The Variation of Animals and Plants under Domestication · Charles Darwin
PART II.
Crossed forms sometimes exhibit the colour or other characters of either parent in stripes or blotches; and this occurs in the first generation, or through reversion in succeeding bud and seminal generations, of which fact several instances were given in the eleventh chapter. In these cases we must follow Naudin[57] and admit that the “essence” or “element” of the two species,—terms which I should translate into the gemmules,—have an affinity for their own kind, and thus separate themselves into distinct stripes or blotches; and reasons were given, when discussing in the fifteenth chapter the incompatibility of certain characters to unite, for believing in such mutual affinity. When two forms are crossed, one is not rarely found to be prepotent in the transmission of its characters over the other; and this we can explain by again assuming that the one form has some advantage over the other in the number, vigour, or affinity of its gemmules. In some cases, however, certain characters are present in the one form and latent in the other; for instance, there is a latent tendency in all pigeons to become blue, and, when a blue pigeon is crossed with one of any other colour, the blue tint is generally prepotent. The explanation of this form of prepotency will be obvious when we come to the consideration of Reversion.
When two distinct species are crossed, it is notorious that they do not yield the full or proper number of offspring; and we can only say on this head that, as the development of each organism depends on such nicely-balanced affinities between a host of gemmules and nascent cells, we need not feel at all surprised that the commixture of gemmules derived from two distinct species should lead to partial or complete failure of development. With respect to the sterility of hybrids produced from the union of two distinct species, it was shown in the nineteenth chapter that this depends exclusively on the reproductive organs being specially affected; but why these organs should be thus affected we do not know, any more than why unnatural conditions of life, though compatible with health, should cause sterility; or why continued close interbreeding, or the illegitimate unions of heterostyled plants, induce the same result. The conclusion that the reproductive organs alone are affected, and not the whole organisation, agrees perfectly with the unimpaired or even increased capacity in hybrid plants for propagation by buds; for this implies, according to our hypothesis, that the cells of the hybrids throw off hybridised gemmules, which become aggregated into buds, but fail to become aggregated within the reproductive organs, so as to form the sexual elements. In a similar manner many plants, when placed under unnatural conditions, fail to produce seed, but can readily be propagated by buds. We shall presently see that pangenesis agrees well with the strong tendency to reversion exhibited by all crossed animals and plants.
Each organism reaches maturity through a longer or shorter course of growth and development: the former term being confined to mere increase of size, and development to changed structure. The changes may be small and insensibly slow, as when a child grows into a man, or many, abrupt, and slight, as in the metamorphoses of certain ephemerous insects, or, again, few and strongly-marked, as with most other insects. Each newly formed part may be moulded within a previously existing and corresponding part, and in this case it will appear, falsely as I believe, to be developed from the old part; or it may be formed within a distinct part of the body, as in the extreme cases of metagenesis. An eye, for instance, may be developed at a spot where no eye previously existed. We have also seen that allied organic beings in the course of their metamorphoses sometimes attain nearly the same structure after passing through widely different forms; or conversely, after passing through nearly the same early forms, arrive at widely different mature forms. In these cases it is very difficult to accept the common view that the first-formed cells or units possess the inherent power, independently of any external agency, of producing new structures wholly different in form, position, and function.
But all these cases become plain on the hypothesis of pangenesis. The units, during each stage of development, throw off gemmules, which, multiplying, are transmitted to the offspring. In the offspring, as soon as any particular cell or unit becomes partially developed, it unites with (or, to speak metaphorically, is fertilised by) the gemmule of the next succeeding cell, and so onwards. But organisms have often been subjected to changed conditions of life at a certain stage of their development, and in consequence have been slightly modified; and the gemmules cast off from such modified parts will tend to reproduce parts modified in the same manner. This process may be repeated until the structure of the part becomes greatly changed at one particular stage of development, but this will not necessarily affect other parts, whether previously or subsequently formed. In this manner we can understand the remarkable independence of structure in the successive metamorphoses, and especially in the successive metageneses of many animals. In the case, however, of diseases which supervene during old age, subsequently to the ordinary period of procreation, and which, nevertheless, are sometimes inherited, as occurs with brain and heart complaints, we must suppose that the organs were affected at an early age and threw off at this period affected gemmules; but that the affection became visible or injurious only after the prolonged growth, in the strict sense of the word, of the part. In all the changes of structure which regularly supervene during old age, we probably see the effects of deteriorated growth, and not of true development.
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[58] 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.[59] When frogs, toads, etc., are born with their limbs doubled, as sometimes happens, the doubling, as Gervais remarks,[60] cannot be due to the complete fusion of two embryos, with the exception of the limbs, for the larvæ are limbless.
The same argument is applicable[61] to certain insects produced with multiple legs or antennæ, for these are metamorphosed from apodal or antennæ-less larvæ. Alphonse Milne-Edwards[62] 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[63] 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[64] 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.”[65] 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[66] 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.”[67]
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.[68]
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 Gärtner—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 re-growth of an amputated limb.
When parts which are homologous and similar in structure, as the vertebræ 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;[69] 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.[70] 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-Séquard 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.
Ideas discussed in this chapter
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