Part 7

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 cells[26].

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 protruded[27].’ 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 months[28]. 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 hunger[29].

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 Organisms[30].

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 some time after separation from the parent by the greater transparency of its younger shell, it is nevertheless impossible to admit that the characteristics of the two animals are in any way different, for just before the separation of the two individuals a circulation of the protoplasm through both shells takes place after the manner described in the text, and there is therefore a complete intermingling of the substance of the two bodies.

The difference between the products is even greater after transverse fission of the _Infusoria_, for a new anus must be formed at the anterior part and a new mouth posteriorly. It is not known whether any circulation of the protoplasm takes place, as in _Euglypha_. But even if this does not occur, there is no reason for believing that the two products of division possess a different duration of life.

The process of fission in the _Diatomaceae_ seems to me to be theoretically important, because here, as in the previously-mentioned _Monothalamia_ (_Euglypha_, etc.), the new silicious skeleton is built up within the primary organism, but not, as in _Euglypha_, for the new individual only, but for both parent and daughter-cell alike[31]. If we compare the diatom shell to a box, then the two halves of the old shell would form two lids, one for each of the products of fission, while a new box is built up afresh for each of them. In this case there is an absolute equality between the products of fission, so far as the shell is concerned.

Note 11. Regeneration.

A number of experiments have been recently undertaken, in connection with a prize thesis at Würzburg, in order to test the powers of regeneration possessed by various animals. In all essential respects the results confirm the statements of the older observers, such as Spallanzani. Carrière has also proved that snails can regenerate not only their horns and eyes, but also part of the head when it has been cut off, although he has shown that Spallanzani's old statement that they can regenerate the whole head, including the nervous system, is erroneous[32].

Note 12. The Duration of Life in Plants.

The title of the work on this subject mentioned in the Text is ‘Die Lebensdauer und Vegetationsweise der Pflanzen, ihre Ursache und ihre Entwicklung,’ F. Hildebrand, Engler’s botanische Jahrbücher, Bd. II. 1. und 2. Heft, Leipzig, 1881.

Note 13.

[Many interesting facts and conclusions upon the subject of this essay will be found in a volume by Professor E. Ray Lankester, ‘On comparative Longevity in Man and the lower Animals,’ Macmillan and Co., 1870.—E. B. P.]

Footnotes for the Appendix to Essay I.

Footnote 1:

Humboldt’s ‘Ausichten der Natur.’

Footnote 2:

This estimate is derived from observation of the time during which these insects are to be seen upon the wing. Direct observations upon the duration of life in this species are unknown to me.

Footnote 3:

[Sir John Lubbock has now kept a queen ant alive for nearly 15 years. See note 2 {note 18 below} on p. 51.—E. B. P.]

Footnote 4:

[After reading these proofs Dr. A. R. Wallace kindly sent me an unpublished note upon the production of death by means of natural selection, written by him some time between 1865 and 1870. The note contains some ideas on the subject, which were jotted down for further elaboration, and were then forgotten until recalled by the argument of this Essay. The note is of great interest in relation to Dr. Weismann’s suggestions, and with Dr. Wallace’s permission I print it in full below.

‘The Action of Natural Selection in Producing Old Age, Decay, and Death.

‘Supposing organisms ever existed that had not the power of natural reproduction, then since the absorptive surface would only increase as the square of the dimensions while the bulk to be nourished and renewed would increase as the cube, there must soon arrive a limit of growth. Now if such an organism did not produce its like, accidental destruction would put an end to the species. Any organism therefore that, by accidental or spontaneous fission, could become two organisms, and thus multiply itself indefinitely without increasing in size beyond the limits most favourable for nourishment and existence, could not be thus exterminated: since the individual only could be accidentally destroyed,—the race would survive. But if individuals did not die they would soon multiply inordinately and would interfere with each other’s healthy existence. Food would become scarce, and hence the larger individuals would probably decompose or diminish in size. The deficiency of nourishment would lead to parts of the organism not being renewed; they would become fixed, and liable to more or less slow decomposition as dead parts within a living body. The smaller organisms would have a better chance of finding food, the larger ones less chance. That one which gave off several small portions to form each a new organism would have a better chance of leaving descendants like itself than one which divided equally or gave off a large part of itself. Hence it would happen that those which gave off very small portions would probably soon after cease to maintain their own existence while they would leave a numerous offspring. This state of things would be in any case for the advantage of the race, and would therefore, by natural selection, soon become established as the regular course of things, and thus we have the origin of _old age_, _decay_, and _death_; for it is evident that when one or more individuals have provided a sufficient number of successors they themselves, as consumers of nourishment in a constantly increasing degree, are an injury to those successors. Natural selection therefore weeds them out, and in many cases favours such races as die almost immediately after they have left successors. Many moths and other insects are in this condition, living only to propagate their kind and then immediately dying, some not even taking any food in the perfect and reproductive state.’—E. B. P.]

Footnote 5:

Johannes Müller, ‘Physiologie,’ Bd. I. p. 31, Berlin, 1840.

Footnote 6:

Oken, ‘Naturgeschichte,’ Stuttgart, 1837, Bd. IV. Abth. 1.

Footnote 7:

Brehm, ‘Leben der Vögel,’ p. 278.

Footnote 8:

‘Naturwissenschaftliche Thatsachen und Probleme,’ Populäre Vorträge, Berlin, 1880; _vide_ Appendix.

Footnote 9:

‘Entomolog. Mag.,’ vol. i. p. 527, 1833.

Footnote 10:

Imhof, ‘Beiträge zur Anatomie der _Perla maxima_,’ Inaug. Diss., Aarau, 1881.

Footnote 11:

Mr. Edwards has meanwhile published these communications in full; cf. ‘On the length of life of Butterflies,’ Canadian Entomologist, 1881, p. 205.

Footnote 12:

When no authority is given, the observations are my own.

Footnote 13:

In the paper quoted above, Edwards, after weighing all the evidence, reduces the length of life from three to four weeks.

Footnote 14:

‘Entomolog. Mag.,’ vol. i. p. 527, 1823.

Footnote 15:

Ibid.

Footnote 16:

Ibid.

Footnote 17:

‘Recherches sur les mœurs des Fourmis indigènes,’ Genève, 1810.

Footnote 18:

These two female ants were still alive on the 25th of September following Sir John Lubbock’s letter, so that they live at least seven years. Cf. ‘Observations on Ants, Bees, and Wasps,’ Part VIII. p. 385; Linn. Soc. Journ. Zool., vol. xv. 1881.

[Sir John Lubbock has kindly given me further information upon the duration of life of these two queen ants. Since the receipt of his letter, the facts have been published in the Journal of the Linnean Society (Zoology), vol. xx. p. 133. I quote in full the passage which refers to these ants:—

‘Longevity.—It may be remembered that my nests have enabled me to keep ants under observation for long periods, and that I have identified workers of _Lasius niger_ and _Formica fusca_ which were at least seven years old, and two queens of _Formica fusca_ which have lived with me ever since December 1874. One of these queens, after ailing for some days, died on the 30th July, 1887. She must then have been more than thirteen years old. I was at first afraid that the other one might be affected by the death of her companion. She lived, however, until the 8th August, 1888, when she must have been nearly fifteen years old, and is therefore by far the oldest insect on record.

‘Moreover, what is very extraordinary, she continued to lay fertile eggs. This remarkable fact is most interesting from a physiological point of view. Fertilization took place in 1874 at the latest. There has been no male in the nest since then, and, moreover, it is, I believe, well established that queen ants and queen bees are fertilized once for all. Hence the spermatozoa of 1874 must have retained their life and energy for thirteen years, a fact, I believe, unparalleled in physiology.’

* * * * *

‘I had another queen of _Formica fusca_ which lived to be thirteen years old, and I have now a queen of _Lasius niger_ which is more than nine years old, and still lays fertile eggs, which produce female ants.’

Both the above-mentioned queens may have been considerably older, for it is impossible to estimate their age at the time of capture. It is only certain (as Sir John Lubbock informs me in his letter) that they must have been at least nine months old (when captured), as the eggs of _F. fusca_ are laid in March or early in April.’ The queens became gradually ‘somewhat lethargic and stiff in their movements (before their death), but there was no loss of any limb nor any abrasion.’ This last observation seems to indicate that queen ants may live for a much longer period in the wild state, for it is stated above that the chitin is often greatly worn, and some of the limbs lost (see pp. 48, 51, and 52).—E. B. P.]

Footnote 19:

A. von Berlepsch, ‘Die Biene und ihre Zucht,’ etc., 3rd ed.; Mannheim, 1872.

Footnote 20:

E. Bevan, ‘Ueber die Honigbiene und die Länge ihres Lebens;’ abstract in Oken’s ‘Isis,’ 1844, p. 506.

Footnote 21:

Dalyell, ‘Rare and Remarkable Animals of Scotland,’ vol. ii. p. 203; London, 1848.

Footnote 22:

[Mr. J. S. Haldane has kindly obtained details of the death of the sea anemone referred to by the author. It died, by a natural death, on August 4, 1887, after having appeared to become gradually weaker for some months previous to this date. It had lived ever since 1828 in the same small glass jar in which it was placed by Sir John Dalyell. It must have been at least 66 years old when it died.—E.B.P.]

Footnote 23:

Bronn, ‘Klassen und Ordnungen des Thierreichs,’ Bd. III. p. 466; Leipzig.

Footnote 24:

Bronn, l. c.

Footnote 25:

Cf. the article ‘Mort’ in the ‘Encyclop. Scienc. Méd.’ vol. M. p. 520.

Footnote 26:

Roux, in his work ‘Der Kampf der Theile im Organismus,’ Jena 1881, has attempted to explain the manner in which division of labour has arisen among the cells of the higher organisms, and to render intelligible the mechanical processes by which the purposeful adaptations of the organism have arisen.

Footnote 27:

von Berlepsch, ‘Die Biene und ihre Zucht,’ etc.

Footnote 28:

Oken, ‘Isis,’ 1844, p. 506.

Footnote 29:

von Berlepsch, l. c., p. 165.

Footnote 30:

Cf. August Gruber, ‘Der Theilungsvorgang bei Euglypha alveolata,’ and ‘Die Theilung der monothalamen Rhizopoden,’ Z. f. W. Z., Bd. XXXV. and XXXVI., p. 104, 1881.

Footnote 31:

Cf. Victor Hensen, ‘Physiologie d. Zeugung,’ p. 152.

Footnote 32:

Cf. J. Carrière, ‘Ueber Regeneration bei Landpulmonaten,’ Tagebl. der 52. Versammlg. deutsch. Naturf. pp. 225-226.

II.

ON HEREDITY.

1883.

ON HEREDITY.

PREFACE.

The following essay was my inaugural lecture as Pro-Rector of the University of Freiburg, and was delivered publicly in the hall of the University, on June 21, 1883; it first appeared in print in the following August. Only a few copies of the first edition were available for the public, and it is therefore now reprinted as a second edition, which only differs from the first in a few not unimportant improvements and additions.

The title which I have chosen requires some explanation. I do not propose to treat of the whole problem of heredity, but only of a certain aspect of it—the transmission of acquired characters which has been hitherto assumed to occur. In taking this course I may say that it was impossible to avoid going back to the foundation of all the phenomena of heredity, and to determine the substance with which they must be connected. In my opinion this can only be the substance of the germ-cells; and this substance transfers its hereditary tendencies from generation to generation, at first unchanged, and always uninfluenced in any corresponding manner, by that which happens during the life of the individual which bears it. If these views, which are indicated rather than elaborated in this paper, be correct, all our ideas upon the transformation of species require thorough modification, for the whole principle of evolution by means of exercise (use and disuse), as proposed by Lamarck, and accepted in some cases by Darwin, entirely collapses.

The nature of the present paper—which is a lecture and not an elaborate treatise—necessitates that only suggestions and not an exhaustive treatment of the subject could be given. I have also abstained from giving further details in the form of an appendix, chiefly because I could hardly have attempted to complete a treatment of the whole range of the subject, and I hope to refer again to these questions in the future, when new experiments and observations have been made.

I am very glad to see that such an important authority as Pflüger[33] has in the meantime come to the same opinion, from an entirely different direction—an opinion which forms the foundation of the views here brought forward, namely, that heredity depends upon the continuity of the molecular substance of the germ from generation to generation.

A. W.

II.

ON HEREDITY.

With your permission I wish to bring before you to-day my views on a problem of general biological interest—the problem of heredity.

Heredity is the process which renders possible that persistence of organic beings throughout successive generations, which is generally thought to be so well understood and to need no special explanation. Nevertheless our minds cannot fail to be much perplexed by the multiplicity of its manifestations, and to be greatly puzzled as to its real nature. A celebrated German physiologist says[34], ‘Although many hands have at all times endeavoured to break the seal which hides the theory of heredity from our view, the results achieved have been but small; and we are in a certain degree justified in looking with little hope upon new efforts undertaken in this direction. We must nevertheless endeavour from time to time to ascertain how far we have advanced towards a complete explanation.’

Such a course is in every way advisable, for we are not dealing with phenomena which from their very nature are incomprehensible by man. The great complexity of the subject has alone rendered it hitherto insuperable, but in the province of heredity we certainly have not reached the limits of attainable knowledge.

From this point of view heredity bears some resemblance to certain anatomical and physiological problems, e. g. the structure and function of the human brain. Its structure—with so many millions of nerve-fibres and nerve-cells—is of such extraordinary complexity that we might well despair of ever completely understanding it. Each fibre is nevertheless distinct in itself, while its connection with the nearest nerve-cell can be frequently traced, and the function of many groups of cell elements is already known. But it would seem to be impossible to unravel the excessively complex network into which the cells and fibres are knit together; and hence to arrive at the function of each single element appears to be also beyond our reach. We have not however commenced to untie this Gordian knot without some hope of success, for who can say how far human perseverance may be able to penetrate into the mechanism of the brain, and to reveal a connected structure and a common principle in its countless elements? But surely this work will be most materially assisted by the simultaneous investigation of the structure and function of the nervous system in the lower forms of life—in the polypes and jelly-fish, worms and Crustacea. In the same way we should not abandon the hope of arriving at a satisfactory knowledge of the processes of heredity, if we consider the simplest processes of the lower animals as well as the more complex processes met with in the higher forms.