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S J Garland

Publications and source records attributed to S J Garland.

At least 19 recordsLinked to original sources

Low-frequency common modulation of soleus motor unit discharge is enhanced during postural control in humans.

The maintenance of quiet stance requires the activation of muscles bilaterally. The soleus muscles in each leg share a common function in standing; that is, each muscle acts to control antero-posterior (AP) sway on its own side. We sought to determine the extent to which oscillations in motor unit discharge were related in motor unit pairs of the soleus muscles during postural and voluntary isometric tasks, both within and between legs. Subjects stood quietly for 5 min or performed a voluntary isometric plantarflexion contraction in a seated position. During the postural tasks, the excursions of AP sway between legs were highly correlated (rho = 0.86 +/- 0.06). The strength of common modulation of motor unit discharge rates was assessed using time- and frequency-domain analyses. The time-domain common drive analysis revealed that the strongest correlation in motor unit discharge modulation occurred in the postural task with unilateral pairs (rho = 0.71 +/- 0.13) being more strongly correlated than bilateral pairs (rho = 0.50 +/- 0.16). Common modulation of motor unit discharge was lowest for the voluntary tasks, with rho = 0.38 +/- 0.11 and 0.16 +/- 0.08 for unilateral and bilateral pairs, respectively. Similarly, the frequency-domain coherence analysis demonstrated an identical ordering effect, with the largest maximum pooled coherence occurring during standing posture in unilateral (0.070 at 1.6 Hz) and bilateral (0.055 at 1.6 Hz) recordings, whereas minimal coherence was observed in the voluntary task in both unilateral and bilateral recordings within the 0-5 Hz range. These results indicate that in the soleus muscle, common modulation of motor unit discharge is greater during postural tasks than during voluntary isometric tasks and can be observed in both bilateral and unilateral motor unit pairs. Differences in the extent of co-modulation of motor unit discharge between tasks may be attributed to either differences in the descending control or differences in the proprioceptive input between postural and isometric tasks.

Action Potentials↗

Synchronization of motor units in human soleus muscle during standing postural tasks.

During standing posture, the soleus muscles acts to control sway in the anteroposterior (AP) direction. The soleus muscles bilaterally share a common function during standing tasks. We sought to determine whether common descending inputs, as evidenced by the synchronization of bilateral motor unit pairs, were employed as a strategy to control this common function. Single motor units were recorded from the soleus muscles in subjects who stood on adjacent force platforms for 5 min with their eyes open or closed. While standing with the eyes open, only 4/39 bilateral motor unit pairs showed significant synchronization. Similarly, only 3/36 motor unit pairs were significantly synchronized during the eyes closed task. The low incidence of synchronization was observed despite a high correlation in the amount of sway in the AP direction between legs in both the eyes open and eyes closed tasks (rho = 0.80 and rho = 0.83, respectively). When the extent of synchronization was assessed between pairs of motor units within the same leg with the eyes open, 10/12 pairs were synchronized. Furthermore, when pairs of soleus motor units were recorded both bilaterally and unilaterally during voluntary isometric ankle plantarflexion, only 4/30 bilateral pairs showed significant synchronization, whereas 19/24 unilateral pairs had significant synchronization. In this study, there was little evidence of the existence of synchronization between bilateral soleus motor unit pairs in either postural tasks or voluntary isometric contractions. In cases in which bilateral synchronization was observed, it was considerably weaker than the synchronization of motor units within a single soleus muscle. The results of this study reveal that it is rather uncommon for bilateral soleus motoneurons to receive common descending synaptic inputs, whereas two motoneurons within a single soleus muscle do.

Action Potentials↗

Modulation of motor unit discharge rate and H-reflex amplitude during submaximal fatigue of the human soleus muscle.

Declining motor unit discharge rates and H-reflex amplitude have been observed in separate experiments during fatiguing submaximal contractions in humans. The purpose of this experiment was to investigate motor unit discharge rate, H-reflex amplitude, and twitch contractile properties concurrently during a fatiguing submaximal isometric contraction of the ankle plantarflexors. Eleven healthy subjects performed fatiguing contractions of low force (25% maximal voluntary contraction (MVC)) or high force (42-66% MVC). Hoffmann (H)-reflexes, muscle compound action potentials (M-waves), twitch contractile properties, and motor unit discharges were recorded from the soleus muscle. In the low-force fatigue task, motor unit firing rate increased gradually over time, whereas the resting H-reflex was significantly depressed at 15% of endurance time and remained quasiconstant for the rest of the task. This suggests that the processes mediating the resting H-reflex depression are relatively independent of those modulating the motor unit firing rate during a low-force fatigue task. In the high-force fatigue task, a decline in the average motor unit discharge rate was accompanied by a decrease in the resting H-reflex amplitude and a prolongation of the twitch half-relaxation time (HRT) at the completion of the fatigue task. Overall, motor unit firing rate was modulated in parallel with changes in the twitch HRT, consistent with the muscle wisdom hypothesis.

Action Potentials↗

Postural muscle activity during bilateral and unilateral arm movements at different speeds.

The muscle activation patterns in anterior and posterior leg muscles were investigated with two types of perturbations to standing balance. Subjects stood with each foot on adjacent force platforms and performed arm flexion movements to shoulder height. Nine subjects performed ten repetitions unilaterally and bilaterally at 100, 75, 50, 25, and 12.5% of maximal acceleration as measured by an accelerometer placed on the dominant hand. Four subjects also performed the fastest movements while leaning forwards and backwards. The area and latency of the EMG activity from the quadriceps (QUAD), hamstrings (BF), soleus (SOL), and tibialis anterior (TA) were measured bilaterally, along with the excursions of the center of pressure (COP) during each movement. In both unilateral and bilateral tasks, subjects showed a scaling of EMG area and COP excursion with the acceleration of the arm movement. Prior to movement onset, significant scaling of EMG area with movement speed occurred in both unilateral and bilateral tasks in most muscles. Following movement onset, EMG areas scaled significantly to movement speed in only the anterior musculature, with the exception of the left BF. The latency of BF was consistent for the four fastest movements. Only the slowest movements resulted in a significant rightward shift of the BF EMG latency. During the unilateral task, the ipsilateral hamstrings were activated significantly earlier than in the bilateral task and the contralateral hamstrings were activated significantly later. It was also observed that subjects utilized one of two different strategies to maintain balance. Five individuals displayed simultaneous anterior/posterior muscle activation while the other four displayed a reciprocal pattern of activation. Regardless of the initial standing position (leaning forwards or backwards), subjects used the same simultaneous or reciprocal activation strategy. The results indicate that muscle activation patterns change with different tasks, but remain the same during variations of the same task.

Adult↗

Changes in motor unit discharge rate are not associated with the amount of twitch potentiation in old men.

This study examined, in nine old men (82 +/- 4 yr), whether there is an association between the magnitude of change in motor unit discharge rate and the amount of twitch potentiation after a conditioning contraction (CC). The evoked twitch force and motor unit discharge rate during isometric ramp-and-hold contractions (10-18 s) of the triceps brachii muscle at 10, 20, and 30% of the maximal voluntary contraction were determined before and 10 s, 2 min, 6 min, and 11 min after a 5-s CC at 75% maximal voluntary contraction. After the CC, there was a potentiation of twitch force (approximately twofold), and the discharge rate of the 47 sampled motor units declined (P < 0.05) an average of 1 Hz 10 s after the CC, compared with the control condition. The CC had no effect on the variability (coefficient of variation) of both force and discharge rate, as well as the electromyographic activity recorded over the triceps brachii and biceps brachii muscles. In contrast to our earlier study of young men (Klein CS, Ivanova TD, Rice CL, and Garland SJ, Neurosci Lett 316: 153-156, 2001), the magnitude of the reduction in discharge rate after the CC was not associated (r = 0.06) with the amount of twitch potentiation. These findings suggest an age-related alteration in the neural strategy for adjusting motor output to a muscle after a CC.

Aged↗

Motor unit discharge rate following twitch potentiation in human triceps brachii muscle.

It has been proposed that during brief voluntary contractions, twitch potentiation may sustain force output despite a decline in motor unit discharge rate. This study examined the evoked twitch force and motor unit discharge rates during submaximal voluntary contractions of the triceps brachii muscle before and after a 5 s conditioning contraction (CC) at 75% of maximal voluntary force. After the CC, twitch force potentiated ( approximately 1.3-2-fold), and the discharge rate in 33 of 35 motor units declined significantly by 1-6 Hz. The increase in twitch force was significantly correlated with the decline in discharge rate (r=-0.74). These findings suggest that the extent of the decrease in motor unit discharge rate following a CC is associated with the magnitude of twitch potentiation.

Action Potentials↗

Muscle vibration sustains motor unit firing rate during submaximal isometric fatigue in humans.

1. In keeping with the 'muscular wisdom hypothesis', many studies have documented that the firing rate of the majority of motor units decreased during fatiguing isometric contractions. The present study investigated whether the application of periodic muscle vibration, which strongly activates muscle spindles, would alter the modulation of motor unit firing rate during submaximal fatiguing isometric contractions. 2. Thirty-three motor units from the lateral head of the triceps brachii muscle were recorded from 10 subjects during a sustained isometric 20 % maximal voluntary contraction (MVC) of the elbow extensors. Vibration was interposed on the contraction for 2 s every 10 s. Twenty-two motor units were recorded from the beginning of the fatigue task. The discharge rate of the majority of motor units remained constant (12/22) or increased (4/22) with fatigue. Six motor units demonstrated a reduction in discharge rate that later returned toward initial values; these motor units had higher initial discharge rates than the other 16 motor units. 3. In a second series of experiments, four subjects held a sustained isometric 20 % MVC for 2 min and then vibration was applied as above for the remainder of the contraction. In this case, motor units initially demonstrated a decrease in firing rate that increased after the vibration was applied. Thus muscle spindle disfacilitation of the motoneurone pool may be associated with the decline of motor unit discharge rate observed during the first 2 min of the contraction. 4. In a third set of experiments, seven subjects performed the main experiment on one occasion and repeated the fatigue task without vibration on a second occasion. Neither the endurance time of the fatiguing contraction nor the MVC torque following fatigue was affected by the application of vibration. This finding calls into question the applicability of the muscular wisdom hypothesis to submaximal contractions.

Adult↗

Blood flow in the triceps brachii muscle in humans during sustained submaximal isometric contractions.

The main purpose of this study was to determine the extent to which blood flow through the profunda artery within the triceps brachii muscle may be compromised during maintained low-force isometric fatiguing contractions. Doppler ultrasound techniques were used to record mean blood velocity and arterial diameter of the profunda brachii artery during sustained isometric contractions of 20% maximal voluntary contraction. The arterial diameter did not change throughout the contraction. Thus, blood velocity was considered to be an indicator of blood flow. The mean blood velocity increased initially and then remained constant during the contraction period. When compared to rest [0.06 (SD 0.03) m s-1] mean blood velocity was significantly larger at the start of the contraction [0.13 (SD 0.07) m s-1] and larger yet during recovery following the contraction [0.30 (SD 0.14) m s-1]. Although blood flow through the conduit artery did not drop during the contraction, the post-contraction hyperaemia suggested that circulatory compromise might have occurred at the level of the capillary beds.

Adult↗

Role of limb movement in the modulation of motor unit discharge rate during fatiguing contractions.

Motor unit firing rates of the triceps brachii muscle have been shown to decline during sustained isometric contractions, but not if the fatiguing contraction incorporates arm movements. The purpose of this study was to determine the impact of the actual physical displacement of the limb on the maintenance of motor unit discharge rate during dynamic muscle fatigue. An isometric force pulse paradigm was used to recreate the motor unit activity patterns that occur during a dynamic contraction. With this paradigm, the variable force output that would occur during a dynamic contraction remained intact, but the movement of the limb was eliminated. Motor unit firing rates declined in the isometric force pulse protocol. Thus, factors related to the actual movement of the limb appear to enable the maintenance of motor unit discharge rates during fatigue.

Adult↗

Motor unit double discharges: statistical anomaly or functional entity?

Motor unit double discharges, or doublets, have been described as two consecutive motor unit discharges that occur with a short interspike interval of 2.5 - 20 ms (Simpson, 1969). Double discharges have been reported in the literature for over 70 years. For instance, Eccles and Hoff (1932) found that double discharges were elicited occasionally at the onset of a crossed extension reflex in the soleus muscle of the anaesthetized cat. With the use of electrical stimulation protocols, short interspike intervals inserted at the beginning of a stimulation train have been shown to increase both the peak force and rate of rise of force production, and also decrease the range of fatigue. The extent to which double discharges occur in naturally-occurring voluntary behaviours remains relatively unexplored. This review examines the issue of whether double discharges occur solely because of an intrinsic property of motoneurones, thereby representing a "statistical anomaly," or whether they may result from a neural control strategy to augment force production, i.e., a "functional entity."

Animals↗

Discharge patterns in human motor units during fatiguing arm movements.

The purpose of this study was to determine whether short interspike intervals (ISIs of <20 ms) would occur naturally during voluntary movement and would increase in number with fatigue. Thirty-four triceps brachii motor units from nine subjects were assessed during a fatigue task consisting of fifty extension and fifty flexion elbow movements against a constant-load opposing extension. Nineteen motor units were recorded from the beginning of the fatigue task; the number of short ISIs was 7.1 +/- 4.1% of the total number of ISIs in the first one-third of the task (unfatigued state). This value increased to 11.8 +/- 5.9% for the last one-third of the task (fatigued state). Fifteen motor units were recruited during the fatigue task and discharged, with 16.4 +/- 6.0% of short ISIs in the fatigued state. For all motor units, the number of short ISIs was positively correlated (r2 = 0.85) with the recruitment threshold torque. Short ISIs occurred most frequently at movement initiation but also occurred throughout the movement. These results document the presence of short ISIs during voluntary movement and their increase in number during fatigue.

Adult↗

Motor unit discharge rate is not associated with muscle relaxation time in sustained submaximal contractions in humans.

The muscle wisdom hypothesis suggests that motor unit discharge rates decrease in parallel with the slowing of muscle contractile properties during fatigue. In this study, the discharge rates of single motor units and the muscle contractile properties were measured during a sustained submaximal contraction. The majority of motor units that were active from the beginning of the task demonstrated a decline in discharge rate in the absence of any slowing of muscle relaxation time. These findings suggest that the muscle wisdom hypothesis may not apply to sustained submaximal contractions.

Adult↗

Control of motor units in human flexor digitorum profundus under different proprioceptive conditions.

1. Changing the posture of the human fingers can functionally 'disengage' the deep finger flexor muscle from its normal action on the terminal phalanx of the fourth (or third) finger. This enables the activity of the muscle to be studied both with and without its normal proprioceptive inputs. 2. Spike trains of long duration from pairs of concurrently active motor units in this muscle were recorded in both the engaged and disengaged hand postures. Subjects voluntarily kept one of the motor units (the 'controlled' unit) discharging at the same target frequency in both postures. The strength of short-term synchrony, the strength of common drive, and the variability of discharge of these pairs of motor units were determined in both postures. 3. All subjects reported that the effort required to activate the motor units in the disengaged hand posture was substantially greater than in the normal engaged posture. 4. Short-term synchrony, which is a function of common corticospinal inputs to pairs of motor units, was similar in both hand postures. However, the strength of common drive was significantly decreased when the muscle was disengaged. Although the neural substrate for common drive is not known, this observation suggests that proprioceptive feedback is involved either directly or indirectly. 5. Although the discharge rate of the 'uncontrolled' motor units increased when the muscle was disengaged, the variability of discharge of these and the 'controlled' motor units increased significantly. This supports the idea that the precision with which fine motor tasks can be performed is improved when proprioceptive feedback is intact.

Action Potentials↗

Motor unit recruitment and discharge behavior in movements and isometric contractions.

The purpose of this study was to contrast the discharge patterns of the same motor units during movements and during isometric contractions that were produced with comparable torque-time characteristics. Subjects performed elbow flexion and extension movements with predetermined acceleration characteristics. The average acceleration and deceleration profiles for the movements were reproduced in the isometric setting by presenting the kinematic profiles as templates for torque production. Trained subjects were able to match the first agonist (AG1) and antagonist (ANT) electromyographic (EMG) bursts, but tended to produce a smaller second agonist burst (AG2) in the isometric contraction. Twenty-five motor units from triceps brachii were studied. The same motor units (with one exception) were recruited and subsequently discharged in a similar fashion in both the isometric and movement tasks in the AG1 and ANT EMG bursts, with fewer motor unit discharges in the AG2 burst in the isometric contraction. The central control mechanisms appear to be the same for the acceleration phase of movement and isometric contraction, but differ during the deceleration phase.

Adult↗

Postural responses to unilateral arm perturbation in young, elderly, and hemiplegic subjects.

OBJECTIVE: To compare postural responses during standing associated with self-paced unilateral arm flexion exhibited by young and elderly healthy subjects and subjects with hemiplegia from stroke. DESIGN AND SETTING: Descriptive cross-sectional study in a laboratory setting. PATIENTS: Ten young, 10 elderly healthy subjects, and 12 volunteers with longstanding hemiparesis following stroke were tested. The hemiplegic group had good functional balance scores on the Berg Balance Scale (BBS). MAIN OUTCOME MEASURES: The peak arm acceleration, center of pressure (CP) excursion speed, and the electromyographic activity in the posterior leg muscles (bilateral hamstrings and triceps surae) were monitored during self-initiated rapid unilateral arm flexion and during quiet stance. RESULTS: During unilateral arm flexion, the young group showed the highest arm accelerations and lowest CP excursion speeds. The variability of postural responses was largest in the elderly and hemiplegic groups with hemiplegic subjects showing the greatest CP excursion speeds for the lowest arm accelerations. The first burst of activity in ipsilateral hamstrings muscle was the same in all subjects. However, the hemiplegic group had less activation (latency and amplitude) of other posterior leg muscles. CONCLUSION: The elderly subjects had more variable responses to perturbation than the young subjects despite similar BBS scores. This may indicate that the BBS fails to detect differences in balance at the high end of the scale. Although the hemiplegic subjects demonstrated some anticipatory control of standing balance, they consistently performed poorer than elderly and young controls. The failure to coordinate postural leg muscle activity with focal movement may contribute to the instability of subjects with hemiplegia.

Adult↗

Responses of human single motor units to transcranial magnetic stimulation.

Transcranial electromagnetic brain stimuli elicit a complex response in the electromyogram of active human hand muscles. Relatively weak stimuli evoke a short-latency primary response via a presumably monosynaptic corticospinal path. This is followed by a silent period that is terminated by a second peak at a latency of 50-80 ms. The responses evoked in single motor units in flexor digitorum profundus (FDP) were recorded. Responses were elicited at the second-peak latency only in trials in which no primary response was elicited in that unit, and only when the stimulus was given during the first half of the interspike interval (ISI). When given during the second half of the ISI, the same stimulus evoked a primary response but no second peak response. Stronger stimuli suppressed the second peak by evoking a primary response in more trials. Having discharged at about 20 ms latency, the parent motoneurone was unable to discharge again at second-peak latency, 30-60 ms later. The response at second-peak latency was not modified by disengaging both FDP and the extensors of the distal interphalangeal joint. Hence, this response is not secondary to a stretch reflex provoked by activation of the finger extensors, nor is it the result of a cutaneous signal resulting from movement of the finger. The latencies suggest that the corticospinal volley evokes a beta-motoneurone-mediated twitch in FDP muscle spindles, which elicits an afferent volley that activates the motoneurone reflexly. The first 100 ms or so of the silent period is due to the realignment of the first post-stimulus spike in most trials to corticospinal latency; i.e. this is not necessarily the result of an inhibitory or disfacilitatory process. Still stronger stimuli increase the duration of the ISI in which the stimulus is given, indicating the presence of an inhibitory/disfacilitatory process.

Adult↗

Standing balance during internally produced perturbations in subjects with hemiplegia: validation of the balance scale.

OBJECTIVE: To examine the concurrent validity of the Berg Balance Scale (BBS) in subjects with hemiparesis after stroke by correlating functional balance performance on the BBS with laboratory measures of balance (center of pressure [CP] and electromyographic [EMG] activity of leg musculature during rapid arm flexion and during quiet stance). DESIGN: Concurrent validation study. Consistency of performance was determined through repeat measures obtained from 13 subjects. SETTING: Laboratory. PATIENTS: Twenty-four volunteers with longstanding hemiparesis from cerebrovascular accident (CVA) were tested. MAIN OUTCOME MEASURES: BBS scores were compared with CP excursion speed during (1) self-initiated rapid unilateral arm flexion with the nonhemiplegic arm and (2) quiet stance. The peak tangential arm acceleration and the sequence of EMG events in the posterior leg muscles were monitored during the arm flexion movement. RESULTS: In the arm flexion condition, BBS performance was significantly correlated with arm acceleration and CP excursion speed (multiple regression R = .81). BBS performance was also related to the presence of anticipatory activation of ipsilateral hamstrings during the rapid arm flexion movement. BBS performance and CP excursion speed during quiet stance were also significantly related (r = .76). The initial and repeat measures were not significantly different in magnitude (paired tests of difference, p < .01). CONCLUSION: The BBS appears to reflect differing abilities to tolerate the internally produced perturbation to standing balance associated with forward arm flexion in individuals with hemiparesis from CVA.

Aged↗