Adaptations in motor unit discharge activity with force control training in young and older human adults

Six young (aged 18–22 years) and six older (aged 66–76 years) healthy humans participated in a visually guided isometric force modulation training program designed to improve accurate control of force during ankle dorsiflexion. Isometric force and the discharge activity of motor units (MU) supplying the tibialis anterior muscle were sampled concurrently at the beginning of the study, following 2 weeks of force modulation training and again after a 4 week retention period which followed immediately. The initial maximal voluntary force (MVC) and MU discharge rates were similar between young and older adults at 10–60% MVC while MU discharge rates during maximal effort were significantly reduced in older adults. Following the 2 weeks of force modulation training, both young and older adults demonstrated significant improvements in force accuracy (44% young, 48% older) and significantly reduced MU discharge rates at 30%, 40%, and 60% MVC. Young adults also demonstrated increased MVC force (11%), while older adults demonstrated significantly increased (30%) maximal MU discharge rate. Thus, following 2 weeks of force modulation training, young and older individuals demonstrated similar MU discharge rates at all force levels. The MU discharge rate adaptations were retained after the 4 week retention period. In young adults, improved force accuracy and increased MVC force were accompanied by significantly reduced MU recruitment thresholds. In the older subjects, improved force accuracy was accompanied by an increase in the difference between the recruitment-derecruitment force threshold and significantly reduced antagonist co-contraction. Age-related alterations in force regulation and MU discharge activity cannot be explained solely on the basis of contractile changes in senescent muscle. Rather, reliance on compensatory neuromuscular changes including antagonist muscle co-contraction is suggested. Accepted: 27 June 2000     

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.     

Effects of aging on force variability,single motor unit discharge patterns,and the structure of 10, 20, and 40 Hz EMG activity

The purpose of this investigation was to examine the discharge properties of single motor units and the structure of the rectified 10, 20, and 40 Hz electromyographic (EMG) activity to determine a physiological correlate for the greater force variability with aging. Young (n=10; mean: 22+/-1 years), old (n=10; mean: 67+/-2 years), and older-old (n=10; mean: 82+/-5 years) adult humans produced isometric second finger abduction force in both constant and sine-wave tasks at 5, 10, 20, and 40% of their maximal voluntary contraction. Force and fine-wire intramuscular electromyography were recorded from the first dorsal interosseous muscle. The amount and time-dependent structure of the discharge rate variability of single motor units and Fourier analysis of the rectified intramuscular EMG was performed. Force output variability increased across the young, old, and older-old groups. The amount and time-dependent structure of the discharge rate variability of single motor units did not differ between the young and aging groups. There was a progressive decrease in the relative power of approximately 40 Hz EMG activity from the young>old>older-old subjects across the 5, 10, 20, and 40% maximum voluntary contraction (MVC) force levels. There was also a progressive increase in the relative power of the approximately 10 Hz EMG activity from young相似文献   

Maximal force,force/time and activation/coactivation characteristics of the neck muscles in extension and flexion in healthy men and women at different ages

This study examined the force production characteristics, activation/coactivation and endurance capacity of the neck extension and flexion muscles in healthy men (n=29) and women (n=28) divided into three age groups (18–26 years, 30–37 years and 45–55 years). Force and electromyography (EMG) measurements were performed during the maximal voluntary isometric extension and flexion actions. This was followed by an endurance test (ET; 60% force level of maximal voluntary contraction sustained until exhaustion), after which the force and EMG recordings were repeated. Men were both stronger and had higher values (P<0.001) for explosive force (rate of force development, RFD) than women in both actions. Younger subjects of both genders exhibited larger (P<0.05 in women) RFD values than older subjects in extension. The coactivation of the antagonist muscles during the maximal extension or flexion did not differ significantly between men and women, but the coactivation of the antagonists was larger (P<0.05) in the older age groups than in the youngest group. Women maintained the 60% force level longer than men in both actions (extension, P<0.001; flexion, not significant). The fatiguing loading led to significant decreases in maximal isometric force (P<0.001) and RFD (P<0.01–0.001), but these relative decreases did not differ between the groups. In conclusion, large gender differences in the voluntary extension and flexion force production characteristics of the neck muscles did exist, as reported earlier for other muscles of the body. No age-related differences were observed in maximal force of the extension and flexion actions within the age ranges of the subject groups studied here, but the older subjects exhibited greater coactivation and produced lower force values in the early portions of the force/time curve of the extension than the youngest group. The data indicate that explosive force production may be sensitive to aging earlier than maximal strength in the case of the neck extensor muscles. Electronic Publication     

Independence between the amount and structure of variability at low force levels

The purpose of the current experiment was to investigate the amount (standard deviation (S.D.) and coefficient of variation (CV)) and structure (approximate entropy (ApEn)) of force variability at very low force levels. Participants produced isometric force output of index finger abduction at five levels (0.4, 0.8, 1.0, 2.0, and 4.0 N) with high and low visual feedback gain. The findings showed that: subjects scaled their force output to the targets; S.D. increased non-linearly with force level and decreased with visual gain; and CV decreased with force level as well as visual gain. ApEn of the force output did not change as a function of force level, although the high gain increased ApEn in contrast to low gain. It is proposed that the recruitment of additional motor units at very low force levels does not significantly alter the structure of the force output, although it does increase the magnitude of force and its amount of variability. Overall, the findings provide evidence that the amount and structure of motor variability can be influenced by separate control processes at low force levels.     

Role of across-muscle motor unit synchrony for the coordination of forces

Evidence from five-digit grasping studies indicates that grip forces exerted by pairs of digits tend to be synchronized. It has been suggested that motor unit synchronization might be a mechanism responsible for constraining the temporal relationships between grip forces. To evaluate this possibility and quantify the effect of motor unit synchrony on force relationships, we used a motor unit model to simulate force produced by two muscles using three physiological levels of motor unit synchrony across the two muscles. In one condition, motor units in the two muscles discharged independently of one another. In the other two conditions, the timing of randomly selected motor unit discharges in one muscle was adjusted to impose low or high levels of synchrony with motor units in the other muscle. Fast Fourier transform analysis was performed to compute the phase differences between forces from 0.5 to 17 Hz. We used circular statistics to assess whether the phase differences at each frequency were randomly or non-randomly distributed (Rayleigh test). The mean phase difference was then computed on the non-random distributions. We found that the number of significant phase-difference distributions increased markedly with increasing synchronization strength from 18% for no synchrony to 65% and 82% for modest and strong synchrony conditions, respectively. Importantly, most of the mean angles clustered at very small phase difference values (~0 to 10°), indicating a strong tendency for forces to be exerted in a synchronous fashion. These results suggest that motor unit synchronization could play a significant functional role in the coordination of grip forces.     

Fluctuations of isometric force after eccentric exercise of the elbow flexors of young,middle-aged,and old men

This study compared force fluctuations during isometric contraction following eccentric exercise of the elbow flexors between young, middle-aged, and old subjects. Ten young (20 ± 2.0 years), 12 middle-aged (48 ± 7.3 years), and 10 old (71 ± 4.1 years) men performed six sets of five eccentric actions of the elbow flexors using a dumbbell weighing 40% of maximal voluntary isometric contraction strength (MVC) at an elbow joint angle of 90° (1.57 rad). MVC was measured before, immediately after, and 1–5 days following exercise, and the force fluctuations were assessed at 30, 50, and 80% of the corresponding time point MVC using coefficient of variation (CV) of force data collected at a frequency of 100 Hz for 4 s. Changes in MVC and CV over time were compared between groups by a two-way repeated measures ANOVA. Changes in MVC following exercise were not significantly different between the young and middle-aged groups, but the old group showed significantly (P < 0.05) smaller decreases in MVC compared with other groups. CV increased significantly (P < 0.05) only immediately after exercise without a significant difference among the three intensities, and no significant differences between groups were evident. It was concluded that force fluctuations during submaximal isometric tasks after eccentric exercise were not affected by age.     

Contraction characteristics of the human quadriceps muscle during percutaneous electrical stimulation

Percutaneous electrical stimulation was used to study the force response of the quadriceps muscle. The normal frequency dependence of force was investigated in muscles at rest and after fatiguing contractions. A comparison between force response during fatigue induced by electrical stimulation at different frequencies and by voluntary work suggested equal changes in contractility, irrespective of the fatigue-inducing procedure. In fresh muscle we found a linear relation between stimulation period (10–100 ms) and force. At fatigue the relation changes with maximal deviation from linearity at a 50-ms period (20 Hz). There is a rapid recovery of high frequency force whereas the low frequency response remains low even after 30 min rest. At very low frequencies there is initially unexpectedly high force in fatigued muscle. This could be a result of increased fusion of twitches with initially prolonged relaxation time. To study the twitch summation we compared experimental results in a wide frequency range with computer-simulated twitch summations and present the frequency dependence of summation processes in human quadriceps muscle.     

Combined effect of motor imagery and peripheral nerve electrical stimulation on the motor cortex

Although motor imagery enhances the excitability of the corticospinal tract, there are no peripheral afferent inputs during motor imagery. In contrast, peripheral nerve electrical stimulation (ES) can induce peripheral afferent inputs; thus, a combination of motor imagery and ES may enhance the excitability of the corticospinal tract compared with motor imagery alone. Moreover, the level of stimulation intensity may also be related to the modulation of the excitability of the corticospinal tract during motor imagery. Here, we evaluated whether a combination of motor imagery and peripheral nerve ES influences the excitability of the corticospinal tract and measured the effect of ES intensity on the excitability induced during motor imagery. The imagined task was a movement that involved touching the thumb to the little finger, whereas ES involved simultaneous stimulation of the ulnar and median nerves at the wrist. Two different ES intensities were used, one above the motor threshold and another above the sensory threshold. Further, we evaluated whether actual movement with afferent input induced by ES modulates the excitability of the corticospinal tract as well as motor imagery. We found that a combination of motor imagery and ES enhanced the excitability of the motor cortex in the thenar muscle compared with the other condition. Furthermore, we established that the modulation of the corticospinal tract was related to ES intensity. However, we found that the excitability of the corticospinal tract induced by actual movement was enhanced by peripheral nerve ES above the sensory threshold.     

Oscillations in motor unit discharge are reflected in the low-frequency component of rectified surface EMG and the rate of change in force

Common drive to a motor unit (MU) pool manifests as low-frequency oscillations in MU discharge rate, producing fluctuations in muscle force. The aim of the study was to examine the temporal correlation between instantaneous MU discharge rate and rectified EMG in low frequencies. Additionally, we attempted to examine whether there is a temporal correlation between the low-frequency oscillations in MU discharge rate and the first derivative of force (dF/dt). Healthy young subjects produced steady submaximal force with their right finger as a single task or while maintaining a pinch-grip force with the left hand as a dual task. Surface EMG and fine-wire MU potentials were recorded from the first dorsal interosseous muscle in the right hand. Surface EMG was band-pass filtered (5–1,000 Hz) and full-wave rectified. Rectified surface EMG and the instantaneous discharge rate of MUs were smoothed by a Hann-window of 400 ms duration (equivalent to 2 Hz low-pass filtering). In each of the identified MUs, the smoothed MU discharge rate was positively correlated with the rectified-and-smoothed EMG as confirmed by the distinct peak in cross-correlation function with greater values in the dual task compared with the single task. Additionally, the smoothed MU discharge rate was temporally correlated with dF/dt more than with force and with rectified-and-smoothed EMG. The results indicated that the low-frequency component of rectified surface EMG and the first derivative of force provide temporal information on the low-frequency oscillations in the MU discharge rate.     

Comparison of fluctuations of motor unit recruitment and de-recruitment thresholds in man

Recruitment and de-recruitment thresholds of motor units in the wrist extensor muscles can undergo important random fluctuations, even when they are measured during stereotyped contractions and relaxations. These fluctuations were statistically quantified and compared. The statistical analysis indicated that recruitment and de-recruitment thresholds display the same kind of fluctuations, and that the successive measurements are randomly distributed following a quasi-normal law. We suggest that the notion of force threshold for motor unit recruitment and de-recruitment might be oversimplified and that a motor unit seems to have a range of force in which it can be recruited or de-recruited. Comparison of the mean values of recruitment and de-recruitment thresholds of the motor units in the extensor carpi radialis muscles showed that de-recruitment thresholds were significantly lower than recruitment thresholds. This difference in the thresholds, together with the difference in the motor unit discharge frequency during a contraction and a relaxation, suggests a differential control of the motoneurone activity during contractions and relaxations.     

Motor variability within a multi-effector system: experimental and analytical studies of multi-finger production of quick force pulses

The purpose of the study was to develop a model of force variability for a fast action performed by a multi-effector system and to verify it for multi-finger quick force production. The experiments involved quick isometric contractions to different target force levels using different finger combinations. Force variance calculated over sets of trials for a multi-finger force production task showed non-monotonic single-peak profiles of force variance with a peak at a time between the times of the maxima of the force rate and of the total force. When analyzed in the four-dimensional space of finger forces, the variance peak was mostly expressed in the direction of the force rate, and was absent in the directions orthogonal to it. The non-monotonic time profile of the force variance could be reproduced by a model of force production, which assumes that each finger force profile is based on a template function scaled in duration and magnitude with two parameters assigned prior to each trial with some variability. The model allows decomposition of the force variance into two fractions related to variability in setting the magnitude and duration scaling parameters. The former fraction changes monotonically with time, while the latter shows a transient peak in the middle of the action. The model was able to reproduce experimental variance time profiles across conditions with the total error of under 8%. The results demonstrate, in particular, that fast multi-finger actions may show transient changes in motor variability in certain directions of the finger force space, particularly in the direction of the first force derivative, without any task-specific coordinating action by the controller. These findings require a reconsideration of some of the conclusions drawn in recent studies on the structure of motor variability in redundant multi-effector systems.     

Comparison of neural damage induced by electrical stimulation with faradaic and capacitor electrodes

Arrays of platinum (faradaic) and anodized, sintered tantalum pentoxide (capacitor) electrodes were implanted bilaterally in the subdural space of the parietal cortex of the cat. Two weeks after implantation both types of electrodes were pulsed for seven hours with identical waveforms consisting of controlled-current, chargebalanced, symmetric, anodic-first pulse pairs, 400 μsec/phase and a charge density of 80–100 μC/cm² (microcoulombs per square cm) at 50 pps (pulses per second). One group of animals was sacrificed immediately following stimulation and a second smaller group one week after stimulation. Tissues beneath both types of pulsed electrodes were damaged, but the difference in damage for the two electrode types was not statistically significant. Tissue beneath unpulsed electrodes was normal. At the ultrastructural level, in animals killed immediately after stimulation, shrunken and hyperchromic neurons were intermixed with neurons showing early intracellular edema. Glial cells appeared essentially normal. In animals killed one week after stimulation most of the damaged neurons had recovered, but the presence of shrunken, vacuolated and degenerating neurons showed that some of the cells were damaged irreversibly. It is concluded that most of the neural damage from stimulations of the brain surface at the level used in this study derives from processes associated with passage of the stimulus current through tissue, such as neuronal hyperactivity rather than electrochemical reactions associated with current injection across the electrode-tissue interface, since such reactions occur only with the faradaic electrodes.     

Area 4 and area 5: differences between the load direction-dependent discharge variability of cells during active postural fixation

Summary Previous experiments have shown that the activity of cells in both areas 4 and 5 is strongly modulated in a continuously-graded manner for movements of the arm in 8 different directions away from a central starting position. However, it is reported here that there is a clear difference between these two areas during postural fixation of the arm in one position against loads applied in 8 different directions. Area 4 cell activity shows large variations when the monkey holds its arm at the central position by exerting forces in different directions to counteract the loads. The activity of cells in area 5 is much less strongly modulated under the same isometric conditions. These observations suggest that cell activity in area 5 is less related to the level and pattern of force output than are cells in area 4. Area 5 may produce instead a signal related more to the limb position, per se, during actively maintained posture.     

On the relations between single cell activity in the motor cortex and the direction and magnitude of three-dimensional dynamic isometric force

The role of the motor cortex in the control of both the direction and magnitude of dynamic force, when both are allowed to vary in 3D, is not known. We recorded the activity of 504 cells in the motor cortex of two monkeys during a behavioral task in which the subjects used a manipulandum to vary both the direction and magnitude of isometric force in 3D space. The majority (86%) of cells active in the task related to the direction, a tiny number (2.5%) to the magnitude, and a moderate number (11.5%) to both the direction and magnitude of dynamic force output. Finally, we compared neural activity in the same population of neurons during dynamic and static force output and found that the relations to direction and magnitude were very similar in both epochs. Our results indicate that during dynamic force production, cells in the motor cortex are primarily concerned with specifying the direction of force. The magnitude signal is not prominent in motor cortex neurons, and in general, magnitude and direction of force are specified together. Furthermore, the data suggest that the control of static and dynamic motor systems is based, to a great extent, on a common control process. The work was supported in part by a Merit Review award from the Department of Veterans Affairs and by the American Legion Brain Sciences Chair.     

Variability of quadriceps femoris motor neuron discharge and muscle force in human aging

The purpose was to determine the contribution of visual feedback and the effect of aging on the variability of knee extensor (KE) muscle force and motor unit (MU) discharge. Single MUs were recorded during two types of isometric trials, (1) visual feedback provided (VIS) and then removed (NOVIS) during the trial (34 MUs from young, 32 from elderly), and (2) only NOVIS (66 MUs from young, 77 from elderly) during the trial. Recruitment threshold (RT) ranged from 0–37% MVC. Standard deviation (SD) and coefficient of variation (CV) of muscle force and MU interspike interval (ISI) was measured during steady contractions at target forces ranging from 0.3 to 54% MVC. Force drift (<0.5 Hz) was removed before analysis. VIS/NOVIS trials: the decrease in the CV of ISI from VIS to NOVIS was greater for MUs from elderly (12.5 ± 4.1 to 9.94 ± 2.6%) than young (10.6 ± 3.3 to 10.3 ± 2.8%, age group × vision interaction, P = 0.006). The change in CV of force from VIS to NOVIS was significantly greater for elderly (1.45 to 1.05%) than young (1.42 to 1.41%). NOVIS only trials: for all MUs, the average RT (6.6 ± 7.7 % MVC), target force above RT (1.20 ± 2.7% MVC), SD of ISI (0.012 ± 0.005 s), and CV of ISI (11.1 ± 3.3%) were similar for young and elderly MUs. The CV of force was similar between age groups for trials between 0 and 3% MVC (1.74 ± 0.74%) and was greater for young subjects from 3 to 10% MVC (1.47 ± 0.5 vs. 1.21 ± 0.4%) and >10% MVC (1.44 ± 0.6 vs. 1.01 ± 0.3%). The CV of ISI was similar between age groups for MUs in 0–3, 3–10, and >10% bins of RT. Thus, the contribution of visuomotor correction to the variability of motor unit discharge and force is greater for elderly adults. The presence of visual feedback appears to be necessary to find greater discharge variability in motor units from the knee extensors of elderly adults.