The Lessening of Neuro-muscular Irritability by Fatigue
The threshold for the peroneus communis nerve in decerebrate animals varied from 0.319 to 2.96 units, with an average in sixteen experiments of 1.179.[*] This average is the same as that found by E. L. Porter[2] for the radial nerve in the spinal cat. For animals under urethane anesthesia a higher average was obtained. In these it varied from .644 to 7.05, or an average in ten experiments of 3.081.
[*] For the detailed data of these and other quantitative experiments, the reader should consult the tables in the original papers.
The threshold for the tibialis anticus muscle varied in the decerebrate animals from 6.75 units to 33.07, or an average in fifteen experiments of 18.8. Ten experiments were performed under urethane anesthesia and the threshold varied from 12.53 to 54.9, with an average of 29.84 β units. From these results it is evident that anesthesia notably affects the threshold.
E. L. Porter proved, by experiments carried on in the Harvard Physiological Laboratory, that the threshold of an undisturbed nerve-muscle remains constant for hours, and his observation was confirmed by Gruber (see [Fig. 19]). If, therefore, after fatigue, a change exists in the threshold, this change is necessarily the result of alterations set up by the fatigue process in the nerve-muscle or muscle.
After fatigue the threshold of the nerve-muscle, in sixteen decerebrate animals, increased from an average of 1.179 to 3.34—an increase of 183 per cent. In ten animals under urethane anesthesia the threshold after fatigue increased from a normal average of 3.08 to 9.408—an increase of 208 per cent.
An equal increase in the threshold stimulus was obtained from the normal muscle directly. In decerebrate animals the normal threshold of 18.8 units was increased by fatigue to 69.54, or an increase of 274 per cent. With urethane anesthesia the threshold increased from 29.849 to 66.238, or an increase of 122 per cent.
[Fig. 18], plotted from the data of one of the many experiments, shows the relative heights of the threshold before and after fatigue. The correspondence of the two readings of the threshold, one from the nerve supplying the muscle and the other from the muscle directly, served as a check on the electrodes. The broken line in the figure represents the threshold (in units) of the nerve-muscle, and the continuous line that of the muscle. The threshold values of the nerve-muscle have been magnified ten times in order to bring the two records close together. In this experiment the threshold of the muscle after fatigue (i. e., at 2) is 167 per cent higher than the normal threshold (at 1), while that of the nerve-muscle after fatigue is 30.5 per cent higher than its normal.
Figure 18.—A record plotted from the data of one experiment. The time intervals in minutes are registered on the abscissa; the value of the threshold in units is registered on the ordinate. The continuous line is the record of the muscle, the broken line that of the nerve-muscle. The values for the nerve-muscle have been magnified ten times, those for the muscle are normal.
(1) Normal values of the threshold.
(2) Fatigue thresholds after one hour’s work, lifting 120 grams 240 times a minute.
(3 and 4) The threshold after rest.
Evidently a direct relation exists between the duration of work and the increase of threshold. For instance, the threshold is higher after a muscle is fatigued for two hours than it is at the end of the first hour. The relation between the work done and the threshold is not so clear. In some animals the thresholds were higher after 120 grams had been lifted 120 times a minute for 30 minutes than they were in others in which 200 grams had been lifted 240 times a minute for the same period. The muscle in the latter instances did almost four times as much work, yet the threshold was lower. The difference may be due to the general condition of the animal.
A few experiments were performed on animals in which the nerve supplying the muscle was cut seven to fourteen days previous to the experiment. The muscle, therefore, had within it no living nerve fibres. The average normal threshold for the denervated muscle in 6 animals was 61.28 units. As in the normal muscle, the percentage increase due to fatigue was large.