The factors limiting force production and exercise endurance time have been briefly described, together with some of the changes occurring at various sites within the muscle and central nervous system. Evidence is presented that, in fatigue of sustained maximal voluntary contractions (MVC) executed by well-motivated subjects, the reduction in force generating capacity need not be due to a decline in central nervous system (CNS) motor drive or to failing neuromuscular transmission, but can be attributed solely to contractile failure of the muscles involved. However, despite this conclusion, both the integrated electromyogram (EMG) and the mean firing rate of individual motor units do decline progressively during sustained MVC. This, however, does not necessarily result in loss of force since the parallel slowing of muscle contractile speed reduces tetanic fusion frequency. It is suggested that the range of motoneuron firing rates elicited by voluntary effort is regulated and limited for each muscle to the minimum required for maximum force generation, thus preventing neuromuscular transmission failure and optimizing motor control. Such a CNS regulating mechanism would probably require some reflex feedback from the muscle.

SUMMARY1. During fatigue from a sustained maximal voluntary contraction (m.v.c.) the mean motoneurone discharge rates decline. In the present experiments we found no recovery of firing rates after 3 min of rest if the fatigued muscle was kept ischaemic, but near full recovery 3 min after the blood supply was restored.2. Since 3 min is thus sufficient time for recovery of any central changes in excitability, the results support the hypothesis that, during fatigue, motoneurone firing rates may be regulated by a peripheral reflex originating in response to fatigue-induced changes within the muscle.

Central and peripheral factors were studied in fatigue of submaximal intermittent isometric contractions of the human quadriceps and soleus muscles. Subjects made repeated 6 s, 50% maximal voluntary contractions (MVC) followed by 4 s rest until the limit of endurance (Tlim). Periodically, a fatigue test was performed. This included a brief MVC, either a single shock or 8 pulses at 50 Hz during a rest period and a shock superimposed on a target force voluntary contraction. At Tlim, the MVC force had declined by 50%, usually in parallel with the force from stimulation at 50 Hz. The twitches superimposed on the target forces declined more rapidly, disappearing entirely at Tlim. In similar experiments on adductor pollicis, no reduction of the evoked M wave was seen. The results suggest that, during fatigue of quadriceps and adductor pollicis induced by this protocol, no central fatigue was apparent, but some was seen in soleus. Thus the reduced force-generating capacity could result mainly or entirely from failure of the muscle contractile apparatus.

SUMMARY1. Tungsten micro-electrodes have been used to record the electrical activity of single motor units in the human adductor pollicis during maximal voluntary contractions. The potentials were characteristic of those from single muscle fibres.2. In brief maximal contractions, the firing rates of over 200 motor units were obtained from five normal subjects. Four subjects had a similar range (mean 26-4 + 6-5 Hz) while the fifth was slightly higher (35 + 7 4 Hz).3. When maximal voluntary force was sustained for 40-120 s, there was a progressive decline in the range and mean rate of motor-unit discharge. In the first 60 s, mean rates fell from about 27 Hz to 15 Hz. There was some evidence to suggest that those units with the highest initial frequencies changed rate most rapidly.4. It is suggested that this decline in motor unit discharge rates is not responsible for force loss, but that it may enable effective modulation of voluntary strength by rate coding to continue during fatigue.

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Measurements were made from the human adductor pollicis muscle of force, contractile speed, and electromyographic activity (EMG) before, during, and after maximal isometric voluntary contractions sustained for 60 s. The use of brief test periods of maximal nerve stimulation with single shocks or trains of shocks enabled various muscle mechanical properties to be studied throughout each contraction. Electrical activity was measured after rectification and smoothing of the surface potentials and also by counting the total number of potentials per unit time from a population of motor units using fine wire intramuscular electrodes. During a 60-s maximal voluntary contraction, the force fell by 30-50%. Throughout the experiment the voluntary force matched that produced by supramaximal tetanic nerve stimulation. This indicated that, with sufficient practice, full muscle activation could be maintained by voluntary effort. However, the amplitude of the smoothed, rectifed EMG and the rate of spike counts declined. Since no evidence for neuromuscular block was found, the decline in EMG and spike counts was attributed to a progressive reduction of the neural drive from the central nervous system, despite maintained maximum effort. After the prolonged voluntary contractions twitch duration was prolonged, mainly as a result of slowing in relaxation rate. Twitch summation in unfused tetani increased. Both the maximum rate of relaxation and the time course of force decay declined by 50-70%. Similar changes were seen in both voluntary contractions and in test periods of stimulation. The percentage change in muscle contractile speed measured by these parameters approximately equaled the percentage change in the surface EMG measured simultaneously. It is concluded that 1) during a 60-s sustained maximal voluntary contraction there is a progressive slowing of contraction speed such that the excitation rate required to give maximal force generation is reduced, 2) the simultaneous decline in EMG may be due to a continuous reduction in motoneuron discharge rate, and 3) the EMG decline may not necessarily contribute to force loss.

The relationship between the electromyographic (EMG) power spectrum and muscle conduction velocity was investigated during both fatiguing and nonfatiguing contractions of the adductor pollicis muscle. Changes in the EMG power spectrum were measured by Fourier transform analysis and by comparing the power in the high (130-238 Hz) and low (20--40 Hz) frequency bands. Changes in conduction velocity were measured during voluntary activity from changes in the muscle mass action potential evoked by periodic maximal shocks to the nerve. This was varied independently either by maintaining a 60-s fatiguing maximal voluntary contraction involving 30--50% loss of force or by changing muscle temperature in the absence of fatigue. Both procedures resulted in similar changes in the power spectrum. However, the change in conduction velocity required to generate equal changes in the EMG was about 10 times greater in the absence of fatigue than those observed during a 60-s maximum contraction initiated at any initial muscle temperature. This suggests that during fatigue of maximal voluntary contractions, factors other than changes in the wave form of individual muscle fiber action potentials must contribute to the observed shift in the total surface EMG frequency components.

Single motor-unit firing rates have been recorded during maximal voluntary contractions using tungsten microelectrodes. Over 300 units from four subjects were sampled from each of three muscles. These were the biceps brachii, adductor pollicis, and soleus, chosen because of known differences in their fiber-type composition and contractile properties. In all cases the contraction maximality was assured by delivering single supramaximal shocks during the voluntary contractions. All motor units were deemed to have already been fully activated if no additional force resulted. Thus for each muscle, the firing rates elicited by a maximal voluntary effort are sufficient to generate a fully fused tetanus in each motor unit. For the biceps brachii and adductor pollicis muscles, the mean firing rates (+/- SD) were 31.1 +/- 10.1 and 29.9 +/- 8.6 Hz, respectively, while for soleus they were only 10.7 +/- 2.9 HZ. For each muscle the firing rates distribution covered approximately a four-fold range about the mean value. The mean firing rates for each muscle varied roughly in proportion to their respective twitch contraction and half relaxation times. These contractile time measurements for both biceps brachii and adductor pollicis agreed well with the mean values reported for human fast-twitch motor units, while those for soleus fell in the range observed for human slow-twitch units. An argument is presented that suggests that, in response to voluntary effort, the range of discharge rates of each motor-unit pool is limited to those only just sufficient to produce maximum force in each motor unit. This suggestion is based on the relationship between the range of motor-unit firing frequencies observed during maximum voluntary contractions, their range of contraction times, and the stimulation frequencies required for maximum force generation. The implications of this hypothesis for motor control are discussed.