-dimensional finger force space both before and during fatigue conditions. Based on this finding, quadratic cost functions with linear terms were estimated from the experimental data. The dihedral angle between the plane of optimal solutions and the plane of experimental observations (D _ANGLE) was very small (a few degrees); it increased during fatigue. There was an increase in fatigue of the coefficient at the quadratic term for the index finger force balanced by a drop in the coefficients for the ring and middle fingers. Within each finger pair (index–middle and ring–little), the contribution of the “central” fingers to moment production increased during fatigue. An index of antagonist moment production dropped with fatigue. Fatigue led to higher co-variation indices during pronation tasks (index finger is an agonist) but opposite effects during supination tasks. The results suggest that adaptive changes in co-variation indices that help stabilize performance may depend on the role of the fatigued element, agonist or antagonist.     

Position sense at the elbow after fatiguing contractions

Fatigue is often associated with increased clumsiness. One possible explanation for this is that the proprioceptive signals from receptors in and around muscles change during muscle fatigue. Thirteen human subjects were tested for their ability to match the elbow angle of one arm with the contralateral arm, before and after a fatiguing contraction of one arm. Contractile fatigue was induced by a series of maximal voluntary contractions of the elbow flexors of the dominant arm. While fatigue of either the target arm or the matching arm usually changed the ability of individual subjects to match arm position, this effect varied markedly from one subject to another and no consistent pattern was discerned. In particular, there was no reciprocal change when the fatigued arm was the matching arm compared with when the nonfatigued arm was the matching arm. The absence of a consistent reciprocal effect indicates that the fatigue-related changes in the ability to match arm position are not solely due to changes in proprioceptive signals and that central fatigue processes are probably involved.     

Older adults use a unique strategy to lift inertial loads with the elbow flexor muscles

The purpose of this study was to determine the effect of age on the ability to exert steady forces and to perform steady flexion movements with the muscles that cross the elbow joint. An isometric task required subjects to exert a steady force to match a target force that was displayed on a monitor. An anisometric task required subjects to raise and lower inertial loads so that the angular displacement around the elbow joint matched a template displayed on a monitor. Steadiness was measured as the coefficient of variation of force and as the normalized standard deviation of wrist acceleration. For the isometric task, steadiness as a function of target force decreased similarly for old adults and young adults. For the anisometric task, steadiness increased as a function of the inertial load and there were significant differences caused by age. Old adults were less steady than young adults during both shortening and lengthening contractions with the lightest loads. Furthermore, old adults were least steady when performing lengthening contractions. These behaviors appear to be associated with the patterns of muscle activation. These results suggest that different neural strategies are used to control isometric and anisometric contractions performed with the elbow flexor muscles and that these strategies do not change in parallel with advancing age.     

Bilateral interactions during contractions of intrinsic hand muscles

During demanding voluntary contractions (e.g., high force or fatigue), activation is not restricted to the target muscle but extends to other ipsilateral muscles; even contralateral muscles become activated. The contralateral "irradiation" of activity was measured in five subjects during submaximal and maximal voluntary contractions (MVCs) of the first dorsal interosseous (FDI) (index finger abduction) and during unfatigued and fatigued conditions. All subjects were tested five times with at least one week between tests. Unilateral MVCs were associated with a substantial amount of contralateral FDI activation [mean = 7.9 +/- 6.7% (SD) MVC prior to fatigue]. The amount of such contralateral irradiation was significantly different between different individuals and was positively correlated between dominant and nondominant hands. During fatigue tests, the contractile activity of the contralateral "nontarget" index finger showed progressive increase (force, electromyogram) as was measured during both the submaximal task and interspersed MVCs of the target finger. In addition, a superimposed saw-tooth pattern of intermittently waxing and waning contractions commonly appeared contralaterally. The expression of contralateral irradiation force was itself fatigue-sensitive: less irradiation was seen in a recently fatigued muscle than was seen before the fatigue test. These fatigue effects could not be explained as having been caused by changes in muscle properties. Possible anatomical sites of contralateral irradiation are briefly discussed.