Topics: Relation of muscle work to energy exchange. Views of Liebig. Experimental evidence. Relation of nitrogen excretion to muscle work. Significance of the respiratory quotient in determining nature of the material oxidized. Fats and carbohydrates as source of energy by muscles. Utilization of proteid as a source of energy. Formation of carbohydrate from proteid. Significance of proteid metabolism. Theories of Carl Voit. Morphotic proteid. Circulating proteid. General conception of proteid metabolism on the basis of Voit's theories. Pfüger'a views of proteid metabolism. Rapidity of elimination of food nitrogen. Methods by which nitrogen is split off from proteid. Theories of Folin. Significance of creatinin and of the percentage distribution of excreted nitrogen. Endogenous or tissue metabolism. Exogenous or intermediate metabolism. Needs of tbe body for proteid food possibly satisfied by quantity sufficient to meet tbe demands of tissue or endogenous metabolism. Bearings of Folin's views on current theories and general facta of proteid metabolism. Large proteid reserve and voluminous exogenous metabolism probably not needed. Importance of feeding experiments in determining the true value of different views.

AS we have already seen, every form of muscular activity begets an increase in the energy exchange of the body. Between the two extremes of absolute rest and excessive muscular exertion, we find differences of 2000 calories or more per day as representing the degree of chemical decomposition corresponding to the particular state of muscular activity. The work of the involuntary muscles, such as have to do with peristalsis, respiration, rythmical beat of the heart, eta., is a relatively constant factor, though naturally subject to some variation, as has been pointed out in other connections. External muscular activity, however, is the one factor above all others that modifies the rate of energy exchange. A little longer walk, a heavier load to carry, a steeper hill to climb, any increase great or small in the activity of the muscles of the body, means a corresponding increase in chemical decomposition, with increased output of the ordinary products of tissue oxidation. The material so consumed, or oxidized, must be made good to hold the body in equilibrium; the supplies drawn upon are to be replaced, if the tissues of the body are to be kept in a proper state of efficiency.

What is the nature of the material used up in connection with muscle work? As can readily be seen, this is an important question, for on its answer depends, in some measure at least, the character of the proper intake, or food, to be supplied in order to make good the loss. If the energy of mechanical work, the energy of muscle contraction, comes from the breaking down of proteid matter alone, then obviously excessive muscular work would need to be accompanied, or followed, by a generous supply of proteid food. If, on the other hand, external work means liberation of energy solely from non-nitrogenous materials, then it is equally clear that fats and carbohydrates are the proper foods to offset the drain incidental to vigorous muscular action.

The views of Liebig, briefly referred to in a previous chapter, held sway over physiologists for many years. His dictum that proteid foods were true plastic foods, entering into the structure of the tissues of the body, and that they alone were the real sources of muscular energy, met for a time with no opposition. It was not until the advent of a more critical spirit, accompanied by a fuller appreciation of the necessity of experimental evidence, that physiologists began to test with scientific accuracy the validity of the current views. It is worthy of note that long prior to this time, even before oxygen was discovered, the far-sighted and resourceful John Mayow, in his work with the various "spirits" of the body and their relation to respiration, etc., evolved the view that muscular power has its origin in the combustion of fat brought to the muscles by the blood and burned there by aid of a gas or "spirit" taken from the air by the lungs, and likewise carried to the muscles by the circulating blood. Considering the time when Mayow lived and the dearth of true scientific knowledge as we measure it to-day, his hypothesis was a wonderful forestalling of present views.

It is quite obvious that the views of Liebig, if true, admit of easy proof; since, if the energy of muscular power comes from the breaking down of proteid, there should be a certain parallelism between the output of nitrogen from the body and the amount of muscular work accomplished, everything else being equal. As stated in a previous chapter, such study of this question as was made soon disclosed the fact that the one element above all others that Beemed to influence the output of nitrogen was the intake of proteid food. Thus, the English investigators, Lawes and Gilbert, found by experimenting with animals that when the latter were kept under uniform conditions of muscular work, the amount of nitrogen excreted ran parallel with the intake of nitrogen. Further, in the early experiments of Voit, the results obtained clearly showed that variations in the amount of work performed were practically without influence on the excretion of nitrogenous waste products.

The experiment, however, that came as a death blow to the theories of Liebig was that of Pidr-tid. Wislicenus,1 who in 1865 made an ascent of the Faulhorn, 6500 feet high, using a diet wholly non-nitrogenous. From the nitrogen excreted they were able, of course, to calculate the amount of proteid oxidized in the body during the period of work, and found that the proteid consumed could not have furnished, at the most, more than one-half the energy required to lift the weights of their bodies to the top of the high peak. Further, they observed that neither during the work period, nor immediately after, was there any noticeable increase in the excretion of nitrogen. Obviously, as they state, the oxidation of proteid matter in the body cannot be the exclusive source of the energy of muscular contraction, since the measurable amount of external work performed in the ascent of the mountain was far greater than the equivalent of the energy capable of being furnished by the proteid actually burned. To which may be added the fact that considerable energy, not measurable in their experiment, must have been employed in the work of the involuntary muscles of the body; thus increasing by so much the difference between the muscular work actually accomplished and the available energy from proteid consumed. lb is true that minor criticisms regarding certain details of the experiment can be offered to-day, such as the fact that the men were, in a measure, in a state of "nitrogen starvation," etc., but these criticisms do not in any degree militate against the main thesis that the energy of muscular contraction does not come exclusively from the consumption or breaking down of proteid, either of food or tissue. Vigojjjug and even severe muscular work does not necessarily increase the decomposi-tinn of proteid material. Dogs made to run in large tread-mills, with the same diet as on resting days, were found to excrete practically no more nitrogen than during the days of rest. Occasionally, however, in some one experiment the output of nitrogen would show an increase over the output on resting days. Further, experiments made with horses led to essentially the same result, except that greater increase in the excretion of nitrogen was observed than with dogs. This increase in nitrogen output, however, as a concomitant of increased muscular activity, could be prevented by adding to the amount of carbohydrate food.

1 See Geeammelte Schriften von Adolf Fick. Ueber die Entitehung der Miukelkraft. Band 2, p. 86. Wurzburg, 1903.