In the first place, I think we may conclude with certainty from the unending duration of unicellular organisms, that such wearing out of tissue cells is a secondary adaptation, that the death of the cell, like general death, has arisen with the complex, higher organisms. Waste does not depend upon the intrinsic nature of the cells, as the primitive organisms prove to us, but it has appeared as an adaptation of the cells to the new conditions by which they are surrounded when they come into combination, and thus form the cell-republic of the metazoan body. The replacement of cells in the tissues must be more advantageous for the functions of the whole organism than the unlimited activity of the same cells, inasmuch as the power of single cells would be much increased by this means. In certain cases, these advantages are obvious, as for example in many glands of which the secretions are made up of cast-off cells. Such cells must die and be separated from the organism, or the secretion would come to an end. In many cases, however, the facts are obscure, and await physiological investigation. But in the meantime we may draw some conclusions from the effects of growth, which are necessarily bound up with a certain rate of production of new cells. In the process of growth a certain degree of choice between the old cells which have performed their functions up to any particular time, and the new ones which have appeared between them, is as it were left to the organism.
The organism may thus, figuratively speaking, venture to demand from the various specific cells of tissues a greater amount of work than they are able to bear, during the normal length of their life, and with the normal amount of their strength. The advantages gained by the whole organism might more than compensate for the disadvantages which follow from the disappearance of single cells. The glandular secretions which are composed of cell-detritus, prove that the cells of a complex organism may acquire functions which result in the loosening of their connexion with the living cell-community of the body, and their final separation from it. And the same facts hold with the blood corpuscles, for the exercise of their function results in ultimate dissolution. Hence it is not only conceivable, but in every way probable, that many other functions in the higher organisms involve the death of the cells which perform them, not because the living cell is necessarily worn out and finally killed by the exercise of any ordinary vital process, but because the specific functions in the economy of the cell community which such cells undertake to perform, involve the death of the cells themselves. But the fact that such functions have appeared,—involving as they do the sacrifice of a great number of cells,—entirely depends upon the replacement of the old by newly formed cells, that is by the process of reproduction in cells26.
We cannot a priori dispute the possibility of the existence of tissues in which the cells are not worn out by the performance of function, but such an occurrence appears to be improbable when we recollect that the cells of all tissues owe their constitution to a very far-reaching process of division of labour, which leaves them comparatively one-sided, and involves the loss of many properties of the unicellular, self-sufficient organism. At any rate we only know of potential immortality in the cells which constitute independent unicellular organisms, and the nature of these is such that they are continually undergoing a complete process of reformation.
If we did not find any replacement of cells in the higher organism, we should be induced to look upon death itself as the direct result of the division of labour among the cells, and to conclude that the specific cells of tissues have lost, as a consequence of the one-sided development of their activities, the power of unending life, which belongs to all independent primitive cells. We should argue that they could only perform their functions for a certain time, and would then die, and with them the organism whose life is dependent upon their activity. The longer they are occupied with the performance of special functions, the less completely do they carry out the phenomena of life, and hence they lead to the appearance of retrogressive changes. But the replacement of cells is certain in many tissues (in glands, blood, etc.), so that we can never seek a satisfactory explanation in the train of reasoning indicated above, but we must assume the existence of limits to the replacement of cells. In my opinion, we can find an explanation of this in the general relations of the single individual to its species, and to the whole of the external conditions of life; and this is the explanation which I have suggested and have attempted to work out in the text.
Note 9. Death by Sudden Shock
The most remarkable example of this kind of death known to me, is that of the male bees. It has been long known that the drone perishes while pairing, and it was usually believed that the queen bites it to death. Later observations have however shown that this is not the case, but that the male suddenly dies during copulation, and that the queen afterwards bites through the male intromittent organ, in order to free herself from the dead body. In this case death is obviously due to sudden excitement, for when the latter is artificially induced, death immediately follows. Von Berlepsch made some very interesting observations on this point, ‘If one catches a drone by the wings, during the nuptial flight, and holds it free in the air without touching any other part, the penis is protruded and the animal instantly dies, becoming motionless as though killed by a shock. The same thing happens if one gently stimulates the dorsal surface of the drone on a similar occasion. The male is in such an excited and irritable condition that the slightest muscular movement or disturbance causes the penis to be protruded27.’ In this case death is caused by the so-called nervous shock. The humble-bees are not similarly constituted, for the male does not die after fertilizing the female, ‘but withdraws its penis and flies away.’ But the death of male bees, during pairing, must not be regarded as normal death. Experiment has shown that these insects can live for more than four months28. They do not, as a matter of fact, generally live so long; for—although the workers do not, as was formerly believed, kill them after the fertilization of the queen, by direct means—they prevent them from eating the honey and drive them from the hive, so that they die of hunger29.
We must also look upon death which immediately, or very quickly, follows upon the deposition of eggs as death by sudden shock. The females of certain species of Psychidae, when they reproduce sexually, may remain alive for more than a week waiting for a male: after fertilization, however, they lay their eggs and die, while the parthenogenetic females of the same species lay their eggs and die immediately after leaving the cocoon; so that while the former live for many days, the latter do not last for more than twenty-four hours. ‘The parthenogenetic form of Solenobia triquetrella, soon after emergence, lays all her eggs together in the empty case, becomes much shrunken, and dies in a few hours.’ (Letter from Dr. Speyer, Rhoden.)
Note 10. Intermingling during the Fission of Unicellular Organisms30
Fission is quite symmetrical in Amoebae, so that it is impossible to recognise mother and daughter in the two resulting organisms. But in Euglypha and allied forms the existence of a shell introduces a distinguishing mark by which it is possible to discriminate between the products of fission; so that the offspring can be differentiated from the parent. The parent organism, before division, builds the parts of the shell for the daughter form. These parts are arranged on the surface of that part of the protoplasm, external to the old shell, which will be subsequently separated as the daughter-cell. On this part the spicules are arranged and unite to form the new shell. The division of the nucleus takes place after that of the protoplasm, so that the daughter-cell is for some time without a nucleus. Although we can in this species recognise the daughter-cell for