Max Verworn and Max Kassowitz, have pointed out, in opposition to modern vitalism, that "life consists in a continuous alternation between the upbuild and the decay of the highly complicated chemical unities of the protoplasm. And if this conception is admitted, we may rightly say that we know what we mean by death. If death is the cessation of life, we must mean by that the cessation of the alternation between the upbuild and the dissolution of the molecules of protoplasm; and as each of the molecules of protoplasm must break up again shortly after its formation, we have in death to deal only with the definite cessation of reconstruction in the destroyed plasma-molecules. Hence a living thing is not finally dead—that is to say, absolutely incompetent to discharge any further vital function—until the whole of its plasma-molecules are destroyed." In the exhaustive justification with which Kassowitz follows up this definition in the fifteenth chapter of his General Biology, the natural causes of physiological death are fully described.
Among the numerous and contradictory views of recent biologists on the nature of death we find many errors and misunderstandings, due to a lack of clear distinction between the duration of the living matter in general and that of the individual life-form. This is particularly noticeable in the contradictory views which have been elicited by August Weismann's theory (1882) of the immortality of the unicellulars. I have shown in the eleventh chapter of the Riddle that it is untenable. But as the distinguished zoologist has again taken up his theory with energy in his instructive Lectures on the Theory of the Descent (1902), and has added to it erroneous observations on the nature of death, I am obliged to return to the point. Precisely because this interesting work gives most valuable support to the theory of evolution, and maintains Darwin's theory of selection and its consequences with great effect, I feel it is necessary to point out considerable weaknesses and dangerous errors in it. The chief of these is the important theory of the germ-plasm and the consequent opposition to the inheritance of acquired characteristics. Weismann deduces from this a radical distinction between the unicellular and the multicellular organisms. The latter alone are mortal, the former immortal; "between the unicellular and the multicellular lies the introduction of physiological—that is to say, normal—death." We must say, in opposition to this, that the physiological individuals (bionta) among the protista are just as limited in their duration as among the histona. But if the chief stress in the question is laid, not on the individuality of the living matter, but on the continuity of the metabolic life-movement through a series of generations, it is just as correct to affirm a partial immortality of the plasm for the multicellulars as for the unicellulars.
The immortality of the unicellulars, on which Weismann has laid so much stress, can only be sustained for a small part of the protists even in his own sense—namely, for those which simply propagate by cleavage, the chromacea and bacteria among the monera (chapter ix.), the diatomes and paulotomes among the protophyta, and a part of the infusoria and rhizopods among the protozoa. Strictly speaking, the individual life is destroyed when a cell splits into two daughter-cells. One might reply with Weismann that in this case the dividing unicellular organism lives on as a whole in its offspring, and that we have no corpse, no dead remains of the living matter, left behind. But that is not true of the majority of the protozoa. In the highly developed ciliata the chief nucleus is lost, and there must be from time to time a conjugation of two cells and a mutual fertilization of their secondary nuclei, before there can be any further multiplication by simple cleavage. However, in most of the sporozoa and rhizopoda, which generally propagate by spore formation, only one portion of the unicellular organism is used for this; the other portion dies, and forms a "corpse." In the large rhizopods (thalamophora and radiolaria) the spore-forming inner part, which lives on in the offspring, is smaller than the decaying outer portion, which becomes the corpse.
Weismann's view of the secondary "introduction of physiological death in the multicellulars" is just as untenable as his theory of the immortality of the unicellulars. According to this opinion, the death of the histona—both the metaphyta and metazoa—is a purposive outcome of adaptation, only introduced by selection when the multicellular organism has reached a certain stage of complexity of structure, which is incompatible with its original immortality. Natural selection would thus kill the immortal and preserve only the mortal; it would interfere with the multiplication of the immortals in the bloom of their years, and only use the mortal for rearing posterity. The curious conclusions which Weismann reached in developing this theory of death, and the striking contradictions to his own theory of the germ-plasm which he fell into, have been pointed out by Kassowitz in the forty-ninth chapter of his General Biology. In my opinion, this paradoxical theory of death has no more basis than the germ-plasm theory he has ingeniously connected with it. We may admire the subtlety and depth of the speculations with which Weismann has worked out his elaborate molecular theory. But the nearer we get to its foundations the less solid we find them. Moreover, not one of the many supporters of the theory of germ-plasm has been able to make profitable use of it in the twenty years since it was first published. On the other hand, it has had an evil influence in so far as it denied the inheriting of acquired characters, which I hold, with Lamarck and Darwin, to be one of the soundest and most indispensable supports of the theory of descent.
In discussing the question of the real causes of death, we confine our attention to normal or physiological death without considering the innumerable causes of accidental or pathological death, by illness, parasites, mishaps, etc. Normal death takes place in all organisms when the limit of the hereditary term of life is reached. This limit varies enormously in different classes of organisms. Many of the unicellular protophyta and protozoa live only a few hours, others several months or years; many one-year plants and lower animals live only a summer in our temperate climate, and only a few weeks or months in the arctic circle or on the snow-covered Alps. On the other hand, the larger vertebrates are not uncommonly a hundred years old, and many trees live for a thousand years. The normal span of life has been determined in all species in the course of their evolution by adaptation to special conditions, and has then been transmitted to offspring by heredity. In the latter, however, it is often subject to considerable modifications.
The organism has been compared, on the modern "machine theory" of life, to an artificially constructed mechanism, or an apparatus in which the human intelligence has put together various parts for the attainment of a certain end. This comparison is inapplicable to the lowest organisms, the monera, which are devoid of such a mechanical structure. In these primitive "organisms without organs" (chromacea and bacteria) the sole cause of life is the invisible chemical structure of the plasm and the metabolism effected by this. As soon as this ceases death takes place (cf. chapter ix.). In the case of all other organisms the comparison is useful in so far as the orderly co-operation of the various organs or parts accomplishes a certain task by the conversion of virtual into active force. But the great difference between the two is that in the case of the machine the regularity is due to the purposive and consciously acting will of man, whereas in the case of the organism it is produced by unconscious natural selection without any design. On the other hand, the two have another important feature in common in the limited span of life which is involved in their being used up. A locomotive, ship, telegraph, or piano, will last only a certain number of years. All their parts are worn out by long use, and, in spite of all repairing, become at last useless. So in the case of all organisms, the various parts are sooner or later worn out and rendered useless; this is equally true of the organella of the protist and the organs of the histon. It is true that the parts may be repaired or regenerated; but sooner or later they cease to be of service, and become the cause of death.
When we take the idea of regeneration, or the recuperation of parts that have been rendered useless, in the widest sense, we find it to be a universal vital function of the greatest importance. The whole metabolism of the living organism consists in the assimilation of plasm, or the replacing of the plasma-particles which are constantly used up by dissimilation (cf. chapter x.). Verworn has given the name of biogens to the hypothetical molecules of living matter—which I regard with Hering as endowed with memory, and (1875) have called plastidules. He says: "The biogens are the real vehicles of life. In their constant decay and reconstruction consists the process of life, which expresses itself in the great variety of vital phenomena." The relation of assimilation (the building-up of the biogens) to dissimilation (the decay of the biogens) may be expressed by a fraction to which the name biotonus is given A/D. It is of