The Variation of Animals and Plants under Domestication — Volume 2

The principle of the independent formation of each part, owing to the union of the proper gemmules with certain nascent cells, together with the superabundance of the gemmules derived from both parents, and the subsequent self-multiplication of the gemmules, throws light on a widely different group of facts, which on any ordinary view of development appears very strange. I allude to organs which are abnormally transposed or multiplied. For instance, a curious case has been recorded by Dr. Elliott Coues (27/58. 'Proc. Boston Soc. of Nat. Hist.' republished in 'Scientific Opinion' November 10, 1869 page 488.) of a monstrous chicken with a perfect additional RIGHT leg articulated to the LEFT side of the pelvis. Gold-fish often have supernumerary fins placed on various parts of their bodies. When the tail of a lizard is broken off, a double tail is sometimes reproduced; and when the foot of the salamander was divided longitudinally by Bonnet, additional digits were occasionally formed. Valentin injured the caudal extremity of an embryo, and three days afterwards it produced rudiments of a double pelvis and of double hind-limbs. (27/59. Todd 'Cyclop. of Anat. and Phys.' volume 4 1849-52 page 975.) When frogs, toads, etc., are born with their limbs doubled, as sometimes happens, the doubling, as Gervais remarks (27/60. 'Compte Rendus' November 14, 1865 page 800.), cannot be due to the complete fusion of two embryos, with the exception of the limbs, for the larvae are limbless. The same argument is applicable (27/61. As previously remarked by Quatrefages in his 'Metamorphoses de l'Homme' etc. 1862 page 129.) to certain insects produced with multiple legs or antennae, for these are metamorphosed from apodal or antennae-less larvae. Alphonse Milne-Edwards (27/62. Gunther 'Zoological Record' 1864 page 279.) has described the curious case of a crustacean in which one eye-peduncle supported, instead of a complete eye, only an imperfect cornea, and out of the centre of this a portion of an antenna was developed. A case has been recorded (27/63. Sedgwick 'Medico-Chirurg. Review' April 1863 page 454.) of a man who had during both dentitions a double tooth in place of the left second incisor, and he inherited this peculiarity from his paternal grandfather. Several cases are known (27/64. Isid. Geoffroy Saint-Hilaire 'Hist. des Anomalies' tome 1 1832 pages 435, 657; and tome 2 page 560.) of additional teeth having been developed in the orbit of the eye, and, more especially with horses, in the palate. Hairs occasionally appear in strange situations, as "within the substance of the brain." (27/65. Virchow 'Cellular Pathology' 1860 page 66.) Certain breeds of sheep bear a whole crowd of horns on their foreheads. As many as five spurs have been seen on both legs of certain Game-fowls. In the Polish fowl the male is ornamented with a topknot of hackles like those on his neck, whilst the female has a top-knot formed of common feathers. In feather- footed pigeons and fowls, feathers like those on the wing arise from the outer side of the legs and toes. Even the elemental parts of the same feather may be transposed; for in the Sebastopol goose, barbules are developed on the divided filaments of the shaft. Imperfect nails sometimes appear on the stumps of the amputated fingers of man (27/66. Muller 'Phys.' English Translation volume 1 1833 page 407. A case of this kind has lately been communicated to me.) and it is an interesting fact that with the snake-like Saurians, which present a series with more and more imperfect limbs, the terminations of the phalanges first disappear, "the nails becoming transferred to their proximal remnants, or even to parts which are not phalanges." (27/67. Dr. Furbringer 'Die Knochen etc. bei den schlangenahnlichen Sauriern' as reviewed in 'Journal of Anat. and Phys.' May 1870 page 286.)

Analogous cases are of such frequent occurrence with plants that they do not strike us with sufficient surprise. Supernumerary petals, stamens, and pistils, are often produced. I have seen a leaflet low down in the compound leaf of Vicia sativa replaced by a tendril; and a tendril possesses many peculiar properties, such as spontaneous movement and irritability. The calyx sometimes assumes, either wholly or by stripes, the colour and texture of the corolla. Stamens are so frequently converted into petals, more or less completely, that such cases are passed over as not deserving notice; but as petals have special functions to perform, namely, to protect the included organs, to attract insects, and in not a few cases to guide their entrance by well-adapted contrivances, we can hardly account for the conversion of stamens into petals merely by unnatural or superfluous nourishment. Again, the edge of a petal may occasionally be found including one of the highest products of the plant, namely, pollen; for instance, I have seen the pollen-mass of an Ophrys, which is a very complex structure, developed in the edge of an upper petal. The segments of the calyx of the common pea have been observed partially converted into carpels, including ovules, and with their tips converted into stigmas. Mr. Salter and Dr. Maxwell Masters have found pollen within the ovules of the passion-flower and of the rose. Buds may be developed in the most unnatural positions, as on the petal of a flower. Numerous analogous facts could be given. (27/68. Moquin-Tandon 'Teratologie Veg.' 1841 pages 218, 220, 353. For the case of the pea see 'Gardener's Chronicle' 1866 page 897. With respect to pollen within ovules see Dr. Masters in 'Science Review' October 1873 page 369. The Rev. J.M. Berkeley describes a bud developed on a petal of a Clarkia in 'Gardener's Chronicle' April 28, 1866.)

I do not know how physiologists look at such facts as the foregoing. According to the doctrine of pangenesis, the gemmules of the transposed organs become developed in the wrong place, from uniting with wrong cells or aggregates of cells during their nascent state; and this would follow from a slight modification in their elective affinities. Nor ought we to feel much surprise at the affinities of cells and gemmules varying, when we remember the many curious cases given in the seventeenth chapter, of plants which absolutely refuse to be fertilised by their own pollen, though abundantly fertile with that of any other individual of the same species, and in some cases only with that of a distinct species. It is manifest that the sexual elective affinities of such plants — to use the term employed by Gartner — have been modified. As the cells of adjoining or homologous parts will have nearly the same nature, they will be particularly liable to acquire by variation each other's elective affinities; and we can thus understand to a certain extent such cases as a crowd of horns on the heads of certain sheep, of several spurs on the legs of fowls, hackle-like feathers on the heads of the males of other fowls, and with the pigeon wing-like feathers on their legs and membrane between their toes, for the leg is the homologue of the wing. As all the organs of plants are homologous and spring from a common axis, it is natural that they should be eminently liable to transposition. It ought to be observed that when any compound part, such as an additional limb or an antenna, springs from a false position, it is only necessary that the few first gemmules should be wrongly attached; for these whilst developing would attract other gemmules in due succession, as in the regrowth of an amputated limb. When parts which are homologous and similar in structure, as the vertebrae of snakes or the stamens of polyandrous flowers, etc., are repeated many times in the same organism, closely allied gemmules must be extremely numerous, as well as the points to which they ought to become united; and, in accordance with the foregoing views, we can to a certain extent understand Isid. Geoffroy Saint-Hilaire's law, that parts, which are already multiple, are extremely liable to vary in number.

Variability often depends, as I have attempted to show, on the reproductive organs being injuriously affected by changed conditions; and in this case the gemmules derived from the various parts of the body are probably aggregated in an irregular manner, some superfluous and others deficient. Whether a superabundance of gemmules would lead to the increased size of any part cannot be told; but we can see that their partial deficiency, without necessarily leading to the entire abortion of the part, might cause considerable modifications; for in the same manner as plants, if their own pollen be excluded, are easily hybridised, so, in the case of cells, if the properly succeeding gemmules were absent, they would probably combine easily with other and allied gemmules, as we have just seen with transposed parts.

In variations caused by the direct action of changed conditions, of which several instances have been given, certain parts of the body are directly affected by the new conditions, and consequently throw off modified gemmules, which are transmitted to the offspring. On any ordinary view it is unintelligible how changed conditions, whether acting on the embryo, the young or the adult, can cause inherited modifications. It is equally or even more unintelligible on any ordinary view, how the effects of the long-continued use or disuse of a part, or of changed habits of body or mind, can be inherited. A more perplexing problem can hardly be proposed; but on our view we have only to suppose that certain cells become at last structurally modified; and that these throw off similarly modified gemmules. This may occur at any period of development, and the modification will be inherited at a corresponding period; for the modified gemmules will unite in all ordinary cases with the proper preceding cells, and will consequently be developed at the same period at which the modification first arose. With respect to mental habits or instincts, we are so profoundly ignorant of the relation between the brain and the power of thought that we do not know positively whether a fixed habit induces any change in the nervous system, though this seems highly probable; but when such habit or other mental attribute, or insanity, is inherited, we must believe that some actual modification is transmitted (27/69. See some remarks to this effect by Sir H. Holland in his 'Medical Notes' 1839 page 32.); and this implies, according to our hypothesis, that gemmules derived from modified nerve-cells are transmitted to the offspring.

It is generally necessary that an organism should be exposed during several generations to changed conditions or habits, in order that any modification thus acquired should appear in the offspring. This may be partly due to the changes not being at first marked enough to catch attention, but this explanation is insufficient; and I can account for the fact only by the assumption, which we shall see under the head of reversion is strongly supported, that gemmules derived from each unmodified unit or part are transmitted in large numbers to successive generations, and that the gemmules derived from the same unit after it has been modified go on multiplying under the same favourable conditions which first caused the modification, until at last they become sufficiently numerous to overpower and supplant the old gemmules.

A difficulty may be here noticed; we have seen that there is an important difference in the frequency, though not in the nature, of the variations in plants propagated by sexual and asexual generation. As far as variability depends on the imperfect action of the reproductive organs under changed conditions, we can at once see why plants propagated asexually should be far less variable than those propagated sexually. With respect to the direct action of changed conditions, we know that organisms produced from buds do not pass through the earlier phases of development; they will therefore not be exposed, at that period of life when structure is most readily modified, to the various causes inducing variability in the same manner as are embryos and young larval forms; but whether this is a sufficient explanation I know not.

With respect to variations due to reversion, there is a similar difference between plants propagated from buds and seeds. Many varieties can be propagated securely by buds, but generally or invariably revert to their parent-forms by seed. So, also, hybridised plants can be multiplied to any extent by buds, but are continually liable to reversion by seed, — that is, to the loss of their hybrid or intermediate character. I can offer no satisfactory explanation of these facts. Plants with variegated leaves, phloxes with striped flowers, barberries with seedless fruit, can all be securely propagated by buds taken from the stem or branches; but buds from the roots of these plants almost invariably lose their character and revert to their former condition. This latter fact is also inexplicable, unless buds developed from the roots are as distinct from those on the stem, as is one bud on the stem from another, and we know that these latter behave like independent organisms.

Finally, we see that on the hypothesis of pangenesis variability depends on at least two distinct groups of causes. Firstly, the deficiency, superabundance, and transposition of gemmules, and the redevelopment of those which have long been dormant; the gemmules themselves not having undergone any modification; and such changes will amply account for much fluctuating variability. Secondly, the direct action of changed conditions on the organisation, and of the increased use or disuse of parts; and in this case the gemmules from the modified units will be themselves modified, and, when sufficiently multiplied, will supplant the old gemmules and be developed into new structures.

Turning now to the laws of Inheritance. If we suppose a homogeneous gelatinous protozoon to vary and assume a reddish colour, a minute separated particle would naturally, as it grew to full size, retain the same colour; and we should have the simplest form of inheritance. (27/70. This is the view taken by Prof. Hackel in his 'Generelle Morphologie' b. 2 s. 171, who says: "Lediglich die partielle Identitat der specifisch constituirten Materie im elterlichen und im kindlichen Organismus, die Theilung dieser Materie bei der Fortpflanzung, ist die Ursache der Erblichkeit.") Precisely the same view may be extended to the infinitely numerous and diversified units of which the whole body of one of the higher animals is composed; the separated particles being our gemmules. We have already sufficiently discussed by implication, the important principle of inheritance at corresponding ages. Inheritance as limited by sex and by the season of the year (for instance with animals becoming white in winter) is intelligible if we may believe that the elective affinities of the units of the body are slightly different in the two sexes, especially at maturity, and in one or both sexes at different seasons, so that they unite with different gemmules. It should be remembered that, in the discussion on the abnormal transposition of organs, we have seen reason to believe that such elective affinities are readily modified. But I shall soon have to recur to sexual and seasonal inheritance. These several laws are therefore explicable to a large extent through pangenesis, and on no other hypothesis which has as yet been advanced.

But it appears at first sight a fatal objection to our hypothesis that a part or organ may be removed during several successive generations, and if the operation be not followed by disease, the lost part reappears in the offspring. Dogs and horses formerly had their tails docked during many generations without any inherited effect; although, as we have seen, there is some reason to believe that the tailless condition of certain sheep-dogs is due to such inheritance. Circumcision has been practised by the Jews from a remote period, and in most cases the effects of the operation are not visible in the offspring; though some maintain that an inherited effect does occasionally appear. If inheritance depends on the presence of disseminated gemmules derived from all the units of the body, why does not the amputation or mutilation of a part, especially if effected on both sexes, invariably affect the offspring? The answer in accordance with our hypothesis probably is that gemmules multiply and are transmitted during a long series of generations — as we see in the reappearance of zebrine stripes on the horse — in the reappearance of muscles and other structures in man which are proper to his lowly organised progenitors, and in many other such cases. Therefore the long-continued inheritance of a part which has been removed during many generations is no real anomaly, for gemmules formerly derived from the part are multiplied and transmitted from generation to generation.

We have as yet spoken only of the removal of parts, when not followed by morbid action: but when the operation is thus followed, it is certain that the deficiency is sometimes inherited. In a former chapter instances were given, as of a cow, the loss of whose horn was followed by suppuration, and her calves were destitute of a horn on the same side of their heads. But the evidence which admits of no doubt is that given by Brown-Sequard with respect to guinea-pigs, which after their sciatic nerves had been divided, gnawed off their own gangrenous toes, and the toes of their offspring were deficient in at least thirteen instances on the corresponding feet. The inheritance of the lost part in several of these cases is all the more remarkable as only one parent was affected; but we know that a congenital deficiency is often transmitted from one parent alone — for instance, the offspring of hornless cattle of either sex, when crossed with perfect animals, are often hornless. How, then, in accordance with our hypothesis can we account for mutilations being sometimes strongly inherited, if they are followed by diseased action? The answer probably is that all the gemmules of the mutilated or amputated part are gradually attracted to the diseased surface during the reparative process, and are there destroyed by the morbid action.

A few words must be added on the complete abortion of organs. When a part becomes diminished by disuse prolonged during many generations, the principle of economy of growth, together with intercrossing, will tend to reduce it still further as previously explained, but this will not account for the complete or almost complete obliteration of, for instance, a minute papilla of cellular tissue representing a pistil, or of a microscopically minute nodule of bone representing a tooth. In certain cases of suppression not yet completed, in which a rudiment occasionally reappears through reversion, dispersed gemmules derived from this part must, according to our view, still exist; we must therefore suppose that the cells, in union with which the rudiment was formerly developed, fail in their affinity for such gemmules, except in the occasional cases of reversion. But when the abortion is complete and final, the gemmules themselves no doubt perish; nor is this in any way improbable, for, though a vast number of active and long-dormant gemmules are nourished in each living creature, yet there must be some limit to their number; and it appears natural that gemmules derived from reduced and useless parts would be more liable to perish than those freshly derived from other parts which are still in full functional activity.

The last subject that need be discussed, namely, Reversion, rests on the principle that transmission and development, though generally acting in conjunction, are distinct powers; and the transmission of gemmules with their subsequent development shows us how this is possible. We plainly see the distinction in the many cases in which a grandfather transmits to his grandson, through his daughter, characters which she does not, or cannot, possess. But before proceeding, it will be advisable to say a few words about latent or dormant characters. Most, or perhaps all, of the secondary characters, which appertain to one sex, lie dormant in the other sex; that is, gemmules capable of development into the secondary male sexual characters are included within the female; and conversely female characters in the male: we have evidence of this in certain masculine characters, both corporeal and mental, appearing in the female, when her ovaria are diseased or when they fail to act from old age. In like manner female characters appear in castrated males, as in the shape of the horns of the ox, and in the absence of horns in castrated stags. Even a slight change in the conditions of life due to confinement sometimes suffices to prevent the development of masculine characters in male animals, although their reproductive organs are not permanently injured. In the many cases in which masculine characters are periodically renewed, these are latent at other seasons; inheritance as limited by sex and season being here combined. Again, masculine characters generally lie dormant in male animals until they arrive at the proper age for reproduction. The curious case formerly given of a Hen which assumed the masculine characters, not of her own breed but of a remote progenitor, illustrates the close connection between latent sexual characters and ordinary reversion.

With those animals and plants which habitually produce several forms, as with certain butterflies described by Mr. Wallace, in which three female forms and one male form co-exist, or, as with the trimorphic species of Lythrum and Oxalis, gemmules capable of reproducing these different forms must be latent in each individual.

Insects are occasionally produced with one side or one quarter of their bodies like that of the male, with the other half or three-quarters like that of the female. In such cases the two sides are sometimes wonderfully different in structure, and are separated from each other by a sharp line. As gemmules derived from every part are present in each individual of both sexes, it must be the elective affinities of the nascent cells which in these cases differ abnormally on the two sides of the body. Almost the same principle comes into play with those animals, for instance, certain gasteropods and Verruca amongst cirripedes, which normally have the two sides of the body constructed on a very different plan; and yet a nearly equal number of individuals have either side modified in the same remarkable manner.

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. (27/71. In these remarks I, in fact, follow Naudin, who speaks of the elements or essences of the two species which are crossed. See his excellent memoir in the 'Nouvelles Archives du Museum' tome 1 page 151.) 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