scholarly journals Neural control of muscle force: indications from a simulation model

2013 ◽  
Vol 109 (6) ◽  
pp. 1548-1570 ◽  
Author(s):  
Paola Contessa ◽  
Carlo J. De Luca
Keyword(s):  
Motor Unit ◽  
Neural Control ◽  
Muscle Force ◽  
Force Production ◽  
Motor Units ◽  
Isometric Contractions ◽  
Alternative Mechanism ◽  
Firing Behavior ◽  
Firing Rates ◽  
Muscle Force Production

We developed a model to investigate the influence of the muscle force twitch on the simulated firing behavior of motoneurons and muscle force production during voluntary isometric contractions. The input consists of an excitatory signal common to all the motor units in the pool of a muscle, consistent with the “common drive” property. Motor units respond with a hierarchically structured firing behavior wherein at any time and force, firing rates are inversely proportional to recruitment threshold, as described by the “onion skin” property. Time- and force-dependent changes in muscle force production are introduced by varying the motor unit force twitches as a function of time or by varying the number of active motor units. A force feedback adjusts the input excitation, maintaining the simulated force at a target level. The simulations replicate motor unit behavior characteristics similar to those reported in previous empirical studies of sustained contractions: 1) the initial decrease and subsequent increase of firing rates, 2) the derecruitment and recruitment of motor units throughout sustained contractions, and 3) the continual increase in the force fluctuation caused by the progressive recruitment of larger motor units. The model cautions the use of motor unit behavior at recruitment and derecruitment without consideration of changes in the muscle force generation capacity. It describes an alternative mechanism for the reserve capacity of motor units to generate extraordinary force. It supports the hypothesis that the control of motoneurons remains invariant during force-varying and sustained isometric contractions.

Neuromuscular drive and force production are not altered during bilateral contractions

1998 ◽  
Vol 84 (1) ◽  
pp. 200-206 ◽  
Author(s):  
J. M. Jakobi ◽  
E. Cafarelli
Keyword(s):  
Motor Unit ◽  
Force Production ◽  
Young Male ◽  
Neuromuscular Control ◽  
Force Generation ◽  
Isometric Contractions ◽  
Firing Rates ◽  
Significant Limitation ◽  
Motor Unit Firing ◽  
Male Subjects

Jakobi, J. M., and E. Cafarelli. Neuromuscular drive and force production are not altered during bilateral contractions. J. Appl. Physiol. 84(1): 200–206, 1998.—Several investigators have studied the deficit in maximal voluntary force that is said to occur when bilateral muscle groups contract simultaneously. A true bilateral deficit (BLD) would suggest a significant limitation of neuromuscular control; however, some of the data from studies in the literature are equivocal. Our purpose was to determine whether there is a BLD in the knee extensors of untrained young male subjects during isometric contractions and whether this deficit is associated with a decreased activation of the quadriceps, increased activation of the antagonist muscle, or an alteration in motor unit firing rates. Twenty subjects performed unilateral (UL) and bilateral (BL) isometric knee extensions at 25, 50, 75, and 100% maximal voluntary contraction. Total UL and BL force (Δ3%) and maximal rate of force generation (Δ2.5%) were not significantly different. Total UL and BL maximal vastus lateralis electromyographic activity (EMG; 2.7 ± 0.28 vs. 2.6 ± 0.24 mV) and coactivation (0.17 ± 0.02 vs. 0.20 ± 0.02 mV) were also not different. Similarly, the ratio of force to EMG during submaximal UL and BL contractions was not different. Analysis of force production by each leg in UL and BL conditions showed no differences in force, rate of force generation, EMG, motor unit firing rates, and coactivation. Finally, assessment of quadriceps activity with the twitch interpolation technique indicated no differences in the degree of voluntary muscle activation (UL: 93.6 ± 2.51 Hz, BL: 90.1 ± 2.43 Hz). These results provide no evidence of a significant limitation in neuromuscular control between BL and UL isometric contractions of the knee extensor muscles in young male subjects.

Activation of type-identified motor units during centrally evoked contractions in the cat medial gastrocnemius muscle. II. Motoneuron firing-rate modulation

1996 ◽  
Vol 75 (1) ◽  
pp. 38-50 ◽  
Author(s):  
K. E. Tansey ◽  
B. R. Botterman
Keyword(s):  
Firing Rate ◽  
Motor Unit ◽  
Muscle Force ◽  
Motor Units ◽  
Medial Gastrocnemius ◽  
Firing Rates ◽  
Rate Modulation ◽  
The Mean ◽  
Fast Twitch ◽  
The Relationship

1. The aim of this study was to examine the nature of motoneuron firing-rate modulation in type-identified motor units during smoothly graded contractions of the cat medial gastrocnemius (MG) muscle evoked by stimulation of the mesencephalic locomotor region (MLR). Motoneuron discharge patterns, firing rates, and the extent of firing-rate modulation in individual units were studied, as was the extent of concomitant changes in firing rates within pairs of simultaneously active units. 2. In 21 pairs of simultaneously active motor units, studied during 41 evoked contractions, the motoneurons' discharge rates and patterns were measured by processing the cells' recorded action potentials through windowing devices and storing their timing in computer memory. Once recruited, most motoneurons increased their firing rates over a limited range of increasing muscle tension and then maintained a fairly constant firing rate as muscle force continued to rise. Most motoneurons also decreased their firing rates over a slightly larger, but still limited, range of declining muscle force before they were derecruited. Although this was the most common discharge pattern recorded, several other interesting patterns were also seen. 3. The mean firing rate for slow twitch (type S) motor units (27.8 imp/s, 5,092 activations) was found to be significantly different from the mean firing rate for fast twitch (type F) motor units (48.4 imp/s, 11,272 activations; Student's t-test, P < 0.001). There was no significant difference between the mean firing rates of fast twitch, fatigue-resistant (type FR) and fast twitch, fatigable (type FF) motor units. When the relationship between motoneuron firing rate and whole-muscle force was analyzed, it was noted that, in general, smaller, lower threshold motor units began firing at lower rates and reached lower peak firing rates than did larger, higher threshold motor units. These results confirm both earlier experimental observations and predictions made by other investigators on the basis of computer simulations of the cat MG motor pool, but are in contrast to motor-unit discharge behavior recorded in some human motor-unit studies. 4. The extent of concomitant changes in firing rate within pairs of simultaneously active motor units was examined to estimate the extent of simultaneous motoneuron firing-rate modulation across the motoneuron pool. A smoothed (5 point sliding average) version of the two motoneurons' instantaneous firing rates was plotted against each other, and the slope and statistical significance of the relationship was determined. In 16 motor-unit pairs, the slope of the motoneurons' firing-rate relationship was significantly distinct from 0. Parallel firing-rate modulation (< 10-fold difference in firing rate change reflected by a slope of > 0.1) was noted only in pairs containing motor units of like physiological type and then only if they were of similar recruitment threshold. 5. Other investigators have demonstrated that changes in a motoneuron's "steady-state" firing rate predictably reflect changes in the amount of effective synaptic current that cell is receiving. The finding in the present study of limited parallel firing-rate modulation between simultaneously active motoneurons would suggest that changes in the synaptic drive to the various motoneurons of the pool is unevenly distributed. This finding, in addition to the findings of orderly motor-unit recruitment and the relationship between motor-unit recruitment threshold and motoneuron firing rate, cannot be adequately accommodated for by the existing models of the synaptic organization in motoneuron pools. Therefore a new model of the synaptic organization within the motoneuron pool has been proposed.

Effects of Aging on Motor-Unit Control Properties

1999 ◽  
Vol 82 (5) ◽  
pp. 2081-2091 ◽  
Author(s):  
Zeynep Erim ◽  
M. Faisal Beg ◽  
David T. Burke ◽  
Carlo J. de Luca
Keyword(s):  
Firing Rate ◽  
Motor Unit ◽  
Motor Units ◽  
Elderly Subjects ◽  
Compensatory Mechanism ◽  
Type I ◽  
Firing Behavior ◽  
Firing Rates ◽  
Age Related ◽  
Young Subjects

It was hypothesized that the age-related alterations in the morphological properties of a motor unit would be accompanied by modifications in the control aspects of the motor unit, as either an adaptive or compensatory mechanism to preserve smooth force production. In specific, the objective of the study was to investigate the age-related alterations in the concurrent firing behavior of multiple motor units in the first dorsal interosseous (FDI) muscle in isometric contractions at 20 and 50% of the subject's voluntary contraction level. Analysis of the data collected from 10 young (24–37 yr of age) and 10 elderly (65–88 yr of age) subjects led to three novel observations regarding the firing behavior of aged motor units. 1) Among elderly subjects, there is a decrease in the common fluctuations that are observed among the firing rates of motor units in the young. 2) The relationship observed between the firing rate and recruitment threshold of young subjects is disturbed in the elderly. Although in young subjects, at any point in a given submaximal contraction, earlier recruited motor units have higher firing rates than later-recruited units; in aged subjects this dependency of firing rate on recruitment rank is compromised. 3) The progressive decrease observed in the firing rates of concurrently active motor units in constant-force contractions in the young is not seen in the aged. In addition to these original findings, this study provided support for earlier reports of 1) decreased average firing rates probably reflecting the slowing of the muscle, 2) a shift in recruitment thresholds toward lower force levels in line with the shift toward type I fibers, and 3) multiphasic action potential shapes indicative of the reinnervation process that takes place during aging. Taken as a whole, these findings indicate significant age-related modifications in the control properties of human motor units.

scholarly journals The compensatory interaction between motor unit firing behavior and muscle force during fatigue

2016 ◽  
Vol 116 (4) ◽  
pp. 1579-1585 ◽  
Author(s):  
Paola Contessa ◽  
Carlo J. De Luca ◽  
Joshua C. Kline
Keyword(s):  
Motor Unit ◽  
Muscle Force ◽  
Vastus Lateralis ◽  
Motor Units ◽  
Vastus Lateralis Muscle ◽  
Clear Understanding ◽  
Contraction Force ◽  
Firing Behavior ◽  
Motoneuron Pool ◽  
Motor Unit Firing

Throughout the literature, different observations of motor unit firing behavior during muscle fatigue have been reported and explained with varieties of conjectures. The disagreement amongst previous studies has resulted, in part, from the limited number of available motor units and from the misleading practice of grouping motor unit data across different subjects, contractions, and force levels. To establish a more clear understanding of motor unit control during fatigue, we investigated the firing behavior of motor units from the vastus lateralis muscle of individual subjects during a fatigue protocol of repeated voluntary constant force isometric contractions. Surface electromyographic decomposition technology provided the firings of 1,890 motor unit firing trains. These data revealed that to sustain the contraction force as the muscle fatigued, the following occurred: 1) motor unit firing rates increased; 2) new motor units were recruited; and 3) motor unit recruitment thresholds decreased. Although the degree of these adaptations was subject specific, the behavior was consistent in all subjects. When we compared our empirical observations with those obtained from simulation, we found that the fatigue-induced changes in motor unit firing behavior can be explained by increasing excitation to the motoneuron pool that compensates for the fatigue-induced decrease in muscle force twitch reported in empirical studies. Yet, the fundamental motor unit control scheme remains invariant throughout the development of fatigue. These findings indicate that the central nervous system regulates motor unit firing behavior by adjusting the operating point of the excitation to the motoneuron pool to sustain the contraction force as the muscle fatigues.

Motor-unit behavior in humans during fatiguing arm movements

1996 ◽  
Vol 75 (4) ◽  
pp. 1629-1636 ◽  
Author(s):  
K. J. Miller ◽  
S. J. Garland ◽  
T. Ivanova ◽  
T. Ohtsuki
Keyword(s):  
Motor Unit ◽  
Force Production ◽  
Motor Units ◽  
Isometric Contractions ◽  
Triceps Brachii ◽  
Unit Discharge ◽  
Recruitment Threshold ◽  
Motor Unit Discharge ◽  
Discharge Behavior ◽  
The Mean

1. The activity of 40 triceps brachii motor units was recorded from the dominant arms of 9 healthy adult volunteers (age 27.8 +/- 4.4 yr, mean +/- SD) during a fatigue task that included both isometric and anisometric contractions. The fatigue task lasted 8.3 min and consisted of 50 extension and 50 flexion movements of the elbow. Each movement (40 degrees in 0.8s) was separated by an isometric contraction. A constant load resisting extension of 17.7 +/- 3.0% of maximal voluntary contractions (MVC) was applied throughout the task. This paradigm enabled the direct contrast of motor-unit discharge behavior during the different types of fatiguing contractions. 2. Motor-unit behavior was examined to determine the relative contribution of two mechanisms for optimizing force production under fatiguing conditions: recruitment of motor units and modulation of motor-unit discharge following recruitment. Threshold torques for motor-unit recruitment thresholds were determined by ramp-and-hold isometric contractions. Motor-unit discharge was evaluated during the fatigue task by contrasting the number of motor-unit potentials (spikes) per contraction for concentric eccentric, and isometric contractions. 3. The fatigue task resulted in a 30 +/- 12% decline in the mean MVC of elbow extension. Recruitment of nine new motor units (23%) was evident during the fatiguing extension movements, often within five to seven movements (i.e., within 25-35 s). Each newly recruited motor unit had the largest recruitment threshold torque in that experiment. 4. Analysis of the motor units that were active from the beginning of the fatigue task revealed that the mean number of motor-unit spikes per contraction increased, or remained constant as fatigue ensued, yet for the majority of motor units it increased or remained constant. None of the newly recruited motor units demonstrated decreased number of mean spikes per contraction after recruitment. Further, concurrently active motor units displayed different discharge behavior in two-thirds of the subjects. It is proposed that if the neural drive to the muscle is distributed uniformly upon the motoneuron pool, peripheral feedback from the exercising muscle may modulate specific motoneuron discharge levels during fatigue.

scholarly journals Contribution from motor unit firing adaptations and muscle coactivation during fatigue

2018 ◽  
Vol 119 (6) ◽  
pp. 2186-2193 ◽  
Author(s):  
Paola Contessa ◽  
John Letizi ◽  
Gianluca De Luca ◽  
Joshua C. Kline
Keyword(s):  
Motor Unit ◽  
Late Phase ◽  
Motor Units ◽  
Force Output ◽  
Firing Behavior ◽  
Firing Rates ◽  
Pronator Teres ◽  
Motor Unit Firing ◽  
The Central Nervous System ◽  
Extensor Carpi Radialis

The control of motor unit firing behavior during fatigue is still debated in the literature. Most studies agree that the central nervous system increases the excitation to the motoneuron pool to compensate for decreased force contributions of individual motor units and sustain muscle force output during fatigue. However, some studies claim that motor units may decrease their firing rates despite increased excitation, contradicting the direct relationship between firing rates and excitation that governs the voluntary control of motor units. To investigate whether the control of motor units in fact changes with fatigue, we measured motor unit firing behavior during repeated contractions of the first dorsal interosseous (FDI) muscle while concurrently monitoring the activation of surrounding muscles, including the flexor carpi radialis, extensor carpi radialis, and pronator teres. Across all subjects, we observed an overall increase in FDI activation and motor unit firing rates by the end of the fatigue task. However, in some subjects we observed increases in FDI activation and motor unit firing rates only during the initial phase of the fatigue task, followed by subsequent decreases during the late phase of the fatigue task while the coactivation of surrounding muscles increased. These findings indicate that the strategy for sustaining force output may occasionally change, leading to increases in the relative activation of surrounding muscles while the excitation to the fatiguing muscle decreases. Importantly, irrespective of changes in the strategy for sustaining force output, the control properties regulating motor unit firing behavior remain unchanged during fatigue. NEW & NOTEWORTHY This work addresses sources of debate surrounding the manner in which motor unit firing behavior is controlled during fatigue. We found that decreases in the motor unit firing rates of the fatiguing muscle may occasionally be observed when the contribution of coactive muscles increases. Despite changes in the strategy employed to sustain the force output, the underlying control properties regulating motor unit firing behavior remain unchanged during muscle fatigue.

scholarly journals Respiration-related discharge of hyoglossus muscle motor units in the rat

2014 ◽  
Vol 111 (2) ◽  
pp. 361-368 ◽  
Author(s):  
Gregory L. Powell ◽  
Amber Rice ◽  
Seres J. Bennett-Cross ◽  
Ralph F. Fregosi
Keyword(s):  
Motor Unit ◽  
Muscle Force ◽  
Critical Role ◽  
Motor Units ◽  
Respiratory Cycle ◽  
Firing Rates ◽  
Rate Modulation ◽  
Inspiratory Phase ◽  
Anesthetized Rats ◽  
Motor Unit Firing

Although respiratory muscle motor units have been studied during natural breathing, simultaneous measures of muscle force have never been obtained. Tongue retractor muscles, such as the hyoglossus (HG), play an important role in swallowing, licking, chewing, breathing, and, in humans, speech. The HG is phasically recruited during the inspiratory phase of the respiratory cycle. Moreover, in urethane anesthetized rats the drive to the HG waxes and wanes spontaneously, providing a unique opportunity to study motor unit firing patterns as the muscle is driven naturally by the central pattern generator for breathing. We recorded tongue retraction force, the whole HG muscle EMG and the activity of 38 HG motor units in spontaneously breathing anesthetized rats under low-force and high-force conditions. Activity in all cases was confined to the inspiratory phase of the respiratory cycle. Changes in the EMG were correlated significantly with corresponding changes in force, with the change in EMG able to predict 53–68% of the force variation. Mean and peak motor unit firing rates were greater under high-force conditions, although the magnitude of discharge rate modulation varied widely across the population. Changes in mean and peak firing rates were significantly correlated with the corresponding changes in force, but the correlations were weak ( r2 = 0.27 and 0.25, respectively). These data indicate that, during spontaneous breathing, recruitment of HG motor units plays a critical role in the control of muscle force, with firing rate modulation playing an important but lesser role.

Does Raising Morning Rectal Temperature to Evening Levels Offset the Diurnal Variation in Muscle Force Production?

2013 ◽  
Vol 30 (4) ◽  
pp. 486-501 ◽  
Author(s):  
Ben J. Edwards ◽  
Samuel A. Pullinger ◽  
Jonathan W. Kerry ◽  
William R. Robinson ◽  
Tom P. Reilly ◽  
...  
Keyword(s):  
Diurnal Variation ◽  
Rectal Temperature ◽  
Muscle Force ◽  
Force Production ◽  
Muscle Force Production

scholarly journals Stroke increases ischemia-related decreases in motor unit discharge rates

2018 ◽  
Vol 120 (6) ◽  
pp. 3246-3256 ◽  
Author(s):  
Spencer A. Murphy ◽  
Francesco Negro ◽  
Dario Farina ◽  
Tanya Onushko ◽  
Matthew Durand ◽  
...  
Keyword(s):  
Motor Unit ◽  
Time Constant ◽  
Motor Units ◽  
Unit Discharge ◽  
Firing Rates ◽  
Sensory Pathways ◽  
Discharge Rates ◽  
Motor Unit Firing ◽  
Motor Unit Discharge ◽  
Density Surface

Following stroke, hyperexcitable sensory pathways, such as the group III/IV afferents that are sensitive to ischemia, may inhibit paretic motor neurons during exercise. We quantified the effects of whole leg ischemia on paretic vastus lateralis motor unit firing rates during submaximal isometric contractions. Ten chronic stroke survivors (>1 yr poststroke) and 10 controls participated. During conditions of whole leg occlusion, the discharge timings of motor units were identified from decomposition of high-density surface electromyography signals during repeated submaximal knee extensor contractions. Quadriceps resting twitch responses and near-infrared spectroscopy measurements of oxygen saturation as an indirect measure of blood flow were made. There was a greater decrease in paretic motor unit discharge rates during the occlusion compared with the controls (average decrease for stroke and controls, 12.3 ± 10.0% and 0.1 ± 12.4%, respectively; P < 0.001). The motor unit recruitment thresholds did not change with the occlusion (stroke: without occlusion, 11.68 ± 5.83%MVC vs. with occlusion, 11.11 ± 5.26%MVC; control: 11.87 ± 5.63 vs. 11.28 ± 5.29%MVC). Resting twitch amplitudes declined similarly for both groups in response to whole leg occlusion (stroke: 29.16 ± 6.88 vs. 25.75 ± 6.78 Nm; control: 38.80 ± 13.23 vs 30.14 ± 9.64 Nm). Controls had a greater exponential decline (lower time constant) in oxygen saturation compared with the stroke group (stroke time constant, 22.90 ± 10.26 min vs. control time constant, 5.46 ± 4.09 min; P < 0.001). Ischemia of the muscle resulted in greater neural inhibition of paretic motor units compared with controls and may contribute to deficient muscle activation poststroke. NEW & NOTEWORTHY Hyperexcitable inhibitory sensory pathways sensitive to ischemia may play a role in deficient motor unit activation post stroke. Using high-density surface electromyography recordings to detect motor unit firing instances, we show that ischemia of the exercising muscle results in greater inhibition of paretic motor unit firing rates compared with controls. These findings are impactful to neurophysiologists and clinicians because they implicate a novel mechanism of force-generating impairment poststroke that likely exacerbates baseline weakness.

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