John G. Semmler

Mechanisms that contribute to differences in motor performance between young and old adults

This paper examines the physiological mechanisms responsible for differences in the amplitude of force fluctuations between young and old adults. Because muscle force is a consequence of motor unit activity, the potential mechanisms include both motor unit properties and the behavior of motor unit populations. The force fluctuations, however, depend not only on the age of the individual but also on the muscle group performing the task, the type and intensity of the muscle contraction, and the physical activity status of the individual. Computer simulations and experimental findings performed on tasks that involved single agonist and antagonist muscles suggest that differences in force fluctuations are not attributable to motor unit twitch force, motor unit number, or nonuniform activation of the agonist muscle, but that they are influenced by the variability and common modulation of motor unit discharge in both the agonist and antagonist muscles. Because the amplitude of the force fluctuations does not vary linearly with muscle activation, these results suggest that multiple mechanisms contribute to the differences in force fluctuations between young and old adults, although the boundary conditions for each mechanism remain to be determined.

Motor unit discharge and force tremor in skill-and strength-trained individuals

Abstract We examined motor unit (MU) discharge properties (mean interspike interval, ISI, discharge variability, short-term synchronization, common drive) and force tremor in the first dorsal interosseous (FDI) muscle of five musicians (skill-trained), five weight-lifters (strength-trained) and six untrained subjects during low-force isometric abduction of the index finger. Mean MU ISI was slightly shorter in skill-trained subjects than in untrained subjects. Discharge variability of FDI MUs did not differ significantly between groups. The mean strength of MU synchronization (expressed as the frequency of extra synchronous discharges above chance) was different in skill-trained (0.22±0.02 s–1, 162 MU pairs), untrained (0.32±0.02 s–1, 199 MU pairs) and strength-trained subjects (0.44±0.03 s–1, 183 MU pairs). FDI MU synchrony was weak and of equivalent strength in both hands of skill-trained subjects and the dominant (skilled) hand of untrained subjects. The stronger FDI MU synchrony in the non-dominant hand of untrained subjects was equivalent to that found in both hands of strength-trained subjects. The extent of common modulation of firing rates (common drive) was assessed for a subset of MU pairs and was weaker in skill-trained subjects (0.30±0.04, n=14) than untrained (0.43±0.3, n=14) and strength-trained (0.48±0.03, n=21) subjects. Force tremor was quantified for each hand in the same subjects during isometric index finger abduction at target forces of 0.5 N and 3.5 N. Tremor rms amplitude and peak power in the force frequency spectrum were significantly lower in skill-trained subjects than strength-trained subjects with the 3.5-N target force. The peak tremor frequency was similar in the three groups. The relatively more independent discharge of pairs of FDI MUs in skill-trained subjects was not responsible for the reduced tremor amplitudes in these subjects. Correlations between the overall extent of MU synchrony and common drive in FDI muscles and tremor measures obtained during the same experimental session were all non-significant. Differences in the central descending command signals are the most likely explanation for the more independent discharge of FDI MUs in skill-trained hands, while neural or peripheral muscular factors may be responsible for the weaker tremor.

Motor unit synchronization and neuromuscular performance.

SEMMLER, J.G. Motor unit synchronization and neuromuscular performance. Exerc. Sport Sci. Rev., Vol. 30, No. 1, pp 8–14, 2002. The acute and chronic plasticity of motor unit synchronization suggests that it must represent a deliberate strategy for neuromuscular activation. The most likely functional role of motor unit synchronization is to increase the rate of force development during rapid contractions or as a mechanism to coordinate the activity of multiple muscles in synergy.

Corticomotor plasticity and learning of a ballistic thumb training task are diminished in older adults

This study examined changes in corticomotor excitability and plasticity after a thumb abduction training task in young and old adults. Electromyographic (EMG) recordings were obtained from right abductor pollicis brevis (APB, target muscle) and abductor digiti minimi (ADM, control muscle) in 14 young (18-24 yr) and 14 old (61-82 yr) adults. The training task consisted of 300 ballistic abductions of the right thumb to maximize peak thumb abduction acceleration (TAAcc). Transcranial magnetic stimulation (TMS) of the left primary motor cortex was used to assess changes in APB and ADM motor evoked potentials (MEPs) and short-interval intracortical inhibition (SICI) before, immediately after, and 30 min after training. No differences in corticomotor excitability (resting and active TMS thresholds, MEP input-output curves) or SICI were observed in young and old adults before training. Motor training resulted in improvements in peak TAAcc in young (177% improvement, P < 0.001) and old (124%, P = 0.005) subjects, with greater improvements in young subjects (P = 0.002). Different thumb kinematics were observed during task performance, with increases in APB EMG related to improvements in peak TAAcc in young (r(2) = 0.46, P = 0.008) but not old (r(2) = 0.09, P = 0.3) adults. After training, APB MEPs were 50% larger (P < 0.001 compared with before) in young subjects, with no change after training in old subjects (P = 0.49), suggesting reduced use-dependent corticomotor plasticity with advancing age. These changes were specific to APB, because no training-related change in MEP amplitude was observed in ADM. No significant association was observed between change in APB MEP and improvement in TAAcc with training in individual young and old subjects. SICI remained unchanged after training in both groups, suggesting that it was not responsible for the diminished use-dependent corticomotor plasticity for this task in older adults.

Motor unit synchronisation is enhanced during slow lengthening contractions of a hand muscle

This study examined the strength of motor unit synchronisation based on time‐ and frequency‐domain measures during postural, shortening and lengthening contractions of a hand muscle in young adults. Single motor unit activity was recorded with intramuscular electrodes in the left first dorsal interosseus muscle as the subject held the index finger at a constant position while supporting a light load for 2‐5 min. The subject then performed slow (1.7 deg s−1) shortening and lengthening contractions to lift and lower the load. The movement required subjects to perform 10‐25 constant‐velocity contractions with the index finger over a 10 deg range of motion by using 6 s shortening and lengthening contractions. Individual discharge times were obtained from 23 pairs of motor units in 14 subjects to assess the strength of motor unit synchronisation and coherence during the three tasks. The strength of motor unit synchronisation was approximately 50 % greater during the lengthening contractions compared with the postural and shortening contractions, and the width of the central synchronous peak in the cross‐correlation histogram was ≈4 ms narrower during shortening contractions. These findings reveal that there is an increase in common input to motoneurones during lengthening contractions and a greater relative contribution of direct common inputs to motoneurones during shortening contractions compared with postural tasks. Furthermore, the amount of motor unit coherence in the low‐frequency band (2‐12 Hz) was reduced during shortening contractions compared with postural and lengthening contractions. These data indicate that the timing of inputs received by the motoneurones innervating the first dorsal interosseus of young adults differs during postural, shortening and lengthening contractions against a light load.

Motor cortex plasticity induced by paired associative stimulation is enhanced in physically active individuals

Recent evidence indicates that regular physical activity enhances brain plasticity (i.e. the ability to reorganise neural connections) and improves neurocognitive function. However, the effect of regular physical activity on human motor cortex function is unknown. The purpose of this study was to examine motor cortex plasticity for a small hand muscle in highly active and sedentary individuals. Electromyographic recordings were obtained from the left abductor pollicis brevis (APB) muscle of 14 active and 14 sedentary subjects (aged 18–38 yrs). The extent of physical activity was assessed by questionnaire, where the physically active subjects performed >150 min per day moderate‐to‐vigorous aerobic activity on at least 5 days per week, whereas the sedentary group performed <20 min per day of physical activity on no more than 3 days per week. Transcranial magnetic stimulation (TMS) of the right hemisphere was used to assess changes in APB motor‐evoked potentials (MEPs), input–output curve (IO curve), short‐interval intracortical inhibition (SICI) and cortical silent period (CSP). Neuroplastic changes were induced using paired‐associative stimulation (PAS), which consisted of 90 paired stimuli (0.05 Hz for 30 min) of median nerve electrical stimulation at the wrist followed 25 ms later by TMS to the hand area of motor cortex. The IO curve slope was 35% steeper in individuals with increased physical activity (combined before and after PAS, P < 0.05), suggesting increased motor cortex excitability, although there was no difference in SICI or CSP between groups. PAS induced an increase in MEP amplitude in the physically active subjects (54% increase compared with before, P < 0.01), but no significant facilitation in the sedentary subjects. We conclude that participation in regular physical activity may offer global benefits to motor cortex function that enhances neuroplasticity, which could improve motor learning and neurorehabilitation in physically active individuals.

Neural adaptations to strength training: Moving beyond transcranial magnetic stimulation and reflex studies

It has long been believed that training for increased strength not only affects muscle tissue, but also results in adaptive changes in the central nervous system. However, only in the last 10 years has the use of methods to study the neurophysiological details of putative neural adaptations to training become widespread. There are now many published reports that have used single motor unit recordings, electrical stimulation of peripheral nerves, and non‐invasive stimulation of the human brain [i.e. transcranial magnetic stimulation (TMS)] to study neural responses to strength training. In this review, we aim to summarize what has been learned from single motor unit, reflex and TMS studies, and identify the most promising avenues to advance our conceptual understanding with these methods. We also consider the few strength training studies that have employed alternative neurophysiological techniques such as functional magnetic resonance imaging and electroencephalography. The nature of the information that these techniques can provide, as well as their major technical and conceptual pitfalls, are briefly described. The overall conclusion of the review is that the current evidence regarding neural adaptations to strength training is inconsistent and incomplete. In order to move forward in our understanding, it will be necessary to design studies that are based on a rigorous consideration of the limitations of the available techniques, and that are specifically targeted to address important conceptual questions.

Motor Unit Synchronization Is Increased in Biceps Brachii After Exercise-Induced Damage to Elbow Flexor Muscles

The purpose of this study was to determine the effect of eccentric exercise on correlated motor unit discharge (motor unit synchronization and coherence) during low-force contractions of the human biceps brachii muscle. Eight subjects (age, 25 +/- 7 yr) performed three tasks involving isometric contraction of elbow flexors while EMG (surface and intramuscular) records were obtained from biceps brachii. Tasks were 1) maximum voluntary contraction (MVC); 2) constant-force contraction at various submaximal targets; and 3) sustained discharge of pairs of concurrently active motor units for 2-5 min. These tasks were performed before, immediately after, and 24 h after fatiguing eccentric exercise. MVC force declined 46% immediately after eccentric exercise and remained depressed (31%) 24 h later, which is indicative of muscle damage. For the constant-force task, biceps brachii EMG ( approximately 100% greater) and force fluctuations ( approximately 75% greater) increased immediately after exercise, and both recovered by approximately 50% 24 h later. Motor unit synchronization, quantified by cross-correlation of motor unit pairs during low-force (1-26% MVC) contractions, was 30% greater immediately after (n = 105 pairs) and 24 h after exercise (n = 92 pairs) compared with before exercise (n = 99 pairs). Similarly, motor unit coherence at low (0-10 Hz) frequencies was 20% greater immediately after exercise and 34% greater 24 h later. These results indicate that the series of events leading to muscle damage from eccentric exercise alters the correlated behavior of human motor units in biceps brachii muscle for > or =24 h after the exercise.

Influence of handedness on motor unit discharge properties and force tremor

Discharge properties of motor units (MUs) in the first dorsal interosseous muscle (FDI) were studied in the dominant and non-dominant hands of six right-handed (RH) and six left-handed (LH) individuals. MU discharge rates and variability were similar in each hand in RH (186 MUs) and LH (160 MUs) subjects. MU synchronization was less prominent in the dominant hand of RH subjects, with 51% (45/88) of cross-correlograms of MU discharge having significant central peaks, compared with 81% (90/111) for the non-dominant hand. The strength of MU synchronization (expressed as the frequency of extra synchronous discharges above chance) was weaker in the dominant hand of right-handers (0.23 ± 0.03 s-1 vs 0.39 ± 0.03 s-1), and synchronous peaks from that hand were slightly broader. Four of six RH subjects had significant differences in synchronization between hands (weaker in dominant hand). In contrast, left-handers had similar incidence (80 vs 82%, n = 161) and strength (0.41 ± 0.03 s-1 vs 0.37 ± 0.03 s-1) of MU synchrony in dominant and non-dominant hands. No LH subject had a significant difference in synchronization between hands. Force tremor was quantified in each hand in the same subjects during isometric abduction of FDI at 0.5 N and 3.5 N, and directly correlated with the extent of MU synchronization in the muscle. Tremor root mean square amplitude was similar in dominant and non-dominant hands. Power spectral analysis of the tremor force revealed that the peak frequency in the power spectrum was not influenced by handedness, but power at the peak frequency was higher in the non-dominant hand of RH subjects. Correlations between MU discharge variability and synchrony with measures of tremor amplitude were weak. The reduced MU synchronization in the dominant hand of right-handers may reflect a more restricted distribution of direct projections from motor cortical neurons within the FDI motoneuron pool, or reduced excitability of the cortical neurons during the task. These differences in MU synchronization, however, had an insignificant influence on the magnitude of physiological tremor in the FDI.

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.