We have spoken of the reactions which occur in protoplasm as divisible into two great series—the one ascending, constructive, endothermal; the other descending, destructive, exothermal. In the one series energy is locked up; in the other series it is set free. Synthesis and analysis are names applied to the two series respectively. Synthesis is characteristic of plants, although analysis is also perpetually occurring. Plants fix carbon from the air and liberate oxygen. They also respire, setting free carbonic acid. Analysis is characteristic of animals, although synthesis is not excluded.
Of the chemical processes which occur in plants very little is known. Few halting-places between raw materials and finished products can be marked. The final products are sugars and starches, oils, proteins, and a vast number of other substances—alkaloids, glucosides, etc. Condensation, dehydration, and deoxidation are the methods by which the synthesis of these compounds is accomplished. These methods are adopted simultaneously in varying degree. The large group of bodies known as sugars and starches are, with few exceptions, built on the C₆H₆ model; in fruit-sugar, C₆H₁₂O₆, six atoms of carbon are linked to one another and to six molecules of water. The formula of starch is (C₆H₁₀O₅)ₙ. Not only has water been removed from the molecule, but an unknown number of molecules have been linked together. This condensation and dehydration is effected whenever sugar carried in cell-sap is deposited as starch in seeds or tubers. These compounds are hexatomic. The chemist pictures them as made by the union in the first place of six atoms. As small drops unite to form larger ones, so small molecules, under the direction of the protoplasm of plants, close together.
The reactions which characterize animal protoplasm are of a different kind. They belong to the descending series. Close molecules are unfolded. Water is incorporated with them. Hydrogen and carbon are oxidized into water and carbonic acid. The conversion into sugar of glycogen or of starch may be taken as an illustration of expansion. Starch, (C₆H₁₀O₅)ₙ, becomes maltose, C₁₂H₂₂O₁₁, and then dextrose, C₆H₁₂O₆. The grouped molecule of starch opens out. The breaking of the double molecule of maltose into two molecules of dextrose is a further illustration of progress towards simplicity. Hydration, union with H₂O, accompanies this expansion. Hydrolysis is the secret of almost all digestive acts. Starch is hydrolysed into sugar, fat hydrolysed into glycerin and fatty acid, proteins hydrolysed into peptones.
All the chemical transformations which protoplasm is able to accomplish are of the nature of fermentations. The term fermentation was first applied to the effervescence which occurs in grape-juice when its sugar is being converted into alcohol, carbonic acid gas, and certain substances which appear in relatively small quantities. It was discovered later that the yeast which effects this change is a unicellular plant. The term “fermentation” was extended to the production of vinegar from alcohol, and eventually to all such reactions as are carried out by living organisms, or by the secretions or products of living organisms, without the destruction of the agent which is effective in the process. A ferment is an organic body which brings about changes in other bodies without itself undergoing change. At the end of the process, however prolonged, there is as much ferment as there was at the beginning, and its chemical nature is the same. Rennin has been made to curdle nearly a million times its weight of milk, pepsin to digest half a million times its weight of fibrin. As the ferment is not consumed, there is no relation, except one of speed, between the ferment and the quantity of fermentable substance which it is able to transform. We said that a ferment is an organic body. It is necessary to introduce the qualification “organic,” because certain reactions termed “catalyses” which occur in mineral chemistry resemble fermentations in respect of the non-destruction of the agent which serves as intermediary. If a solution of cane-sugar containing a very small quantity of sulphuric acid is boiled, the cane-sugar is “inverted.” It is changed into a mixture of fruit-sugar and levulose. The ferment invertin of the gastric juice and of intestinal juice produces a similar effect; and just as invertin remains unchanged, so also the sulphuric acid is found in the mixture unchanged in nature and in amount after an unlimited inversion of cane-sugar. Great stress was formerly laid upon the similarity between fermentation and catalysis. It has now been shown that catalytic actions are not necessarily of the same nature as fermentation, although the results and, as far as is visible, the means are similar. For example, finely divided platinum (or, better, palladium) causes an indefinite quantity of oxygen and hydrogen to unite. The reaction comes within the category of catalyses. But it is widely different from a fermentation. The metal causes hydrogen to condense, and actually absorbs it into its surface layer. In the liquid form hydrogen cannot resist combination with oxygen. This may be termed a “physical phenomenon,” adopting the common distinction between chemistry and physics. There is no reason for thinking that fermentations can be explained in so simple a way. They may, however, be grouped under the designation “catalyses.” As the initial conditions and final results are similar, it is inevitable that fermentations and catalyses should obey the same “laws” as to mass action, speed, effect of accumulation of products of action, and the like; but it does not follow that invertin and sulphuric acid produce their effects in the same way. Fermentations are instances of catalysis, but all catalytic actions are not fermentations.
So far from dwelling upon the resemblance between fermentation and the catalysis of mineral chemistry, chemists nowadays incline to regard fermentation as essentially a reaction of life. It is very difficult, when attempting to present ideas which are new to thought, to adapt, without ambiguity, existing words. It would be absurd to talk of a substance removed from yeast or bacteria or blood-corpuscles by a process which involves cooling with liquid air, grinding with powdered glass, solution in water, precipitation with absolute alcohol, and resolution in water, as alive. Yet, unlike any known mineral product, it is easily killed. Ferments are not destroyed by cold, but their activity is arrested. They are most active at about the body temperature. Their activity is annihilated by heating them, in solution, to the temperature at which albumin coagulates—a little over 50° C. Although they are not alive, their behaviour very closely resembles that of living matter. They can be obtained only from living things. They produce their effects even though they are present in almost infinitely small quantity. It is impracticable to make a chemical analysis of a ferment, owing, in the first place, to the very small amount available for analysis, and, in the second place, because of the impossibility, with existing methods, of obtaining a ferment pure. The amount of ferment present in even a great mass of yeast, or in many pounds of salivary gland or pancreas, is extremely small. However prepared, it is always accompanied with proteid substances. It is impossible to say whether ferments, like proteins, have heavy nitrogen-containing molecules. The fact that they are not diffusible suggests that they have.
It would be straining language to term fermentation a phenomenon of life; worse, to define life as a sequence of fermentations. Yet it is safe to say that all the chemical changes carried out by living organisms are fermentations. Fermentation and the chemistry of life are almost synonymous terms.
A very large number of ferments are already known. Each has its own specific work to do: “To every fermentable substance is fitted a ferment, as a key to a lock.” It will be understood, from what has been already said regarding our inability to determine the composition of any ferment, that we cannot say whether or not these various ferments differ one from another in chemical constitution. They are classified according to their action, and not according to their nature. Those which build up are termed “synaptases” (συνάπτω, I unite); those which decompose, or hydrolyse, “diastases” (διάστασις, separation). The termination “ase” is added to the name of the substance upon which the ferment acts, except in cases in which other terms have already become so general as not to be displaceable: amylase, hydrolysing starch; sucrase, inverting cane-sugar; protease, hydrolysing proteins. Unfortunately, there is little uniformity in this nomenclature; amylopsin, invertin, pepsin, are terms used as often as those terminating in “ase.” As a distinguishing termination, “in” or “sin” is less desirable than “ase,” owing to the fact that it has been appropriated already as the termination of the names of albuminoids—e.g., gelatin, chondrin, mucin.
The various ferments are substances which protoplasm sets aside for specific purposes. Primitively, contact with the substance to be fermented determined the nature of the ferment assigned to the task. There are reasons for thinking that protoplasm still retains its power of making a suitable response; cases may be cited in which the lock presented to protoplasm shapes the wards of the key. In such cases the fermentable substance provokes the formation of the ferment. But, for the most part, in situations where particular ferments are regularly needed, protoplasm has acquired the habit of making such ferments and no others. The cells of salivary glands accumulate ptyalin, the cells of gastric glands accumulate pepsin, during the intervals between meals.
The capacity of protoplasm for producing a new ferment when it is needed is shown by such examples as the following: Blood-plasm contains a variety of proteid substances. If a solution of white of egg be added to it, the mixture is clear and uniform. Yet egg-albumin is treated by the blood as a foreign body, a poison. When injected into the veins of a living animal, some of it is excreted by the kidneys, some destroyed in the blood-stream. If several successive doses of egg-albumin are injected into an animal (it is most convenient to inject it into the peritoneal cavity), the power of the blood to destroy the intruder is greatly increased. If now a specimen of blood be taken, and the plasma or serum mixed with egg-albumin, the mixture is no longer clear. The egg-albumin is precipitated. The blood of the animal thus “prepared” has developed a ferment, termed a “precipitin,” which throws down egg-albumin. If instead of egg-albumin, which, although a foreign body, is comparatively innocent, a substance which is distinctly poisonous, toxic, be injected into an animal, the first dose, if a large one, will prove fatal. If, however, the first dose be small, and succeeding doses progressively larger, the animal acquires the power of tolerating a quantity of the poison much larger than would have proved fatal in the first instance. A classical example of this, because it afforded an opportunity of directly observing under the microscope the difference between “unprepared” blood and blood from an immune animal, is the acquisition by a mammal of the power of tolerating the injection of the blood of an eel. Eel’s blood contains a toxin which destroys the red blood-corpuscles of a mammal. The dissolution of the blood-corpuscles may be watched with the microscope. If successively increasing doses of serum of eel’s blood be injected into the body of a rabbit, the rabbit acquires the power of resisting the toxin. Further than this, the serum of the immune rabbit injected into a rabbit which has not been prepared confers immunity upon the latter. If the blood of the prepared animal be mixed with the blood of an unprepared rabbit and with eel’s serum, and the mixture examined under the microscope, it will be seen that red blood-corpuscles are no longer dissolved. The immune serum is able to save the blood-corpuscles of the unprepared blood from destruction. During its course of preparation the rabbit developed an antitoxin.
If germs of diphtheria are injected into the blood of a horse, the first injections give rise to marked febrile symptoms. After a number of injections the horse becomes completely tolerant of the virus. Not only does its blood develop sufficient antitoxin to protect it against the toxin of diphtheria, however large may be the quantity injected into its system, but the serum of the prepared horse, when injected beneath the skin of a child suffering from diphtheria, carries with it sufficient antitoxin to destroy the toxin which has gained admission to the child’s blood.