98 THE PHYSIOLOGY OF MUSCLE AND NERVE. 



pool. A similar phenomenon occurs in muscle fibers when stimu- 

 lated, but the negative condition travels over the muscle fiber at a 

 slower speed, 3 to 4 meters per second in frog's muscle, and with a 

 wave length, according to Bernstein, of only 10 mms. This wave 

 of negativity or of excitation in the muscle precedes the actual 

 wave of contraction. 



This phenomenon of a negative electrical condition traveling 

 over the nerve or muscle and giving us an active current when led 

 off through a galvanometer is of the greatest physiological impor- 

 tance, particularly in the study of nerves. It has been shown 

 that in the nerve this wave of negativity marks the progress of the 

 wave of excitation, and, since we can study its progress by means 

 of the galvanometer or capillary electrometer, we can thus study the 

 excitability and conductivity in nerves when removed from con- 

 nection with their end-organs. That the negative wave, or the 

 action current that it gives rise to, is an invariable sign of the 

 passage of an excitation or nerve impulse is shown by the facts 

 that it is absent in the dead nerve, and that in the living nerve it is 

 produced by mechanical,* chemical,! and reflex t stimulations, as 

 well as by the more usual method of electrical stimulation. 



Herzen has claimed that under certain conditions of local narcosis the 

 nerve fibers when stimulated may give an action current, but no muscle con- 

 traction, a fact which if true would seem to show that the excitation wave 

 or nerve impulse and the wave of negative potential are not associated 

 invariably. This result, however, has been denied by other competent 

 observers (Wedenski, Boruttau). 



Monophasic and Diphasic Action Currents. According to 

 the conception of the action current given above, it is evident that 

 it should be obtained upon stimulation when a living normal nerve, 

 is connected at any two points of its course with a galvanometer or 

 capillary electrometer. The detection of the current under such 

 conditions offers more difficulties, because it is diphasic, as will 

 be seen from the accompanying diagram (Fig. 43). The figure rep- 

 resents a normal nerve led off to the galvanometer from two points, 

 b and c, of its longitudinal surface. As these points in the uninjured 

 nerve have the same potential, no current is shown by the galvanom- 

 eter. If the nerve is stimulated at a by a single stimulus a neg- 

 ative condition or charge passes along the nerve. When it reaches 

 the point b there will be a momentary current through the gal- 

 vanometer from c to b; as the charge passes on to c this point in 

 turn will become negative to b, and there will be a momentary cur- 



* Steinach, "Pfluger's Archiv," 55, 487, 1894. 



tGriitzner, "Pfluger's Archiv," 25, 255, 1881. 



j Boruttau, " Pfluger's Archiv," 84 and 90, 1901-1902. 



