The usual view among physiologists is that the muscle fibre is a thermodynamic apparatus transforming the heat generated during metabolism into mechanical energy. How is this transformation effected? It cannot be said that we have any one hypothesis more convincing than another. It has been suggested that alterations of surface tension play a part, or that the heat produced by oxidation causes the fibre to imbibe water and shorten. Engelmann has devised an artificial muscle consisting of a catgut string and an electrical current passing through a coil of wire, and by means of this he has reproduced the phenomena of simple contraction and tetanus. But it remains for future investigation to verify any one of these hypotheses.
When Huxley published his Physical Basis of Life, probably few physiologists had any doubt that protoplasm was a definite chemical substance, differing from other organic substances only by its much greater complexity. But in 1880 Reinke and Rodewald published the results of an analysis of the substance of a plant protoplasm and these appear to have demonstrated that the substance was really a mixture of a number of true chemical compounds and was not a single definite one. Now all of these substances might exist apart from protoplasm, and in the lifeless form, and a simple mixture of them could hardly bring forth vital reactions. These results were followed by the morphological study of the cell—the discovery of the architecture of the nucleus, and so on, and so opinion began to turn to the hypothesis that the vital manifestations of protoplasm were the result of its structure. Microscopical examination of the cell appeared to disclose a definite arrangement, the “foam” or “froth” of Butschli, for instance. But, again, it was easily shown that the foam, or alveolar structure of protoplasm was merely the expression of physical differences in the substances composing the cell-stuff—they reduced to phenomena of surface tension and the like. Artificial protoplasm and artificial Amœbæ were made—at least mixtures of olive oil and various other substances were made which simulated many of the phenomena of protoplasm in much the same way as crystalline products may be made which simulate the growth of a plant stem with its branches. For instance, one has only to shake up a little soapy water in a flask to see what resembles surprisingly the arrangement of certain kinds of connective tissues in the organism. Obviously these artificial phenomena have nothing to do with living substance.
Yet if we grind up a living muscle with some sand in a mortar we do destroy something. The muscle could be made to contract, but after disintegration this power is lost. We have certainly destroyed a structure, or mechanism, of some kind. But, again, the paste of muscle substance and sand still possesses some kind of vital activity, for with certain precautions it can be made to exhibit many of the phenomena of enzyme activity displayed by the intact muscle fibres, or even the entire organism. Mechanical disintegration, therefore, abolishes some of the activities of the organism, but not all of them. If, however, we heat the muscle paste above a certain temperature, the residue of vital phenomena exhibited by it are irreversibly removed, so that heating destroys the mechanism. This we can hardly imagine to be the case (within ordinary limits of temperature at least) with a physical mechanism, but again a mechanism which is partly chemical might be so destroyed. We see, then, that protoplasm possesses a mechanical structure, but that all of its vital activities do not necessarily depend on this structure. The full manifestation of these activities depends on the protoplasmic substance possessing a certain volume or mass, and also on a certain chemical structure.
If living protoplasm has a structure, and is not simply a mixture of chemical compounds, what is it then? Two or three physico-chemical concepts are at the present time very much in evidence in this connection. When the substances known as colloids were fully investigated by the chemists, much attention was paid to them by the physiologists, so that life was called “the chemistry of the colloids,” just as after the investigation of the enzymes it was called the “chemistry of the enzymes,” and when the discovery of the relative abundance of phosphorus in cell-nuclei and in the brain was discovered, it was called the “chemistry of phosphorus.” Colloids (e.g. glue) are substances that do not readily diffuse through certain membranes, in opposition to crystalloids (e.g. solution of common salt) which do readily so diffuse. They form solutions which easily gelatinise reversibly, that is, can become liquid again (glue); or coagulate irreversibly, that is, cannot become liquid again (albumen); which have no definite saturation point; which have a low osmotic pressure (and derived properties), etc.; and the molecules of which are compound ones consisting of combinations of the molecules of the substance with the molecules of the solvent, or with each other, that is, they are molecular aggregates.
Colloids pass insensibly into crystalloids on the one hand and into coarse suspensions (water shaken up with fine mud, for instance) on the other. We may replace the concept of a colloid by those of “suspensoids” and “emulsoids.” A suspensoid is a liquid containing particles in a fine state of division—if the division is that into the separate molecules we have a solution, if into large aggregates of molecules we have a suspension. If the substance in the liquid is itself liquid, the whole is called an emulsoid. On the one hand this approaches to a mixture of oil in soap and water—an emulsion—and on the other hand to such a mixture as chloroform shaken up with water, when the drops of chloroform readily join together so that two layers of liquid (chloroform and water) form. What we see, then, in protoplasm is a viscid substance possessing a structure of some kind, and containing specialised protoplasmic bodies in its mass (nuclei, nucleoli, granules of various kinds, chlorophyll, and other plastids, etc.). It may contain or exhibit suspensoid or emulsoid parts or substances, or it may contain truly crystalloid solutions. These phases of its constituents are not fixed, but pass into each other during its activity. Nothing that we know about it justifies us in speaking about a “living chemical substance.” On analysis we find that it is a mixture of true chemical substances rather than a substance. It is no use saying that in order to analyse it we must kill it, for what we can observe in it without destroying its structure or activities indicates that it is chemically heterogeneous.
This is not a textbook of general physiology, and the examples of physico-chemical reactions in the organism which we have selected have been quoted in order to show to what extent the chemical and physical methods applied by the physiologists have succeeded in resolving the activities of the organism. The question for our consideration is this: do these results of physico-chemical analysis fully describe organic functioning? Dogmatic mechanism says “yes” without equivocation.
Now it is clear, from even the few typical examples that we have quoted, that physiological analysis shows, indeed, a resolution of the activities of the organism into chemical and physical reactions. How could it do otherwise? How could chemical and physical methods of investigation yield anything else than chemical and physical results? The fact that these methods can be applied to the study of the organism with consistent results shows that their application is valid; that we are justified in seeing physico-chemical activities in life. But are these results all that we have reason to expect?
We turn now to Bergson’s fertile comparison of the physiological analysis of the organism with the action of a cinematograph. If we take a series of photographic snapshots of, e.g., a trotting horse and then superpose these pictures upon each other, we produce all the semblance of the co-ordinated motions of the limbs of the animal. Yet all that is contained in the simulated motion is immobility. From a succession of static conditions we appear to produce a flux. Yet if we could contract our duration of, e.g., a week, into that corresponding to five minutes—if we could speed up our perceptual activity—should we not see the cinematographic pictures as they really are—a series of immovable postures and nothing more: truly an illusion? If, again, we reverse the direction of motion of the film, we integrate our snapshots into something which is absolutely different from the reality which they at first represented; and by such devices the illusions and paradoxical effects of the picture-house farces are made possible. Well, then, in the physiological analysis of the activity of the organism do we not do something very analogous to this? The complexity of even the simplest function of the animal is such that we can only attend to one or two aspects of it at once, arbitrarily neglecting all the rest. We find that the hydrostatic pressure of blood, and lymph, and secretion, the osmotic pressure, the diffusibility, vaso-motor actions, and other things must be investigated when considering the question of how the submaxillary gland secretes saliva. One, or as many as possible, of these reactions are investigated at one time, and then the results are pieced together—integrated—in order to reproduce the full activity of the whole indivisible process. But in doing this do we not introduce something new—a direction or order of happening—into the elements of the dissociated activity of the organism? Each elemental process must occur at just the right time.
What right have we to say that the activity of the organism is made up of physico-chemical elements? Just as much as we have in saying that a curve is made up of infinitesimal straight lines. Let us adopt Bergson’s illustration, with a non-essential modification.