Properties Of Human Motor Units Research Articles

Inappropriate interpretation of surface EMG signals and muscle fiber characteristics impedes understanding of the control of neuromuscular function

as the final common pathway from the nervous system to muscle, the motor unit transmits an activation signal generated by the nervous system to engage the contractile proteins and produce the muscle forces needed for reflex responses, automatic behaviors, and voluntary actions ([11][1]). The net

Limitations of the Spike-Triggered Averaging for Estimating Motor Unit Twitch Force: A Theoretical Analysis

Contractile properties of human motor units provide information on the force capacity and fatigability of muscles. The spike-triggered averaging technique (STA) is a conventional method used to estimate the twitch waveform of single motor units in vivo by averaging the joint force signal. Several limitations of this technique have been previously discussed in an empirical way, using simulated and experimental data. In this study, we provide a theoretical analysis of this technique in the frequency domain and describe its intrinsic limitations. By analyzing the analytical expression of STA, first we show that a certain degree of correlation between the motor unit activities prevents an accurate estimation of the twitch force, even from relatively long recordings. Second, we show that the quality of the twitch estimates by STA is highly related to the relative variability of the inter-spike intervals of motor unit action potentials. Interestingly, if this variability is extremely high, correct estimates could be obtained even for high discharge rates. However, for physiological inter-spike interval variability and discharge rate, the technique performs with relatively low estimation accuracy and high estimation variance. Finally, we show that the selection of the triggers that are most distant from the previous and next, which is often suggested, is not an effective way for improving STA estimates and in some cases can even be detrimental. These results show the intrinsic limitations of the STA technique and provide a theoretical framework for the design of new methods for the measurement of motor unit force twitch.

Commentaries on Viewpoint: Fatigue mechanisms determining exercise performance: Integrative physiology is systems physiology

Commentaries on Viewpoint: Fatigue mechanisms determining exercise performance: Integrative physiology is systems physiology

Properties of human motor units after prolonged activity at a constant firing rate

The primary purpose of this study was to examine if there are changes in the intrinsic properties of spinal motoneurons after prolonged submaximal contractions. To do this, we assessed whether or not the synaptic drive to motoneurons needs to increase in order to maintain a constant firing rate of a motor unit. Recruitment of new units and an increase in total electromyographic (EMG) activity of the muscle of interest were taken as estimates of an increase in synaptic drive. Subjects were asked to maintain a constant firing rate of a clearly identifiable (targeted) motor unit from the first dorsal interosseous muscle for approximately 10 min, while surface EMG and force were recorded simultaneously. For the 60 units studied, the duration of the constant-firing-rate period ranged from 73 to 1,140 s (448 +/- 227 s; mean +/- SD). There was a significant increase ( t-test, p<0.001) in the magnitude of mean surface EMG, and DC force while the targeted motoneuron maintained a constant rate suggesting an increase in the net excitatory input to the motoneuron pool. Changes occurring simultaneously in other parameters, namely, variability in interspike interval, magnitude of force fluctuations, the duration of motor unit action potentials, and the median power frequency of surface EMG were also computed. The firing rates of 16 concurrently firing motoneurons, not controlled by the subject, remained constant. The key finding of this study is that after prolonged activity, a motoneuron requires a stronger excitatory input to maintain its firing rate. Additional results are indicative of significant changes in the characteristics of the synaptic inputs, changes at the neuromuscular junction (both pre- and postsynaptic regions) and the sarcolemma.

Contractile properties of human motor units in health, aging, and disease.

The primary function of skeletal muscle is to produce force for postural control and movement. Although the contractile properties of the whole muscle are useful functional indicators, they do not accurately reflect the heterogeneity of the constituent motor units (MUs) and their changes in health and disease. However, data on the contractile properties of human MUs, in comparison to other animal species, are relatively sparse. This, in part, is due to greater methodological challenges of in vivo studies of MUs in the human. The purpose of this review is to critically appraise the methods used in humans; to describe the normative data from different muscle groups; to discuss differences between data from healthy humans and other animal species; and, last, to characterize changes of the MU contractile properties in aging, disease, and in response to intervention. Because the spike-triggered averaging technique can only be used to study the twitch properties, other methods were subsequently developed to measure a wider range of contractile properties. Although there is general agreement between human data and those from other animal species, major differences do exist. Potential reasons for these discrepancies include true biological differences, but differences in the techniques used may also be responsible. Although limited, measurement of MU contractile properties in humans has provided insight into the changes associated with aging and motoneuronal diseases and provides a means of gauging their adaptive capacity for training and immobilization.

Discharge properties of human motor units during sustained contraction at low level force

The characteristic of discharge behaviors of motor units (MUs) during low level contraction was investigated. The discharge of MUs in the m. vastus medialis was observed during the sustained contraction at 4 different levels below 10% MVC (2, 4, 8 and 10% MVC) for 15 min. The spike interval of all observed MUs gradually elongated during an initial several minutes of the contraction and the characteristic discharge patterns following the elongation were observed. i.e. continuous discharge throughout the contraction (CONT), decruitment (D-N), and re-recruitment following decruitment (D-REC). The relationship between recruitment threshold force ( F th) and discharge pattern was not significant at 2% MVC but, at 10% MVC, there were significant differences in F th between D-N and CONT, and between D-REC and CONT MU populations. In pooled data, the MUs with the shorter mean spike interval at the beginning of the contraction (MSI 0, below 90 ms) tend to discharge continuously, but the MUs with longer MSI 0 showed various discharge patterns. In conclusion, during low level contraction MUs discharge characteristically, and the MU with high excitation levels tend to discharge continuously, but individual MU represents an intrinsic discharge pattern at not a high excitation level.

Effects of Short-Term Training on Physiologic Properties of Human Motor Units

The human neuromuscular system possesses a remarkable ability to adapt to any change in the demands placed upon it. Adaptation to training or disuse is reflected in the activation patterns and physiologic properties of the motor unit (MU) pool in a given muscle group. This review summarizes the studies that have examined such adaptation at the level of the single MU. Disuse, as revealed by electrophysiologic studies, results in impaired MU recruitment and smaller twitch tensions in low and high threshold MUs. Alternatively, shortterm training improves MU recruitment and generally results in larger MU twitch tensions. A method has been developed to physiologically characterize and longitudinally follow single thenar MUs. Studies utilizing these methods have demonstrated that MUs differing in their baseline physiologic characteristics respond very differently to identical short-term training programs. These observations at the single MU level provide insight into training adaptations in whole muscles and muscle groups.

Effects of aging on motor-unit control properties.

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.

Contractile Properties of Human Motor Units: Is Man a Gat?

A major goal in neuroscience is to understand how the CNS controls posture and movement in humans. This requires an understanding of individual human motor unit properties and how they interact within the muscle to perform different tasks. This article describes differences and similarities between the contractile properties of human motor units and those of the cat prototype medial gastrocnemius (MG) muscle, on which so many studies have been conducted. The article describes the methods available for measuring human motor unit properties and their limitations, and it discusses how far the behavior of whole muscles can be predicted from their histochemistry. It questions the extent to which human motor units conform to the conventional criteria by which S (slow, fatigue resistant), FR (fast but fatigue resistant) and FF (fast, fatigable) unit types are usually classified. An important difference between human and cat MG data is that weak human motor units are not necessarily slow, nor strong ones fast; that is, generally, human unit force is not correlated with contractile speed. Also, unlike cat MG, the few human muscles studied so far contain few if any FF units but a high proportion of units with intermediate fatigue resistance (Flnt). These apparently aberrant human properties, however, are also found in other cat and rat muscles. Thus, cat MG may not be the best model for motor unit behavior generally. Finally, the influence of human motor unit properties on force output by recruitment and/or rate coding is discussed.

Computational methods for improving estimates of motor unit twitch contraction properties

Estimates of mechanical properties of human motor units have usually been made indirectly, using the technique of "spike-triggered averaging" (STA). In this method, a single motor unit action potential is used to synchronize the accumulation of an ensemble average of correlated force transients. However, under most realizable conditions, these transients are recorded during periods of sustained motor unit discharge, in which each motor unit is producing a partially fused tetanus. Therefore, the STA technique extracts the characteristics of the unfused force transient, instead of the desired single motor unit twitch. Although the STA method has been widely used, there is as yet no well-established relation between the force transient in the unfused tetanus, and the twitch contraction properties of the motor unit. To evaluate the accuracy of the STA as a measure of the motor unit mechanical properties, we applied two types of muscle models to the force transients recorded in an unfused tetanus, using data derived from experiments in which the response to a single twitch was also recorded. Our objective was to see whether accurate predictions of single motor unit mechanical characteristics are possible, working backward from the STA. The models chosen for this task were a linear second order model, and the distribution-moment (DM) model. These model predictions were then compared with the STA response, and with the twitch properties of the individual motor units. We also evaluated the utility of extrapolating the initial slope of the STA backward to improve the accuracy of the mechanical estimates. The results of our simulation suggest that there is no straightforward relation between the characteristics of the unfused tetanus and the mechanical properties of the single twitch. Although our attempts to predict the properties of the single twitch from the STA were only partly successful, the results of the simulations were far more accurate than those derived from the STA alone. Because the errors in the use of the STA method were so substantial, we would urge that the STA technique be used with great caution as a measure of twitch contraction properties, unless accompanied by appropriate simulations of muscle mechanical behavior.

A comparison of human thenar motor-unit properties studied by intraneural motor-axon stimulation and spike-triggered averaging

1. Measurements of twitch contractile properties of human motor units recorded by spike-triggered averaging may be distorted by partial fusion between twitches, because motor units seldom fire at rates below 8-10 Hz. The effects of this fusion were examined by comparing the responses of 27 human thenar motor units when their motor axons were stimulated at 1, 8, and 10 Hz. 2. Resultant forces were calculated from the abduction and flexion force components, together with various contraction and relaxation rate indexes as reported previously. Values for single twitches were compared to measurements made from the unfused force fluctuations ("apparent twitches") of the same units recorded during 8 and 10 Hz stimulation. 3. For all units, stimulation at 8 and 10 Hz caused partial twitch fusion. At 10 Hz, mean values for "apparent twitch" forces, contraction times (CT), and one-half relaxation times (1/2RT) were reduced to 44, 76, and 52% of the corresponding values measured from separate twitches evoked by 1 Hz stimulation. Similar but smaller reductions were seen at 8 Hz. 4. Slow units, with initial twitch CT greater than 60 ms, showed significantly more distortion of all "apparent twitch" parameters when stimulated at both 8 and 10 Hz, compared to fast units (less than 50 ms). 5. The potentiated abduction force component data were compared with abduction forces obtained previously by spike-triggered averaging from the same muscle group. Mean force obtained by spike-triggered averaging ("STA twitch" force: 21 mN) was significantly larger than that measured in abduction in response to either 1 or 10 Hz motor-axon stimulation (14 mN, 6 mN, respectively).(ABSTRACT TRUNCATED AT 250 WORDS)

The effects of motor activity on the contractile properties of human motor units

The effects of motor activity on the contractile properties of human motor units

Rheobase, input resistance, and motor-unit type in medial gastrocnemius motoneurons in the cat.

Rheobase, input resistance, and motor-unit type in medial gastrocnemius motoneurons in the cat.

Motor units and fiber types of primary ankle extensors of the skunk (Mephitis mephitis)

ArticlesMotor units and fiber types of primary ankle extensors of the skunk (Mephitis mephitis)K. M. Van De Graaff, E. C. Frederick, R. G. Williamson, and G. E. Goslow JrK. M. Van De Graaff, E. C. Frederick, R. G. Williamson, and G. E. Goslow JrPublished Online:01 Nov 1977https://doi.org/10.1152/jn.1977.40.6.1424MoreSectionsPDF (1 MB)Download PDF ToolsExport citationAdd to favoritesGet permissionsTrack citations ShareShare onFacebookTwitterLinkedInWeChat Previous Back to Top Next Download PDF FiguresReferencesRelatedInformation Cited ByForce-Frequency and Fatigue Properties of Motor Units in Muscles That Control Digits of the Human HandAndrew J. Fuglevand, Vaughan G. Macefield, and Brenda Bigland-Ritchie1 April 1999 | Journal of Neurophysiology, Vol. 81, No. 4Contractile Properties of Human Motor Units: Is Man a Gat?1 July 1998 | The Neuroscientist, Vol. 4, No. 4Fiber type composition of the plantarflexors of giraffes (giraffa camelopardalis) at different postnatal stages of developmentComparative Biochemistry and Physiology Part A: Physiology, Vol. 91, No. 2Motor unit types: functional specializations in motor controlTrends in Neurosciences, Vol. 3, No. 11 More from this issue > Volume 40Issue 6November 1977Pages 1424-1431 Copyright & PermissionsCopyright © 1977 the American Physiological Societyhttps://doi.org/10.1152/jn.1977.40.6.1424PubMed925736History Published online 1 November 1977 Published in print 1 November 1977 Metrics

The mechanical properties of human motor units with special reference to their fatiguability and recruitment threshold

The recruitment thresholds, twitch tension, twitch contraction time and fatiguability of human motor units have been studied in the first dorsal interosseous of the hand. Units recruited at contraction strengths <50 g had relatively low twitch tensions (0.18–1.9 g), long contraction times (59–146 msec) and were non-fatiguable. Units recruited at higher contraction strengths (>200 g) had higher twitch tensions (15–26 g), faster contraction times (33–57 msec) and were highly fatiguable. It is concluded that during graded voluntary muscle contractions motor units are recruited in order of increasing contraction strength and diminishing fatigue resistance.

Contractile and electrical properties of human motor units in neuropathies and motor neurone disease.

The contractile and electrical properties of motor units in the first dorsal interosseous muscle of the hand have been studied in 26 patients with ulnar neuropathies and motor neurone disease (amyotrophic lateral sclerosis). Among patients with unilateral pressure or entrapment ulnar neuropathies, there was a tendency for the twitch tensions for single motor units to be smaller, while the surface EMG amplitudes were generally larger in the affected hands. Very large EMG amplitudes but normal size twitch tensions were observed among the motor neurone disease patients, indicating that, in general, motor units enlarged by sprouting are less efficient contractile units than units of normal physiological size.

The orderly recruitment of human motor units during voluntary isometric contractions.

1. The contractile properties of human motor units from the first dorsal interosseus muscle of the hand were studied during voluntary isometric contractions using recently developed techniques.2. The twitch tensions produced by motor units varied widely from about 0.1-10 g. The twitch tension of a motor unit varied nearly linearly as a function of the level of voluntary force at which it was recruited over the entire range of forces studied (0-2 kg).3. The number of additional motor units recruited during a given increment in force declined sharply at high levels of voluntary force. This suggests that even though the high threshold units generate more tension, the contribution of recruitment to increases in voluntary force declines at higher force levels.4. Contraction times for these motor units varied from 30 to 100 msec. Over 80% had contraction times less than 70 msec, and might be classed as fast twitch motor units. The larger motor units, which were recruited at higher threshold forces, tended to have shorter contraction times than the smaller units.

The contractile properties of human motor units during voluntary isometric contractions.

1. The electrical activity of single motor units has been recorded from the first dorsal interosseus muscle of normal human subjects during voluntary, isometric contractions, together with the force generated by the muscle.2. By averaging the force correlated with the impulses from a single motor unit, the contraction time and twitch tension generated by that motor unit could be measured. When the rate of discharge was limited, either voluntarily or by automatic selection of intervals for analysis, the time for the tension to decline to half its maximum value (half-relaxation time) could also be measured for some motor units.3. Under our experimental conditions the trains of impulses from different motor units in most subjects were generated quite independently as tested by (a) measuring the correlation between activity in single units and that in the whole muscle as recorded by the surface electromyogram (e.m.g.), (b) measuring the cross-correlations between pairs of single units and (c) comparing the tension generated by stimulating single motor units with the average tension correlated in time with voluntary activity of single units in the same location.4. In one normal subject evidence of synchronization between separate motor units was obtained. Cross-correlation studies suggested that the cause of the synchronization was the presence of substantial common excitation received by the various motor units in the muscle.5. The frequency response for the contractions of single motor units was well fitted by that for a linear, second-order system with nearly critical damping. However, when stimulation of a few motor units was superimposed on a voluntary contraction, underdamped (oscillatory) responses were seen which were probably of reflex origin.6. The significance of these results in relation to the normal postural tremor in hand muscles is discussed.