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Charles Darwin
The Variation of Animals and Plants under Domestication — Volume 2

CHAPTER 2.XIII

INHERITANCE continued — REVERSION OR ATAVISM.

DIFFERENT FORMS OF REVERSION. IN PURE OR UNCROSSED BREEDS, AS IN PIGEONS, FOWLS, HORNLESS CATTLE AND SHEEP, IN CULTIVATED PLANTS. REVERSION IN FERAL ANIMALS AND PLANTS. REVERSION IN CROSSED VARIETIES AND SPECIES. REVERSION THROUGH BUD-PROPAGATION, AND BY SEGMENTS IN THE SAME FLOWER OR FRUIT. IN DIFFERENT PARTS OF THE BODY IN THE SAME ANIMAL. THE ACT OF CROSSING A DIRECT CAUSE OF REVERSION, VARIOUS CASES OF, WITH INSTINCTS. OTHER PROXIMATE CAUSES OF REVERSION. LATENT CHARACTERS. SECONDARY SEXUAL CHARACTERS. UNEQUAL DEVELOPMENT OF THE TWO SIDES OF THE BODY. APPEARANCE WITH ADVANCING AGE OF CHARACTERS DERIVED FROM A CROSS. THE GERM, WITH ALL ITS LATENT CHARACTERS, A WONDERFUL OBJECT. MONSTROSITIES. PELORIC FLOWERS DUE IN SOME CASES TO REVERSION.

The great principle of inheritance to be discussed in this chapter has been recognised by agriculturists and authors of various nations, as shown by the scientific term ATAVISM, derived from atavus, an ancestor; by the English terms of REVERSION, or THROWING-BACK; by the French PAS-EN-ARRIERE; and by the German RUCKSCHLAG, or RUCKSCHRITT. When the child resembles either grandparent more closely than its immediate parents, our attention is not much arrested, though in truth the fact is highly remarkable; but when the child resembles some remote ancestor or some distant member in a collateral line, — and in the last case we must attribute this to the descent of all the members from a common progenitor, — we feel a just degree of astonishment. When one parent alone displays some newly-acquired and generally inheritable character, and the offspring do not inherit it, the cause may lie in the other parent having the power of prepotent transmission. But when both parents are similarly characterised, and the child does not, whatever the cause may be, inherit the character in question, but resembles its grandparents, we have one of the simplest cases of reversion. We continually see another and even more simple case of atavism, though not generally included under this head, namely, when the son more closely resembles his maternal than his paternal grand-sire in some male attribute, as in any peculiarity in the beard of man, the horns of the bull, the hackles or comb of the cock, or, as in certain diseases necessarily confined to the male sex; for as the mother cannot possess or exhibit such male attributes, the child must inherit them, through her blood, from his maternal grandsire.

The cases of reversion may be divided into two main classes which, however, in some instances, blend into one another; namely, first, those occurring in a variety or race which has not been crossed, but has lost by variation some character that it formerly possessed, and which afterwards reappears. The second class includes all cases in which an individual with some distinguishable character, a race, or species, has at some former period been crossed, and a character derived from this cross, after having disappeared during one or several generations, suddenly reappears. A third class, differing only in the manner of reproduction, might be formed to include all cases of reversion effected by means of buds, and therefore independent of true or seminal generation. Perhaps even a fourth class might be instituted, to include reversions by segments in the same individual flower or fruit, and in different parts of the body in the same individual animal as it grows old. But the two first main classes will be sufficient for our purpose.

REVERSION TO LOST CHARACTERS BY PURE OR UNCROSSED FORMS.

Striking instances of this first class of cases were given in the sixth chapter, namely, of the occasional reappearance, in variously-coloured breeds of the pigeon, of blue birds with all the marks characteristic of the wild Columba livia. Similar cases were given in the case of the fowl. With the common ass, as the legs of the wild progenitor are almost always striped, we may feel assured that the occasional appearance of such stripes in the domestic animal is a case of simple reversion. But I shall be compelled to refer again to these cases, and therefore here pass them over.

The aboriginal species from which our domesticated cattle and sheep are descended, no doubt possessed horns; but several hornless breeds are now well established. Yet in these — for instance, in Southdown sheep — "it is not unusual to find among the male lambs some with small horns." The horns, which thus occasionally reappear in other polled breeds, either "grow to the full size," or are curiously attached to the skin alone and hang "loosely down, or drop off." (13/1. 'Youatt on Sheep' pages 20, 234. The same fact of loose horns occasionally appearing in hornless breeds has been observed in Germany; Bechstein 'Naturgesch. Deutschlands.' b. 1 s. 362.) The Galloways and Suffolk cattle have been hornless for the last 100 or 150 years, but a horned calf, with the horn often loosely attached, is occasionally produced. (13/2. 'Youatt on Cattle' pages 155, 174.)

There is reason to believe that sheep in their early domesticated condition were "brown or dingy black;" but even in the time of David certain flocks were spoken of as white as snow. During the classical period the sheep of Spain are described by several ancient authors as being black, red, or tawny. (13/3. 'Youatt on Sheep' 1838 pages 17, 145.) At the present day, notwithstanding the great care which is taken to prevent it, particoloured lambs and some entirely black are occasionally, or even frequently, dropped by our most highly improved and valued breeds, such as the Southdowns. Since the time of the famous Bakewell, during the last century, the Leicester sheep have been bred with the most scrupulous care; yet occasionally grey-faced, or black-spotted, or wholly black lambs appear. (13/4. I have been informed of this fact through the Rev. W.D. Fox on the excellent authority of Mr. Wilmot: see also remarks on this subject in an article in the 'Quarterly Review' 1849 page 395.) This occurs still more frequently with the less improved breeds, such as the Norfolks. (13/5. Youatt pages 19, 234.) As bearing on this tendency in sheep to revert to dark colours, I may state (though in doing so I trench on the reversion of crossed breeds, and likewise on the subject of prepotency) that the Rev. W.D. Fox was informed that seven white Southdown ewes were put to a so-called Spanish ram, which had two small black spots on his sides, and they produced thirteen lambs, all perfectly black. Mr. Fox believes that this ram belonged to a breed which he has himself kept, and which is always spotted with black and white; and he finds that Leicester sheep crossed by rams of this breed always produce black lambs: he has gone on recrossing these crossed sheep with pure white Leicesters during three successive generations, but always with the same result. Mr. Fox was also told by the friend from whom the spotted breed was procured, that he likewise had gone on for six or seven generations crossing with white sheep, but still black lambs were invariably produced.

Similar facts could be given with respect to tailless breeds of various animals. For instance, Mr. Hewitt (13/6. 'The Poultry Book' by Mr. Tegetmeier 1866 page 231.) states that chickens bred from some rumpless fowls, which were reckoned so good that they won a prize at an exhibition, "in a considerable number of instances were furnished with fully developed tail-feathers." On inquiry, the original breeder of these fowls stated that, from the time when he had first kept them, they had often produced fowls furnished with tails; but that these latter would again reproduce rumpless chickens.

Analogous cases of reversion occur in the vegetable kingdom; thus "from seeds gathered from the finest cultivated varieties of Heartsease (Viola tricolor), plants perfectly wild both in their foliage and their flowers are frequently produced;" (13/7. Loudon's 'Gardener's Mag.' volume 10 1834 page 396: a nurseryman, with much experience on this subject, has likewise assured me that this sometimes occurs.) but the reversion in this instance is not to a very ancient period, for the best existing varieties of the heartsease are of comparatively modern origin. With most of our cultivated vegetables there is some tendency to reversion to what is known to be, or may be presumed to be, their aboriginal state; and this would be more evident if gardeners did not generally look over their beds of seedlings, and pull up the false plants or "rogues" as they are called. It has already been remarked, that some few seedling apples and pears generally resemble, but apparently are not identical with, the wild trees from which they are descended. In our turnip (13/8. 'Gardener's Chronicle' 1855 page 777.) and carrot-beds a few plants often "break " — that is, flower too soon; and their roots are generally hard and stringy, as in the parent-species. By the aid of a little selection, carried on during a few generations, most of our cultivated plants could probably be brought back, without any great change in their conditions of life, to a wild or nearly wild condition: Mr. Buckman has effected this with the parsnip (13/9. Ibid 1862 page 721.); and Mr. Hewett C. Watson, as he informs me, selected, during three generations, "the most diverging plants of Scotch kail, perhaps one of the least modified varieties of the cabbage; and in the third generation some of the plants came very close to the forms now established in England about old castle-walls, and called indigenous."

REVERSION IN ANIMALS AND PLANTS WHICH HAVE RUN WILD.

In the cases hitherto considered, the reverting animals and plants have not been exposed to any great or abrupt change in their conditions of life which could have induced this tendency; but it is very different with animals and plants which have become feral or run wild. It has been repeatedly asserted in the most positive manner by various authors, that feral animals and plants invariably return to their primitive specific type. It is curious on what little evidence this belief rests. Many of our domesticated animals could not subsist in a wild state; thus, the more highly improved breeds of the pigeon will not "field" or search for their own food. Sheep have never become feral, and would be destroyed by almost every beast of prey. (13/10. Mr. Boner speaks ('Chamois-hunting' 2nd edition 1860 page 92) of sheep often running wild in the Bavarian Alps; but, on making further inquiries at my request, he found that they are not able to establish themselves; they generally perish from the frozen snow clinging to their wool, and they have lost the skill necessary to pass over steep icy slopes. On one occasion two ewes survived the winter, but their lambs perished.) In several cases we do not know the aboriginal parent- species, and cannot possibly tell whether or not there has been any close degree of reversion. It is not known in any instance what variety was first turned out; several varieties have probably in some cases run wild, and their crossing alone would tend to obliterate their proper character. Our domesticated animals and plants, when they run wild, must always be exposed to new conditions of life, for, as Mr. Wallace (13/11. See some excellent remarks on this subject by Mr. Wallace 'Journal Proc. Linn. Soc.' 1858 volume 3 page 60.) has well remarked, they have to obtain their own food, and are exposed to competition with the native productions. Under these circumstances, if our domesticated animals did not undergo change of some kind, the result would be quite opposed to the conclusions arrived at in this work. Nevertheless, I do not doubt that the simple fact of animals and plants becoming feral, does cause some tendency to reversion to the primitive state; though this tendency has been much exaggerated by some authors.

[I will briefly run through the recorded cases. With neither horses nor cattle is the primitive stock known; and it has been shown in former chapters that they have assumed different colours in different countries. Thus the horses which have run wild in South America are generally brownish-bay, and in the East dun-coloured; their heads have become larger and coarser, and this may be due to reversion. No careful description has been given of the feral goat. Dogs which have run wild in various countries have hardly anywhere assumed a uniform character; but they are probably descended from several domestic races, and aboriginally from several distinct species. Feral cats, both in Europe and La Plata, are regularly striped; in some cases they have grown to an unusually large size, but do not differ from the domestic animal in any other character. When variously-coloured tame rabbits are turned out in Europe, they generally reacquire the colouring of the wild animal; there can be no doubt that this does really occur, but we should remember that oddly- coloured and conspicuous animals would suffer much from beasts of prey and from being easily shot; this at least was the opinion of a gentleman who tried to stock his woods with a nearly white variety; if thus destroyed, they would be supplanted by, instead of being transformed into, the common rabbit. We have seen that the feral rabbits of Jamaica, and especially of Porto Santo, have assumed new colours and other new characters. The best known case of reversion, and that on which the widely spread belief in its universality apparently rests, is that of pigs. These animals have run wild in the West Indies, South America, and the Falkland Islands, and have everywhere acquired the dark colour, the thick bristles, and great tusks of the wild boar; and the young have reacquired longitudinal stripes. But even in the case of the pig, Roulin describes the half-wild animals in different parts of South America as differing in several respects. In Louisiana the pig (13/12. Dureau de la Malle 'Comptes Rendus' tome 41 1855 page 807. From the statements above given, the author concludes that the wild pigs of Louisiana are not descended from the European Sus scrofa.) has run wild, and is said to differ a little in form, and much in colour, from the domestic animal, yet does not closely resemble the wild boar of Europe. With pigeons and fowls (13/13. Capt. W. Allen, in his 'Expedition to the Niger' states that fowls have run wild on the island of Annobon, and have become modified in form and voice. The account is so meagre and vague that it did not appear to me worth copying; but I now find that Dureau de la Malle ('Comptes Rendus' tome 41 1855 page 690) advances this as a good instance of reversion to the primitive stock, and as confirmatory of a still more vague statement in classical times by Varro.), it is not known what variety was first turned out, nor what character the feral birds have assumed. The guinea-fowl in the West Indies, when feral, seems to vary more than in the domesticated state.

With respect to plants run wild, Dr. Hooker (13/14. 'Flora of Australia' 1859 Introduction page 9.) has strongly insisted on what slight evidence the common belief in their reversion to a primitive state rests. Godron (13/15. 'De l'Espece' tome 2 pages 54, 58, 60.) describes wild turnips, carrots, and celery; but these plants in their cultivated state hardly differ from their wild prototypes, except in the succulency and enlargement of certain parts, — characters which would certainly be lost by plants growing in poor soil and struggling with other plants. No cultivated plant has run wild on so enormous a scale as the cardoon (Cynara cardunculus) in La Plata. Every botanist who has seen it growing there, in vast beds, as high as a horse's back, has been struck with its peculiar appearance; but whether it differs in any important point from the cultivated Spanish form, which is said not to be prickly like its American descendant, or whether it differs from the wild Mediterranean species, which is said not to be social (though this may be due merely to the nature of the conditions), I do not know.]

REVERSION TO CHARACTERS DERIVED FROM A CROSS, IN THE CASE OF SUB-VARIETIES, RACES, AND SPECIES.

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