Essays Upon Heredity and Kindred Biological Problems. Weismann August

end of a line of development tending in the direction of a shortened life. This is made clear by the fact that one of these may-flies (Palingenia) does not even leave its pupa-skin, but reproduces in the so-called sub-imago stage.

      It is therefore obvious that the duration of life is extremely variable, and not only depends upon physiological considerations, but also upon the external conditions of life. With every change in the structure of a species, and with the acquisition of new habits, the length of its life may, and in most cases must, be altered.

      In answering the question as to the means by which the lengthening or shortening of life is brought about, our first appeal must be to the process of natural selection. Duration of life, like every other characteristic of an organism, is subject to individual fluctuations. From our experience with the human species we know that long life is hereditary. As soon as the long-lived individuals in a species obtain some advantage in the struggle for existence, they will gradually become dominant, and those with the shortest lives will be exterminated.

      So far everything is quite simple; but hitherto we have only considered the external mechanism, and we must now further inquire as to the concomitant internal means by which such processes are rendered possible.

      This brings us face to face with one of the most difficult problems in the whole range of physiology,—the question of the origin of death. As soon as we thoroughly understand the circumstances upon which normal death depends in general, we shall be able to make a further inquiry as to the circumstances which influence its earlier or later appearance, as well as to any functional changes in the organism which may produce such a result.

      The changes in the organism which result in normal death,—senility so-called,—have been most accurately studied among men. We know that with advancing age certain alterations take place in the tissues, by which their functional activity is diminished; that these changes gradually increase, and finally either lead to direct or so-called normal death, or produce indirect death by rendering the organism incapable of resisting injuries due to external influences. These senile changes have been so well described from the time of Burdach and Bichat to that of Kussmaul, and are so well known, that I need not enter into further details here.

      In answer to an inquiry as to the causes which induce these changes in the tissues, I can only suggest that the cells which form the vital constituents of tissues are worn out by prolonged use and activity. It is conceivable that the cells might be thus worn out in two ways; either the cells of a tissue remain the same throughout life, or else they are being continually replaced by younger generations of cells, which are themselves cast off in their turn.

      In the present state of our knowledge the former alternative can hardly be maintained. Millions of blood corpuscles are continually dying and being replaced by new ones. On both the internal and external surfaces of the body countless epithelial cells are being incessantly removed, while new ones arise in their place; the activity of many and probably of all glands is accompanied by a change in their cells, for their secretions consist partly of detached and partly of dissolved cells; it is stated that even the cells of bone, connective tissue, and muscle undergo the same changes, and nervous tissue alone remains, in which it is doubtful whether such a renewal of cells takes place. And yet as regards even this tissue, certain facts are known which indicate a normal, though probably a slow renewal of the histological elements. I believe that one might reasonably defend the statement,—in fact, it has already found advocates,—that the vital processes of the higher (i.e. multicellular) animals are accompanied by a renewal of the morphological elements in most tissues.

      This statement leads us to seek the origin of death, not in the waste of single cells, but in the limitation of their powers of reproduction. Death takes place because a worn-out tissue cannot for ever renew itself, and because a capacity for increase by means of cell-division is not everlasting, but finite [See Note 8]. This does not however imply that the immediate cause of death lies in the imperfect renewal of cells, for death would in all cases occur long before the reproductive power of the cells had been completely exhausted. Functional disturbances will appear as soon as the rate at which the worn-out cells are renewed becomes slow and insufficient.

      But it must not be forgotten that death is not always preceded by senility, or a period of old age. For instance, in many of the lower animals death immediately follows the most important deed of the organism, viz. reproduction. Many Lepidoptera, all may-flies, and many other insects die of exhaustion immediately after depositing their eggs. Men have been known to die from the shock of a strong passion. Sulla is said to have died as the result of rage, whilst Leo X succumbed to an excess of joy. Here the psychical shock caused too intense an excitement of the nervous system. In the same manner the exercise of intense effort may also produce a similarly fatal excitement in the above-mentioned insects. At any rate it is certain that when, for some reason, this effort is not made, the insect lives for a somewhat longer period.

      It is clear that in such animals as insects we can only speak figuratively of normal death, if we mean by this an end which is not due to accident. In these animals an accidental end is the rule, and is therefore, strictly speaking, normal [See Note 9].

      Assuming the truth of the above-mentioned hypothesis as to the causes of normal death, it follows that the number of cell-generations which can proceed from the egg-cell is fixed for every species, at least within certain limits; and this number of cell-generations, if attained, corresponds to the maximum duration of life in the individuals of the species concerned. Shortening of life in any species must depend upon a decrease in the number of successive cell-generations, while conversely, the lengthening of life depends upon an increase in the number of cell-generations over those which were previously possible.

      Such changes actually take place in plants. When an annual plant becomes perennial, the change—one in every way possible—can only happen by the production of new shoots, i. e. by an increase in the number of cell-generations. The process is not so obvious in animals, because in them the formation of young cells does not lead to the production of new and visible parts, for the new material is merely deposited in the place of that which is worn out and disappears. Among plants, on the other hand, the old material persists, its cells become lignified, and it is built over by new cells which assume the functions of life.

      It is certainly true that the question as to the necessity of death in general does not seem much clearer from this point of view than from the purely physiological one. This is because we do not know why a cell must divide 10,000 or 100,000 times and then suddenly stop. It must be admitted that we can see no reason why the power of cell-multiplication should not be unlimited, and why the organism should not therefore be endowed with everlasting life. In the same manner, from a physiological point of view, we might admit that we can see no reason why the functions of the organism should ever cease.

      It is only from the point of view of utility that we can understand the necessity of death. The same arguments which were employed to explain the necessity for as short a life as possible, will with but slight modification serve to explain the common necessity of death4.

      Let us imagine that one of the higher animals became immortal; it then becomes perfectly obvious that it would cease to be of value to the species to which it belonged. Suppose that such an immortal individual could escape all fatal accidents, through infinite time,—a supposition which is of course hardly conceivable. The individual would nevertheless be unable to avoid, from time to time, slight injuries to one or another part of its body. The injured parts could not regain their former integrity, and thus the longer the individual lived, the more defective and crippled it would become, and the less perfectly would it fulfil the purpose of its species. Individuals are injured by the operation of external forces, and for this reason alone it is necessary that new and perfect individuals should continually arise and take their place, and this necessity would remain even if the individuals possessed the power of living eternally.

      From this follows, on the one hand, the necessity of reproduction, and, on the other, the utility of death. Worn-out individuals are not only valueless to the species, but they are even harmful, for they take the place of those which are sound. Hence by the operation of natural selection, the life of our hypothetically