Irritability: A Physiological Analysis of the General Effect of Stimuli in Living Substance

by Max Verworn

Contents: Only processes of excitation are conducted, not processes of depression. Conduction of excitation in its two extreme instances. Conduction in undifferentiated pseudopod protoplasm of rhizopoda. Conduction of excitation with decrement of intensity and rapidity. Conduction of excitation in the nerve. Rapidity of conduction of excitation without decrement. Relation between irritability and conductivity. Conduction of excitation with decrement of the nerve after artificial depression of irritability by narcosis. Theory of the decrementless conduction of the normal nerve. Proof of the validity of the “all or none law” in the medullated nerve. Theory of the process of the conductivity of excitation. Theory of core model (Kernleiter). Electrochemical theory of conduction based on the properties of semi-permeable surfaces.

When the response to a stimulus is studied in a living system, whether it be a single cell, a tissue, or a complex organism, the indicator used, either that of movement, current of action, production of certain substances, the development of light, of heat or the alteration of form, is the result of two distinct processes. The first of these is primary excitation, brought about by the stimulus at a local point, and the second is an extension of the excitation to the surrounding tissue. We are not in a position to experimentally bring about a response to stimulation, in which the primary excitation occurs and not the secondary process, that of conductivity. All living substance contains this property, although to a very different degree, as all living substance possesses irritability, and this presents the condition not only for the taking place of the process of excitation but also that of its conduction.

If I here speak only specifically of the conduction of excitation instead of the conductivity of response to stimulation this is not only primarily for the reason that we intend especially to analyze the conductivity of excitation on this occasion, but also because no other effects of stimulation except those of excitation can be conducted from the part affected by the stimulus to the surroundings.

Although considered on theoretical grounds it appears more or less improbable that depression is extended from the place of its origin, it is very easy to convince one’s self experimentally of the fact that depression following a stimulus is invariably localized to that portion directly affected by the stimulus. The nerve furnishes a very favorable object for this purpose. If a nerve muscle preparation of the frog is made and introduced in the glass chamber previously described containing platinum electrodes, and another pair is applied to the nerve between the chamber and the muscle, it is possible to subject the stretch of nerve in the chamber to various agents, producing a paralyzing effect. In this way it may be exposed to an atmosphere of pure nitrogen for example, or to narcosis as by ether, chloroform, carbon dioxide and other gases, to an increase in temperature or to other agents, without these in any way affecting the irritability of the nerve stretch situated over the electrode between the chamber and the muscle. The contractions of the muscle, which are produced by stimulation of the periphery region of the nerve with stimuli of a definite strength, remain unaltered, even when the asphyxiated stretch of nerve in the chamber is already completely degenerated. The central depression of a ganglion cell of a motory neuron is likewise wholly without influence on the degree of excitability of its nerve fiber, as I was able to demonstrate[79] in the reflex inhibition of the motor neurons of the spinal cord of the dog. (Figure [14].) That which is conducted by the nerves is solely the process of excitation.

It is our task to analyze in detail the conditions involved in the conduction of excitation in order to obtain a deeper insight into the physics of this process. A comparative survey of a series of various types of living substance shows us that they differ in respect to the conduction of excitation in the following points: In regard to the rapidity with which the excitation is conducted, the extent of the area over which it spreads, and the intensity with which it extends. These conditions may be best illustrated by citing two extreme examples. The one is formed by the rhizopods, the other by the nerve fibers. Between these two extremes we have manifold gradations in the conditions of conductivity. Not all cell forms are suitable objects for the study of conductivity. There are forms of rhizopods which are as favorable to investigation as the nerve; this is due to the fact that, although they are often of microscopic dimensions, they possess elongated fingerlike or threadlike pseudopods.

Fig. 14.

Contractions of the musculus extensor digitorum communis longus of the dog, brought about by rhythmic stimulation of the nervus peroneus. The muscle is in the condition of tonic excitation which proceeds from the center. The arrows indicate the point where reflex inhibition of the central tonus is produced. The height of the single contraction undergoes no diminution.

Indeed, a rhizopod cell, with its straight, elongated pseudopods, is preëminently fitted as an object of comparison with a neuron. Although the difference in respect to the individual points is so far-reaching, still, based on their outward morphological similarity various physiological parallels in both are forced on our observation. A comparison of the rhizopod cell with the neuron can consequently guard us from many erroneous generalizations which we might be inclined to deduce from a one-sided investigation of the nerve. This is especially the case in regard to the conductivity of excitation, which was formerly studied almost exclusively on the nerve and only occasionally on the muscle, which offers similar conditions. The nerve, in which the function of the conductivity of excitation is particularly highly developed, was considered at the same time as the type in which this process could be most readily analyzed, and from which it was believed general information of the process of the conductivity of excitation could first be gained. This view has led to serious errors, as the nerve, resulting from the high development of its conductive capability, shows quite one-sided specialized conditions, which can by no means be transferred to other forms of living substance.

A very suitable object among rhizopods for the study of conductivity, and which is everywhere easily procured, is Difflugia. This species living in small pools has a delicate urn-shaped, pear-shaped or flask-shaped capsule built up of sand grains, diatomes or material produced by the organism itself. From the opening the protoplasm extends often to a considerable length its finger-shaped hyaline pseudopods. When Difflugia is placed in a flat dish in water and observed under the microscope, it is frequently seen to extend from the opening long pseudopods in exactly opposite directions, which reach for a considerable distance on the bottom. These offer particularly favorable conditions for the study of the conduction of excitation. When this animal is placed under a microscope, the pseudopods are very readily stimulated at any position to a desired extent by means of a sharp needle, to which fat has been previously applied and subsequently the excess removed. The extension of the response from one point toward the other can then be followed with great ease. The pseudopod of the rhizopod has the great advantage over the nerve that its excitation can be directly observed. The excitation following weaker stimulation is manifested by a wrinkling of the previously completely smooth surface; stronger stimulation produces differentiation of the hyaline protoplasm to a strongly refractive strand in the axis and a turbid myelinlike mass at the periphery, the pseudopod at the same time retracting toward the central cell body. In spite of all these occurrences being of microscopic dimensions, still with some practice it is quite possible to experiment on them under the microscope. In this way I found it comparatively simple to study the fundamental principles of conductivity.[80]