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Biomedical subjects

R M Enoka

Publications and source records attributed to R M Enoka.

At least 19 recordsLinked to original sources

Motor unit physiology: some unresolved issues.

The purpose of this review was to examine three issues that limit our understanding of motor unit physiology: (1) the range and distribution of the innervation ratios in a muscle; (2) the association between discharge rate and force; and (3) the variation in motor unit activity across contractions that differ in speed and type. We suggest that if more data were available on these issues, the understanding of neuromuscular function would be enhanced substantially, especially with regard to plasticity in the motor neuron pool, adequacy of the neural drive to muscle, and flexibility of activation patterns across various types of contractions. Current data are limited and these limitations influence our ability to interpret adaptations in muscle function in health and disease.

Animals↗

Activation of the arousal response can impair performance on a simple motor task.

The purpose of this study was to determine the effect of arousal in men and women on the moment-to-moment performance of a simple motor task. We examined the control of a precision task in the presence and absence of imposed stressors. Twenty-nine subjects (14 men, 15 women; 18--44 yr) were randomly assigned to either a control group or one of two stressor groups, Mental Math or Electric Shock. Subjects presented with Math and Shock stressors, which lasted 10 min, experienced significant increases in cognitive and physiological arousal compared with baseline and control subjects. Heart rate, systolic blood pressure, and electrodermal activity were elevated 5--80% with presentation of the stressors, whereas diastolic blood pressure and salivary cortisol were unchanged. The greater levels of cognitive and physiological arousal were associated with reductions in steadiness of a pinch grip for the Shock subjects (approximately 130% reduction from baseline) but not for the subjects in the Math group, who experienced heightened arousal but no change in steadiness (10% reduction from baseline). Although women exhibited more of a reduction in steadiness than men, the effect was largely unrelated to the magnitude of the change in arousal.

Adult↗

Activation of the arousal response and impairment of performance increase with anxiety and stressor intensity.

The purpose of this study was to determine the effect of trait anxiety and stressor intensity on arousal and motor performance during a pinch task. We examined the steadiness of a precision task in the presence and absence of an imposed stressor on subjects with moderate and low trait anxiety. Subjects with the 26 highest and 14 lowest anxiety scores were assigned to one of three groups: a control group (5 women, 5 men), a moderate-anxiety group (8 women, 8 men), or a low-anxiety group (7 women, 7 men). Subjects in the anxiety groups received electric shocks and experienced significant increases in cognitive and physiological arousal compared with baseline and control subjects, especially subjects in the moderate-anxiety group. Heart rate, systolic blood pressure, and electrodermal activity were elevated during the stressor, whereas diastolic blood pressure was unchanged. Cognitive and physiological arousal tended to increase with stressor intensity and was accompanied by changes in steadiness. Although steadiness was markedly reduced with the highest intensity of shock, the average electromyogram activity was unaffected by the stressor. These findings indicate that the increase in arousal and the impairment of steadiness increased with trait anxiety and with the intensity of the noxious stimulus.

Adolescent↗

Long-term activity in upper- and lower-limb muscles of humans.

Despite limited data on humans, previous studies suggest that there is an association between the duration of daily muscle activity and the proportion of type I muscle fibers. We quantified the activity of limb muscles in healthy men and women during normal use and compared these measurements with published reports on fiber-type proportions. Seven men (age range = 21-28 yr) and seven women (age range = 18-26 yr) participated in two 10-h recording sessions. Electromyogram (EMG) activity of four muscles in nondominant upper (first dorsal interosseus and biceps brachii) and lower limbs (vastus medialis and vastus lateralis) was recorded with surface electrodes. Hand and arm muscles were active for 18% of the recording time, whereas leg muscles were active for only 10% of the recording time. On average, upper-limb muscles were activated 67% more often than lower-limb muscles. When lower-limb muscles were activated, however, the mean amplitude of each burst was greater in leg muscles [18 and 17% maximum voluntary contraction (MVC)] compared with hand (8% MVC) and arm (6% MVC) muscles. Temporal association in activity between pairs of muscles was high for the two lower-limb muscles (r2 = 0.7) and relatively weak for the two upper-limb muscles (r2 = 0.09). Long-term muscle activity was only different between men and women for the biceps brachii muscle. We found no relation between duration of muscle activity in 10-h recordings and the reported values of type I fibers in men and women.

Adolescent↗

Sex differences in the fatigability of arm muscles depends on absolute force during isometric contractions.

Women are capable of longer endurance times compared with men for contractions performed at low to moderate intensities. The purpose of the study was 1) to determine the relation between the absolute target force and endurance time for a submaximal isometric contraction and 2) to compare the pressor response and muscle activation patterns of men [26.3 +/- 1.1 (SE) yr] and women (27.5 +/- 2.3 yr) during a fatiguing contraction performed with the elbow flexor muscles. Maximal voluntary contraction (MVC) force was greater for men (393 +/- 23 vs. 177 +/- 7 N), which meant that the average target force (20% of MVC) was greater for men (79.7 +/- 6.5 vs. 36.7 +/- 2.0 N). The endurance time for the fatiguing contractions was 118% longer for women (1,806 +/- 239 vs. 829 +/- 94 s). The average of the rectified electromyogram (%MVC) for the elbow flexor muscles at exhaustion was similar for men (31 +/- 2%) and women (30 +/- 2%). In contrast, the heart rate and mean arterial pressure (MAP) were less at exhaustion for women (94 +/- 6 vs. 111 +/- 7 beats/min and 121 +/- 5 vs. 150 +/- 6 mmHg, respectively). The target force and change in MAP during the fatiguing contraction were exponentially related to endurance time (r(2) = 0.68 and r(2) = 0.64, respectively), whereas the change in MAP was linearly related to target force (r(2) = 0.51). The difference in fatigability of men and women when performing a submaximal contraction was related to the absolute contraction intensity and was limited by mechanisms that were distal to the activation of muscle.

Adult↗

Steadiness is reduced and motor unit discharge is more variable in old adults.

The purpose of this study was to compare the steadiness and discharge rate of motor units during submaximal contractions performed by young and old adults. Subjects performed isometric and slow shortening and lengthening contractions with the first dorsal interosseous muscle. The steadiness of the isometric and slow anisometric contractions was less for the old subjects compared with young subjects, especially at the lower target forces and with the lightest loads. Furthermore, the steadiness of the lengthening contractions was less compared with the shortening contractions for the old subjects. Although the mean discharge rates of motor units were not different for the two groups of subjects, the variability of the discharge rates was greater for the old subjects during the isometric and anisometric contractions. We conclude that a more variable discharge by single motor units probably contributes to the reduced ability of old adults to perform steady muscle contractions.

Adult↗

Strength training can improve steadiness in persons with essential tremor.

We evaluated the effect of a strength-training program on the ability of persons with essential tremor to exert steady forces with the index finger. Thirteen subjects with a diagnosis of essential tremor were assigned to three different groups: one group trained with heavy loads, one with light loads, and one did not perform any training. Subjects attempted to generate steady contractions during both postural and constant-force tasks. Steadiness was quantified by the root mean square amplitude of acceleration during postural tasks and the standard deviation and coefficient of variation of force during the constant-force tasks. Subjects who performed the training program with heavy loads experienced an increase in steadiness around the target force during the constant-force tasks. Subjects in the other two groups did not exhibit any changes. These findings suggest that strength training can decrease the magnitude of tremor. However, we did not observe any associated improvements in functional abilities.

Adult↗

Limb immobilization alters muscle activation patterns during a fatiguing isometric contraction.

The purpose of the study was to determine the role of excitation-contraction coupling in the increased endurance time for low-force contractions after 4 weeks of elbow joint immobilization. Twelve subjects participated in a protocol that required immobilization of the elbow joint in a fiberglass cast for 4 weeks, and 4 subjects acted as controls. Measurements of muscle strength, contractile properties, and fatigability were performed before and after 4 weeks of limb immobilization, and after 4 weeks of recovery. The immobilization intervention produced significant reductions in the daily activity of the elbow flexor muscles, a 21% decline in the maximum voluntary contraction (MVC) force, and a 31% decrease in the maximum load that could be lifted once. Seven of the immobilized subjects exhibited an unusual pattern of muscle activity during the fatiguing contraction after immobilization, which was associated with an increase in the endurance time of the elbow flexor muscles (mean = 220%) in these subjects. The unusual pattern of muscle activity involved lower relative activity of the brachialis muscle, no increase in the amplitude of the electromyogram (EMG) for the elbow flexor muscles, and intermittent rather than continuous EMG. In contrast, the force-frequency relationship of biceps brachii was not altered by immobilization in these subjects, suggesting that adaptations in excitation-contraction coupling were not the primary cause of the prolonged endurance time after immobilization. Rather, the results suggest that the prolonged endurance time exhibited by some subjects after immobilization was largely due to adaptations within the nervous system.

Elbow↗

Coactivation of the antagonist muscle does not covary with steadiness in old adults.

The purpose of the study was to determine the association between steadiness and activation of the agonist and antagonist muscles during isometric and anisometric contractions. Young (n = 14) and old (n = 15) adults used the first dorsal interosseus muscle to perform constant-force and constant-load tasks (2.5, 5, 20, 50, and 75% maximum) with the left index finger. Steadiness was quantified as the coefficient of variation of force and the SD of acceleration normalized to the load lifted. The old adults were less steady at most target forces with isometric contractions (2.5, 5, and 50%) and with most loads during the anisometric contractions (2.5, 5, and 20%). Furthermore, the old adults were less steady when performing lengthening contractions (up to 50%) compared with shortening contractions, whereas there was no difference for young adults. The reduced steadiness exhibited by the old adults during these tasks was not associated with differences in the average level of agonist muscle electromyogram or with coactivation of the antagonist muscle.

Acceleration↗

Motor-unit synchronization increases EMG amplitude and decreases force steadiness of simulated contractions.

The purpose of the study was to determine the effect of motor-unit synchronization on the surface electromyogram (EMG) and isometric force using a computer model of muscle contraction. The EMG and force were simulated by generating muscle fiber action potentials, defining motor-unit mechanical characteristics and territories, estimating motor-unit action potentials, specifying motor-unit discharge times, and imposing various levels of motor-unit synchronization. The output (EMG and force) was simulated at 11 levels of excitation, ranging from 5 to 100% of maximum. To synchronize motor-unit activity, selected motor-unit discharge times were adjusted; however, the number of motor units recruited and the average discharge rate of each unit was constant across synchronization conditions for a given level of excitation. Two levels of synchronization were imposed on the discharge times: a moderate and a high level, which approximated the experimentally observed range of motor-unit synchronization. The moderate level of synchrony caused the average EMG to increase by approximately 65%, whereas the high level caused a 130% increase in the EMG with respect to the no-synchrony condition. Neither synchrony condition influenced the magnitude of the average force. However, motor-unit synchronization did increase the amplitude of the fluctuations in the simulated force, especially at intermediate levels of excitation. In conclusion, motor-unit synchronization increased the amplitude of the average rectified EMG and decreased the steadiness of the force exerted by the muscle in simulated contractions.

Action Potentials↗

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.

Adult↗

Motor-unit synchronization is not responsible for larger motor-unit forces in old adults.

Motor-unit synchronization, which is a measure of the near simultaneous discharge of action potentials by motor units, has the potential to influence spike-triggered average force and the steadiness of a low-force isometric contraction. The purpose of the study was to estimate the contribution of motor-unit synchronization to the larger spike-triggered average forces and the decreased steadiness exhibited by old adults. Eleven young (age 19-30 yr) and 14 old (age 63-81 yr) adults participated in the study. Motor-unit activity was recorded with two fine-wire intramuscular electrodes in the first dorsal interosseus muscle during isometric contractions that caused the index finger to exert an abduction force. In a separate session, steadiness measurements were obtained during constant-force isometric contractions at target forces of 2.5, 5, 7. 5, and 10% of the maximum voluntary contraction (MVC) force. Mean (+/-SD) motor-unit forces measured by spike-triggered averaging were larger in old (15.5 +/- 12.1 mN) compared with young (7.3 +/- 5.7 mN) adults, and the differences were more pronounced between young (8.7 +/- 6.4 mN) and old (19.9 +/- 12.2 mN) men. Furthermore, the old adults had a reduced ability to maintain a steady force during an isometric contraction, particularly at low target forces (2.5 and 5% MVC). Mean (+/-SD) motor-unit synchronization, expressed as the frequency of extra synchronous discharges above chance in the cross-correlogram, was similar in young [0.66 +/- 0.4 impulses/s (imp/s); range, 0.35-1.51 imp/s; 53 pairs) and old adults (0.72 +/- 0.5 imp/s; range, 0.27-1.38 imp/s; 56 pairs). The duration of synchronous peaks in the cross-correlogram was similar for each group (approximately 16 ms). These data suggest that motor-unit synchronization is not responsible for larger spike-triggered average forces in old adults and that motor-unit synchronization does not contribute to the decreased steadiness of low-force isometric contractions observed in old adults.

Action Potentials↗

Strength training improves the steadiness of slow lengthening contractions performed by old adults.

When old adults participate in a strength-training program with heavy loads, they experience an increase in muscle strength and an improvement in the steadiness of submaximal isometric contractions. The purpose of this study was to determine the effect of light- and heavy-load strength training on the ability of old adults to perform steady submaximal isometric and anisometric contractions. Thirty-two old adults (60-91 yr) participated in a 4-wk training program of a hand muscle. Both the light- and heavy-load groups increased one-repetition maximum and maximal voluntary contraction (MVC) strength and experienced similar improvements in the steadiness of the isometric and shortening and lengthening contractions. The increase in MVC strength was greater for the heavy-load group and could not be explained by changes in muscle activation. Before training, the lengthening contractions were less steady than the shortening contractions with the lightest loads (10% MVC). After training, there was no difference in steadiness between the shortening and lengthening contractions, except with the lightest load. These improvements were associated with a reduced level of muscle activation, especially during the lengthening contractions.

Aged↗

Gender differences in the fatigability of human skeletal muscle.

After participating in a 4-wk intervention that reduced normal usage of the elbow flexor muscles, all six women, but only one of six men, experienced a marked increase in the endurance time during a low-force fatiguing contraction. The increase in endurance time was associated with an altered pattern of muscle activation that did not involve the commonly observed progressive increase in muscle activity. Rather, the muscle activity comprised intermittent motor unit activity. In those individuals who exhibited this behavior, the novel pattern of muscle activity was only present immediately after 4 wk of limb immobilization and not before the intervention or after 4 wk of recovery. These findings suggest possible differences between women and men in the adaptations of the neuromuscular system.

Adolescent↗

Older adults can maximally activate the biceps brachii muscle by voluntary command.

Because some of the decline in strength with age may be explained by an impairment of muscle activation, the purpose of this study was to determine the activation level achieved in biceps brachii by older adults during a maximum voluntary contraction (MVC). This capability was assessed with two superimposition techniques: one calculated the activation level that was achieved during an MVC, and the other provided an estimate of the expected MVC force based on extrapolation with submaximal forces. The activation level in biceps brachii was incomplete (< 100%) for the young (n = 16) and elderly (n = 16) subjects, with the elderly subjects exhibiting the greater deficit. In contrast, there was no difference between the measured and expected MVC forces for either group of subjects, whether the extrapolation involved a third-order polynomial or linearization of the data. Because of the lower signal-to-noise ratio associated with the measurement of activation level and the greater number of measurements that contributed to the estimate of the expected MVC force, we conclude that the older adults were able to achieve complete activation of the biceps brachii muscle during an MVC.

Adolescent↗

Neural adaptations with chronic physical activity.

Chronic activity patterns, such as strength training, limb immobilization, and aging, produce marked adaptations in both the muscular and nervous systems. In this brief review, some of the involved mechanisms are examined as they are revealed through studies on the maximality, specificity, and pattern of the neural drive to muscle. The studies on maximality indicate that it is difficult to activate maximally a muscle by voluntary command, the capacity varies across muscles, tasks, and training, and the maximum discharge rates of motor neurons decreases with immobilization and increases with strength training. The data on specificity demonstrate that: strength can be increased by training with imagined contractions; the velocity specificity of isokinetic training is evident with intended contractions; the strength training influences the untrained homologous muscle in the contralateral limb; the bilatral deficit can become a bilateral facilitation with appropriate training; and that eccentric contractions appear to involve a different activation scheme compared to isometric and concentric contractions. Finally, the literature on the pattern of the neural drive suggests that: coactivation varies with training and often decreases as skill level increases; measures of motor-unit synchronization reveal changes in neuronal connectivity with physical training; the reflex potentiation varies across muscles, individuals, and activity patterns; the modulation of the H-reflex amplitude with training involves changes in the motor neuron; and the motor neurons exhibit a bistable, excitability property that may be influenced by exercise. Despite the breadth of this evidence, there remain substantial gaps in our knowledge, particularly regarding the symmetry of adaptations with increased and decreased chronic physical activity.

Adaptation, Physiological↗

Task-dependent effect of limb immobilization on the fatigability of the elbow flexor muscles in humans.

Because short-term limb immobilization produces selective adaptations in the neuromuscular system that probably interact with the task-dependent expression of muscle fatigue, the purpose of this study was to determine the effects of limb immobilization on the ability of human subjects to sustain isometric contractions at low and moderate submaximal forces. Four weeks of elbow joint immobilization caused a substantial decrease in the daily activity of biceps brachii during immobilization, a significant reduction in the cross-sectional area and volume of the elbow flexor muscles as measured by magnetic resonance imaging, and a decline in the maximum voluntary contraction (MVC) activation and force of the elbow flexor muscles. Immobilization had a task-dependent effect on muscle fatigue with a substantially increased endurance time (reduced fatigability) at a low force (20% MVC) and no statistical effect at a moderate force (65% MVC). Despite atrophy of the elbow flexor muscles due to the immobilization, the twitch force elicited in biceps brachii by electrical stimulation was greater after immobilization. The selective improvement of fatigue resistance for the low-force contraction and the absence of a change in the time course of the twitch suggests that the immobilization-induced adaptations included an improved efficacy of some excitation-contraction processes and underscored the major role of these mechanisms in determining the endurance time for low-force, long-duration contractions.

Adult↗