A Bipolar Theory of Living Processes

Two colloidal solutions of different H-ion concentrations separated from each other by a semi-permeable dielectric partition constitute an electric cell, within which the potential difference bears a direct relation to the difference between the H-ion concentrations of the two solutions. The difference of potential created by a cell or battery will pass along conductors to points of lower potential. If there are gaps—synapses—in the conducting paths a sufficient difference of potential between the two ends of the path makes it possible for the electric current to bridge this gap.

Histologic studies by Dr. M. L. Menten and Dr. J. B. Austin have shown that the physical structure of the brain cells of electric fish is changed after the electric organ is discharged. (Fig. 62.) Dahlgren * has found that the electric organ of the electric fish does not store electricity but that it is capable of the instantaneous production of electricity by means of oxidation and that this process of oxidation within the electric organ Fic. 62.—F ffect of the discharge of its electric organ on the brain cells of а torpedo. (From photemicrographs Х 310.)

is not performed if it is disconnected from the brain. This last observation, as stated in Chapter III, was made by Galvani in experiments described by Becquerel.? Finally, of the utmost value in its support of the conception that the organism as a whole is operated by electricity should be cited the biological fact that the application of electricity to — organs or tissues can make them perform all the functions which they normally perform under nervous stimulation.

Although the relation of light energy to the electric energy which operates the bipolar animal organism may not at first be apparent, the following facts are peculiarly significant: (1) that there is increasing evidence that the original unit of living matter was organized from the inorganic elements of the sea by means of electric energy or sunlight; (2) that living organisms, even those whose lives are spent in apparent darkness, live in a world the atmosphere of which is constantly subjected to a bombardment of electrons from the sun; (3) that the energy of heat and light is essential for the production of chemical action; and (4) that the energy transmitted through one of the most delicate and intricate mechanisms in man as the result of the impact of light waves, has been demonstrated to be electric energy. It is therefore fitting that we consider briefly the physical qualities of light and its biological effects in their relation to a bipolar mechanism.

According to the electronic theory “light waves” are electric waves, and the light of the sun is electric energy transmitted by the constant emission of eleetrons from the vast number of electrons constantly vibrating within the body of the sun itself. Light, then, is electricity, and the effects of light are the result of the applieation of its eleetrieal energy and are to be interpreted by the laws which govern electrical energy. That sunlight is electricity is shown by its capacity to induce positive charges. This has been demonstrated by Nodon 3 in his observations of the electric action of the sun and moon, from which he concludes that “the sun induces a positive elec-

tric charge” and that “the full moon produces a positive electric induction analogous to that of the sun.” Various investigators have produced formaldehyde by the action of sunlight on water and carbonic acid in the presence of salts of iron or other inorganic salts; and by the further action of sunlight or of ultraviolet light on formaldehyde, earbohydrates—sugars—have been produced. That is, sunlight—electric energy—ean produce organic compounds from inorganic elements.

The phenomena of heliotropism as demonstrated by Loeb and his conclusions therefrom are strong arguments in favor of the assumption that the animals thus activated are electric mechanisms and that the energy that activates them is electricity. The facts that there is an optimum and a lethal temperature for every type of life; that variations in one of the most fundamental processes of life—oxidation—are accompanied by variations in temperature, make it essential to consider a few of the more important physical properties and biological effects of heat in their relation to electric energy.

Heat in common with electricity and with light is radiant energy, but while the latter forms of energy are electronic phenomena, heat is concerned with the activities of atoms and molecules. As heat is radiant energy, its manifestations are governed by the same primary physical laws as those which govern electric energy and light energy. Moreover, in its manifestations in living organisms and in inorganic matter, heat energy is indissolubly linked with electric energy, light energy and chemical energy.

Of primary importance is the fact that “were there no heat or light, or were the intensity of these below a certain limit, depending on the nature of the substances, we could get no chemical action. Thus inertness would probably be a property of all substances in the dark at the so-called absolute zero of temperature.” (Comstock and Troland. )* sider that chemical action means an alteration of atomic relations, and hence of electronic relations and that the relation between heat and oxidation is reciprocal, that heat promotes oxidation, oxidation producing heat, the direct relation of heat to the electric phenomena of the organism becomes apparent.

An equation has been established by means of which the heat of ionization of water may be related to the ionization constant of water, and this equation is equally “applicable to the ionization process of electrolytes in water. If the ionization values are known at different temperatures, the energy change accompanying the ionization process may be calculated, assuming that the energy change accompanying the process remains constant. . . . It appears, thus, that a knowledge of the thermal properties of electrolytic solutions has a very direct bearing on our interpretation of the phenomena observed in electrolytic solutions. . . . In this connection, it should be noted that a number of investigators, from a study of the temperature coefficient of the electromotive force of concentration cells have obtained values for the energy changes accompanying the transfer of electrolytes from solutions of one concentration to another.” (Kraus.) °

Whether or not this equation and its derivatives can be practieally applied to the biologie concentration cell, these observations throw still further light upon the electro-chemical róle of heat in the organism: “The extraordinarily good heat conductivity of metals is accounted for in terms of the free electrons which they contain. If this is the true explanation, it can be shown to follow that, other things equal, those metals which contain the largest number of free electrons will be the best heat conductors. But such metals will also be the best conductors of electricity, and hence it would appear that some sort of proportionality should exist between the power of a substance to conduct heat and its power to conduct electricity. Accurate measurements and calculations show that a relationship of this kind holds in nature, and that its quantitative character is in remarkable accord with the assumptions of the electronic and molecular theories.” (Comstock and 'lroland.)*

This is strongly in accord with the electro-chemical сопсер- tion, for our researches have shown that the electric conductivity of tissues bears a direct relation to their chemical activity, hence rate of oxidation, hence ionie activity, hence free electrons. That heat is favorable to the growth of plants while cold retards growth, is a common observation; that there is a definite limited range of temperature within which each type of living organism can exist is well known; that all the major activities of life are accompanied by variations in the manifestation of heat is a universal experience. That the chemical reactions which accompany the operation of the mechanism must be accompanied by variations in heat production is the direct corollary of the physico-chemical facts.

The source of heat within the organism has always been the subject of discussion, Lavoisier being the first to prove that it is the result of the combustion of the tissues themselves. Following Lavoisier various investigators have endeavored to locate the principal site of this combustion in some specific organ or tissue—the lungs, the blood, the muscles, ete. It is only within comparatively recent years that it has been generally accepted that the combustion takes place within the living cells themselves.

When atoms, according to the positive or negative preponderance of the charges of which they are composed, are grouped into molecules, the resultant electrical charges of the molecules in turn are positive or negative. The manifestations of these molecular charges produce those types of energy which are designated intermolecular or intramolecular forces. These forces are variously manifested in gases, solids and liquids. In the phenomena of living organisms we are principally concerned with the intermolecular forces of solutions which are manifested as surface tension and osmotic pressure. The phenomena of sound and of heat are the result of molecular activity. In the final analysis, however, as has been stated in our discussion of the different forms of energy, all of these phenomena are due to electrical forces.

Nernst’s application of the osmotie theory to the mechanism of current-production in solutions is of especial interest in this discussion and is given at length in the Appendix, as is Loeb’s description of the relation of the intramolecular forces in biological cells to electrical phenomena. In view of all these facts, which demonstrate the omnipresence of electricity in vital phenomena, its versatility, its relation to thermal, light, chemical and intermolecular energy, its production by oxidation, its identity with sunlight—all these characteristics present overwhelming evidence that electricity is the energy by means of which living organisms operate. In addition, the fact that electrical currents operate in a circuit between points of highest and lowest potential is in accord with our conception that such a circuit exists in all living organisms from the unicellular protozoan to man.

4. Comstock and Твоглмо. The Nature of Matter and Electricity, New York, 1921, p. 29. 5. Kraus. The Properties of Electrically Conducting Systems, New York, 1922, p. 304. Tue ELECTRICAL SIGNIFICANCE OF CERTAIN ESTABLISHED Facts REGARDING THE PRINCIPAL CONSTITUENTS А моха the important constituents of living matter are hydrogen, oxygen, nitrogen and carbon in combination in water, carbohydrates, proteins and lipins; and certain inorganic salts, especially those of potassium, sodium, magnesium, calcium, iron, sulphur, chlorin and phosphorus. Each of these contributes Из share to the complexity of reactions which constitute living processes, but we shall consider here only those elements or compounds the presence of which is primarily essential to the electrical operation of the organism— water, oxygen, hydrogen, potassium and other electrolytes, oils and carbon.

Water constitutes from 70 to 90 per cent of the body weight. Water is therefore the vehicle in which the organism is suspended. Among the various properties of water which render it of value to the organism, there is one which has an essential and vital relationship to a consideration of the organism as a bipolar mechanism: water has an extremely high dielectric constant which means that it has a very high ionizing power as a solvent. Its ionizing power is so high that absolutely pure water is unobtainable, the purest which can be obtained always containing hydrogen and hydroxyl ions in solution. Moreover, water is not, as commonly considered, a simple inert substance, but is complex and unstable in its constitution. The structure of the water molecule is not constant, but is influenced primarily by temperature, and by the substances which it holds

in solution. These variations in molecular structure in turn influence its ionizing power; and variations in ionizing power mean variations in electrical properties. By virtue of its catalyzing power water promotes chemical action, which in turn promotes the production of energy. Water facilitates oxidation. “It is only in the presence of water that oxygen has the power of oxidizing rapidly." (Mathews.)! Therefore, before oxygen can become available for use in the cell it must be dissolved in water.

Water has a high specific heat. Since an elevation of temperature of one degree centigrade increases chemical activity 10 per cent and electric conductivity 2.5 per cent, it is essential that the bipolar units of the organism be protected. against such extraordinary activations or depressions as would result from comparatively slight variations in temperature. This need is met by the fact that each cell is practically suspended in a medium of a high specific heat.

One other property of water has a fundamental bearing on the bipolar character of the cells, namely, the immiscibility of oil and water. This property makes possible the establishment of the semi-permeable lipoid cell membranes. These lipoid films which separate the cells of the body from each other and from the surrounding fluids; which separate the nucleus from the eytoplasm ; which separate the spherules (Mott) within the cells from each other and from the other parts of the cell—these films could not have been established in a medium of an oil or lipoid solvent but only in a water medium.

Electrolytes in solution, especially aqueous solution, conduct the current. According to the electrolytic theory this is explained by their dissociation into ions. Salts as a rule dissociate readily, acids and bases dissociate in varying degrees. When to this statement we add the facts that the protoplasmic contents of the cells consist of water, oils and salts; that in some instances 94 per cent of living matter consists of nothing more unusual or remarkable than water and the com-

monest salts; and that water will hydrolyze any salt until | the product of the hydroxyl and hydrogen ions reaches a value of about 1074 (Bancroft) 2, thus setting free the metal ions, each with its definite electrieal eharge, the vital importance of these electrolytes to the electrie activity of th» cell is at once apparent. Of especial value in relation to the bipolar theory is the influence of electrolytes upon the type of emulsion in which they are dissolved. Studies by Clowes, by Loeb, and by Osterhout are of especial significance in this direction.

“These salts are not mere inert substances, they are not simply absorbed with the water and tolerated, but they are in combination, in part at least, with the organie matter of the protoplasm. They are not simply clinkers clogging the grates of the protoplasmie fires, but they are active in the production of the vital phenomena. Indeed, some have gone so far as to believe, as we shall see, that by means of the electrical charges they bear when in solution they vitalize the colloidal, organie substratum of the cell and make it alive. Any change in their relative proportions at once affects the activity of the cell; thus by increasing or decreasing the proportion of sodium, calcium, or potassium, skeletal muscle may be made to twitch rhythmically or to remain at rest; nerve impulses may be set up in motor nerves, or the irritability of the nerve raised or lowered; chromophores of fish scales may be contracted or expanded; and the activities of all cells increased or diminished. . . . Furthermore, by increasing the total amount of salt in protoplasm many cells may be stimulated and egg cells of some animals caused to develop parthenogenetically without the aid of sperm.” (Mathews.) ?

“This antagonistic action of sodium and calcium salts with reference to emulsions throws light upon some perplexing physiological problems. Jaques Loeb showed that certain marine organisms · died in sodium chlorid or calcium chlorid solutions which were isotonic with sea-water; but flourished when there was a definite ratio of sodium to calcium, a result which could not be explained on the basis of osmotic pressure. If we consider protoplasm as consisting of lipoids (oils) and water, we shall have an emulsion of oil in water in presence of calcium salts. When the sodium and calcium salts are present in a definite ratio, there will be a balancing between these two types of emulsion, and it may well be

that this critical state is the one which is conducive to life and growth. Аз a matter of fact, Clowes found that the ratio of sodium and calcium salts necessary to produce a balancing between the two types of emulsions when working with oil, water and soap, was practically the same as that found in sea-water. This indicates a very close connection between the two sets of phenomena. Osterhout of Harvard has shown that the specific electric conductivity of certain seaweeds is increased by addition of sodium salts and decreased by the addition of calcium salts. If the hypothetical emulsion changed to one of oil in water, the conductivity should increase, and it should decrease if the emulsion changed to one of water in oil.” (Bancroft.)*

Although the validity of Clowes’ application of his discovery of the reversal of phases in oil emulsions by electrolytes to changes in protoplasmic permeability is questioned (Seifriz),? there can be no question that variations in the ionic concentration of the elements within the cell must play an indispensable role in the production of the activities of the cell as an electrical unit. Oxygen, the “acid maker,” is, essential to the life of the cells. Oxidation produces acids. Is the production of acids the essential rôle of oxidation?

If a state of suboxidation of the cells is induced by protracted hemorrhage; by any interference with the intake of oxygen as by gradual asphyxia from any cause; by interference with the absorption of oxygen in the alveoli of the lungs as in pneumonia; or by loss of the oxygen-earrying property of hemoglobin as in carbon monoxid poisoning—whatever the eause of the suboxidation, the differential stainability of the cells is diminished or lost, the diminished energy-transforming function being manifested in the organism as a whole by a diminution of mental and physical energy which is progressive until equilibrium of the acid-alkali balanee—death—-is established.

These facts indicate that the acids produced by oxidation in turn produce the essential alterations in the difference of potential between the nucleus and the cell-body and probably between parts within the cytoplasm (Mott), as a result of which accumulations of acid and an adaptive discharge of energy are provided. Water carries the oxygen in solution to the cells, dissolves the acid by-products of oxidation, and bears them away from the cells, thus constantly restoring the acid-alkali balance which is as constantly altered by the acidulating oxygen. It would appear, therefore, that water and oxygen in a vital and inseparable relationship are essential to the production of the electrical variations within the cells, the manifestations of which, as it would appear, constitute life.

Reduction is as vital a function of the cell as is oxidation. The most active oxidizing agent, as one would suppose, is oxygen; the most active reducing agent is hydrogen. Ву means of oxidation the strongly positive hydrogen ion is released; by means of reduction it is bound, and therefore by the constant interplay of the oxidizing and reducing activities within the cell its electric potential is varied with consequent variations of the phenomena of the cell. “Аз a reservoir of life energy which is liberated by oxidation, hydrogen exceeds any other element in the heat it yields, namely, 34.5 calories per gram, while carbon yields 8.1 calories per gram.” (Os- born.)® It would appear, therefore, that the róles of hydrogen and of oxygen in the activities of the cells are interdependent.

The hydrogen atom is the smallest and most swiftly moving atom, and also the simplest and the most unstable in its construction. It consists of a positive nucleus around which revolves a single negative particle of electricity. Its instability as compared with an atom of helium, the nucleus of which is bound with two electrons, is obvious. By reason of this symmetrically balanced structure the atom of helium is comparatively inert; the unbalanced hydrogen atom is readily disrupted. Because of its instability the hydrogen constituent is readily released from compounds by oxidation and is itself disrupted, setting free the positive nucleus of its atom—the hydrogen ion, the negative electron in combination with oxygen forming the hydroxyl ion.