Reversion, in the ordinary sense of the word, acts so incessantly, that it evidently forms an essential part of the general law of inheritance. It occurs with beings, however propagated, whether by buds or seminal generation, and sometimes may be observed with advancing age even in the same individual. The tendency to reversion is often induced by a change of conditions, and in the plainest manner by crossing. Crossed forms of the first generation are generally nearly intermediate in character between their two parents; but in the next generation the offspring commonly revert to one or both of their grandparents, and occasionally to more remote ancestors. How can we account for these facts? Each unit in a hybrid must throw off, according to the doctrine of pangenesis, an abundance of hybridised gemmules, for crossed plants can be readily and largely propagated by buds; but by the same hypothesis dormant gemmules derived from both pure parent-forms are likewise present; and as these gemmules retain their normal condition, they would, it is probable, be enabled to multiply largely during the lifetime of each hybrid. Consequently the sexual elements of a hybrid will include both pure and hybridised gemmules; and when two hybrids pair, the combination of pure gemmules derived from the one hybrid with the pure gemmules of the same parts derived from the other, would necessarily lead to complete reversion of character; and it is, perhaps, not too bold a supposition that unmodified and undeteriorated gemmules of the same nature would be especially apt to combine. Pure gemmules in combination with hybridised gemmules would lead to partial reversion. And lastly, hybridised gemmules derived from both parent-hybrids would simply reproduce the original hybrid form.[[71]] All these cases and degrees of reversion incessantly occur.

It was shown in the fifteenth chapter that certain characters are antagonistic to each other or do not readily blend; hence, when two animals with antagonistic characters are crossed, it might well happen that a sufficiency of gemmules in the male alone for the reproduction of his peculiar characters, and in the female alone for the reproduction of her peculiar characters, would not be present; and in this case dormant gemmules derived from the same part in some remote progenitor might easily gain the ascendancy, and cause the reappearance of the long-lost character. For instance, when black and white pigeons, or black and white fowls, are crossed,—colours which do not readily blend,—blue plumage in the one case, evidently derived from the rock-pigeon, and red plumage in the other case, derived from the wild jungle-cock, occasionally reappear. With uncrossed breeds the same result follows, under conditions which favour the multiplication and development of certain dormant gemmules, as when animals become feral and revert to their pristine character. A certain number of gemmules being requisite for the development of each character, as is known to be the case from several spermatozoa or pollen-grains being necessary for fertilisation, and time favouring their multiplication, will perhaps account for the curious cases, insisted on by Mr. Sedgwick, of certain diseases which regularly appear in alternate generations. This likewise holds good, more or less strictly, with other weakly inherited modifications. Hence, as I have heard it remarked, certain diseases appear to gain strength by the intermission of a generation. The transmission of dormant gemmules during many successive generations is hardly in itself more improbable, as previously remarked, than the retention during many ages of rudimentary organs, or even only of a tendency to the production of a rudiment; but there is no reason to suppose that dormant gemmules can be transmitted and propagated for ever. Excessively minute and numerous as they are believed to be, an infinite number derived, during a long course of modification and descent, from each unit of each progenitor, could not be supported or nourished by the organism. But it does not seem improbable that certain gemmules, under favourable conditions, should be retained and go on multiplying for a much longer period than others. Finally, on the view here given, we certainly gain some insight into the wonderful fact that the child may depart from the type of both its parents, and resemble its grandparents, or ancestors removed by many hundreds of generations.

Conclusion.

The hypothesis of Pangenesis, as applied to the several great classes of facts just discussed, no doubt is extremely complex, but so are the facts. The chief assumption is that all the units of the body, besides having the universally admitted power of growing by self-division, throw off minute gemmules which are dispersed through the system. Nor can this assumption be considered as too bold, for we know from the cases of graft-hybridisation that formative matter of some kind is present in the tissues of plants, which is capable of combining with that included in another individual, and of reproducing every unit of the whole organism. But we have further to assume that the gemmules grow, multiply, and aggregate themselves into buds and the sexual elements; their development depending on their union with other nascent cells or units. They are also believed to be capable of transmission in a dormant state, like seeds in the ground, to successive generations.

In a highly-organised animal, the gemmules thrown off from each different unit throughout the body must be inconceivably numerous and minute. Each unit of each part, as it changes during development, and we know that some insects undergo at least twenty metamorphoses, must throw off its gemmules. But the same cells may long continue to increase by self-division, and even become modified by absorbing peculiar nutriment, without necessarily throwing off modified gemmules. All organic beings, moreover, include many dormant gemmules derived from their grandparents and more remote progenitors, but not from all their progenitors. These almost infinitely numerous and minute gemmules are contained within each bud, ovule, spermatozoon, and pollen-grain. Such an admission will be declared impossible; but number and size are only relative difficulties. Independent organisms exist which are barely visible under the highest powers of the microscope, and their germs must be excessively minute. Particles of infectious matter, so small as to be wafted by the wind or to adhere to smooth paper, will multiply so rapidly as to infect within a short time the whole body of a large animal. We should also reflect on the admitted number and minuteness of the molecules composing a particle of ordinary matter. The difficulty, therefore, which at first appears insurmountable, of believing in the existence of gemmules so numerous and small as they must be according to our hypothesis, has no great weight.

The units of the body are generally admitted by physiologists to be autonomous. I go one step further and assume that they throw off reproductive gemmules. Thus an organism does not generate its kind as a whole, but each separate unit generates its kind. It has often been said by naturalists that each cell of a plant has the potential capacity of reproducing the whole plant; but it has this power only in virtue of containing gemmules derived from every part. When a cell or unit is from some cause modified, the gemmules derived from it will be in like manner modified. If our hypothesis be provisionally accepted, we must look at all the forms of asexual reproduction, whether occurring at maturity or during youth, as fundamentally the same, and dependent on the mutual aggregation and multiplication of the gemmules. The re-growth of an amputated limb and the healing of a wound is the same process partially carried out. Buds apparently include nascent cells, belonging to that stage of development at which the budding occurs, and these cells are ready to unite with the gemmules derived from the next succeeding cells. The sexual elements, on the other hand, do not include such nascent cells; and the male and female elements taken separately do not contain a sufficient number of gemmules for independent development, except in the cases of parthenogenesis. The development of each being, including all the forms of metamorphosis and metagenesis, depends on the presence of gemmules thrown off at each period of life, and on their development, at a corresponding period, in union with preceding cells. Such cells may be said to be fertilised by the gemmules which come next in due order of development. Thus the act of ordinary impregnation and the development of each part in each being are closely analogous processes. The child, strictly speaking, does not grow into the man, but includes germs which slowly and successively become developed and form the man. In the child, as well as in the adult, each part generates the same part. Inheritance must be looked at as merely a form of growth, like the self-division of a lowly-organised unicellular organism. Reversion depends on the transmission from the forefather to his descendants of dormant gemmules, which occasionally become developed under certain known or unknown conditions. Each animal and plant may be compared with a bed of soil full of seeds, some of which soon germinate, some lie dormant for a period, whilst others perish. When we hear it said that a man carries in his constitution the seeds of an inherited disease, there is much truth in the expression. No other attempt, as far as I am aware, has been made, imperfect as this confessedly is, to connect under one point of view these several grand classes of facts. An organic being is a microcosm—a little universe, formed of a host of self-propagating organisms, inconceivably minute and numerous as the stars in heaven.

REFERENCES

[1] This hypothesis has been severely criticised by many writers, and it will be fair to give references to the more important articles. The best essay which I have seen is by Prof. Delpino, entitled ‘Sulla Darwiniana Teoria della Pangenesi, 1869,’ of which a translation appeared in ‘Scientific Opinion,’ Sept. 29th, 1869, and the succeeding numbers. He rejects the hypothesis, but criticises it fairly, and I have found his criticisms very useful. Mr. Mivart (‘Genesis of Species,’ 1871, chap. x.) follows Delpino, but adds no new objections of any weight. Dr. Bastian (‘The Beginnings of Life,’ 1872, vol. ii. p. 98) says that the hypothesis “looks like a relic of the old rather than a fitting appanage of the new evolution philosophy.” He shows that I ought not to have used the term “pangenesis,” as it had been previously used by Dr. Gros in another sense. Dr. Lionel Beale (‘Nature,’ May 11th, 1871, p. 26) sneers at the whole doctrine with much acerbity and some justice. Prof. Wigand (‘Schriften der Gesell. der gesammt. Naturwissen. zu Marburg,’ B. ix. 1870) considers the hypothesis as unscientific and worthless. Mr. G. H. Lewes (‘Fortnightly Review,’ Nov. 1st, 1868, p. 503) seems to consider that it may be useful: he makes many good criticisms in a perfectly fair spirit. Mr. F. Galton, after describing his valuable experiments (‘Proc. Royal Soc.,’ vol. xix. p. 393) on the intertransfusion of the blood of distinct varieties of the rabbit, concludes by saying that in his opinion the results negative beyond all doubt the doctrine of Pangenesis. He informs me that subsequently to the publication of his paper he continued his experiments on a still larger scale for two more generations, without any sign of mongrelism showing itself in the very numerous offspring. I certainly should have expected that gemmules would have been present in the blood, but this is no necessary part of the hypothesis, which manifestly applies to plants and the lowest animals. Mr. Galton, in a letter to ‘Nature’ (April 27th, 1871, p. 502), also criticises various incorrect expressions used by me. On the other hand, several writers have spoken favourably of the hypothesis, but there would be no use in giving references to their articles. I may, however, refer to Dr. Ross’ work, ‘The Graft Theory of Disease; being an application of Mr. Darwin’s hypothesis of Pangenesis,’ 1872, as he gives several original and ingenious discussions.

[2] Quoted by Paget, ‘Lectures on Pathology,’ 1853, p. 159.

[3] Dr. Lachmann, also, observes (‘Annals and Mag. of Nat. History,’ 2nd series, vol. xix. 1857, p. 231) with respect to infusoria, that “fissation and gemmation pass into each other almost imperceptibly.” Again, Mr. W. C. Minor (‘Annals and Mag. of Nat. Hist.,’ 3rd series, vol. xi. p. 328) shows that with Annelids the distinction that has been made between fission and budding is not a fundamental one. See also Professor Clark’s work ‘Mind in Nature,’ New York, 1865, pp. 62, 94.