SUMMARY1. 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 041-10 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.

SUMMARY1. Motor units in the first dorsal interosseus muscle of normal human subjects were recorded by needle electrodes, together with the surface electromyogram (e.m.g.). The wave form contributed by each motor unit to the surface e.m.g. was determined by signal averaging.2. The peak-to-peak amplitude of the wave form contributed to the surface e.m.g. by a motor unit increased approximately as the square root of the threshold force at which the unit was recruited. The peak-to-peak duration of the wave form was independent of the threshold force.3. Large and small motor units are uniformly distributed throughout this muscle, and the muscle fibres making up a motor unit may be widely dispersed.4. The rectified surface e.m.g. was computed as a function of force, based on the sample of motor units recorded. The largest contribution of motor unit recruitment occurs at low force levels, while the contribution of increased firing rate becomes more important at higher force levels.5. Possible bases for the common experimental observation that the mean rectified surface e.m.g. varies linearly with the force generated by a muscle are discussed. E.m.g. potentials and contractile responses may both sum non-linearly at moderate to high force levels, but in such a way that the rectified surface e.m.g. is still approximately linearly related to the force produced by the muscle.

SUMMARY1. Human subjects generated approximately linearly increasing or decreasing voluntary, isometric contractions using the first dorsal interosseus muscle of the hand.2. Single motor units began firing at 8-4+ 13 impulses/sec (mean + S.D. of an observation) and increased their firing rate 1*4 + 0-6 impulses/ sec for each change of 100 g in voluntary force. These values were independent of the threshold force for recruiting motor units.3. At intermediate rates of increasing and decreasing voluntary force (one complete cycle every 10 sec) the firing rate of single motor units varied linearly with force over the entire range of forces studied. However, during slow increases in voluntary force, the firing rate tended to reach a plateau, while during rapid increases an initial train of impulses at a roughly constant rate was observed.4. The relative importance of recruitment and increased firing rate, as mechanisms for increasing the force of voluntary contraction, was determined. Only at low levels of force is recuitment the major mechanism. Increased firing rate becomes the more important mechanism at intermediate force levels and contributes the large majority of force if the entire physiological range is considered.

SUMMARY1. 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.

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Experiments were designed to evaluate the relative contribution of impulse propagation failure, high-energy phosphate depletion, lowered pH, and impaired excitation-contraction coupling to human muscle fatigue and recovery. 31P nuclear magnetic resonance spectroscopy measurements were made on adductor pollicis muscle, together with simultaneous measurements of M-wave, force, and rectified integrated EMG (RIEMG). During fatigue, maximum voluntary contraction force (MVC) fell by 90%, pH fell from 7.1 to 6.4, and phosphocreatine was almost totally depleted. Neuromuscular efficiency (NME = force/RIEMG) was reduced to 40% of control at the end of the fatiguing contraction, and the M wave was reduced in amplitude and prolonged in duration. Following exercise, the M wave returned to normal within 4 minutes. pH, high-energy phosphates, and MVC recovered within 20 minutes. By contrast, neuromuscular efficiency did not recover within 60 minutes. These findings indicate three different components of fatigue. The first is reflected by the altered M wave and indicates impaired muscle membrane excitation and impulse propagation. The second, associated with reduced MVC, correlates with the metabolic state of the muscle (PCr and pH). The third, indicated by reduced NME, is independent of changes in high-energy phosphates and pH and is probably due to impaired excitation-contraction coupling.

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In order to determine whether or not impulse propagation was impaired during muscle fatigue, evoked muscle compound potentials (MCP) and twitches were recorded, both before and after fatigue, from the first dorsal interosseus (FDI), adductor pollicis (AP), and anterior tibialis (AT) muscles following supramaximal ulnar and peroneal nerve stimulation, respectively. The muscles were fatigued by maintaining maximum voluntary isometric, index finger abduction, thumb adduction, or ankle dorsiflexion for 1-5 minutes. FDI was most markedly altered, with reduced MCP amplitude (mean 32%) and increased MCP duration (mean 47%) after only 1 minute. After fatigue of longer duration (3-5 minutes), there were corresponding reductions in both the MCP amplitudes and the twitch tensions recorded from both the FDI and ankle dorsiflexors. We conclude that (1) a reduction in both the level of excitation and impulse propagation velocity of muscle membranes occurs during muscle fatigue, and (2) the magnitude of this reduced membrane function and its contribution to the mechanisms underlying fatigue depend both on the duration and degree of fatigue, as well as on the intrinsic properties of the particular muscle.