The contradiction between the theories of the peripheral and of the central origin of the dicrotic, however, is only apparent, and neither may be true, because it might be that this elevation is not due to a wave which travels in the blood. The experiments of the previous investigators seem to point in this direction. The disappearance of the secondary elevations when the arterial wall has lost the properties of an elastic body, the above-mentioned experiments of v. Kries, and the observations of Grashey and Marey on the movements of the walls of an elastic tube indicate clearly that nothing but elasticity is needed to produce these secondary or dicrotic elevations, for, in the different experiments, they are produced as well when the heart and its valves are replaced by a valveless bag as when the function of the valves is unimpaired; as well with resistance at the periphery as without, the only condition being that the walls are elastic. This proves the importance of the elasticity of the arterial wall. The experiments of the graphic registration of the movements of the walls of an elastic tube, furthermore, indicate that the conditions of this experiment are a close imitation of the mechanical conditions which prevail in the arteries. It may be expected that the analysis of the conditions of the experiment will give an insight into the origin of the sphygmographic curves, because the tracings which Grashey and Marey took from the walls of a rubber tube resemble closely the tracings of the human pulse. This experiment, first, proves that the form of the curve depends merely on physical conditions. The movement of a point of the wall of the tube depends on the following four factors: (1) The elasticity of the wall; (2) the incompressibility of the liquid; (3) the form of the original wave, i. e., the way in which the liquid is pumped into the tube; (4) the rate of outflow. If the process of pressing liquid into the tube is repeated regularly, a stationary form of movement will be obtained eventually; the amount of outflow for one interval is constant in this case. This means that eventually a state is attained where the same quantity of liquid which is pumped into the tube at one end flows out from the tube at the other. The physiological bearing of this result is that the turgor of an artery does not change without a cause. Such a change would be indicated by the going up or down of the base-line of the tracing.
The first two factors are, in physiology, studied with relative ease. The elastic qualities of the arteries have been studied since Poiseuille and John Hunter by Wertheim, Zwardemaaker, Marey, and others, and they are more or less well known. The physical properties of the blood are very nearly those of an incompressible liquid, and this is certainly true for the small pressure to which the blood is exposed in the arteries.
As to the initial form of the wave which the action of the left ventricle produces in the arterial system, we get a hint from the experiments of Grashey, v. Kries, and Marey, where the sudden compression of a bag furnished the initial shock.[64] These changes of pressure can be represented by a curve like that in Fig. 1.
So long as the contraction of the left ventricle lasts and the valves are open, the action of the heart produces a certain pressure in the aorta, but the influence of the intraventricular pressure is zero when the valves are closed. The second phase of the curve Fig. 1, where the pressure is zero, certainly gives the influence of the intraventricular pressure during the diastole, because there is no communication between the ventricle and the arterial system when the valves are closed. The question is whether the rest of the curve can represent the changes of the intraventricular pressure when the valves are open.
Fig. 1. Changes of pressure produced in a bag by sudden compression.
Fig 2. Decreasing amount of liquid in a tube when the outflow is uniform.
The first curves of intraventricular pressure were traced by Chauveau and Marey. These experiments were made on a horse, and they have been repeated since it was discovered that they can be performed also on smaller animals. Besides Chauveau and Marey may be mentioned the names of Fick, Huerthle, v. Frey, Rolleston, Bayliss and Starling. The curves obtained by various observers belong to two types; one shows the so called "plateau," the other does not. Recent experiments have proved that this difference of results is due to a difference in methods. This also is suggested by the fact that different curves have been obtained from animals of the same species. Two methods have been applied lately for testing these curves of intraventricular pressure. The first was devised by Bayliss and Starling. It consisted chiefly in the photographic registration of the movement of the liquid in a manometer tube. The photographic registration is frictionless, and the mass of the moving liquid was so small that vibrations by inertia were fairly excluded for pressures which are not greater than the intraventricular pressure.[65] The second method was used by Porter. The idea of this method was to trace only a part of the curve, not the whole. The writing lever, thus, has in the beginning of the tracing no inertia at all, and the tracing may be overdrawn but is certainly correct in form up to the next point of inflexion of the curve.[66] These tests and the repeated experiments of Chauveau leave no doubt as to the existence of the plateau.
The varying pressure from the heart which produces the pulse wave may be described in this way: The pressure suddenly rises to a maximum and maintains it for a certain time; when the semilunar valves close, the pressure drops as suddenly as it rose, and remains at zero until the valves open again. Such a function can be represented by a curve like Fig. 1, and this is the reason why the complicated action of the heart can be superseded by the compression of a bag without changing the mechanical conditions of the problem. Of course it can not be expected that a schematic curve will show all the details of the real tracing. It is suggested, however, by Frank[67] that many of the small irregularities of the curve of intraventricular pressure are due to vibrations caused by the inertia of the apparatus and that the true form of the curve of intraventricular pressure is very simple. This remark is supported by Huerthle,[68] who tested the apparatus of Marey, Knoll and Grunmach. Marey's tambour was found to be the most exact, but even this instrument produces deformities in the tracings, though the general outlines are exact. This would indicate that the schematic representation of Fig. 1 is a very close imitation of the real form of the curve of intraventricular pressure, although empirical tracings do not show right angles and straight lines. It seems, however, that the undulations of the plateau are genuine, since they are found in the most reliable tracings, and it may be possible to explain them merely on the basis of the physical conditions of the experiment.