Чарльз Дарвин
The Variation of Animals and Plants under Domestication — Volume 2
In all these respects the forms of the same undoubted species, when illegitimately united, behave in exactly the same manner as do two distinct species when crossed. This led me carefully to observe during four years many seedlings, raised from several illegitimate unions. The chief result is that these illegitimate plants, as they may be called, are not fully fertile. It is possible to raise from dimorphic species, both long-styled and short-styled illegitimate plants, and from trimorphic plants all three illegitimate forms. These can then be properly united in a legitimate manner. When this is done, there is no apparent reason why they should not yield as many seeds as did their parents when legitimately fertilised. But such is not the case; they are all infertile, but in various degrees; some being so utterly and incurably sterile that they did not yield during four seasons a single seed or even seed-capsule. These illegitimate plants, which are so sterile, although united with each other in a legitimate manner, may be strictly compared with hybrids when crossed inter se, and it is well known how sterile these latter generally are. When, on the other hand, a hybrid is crossed with either pure parent- species, the sterility is usually much lessened: and so it is when an illegitimate plant is fertilised by a legitimate plant. In the same manner as the sterility of hybrids does not always run parallel with the difficulty of making the first cross between the two parent-species, so the sterility of certain illegitimate plants was unusually great, whilst the sterility of the union from which they were derived was by no means great. With hybrids raised from the same seed-capsule the degree of sterility is innately variable, so it is in a marked manner with illegitimate plants. Lastly, many hybrids are profuse and persistent flowerers, whilst other and more sterile hybrids produce few flowers, and are weak, miserable dwarfs; exactly similar cases occur with the illegitimate offspring of various dimorphic and trimorphic plants.
Although there is the closest identity in character and behaviour between illegitimate plants and hybrids, it is hardly an exaggeration to maintain that the former are hybrids, but produced within the limits of the same species by the improper union of certain forms, whilst ordinary hybrids are produced from an improper union between so-called distinct species. We have already seen that there is the closest similarity in all respects between first illegitimate unions, and first crosses between distinct species. This will perhaps be made more fully apparent by an illustration: — we may suppose that a botanist found two well-marked varieties (and such occur) of the long-styled form of the trimorphic Lithrum salicaria, and that he determined to try by crossing whether they were specifically distinct. He would find that they yielded only about one-fifth of the proper number of seed, and that they behaved in all the other above-specified respects as if they had been two distinct species. But to make the case sure, he would raise plants from his supposed hybridised seed, and he would find that the seedlings were miserably dwarfed and utterly sterile, and that they behaved in all other respects like ordinary hybrids, he might then maintain that he had actually proved, in accordance with the common view, that his two varieties were as good and as distinct species as any in the world; but he would be completely mistaken.
The facts now given on dimorphic and trimorphic plants are important, because they show us, first, that the physiological test of lessened fertility, both in first crosses and in hybrids, is no criterion of specific distinction; secondly, because we may conclude that there is some unknown bond which connects the infertility of illegitimate unions with that of their illegitimate offspring, and we are led to extend the same view to first crosses and hybrids; thirdly, because we find, and this seems to me of especial importance, that two or three forms of the same species may exist and may differ in no respect whatever, either in structure or in constitution, relatively to external conditions, and yet be sterile when united in certain ways. For we must remember that it is the union of the sexual elements of individuals of the same form, for instance, of two long-styled forms, which results in sterility; whilst it is the union of the sexual element proper to two distinct forms which is fertile. Hence the case appears at first sight exactly the reverse of what occurs in the ordinary unions of the individuals of the same species, and with crosses between distinct species. It is, however, doubtful whether this is really so; but I will not enlarge on this obscure subject.
We may, however, infer as probable from the consideration of dimorphic and trimorphic plants, that the sterility of distinct species when crossed, and of their hybrid progeny, depends exclusively on the nature of their sexual elements, and not on any difference in their structure or general constitution. We are also led to this same conclusion by considering reciprocal crosses, in which the male of one species cannot be united, or only with great difficulty, with the female of a second species, whilst the converse cross can be effected with perfect facility. That excellent observer, Gartner, likewise concluded that species when crossed are sterile owing to differences confined to their reproductive systems.
On the principle which makes it necessary for man, whilst he is selecting and improving his domestic varieties, to keep them separate, it would clearly be advantageous to varieties in a state of nature, that is to incipient species, if they could be kept from blending, either through sexual aversion, or by becoming mutually sterile. Hence it at one time appeared to me probable, as it has to others, that this sterility might have been acquired through natural selection. On this view we must suppose that a shade of lessened fertility first spontaneously appeared, like any other modification, in certain individuals of a species when crossed with other individuals of the same species; and that successive slight degrees of infertility, from being advantageous, were slowly accumulated. This appears all the more probable, if we admit that the structural differences between the forms of dimorphic and trimorphic plants, as the length and curvature of the pistil, etc., have been co-adapted through natural selection; for if this be admitted, we can hardly avoid extending the same conclusion to their mutual infertility. Sterility, moreover, has been acquired through natural selection for other and widely different purposes, as with neuter insects in reference to their social economy. In the case of plants, the flowers on the circumference of the truss in the guelder rose (Viburnum opulus) and those on the summit of the spike in the feather-hyacinth (Muscari comosum) have been rendered conspicuous, and apparently in consequence sterile, in order that insects might easily discover and visit the perfect flowers. But when we endeavour to apply the principle of natural selection to the acquirement by distinct species of mutual sterility, we meet with great difficulties. In the first place, it may be remarked that separate regions are often inhabited by groups of species or by single species, which when brought together and crossed are found to be more or less sterile; now it could clearly have been no advantage to such separated species to have been rendered mutually sterile, and consequently this could not have been effected through natural selection; but it may perhaps be argued, that, if a species were rendered sterile with some one compatriot, sterility with other species would follow as a necessary consequence. In the second place, it is as much opposed to the theory of natural selection, as to the theory of special creation, that in reciprocal crosses the male element of one form should have been rendered utterly impotent on a second form, whilst at the same time the male element of this second form is enabled freely to fertilise the first form; for this peculiar state of the reproductive system could not possibly have been advantageous to either species.
In considering the probability of natural selection having come into action in rendering species mutually sterile, one of the greatest difficulties will be found to lie in the existence of many graduated steps from slightly lessened fertility to absolute sterility. It may be admitted, on the principle above explained, that it would profit an incipient species if it were rendered in some slight degree sterile when crossed with its parent-form or with some other variety; for thus fewer bastardised and deteriorated offspring would be produced to commingle their blood with the new species in process of formation. But he who will take the trouble to reflect on the steps by which this first degree of sterility could be increased through natural selection to that higher degree which is common to so many species, and which is universal with species which have been differentiated to a generic or family rank, will find the subject extraordinarily complex. After mature reflection it seems to me that this could not have been effected through natural selection. Take the case of any two species which, when crossed, produce few and sterile offspring; now, what is there which could favour the survival of those individuals which happened to be endowed in a slightly higher degree with mutual infertility, and which thus approached by one small step towards absolute sterility? Yet an advance of this kind, if the theory of natural selection be brought to bear, must have incessantly occurred with many species, for a multitude are mutually quite barren. With sterile neuter insects we have reason to believe that modifications in their structure and fertility have been slowly accumulated by natural selection, from an advantage having been thus indirectly given to the community to which they belonged over other communities of the same species; but an individual animal not belonging to a social community, if rendered slightly sterile when crossed with some other variety, would not thus itself gain any advantage or indirectly give any advantage to the other individuals of the same variety, thus leading to their preservation.
But it would be superfluous to discuss this question in detail; for with plants we have conclusive evidence that the sterility of crossed species must be due to some principle, quite independent of natural selection. Both Gartner and Kolreuter have proved that in general including numerous species, a series can be formed from species which when crossed yield fewer and fewer seeds, to species which never produce a single seed, but yet are affected by the pollen of certain other species, for the germen swells. It is here manifestly impossible to select the more sterile individuals, which have already ceased to yield seeds; so that this acme of sterility, when the germen alone is affected, cannot have been gained through selection; and from the laws governing the various grades of sterility being so uniform throughout the animal and vegetable kingdoms, we may infer that the cause, whatever it may be, is the same or nearly the same in all cases.
As species have not been rendered mutually infertile through the accumulative action of natural selection, and as we may safely conclude, from the previous as well as from other and more general considerations, that they have not been endowed through an act of creation with this quality, we must infer that it has arisen incidentally during their slow formation in connection with other and unknown changes in their organisation. By a quality arising incidentally, I refer to such cases as different species of animals and plants being differently affected by poisons to which they are not naturally exposed; and this difference in susceptibility is clearly incidental on other and unknown differences in their organisation. So again the capacity in different kinds of trees to be grafted on each other, or on a third species, differs much, and is of no advantage to these trees, but is incidental on structural or functional differences in their woody tissues. We need not feel surprise at sterility incidentally resulting from crosses between distinct species, — the modified descendants of a common progenitor, — when we bear in mind how easily the reproductive system is affected by various causes — often by extremely slight changes in the conditions of life, by too close interbreeding, and by other agencies. It is well to bear in mind such cases as that of the Passiflora alata, which recovered its self-fertility from being grafted on a distinct species — the cases of plants which normally or abnormally are self-impotent, but can readily be fertilised by the pollen of a distinct species — and lastly the cases of individual domesticated animals which evince towards each other sexual incompatibility.
We now at last come to the immediate point under discussion: how is it that, with some few exceptions in the case of plants, domesticated varieties, such as those of the dog, fowl, pigeon, several fruit-trees, and culinary vegetables, which differ from each other in external characters more than many species, are perfectly fertile when crossed, or even fertile in excess, whilst closely allied species are almost invariably in some degree sterile? We can, to a certain extent, give a satisfactory answer to this question. Passing over the fact that the amount of external difference between two species is no sure guide to their degree of mutual sterility, so that similar differences in the case of varieties would be no sure guide, we know that with species the cause lies exclusively in differences in their sexual constitution. Now the conditions to which domesticated animals and cultivated plants have been subjected have had so little tendency towards modifying the reproductive system in a manner leading to mutual sterility, that we have very good grounds for admitting the directly opposite doctrine of Pallas, namely, that such conditions generally eliminate this tendency; so that the domesticated descendants of species, which in their natural state would have been in some degree sterile when crossed, become perfectly fertile together. With plants, so far is cultivation from giving a tendency towards mutual sterility, that in several well-authenticated cases, already often alluded to, certain species have been affected in a very different manner, for they have become self- impotent, whilst still retaining the capacity of fertilising, and being fertilised by, distinct species. If the Pallasian doctrine of the elimination of sterility through long-continued domestication be admitted, and it can hardly be rejected, it becomes in the highest degree improbable that similar circumstances should commonly both induce and eliminate the same tendency; though in certain cases, with species having a peculiar constitution, sterility might occasionally be thus induced. Thus, as I believe, we can understand why with domesticated animals varieties have not been produced which are mutually sterile; and why with plants only a few such cases have been observed, namely, by Gartner, with certain varieties of maize and verbascum, by other experimentalists with varieties of the gourd and melon, and by Kolreuter with one kind of tobacco.
With respect to varieties which have originated in a state of nature, it is almost hopeless to expect to prove by direct evidence that they have been rendered mutually sterile; for if even a trace of sterility could be detected, such varieties would at once be raised by almost every naturalist to the rank of distinct species. If, for instance, Gartner's statement were fully confirmed, that the blue and red flowered forms of the pimpernel (Anagallis arvensis) are sterile when crossed, I presume that all the botanists who now maintain on various grounds that these two forms are merely fleeting varieties, would at once admit that they were specifically distinct.
The real difficulty in our present subject is not, as it appears to me, why domestic varieties have not become mutually infertile when crossed, but why this has so generally occurred with natural varieties as soon as they have been modified in a sufficient and permanent degree to take rank as species. We are far from precisely knowing the cause; but we can see that the species, owing to their struggle for existence with numerous competitors, must have been exposed to more uniform conditions of life during long periods of time than domestic varieties have been, and this may well make a wide difference in the result. For we know how commonly wild animals and plants, when taken from their natural conditions and subjected to captivity, are rendered sterile; and the reproductive functions of organic beings which have always lived and been slowly modified under natural conditions would probably in like manner be eminently sensitive to the influence of an unnatural cross. Domesticated productions, on the other hand, which, as shown by the mere fact of their domestication, were not originally highly sensitive to changes in their conditions of life, and which can now generally resist with undiminished fertility repeated changes of conditions, might be expected to produce varieties, which would be little liable to have their reproductive powers injuriously affected by the act of crossing with other varieties which had originated in a like manner.
Certain naturalists have recently laid too great stress, as it appears to me, on the difference in fertility between varieties and species when crossed. Some allied species of trees cannot be grafted on one another, whilst all varieties can be so grafted. Some allied animals are affected in a very different manner by the same poison, but with varieties no such case until recently was known; whilst now it has been proved that immunity from certain poisons sometimes stands in correlation with the colour of the individuals of the same species. The period of gestation generally differs much in distinct species, but with varieties until lately no such difference had been observed. Here we have various physiological differences, and no doubt others could be added, between one species and another of the same genus, which do not occur, or occur with extreme rarity, in the case of varieties; and these differences are apparently wholly or in chief part incidental on other constitutional differences, just in the same manner as the sterility of crossed species is incidental on differences confined to the sexual system. Why, then, should these latter differences, however serviceable they may indirectly be in keeping the inhabitants of the same country distinct, be thought of such paramount importance, in comparison with other incidental and functional differences? No sufficient answer to this question can be given. Hence the fact that widely distinct domestic varieties are, with rare exceptions, perfectly fertile when crossed, and produce fertile offspring, whilst closely allied species are, with rare exceptions, more or less sterile, is not nearly so formidable an objection as it appears at first to the theory of the common descent of allied species.
CHAPTER 2.XX
SELECTION BY MAN.
SELECTION A DIFFICULT ART. METHODICAL, UNCONSCIOUS, AND NATURAL SELECTION. RESULTS OF METHODICAL SELECTION. CARE TAKEN IN SELECTION. SELECTION WITH PLANTS. SELECTION CARRIED ON BY THE ANCIENTS AND BY SEMI-CIVILISED PEOPLE. UNIMPORTANT CHARACTERS OFTEN ATTENDED TO. UNCONSCIOUS SELECTION. AS CIRCUMSTANCES SLOWLY CHANGE, SO HAVE OUR DOMESTICATED ANIMALS CHANGED THROUGH THE ACTION OF UNCONSCIOUS SELECTION. INFLUENCE OF DIFFERENT BREEDERS ON THE SAME SUB-VARIETY. PLANTS AS AFFECTED BY UNCONSCIOUS SELECTION. EFFECTS OF SELECTION AS SHOWN BY THE GREAT AMOUNT OF DIFFERENCE IN THE PARTS MOST VALUED BY MAN.
The power of Selection, whether exercised by man, or brought into play under nature through the struggle for existence and the consequent survival of the fittest, absolutely depends on the variability of organic beings. Without variability nothing can be effected; slight individual differences, however, suffice for the work, and are probably the chief or sole means in the production of new species. Hence our discussion on the causes and laws of variability ought in strict order to have preceded the present subject, as well as inheritance, crossing, etc.; but practically the present arrangement has been found the most convenient. Man does not attempt to cause variability; though he unintentionally effects this by exposing organisms to new conditions of life, and by crossing breeds already formed. But variability being granted, he works wonders. Unless some degree of selection be exercised, the free commingling of the individuals of the same variety soon obliterates, as we have previously seen, the slight differences which arise, and gives uniformity of character to the whole body of individuals. In separated districts, long- continued exposure to different conditions of life may produce new races without the aid of selection; but to this subject of the direct action of the conditions of life I shall recur in a future chapter.
When animals or plants are born with some conspicuous and firmly inherited new character, selection is reduced to the preservation of such individuals, and to the subsequent prevention of crosses; so that nothing more need be said on the subject. But in the great majority of cases a new character, or some superiority in an old character, is at first faintly pronounced, and is not strongly inherited; and then the full difficulty of selection is experienced. Indomitable patience, the finest powers of discrimination, and sound judgment must be exercised during many years. A clearly predetermined object must be kept steadily in view. Few men are endowed with all these qualities, especially with that of discriminating very slight differences; judgment can be acquired only by long experience; but if any of these qualities be wanting, the labour of a life may be thrown away. I have been astonished when celebrated breeders, whose skill and judgment have been proved by their success at exhibitions, have shown me their animals, which appeared all alike, and have assigned their reasons for matching this and that individual. The importance of the great principle of Selection mainly lies in this power of selecting scarcely appreciable differences, which nevertheless are found to be transmissible, and which can be accumulated until the result is made manifest to the eyes of every beholder.
The principle of selection may be conveniently divided into three kinds. METHODICAL SELECTION is that which guides a man who systematically endeavours to modify a breed according to some predetermined standard. UNCONSCIOUS SELECTION is that which follows from men naturally preserving the most valued and destroying the less valued individuals, without any thought of altering the breed; and undoubtedly this process slowly works great changes. Unconscious selection graduates into methodical, and only extreme cases can be distinctly separated; for he who preserves a useful or perfect animal will generally breed from it with the hope of getting offspring of the same character; but as long as he has not a predetermined purpose to improve the breed, he may be said to be selecting unconsciously. (20/1. The term "unconscious selection" has been objected to as a contradiction; but see some excellent observations on this head by Prof. Huxley ('Nat. Hist. Review' October 1864 page 578) who remarks that when the wind heaps up sand-dunes it sifts and UNCONSCIOUSLY SELECTS from the gravel on the beach grains of sand of equal size.) Lastly, we have NATURAL SELECTION, which implies that the individuals which are best fitted for the complex, and in the course of ages changing conditions to which they are exposed, generally survive and procreate their kind. With domestic productions, natural selection comes to a certain extent into action, independently of, and even in opposition to, the will of man.
METHODICAL SELECTION.
What man has effected within recent times in England by methodical selection is clearly shown by our exhibitions of improved quadrupeds and fancy birds. With respect to cattle, sheep, and pigs, we owe their great improvement to a long series of well-known names — Bakewell, Coiling, Ellman, Bates, Jonas Webb, Lords Leicester and Western, Fisher Hobbs, and others. Agricultural writers are unanimous on the power of selection: any number of statements to this effect could be quoted; a few will suffice. Youatt, a sagacious and experienced observer, writes (20/2. 'On Sheep' 1838 page 60.) the principle of selection is "that which enables the agriculturist, not only to modify the character of his flock, but to change it altogether." A great breeder of Shorthorns (20/3. Mr. J. Wright on Shorthorn Cattle in 'Journal of Royal Agricult. Soc.' volume 7 pages 208, 209.) says, "In the anatomy of the shoulder modern breeders have made great improvement on the Ketton shorthorns by correcting the defect in the knuckle or shoulder-joint, and by laying the top of the shoulder more snugly in the crop, and thereby filling up the hollow behind it...The eye has its fashion at different periods: at one time the eye high and outstanding from the head, and at another time the sleepy eye sunk into the head; but these extremes have merged into the medium of a full, clear and prominent eye with a placid look."
Again, hear what an excellent judge of pigs (20/4. H.D. Richardson 'On Pigs' 1847 page 44.) says: "The legs should be no longer than just to prevent the animal's belly from trailing on the ground. The leg is the least profitable portion of the hog, and we therefore require no more of it than is absolutely necessary for the support of the rest." Let any one compare the wild-boar with any improved breed, and he will see how effectually the legs have been shortened.
Few persons, except breeders, are aware of the systematic care taken in selecting animals, and of the necessity of having a clear and almost prophetic vision into futurity. Lord Spencer's skill and judgment were well known; and he writes (20/5. 'Journal of Royal Agricult. Soc.' volume 1 page 24.), "It is therefore very desirable, before any man commences to breed either cattle or sheep, that he should make up his mind to the shape and qualities he wishes to obtain, and steadily pursue this object." Lord Somerville, in speaking of the marvellous improvement of the New Leicester sheep, effected by Bakewell and his successors, says, "It would seem as if they had first drawn a perfect form, and then given it life." Youatt (20/6. 'On Sheep' pages 520, 319.) urges the necessity of annually drafting each flock, as many animals will certainly degenerate "from the standard of excellence which the breeder has established in his own mind." Even with a bird of such little importance as the canary, long ago (1780-1790) rules were established, and a standard of perfection was fixed according to which the London fanciers tried to breed the several sub- varieties. (20/7. Loudon's 'Mag. of Nat. Hist.' volume 8 1835 page 618.) A great winner of prizes at the Pigeon-shows (20/8. 'A treatise on the Art of Breeding the Almond Tumbler' 1851 page 9.), in describing the short-faced Almond Tumbler, says, "There are many first-rate fanciers who are particularly partial to what is called the goldfinch-beak, which is very beautiful; others say, take a full-size round cherry then take a barleycorn, and judiciously placing and thrusting it into the cherry, form as it were your beak; and that is not all, for it will form a good head and beak, provided, as I said before, it is judiciously done; others take an oat; but as I think the goldfinch-beak the handsomest, I would advise the inexperienced fancier to get the head of a goldfinch, and keep it by him for his observation." Wonderfully different as are the beaks of the rock pigeon and goldfinch, the end has undoubtedly been nearly gained, as far as external shape and proportions are concerned.
Not only should our animals be examined with the greatest care whilst alive, but, as Anderson remarks (20/9. 'Recreations in Agriculture' volume 2 page 409.) their carcases should be scrutinised, "so as to breed from the descendants of such only as, in the language of the butcher, cut up well." The "grain of the meat" in cattle, and its being well marbled with fat (20/10. 'Youatt on Cattle' pages 191, 227.), and the greater or less accumulation of fat in the abdomen of our sheep, have been attended to with success. So with poultry, a writer (20/11. Ferguson 'Prize Poultry' 1854 page 208.), speaking of Cochin-China fowls, which are said to differ much in the quality of their flesh, says, "the best mode is to purchase two young brother-cocks, kill, dress, and serve up one; if he be indifferent, similarly dispose of the other, and try again; if, however, he be fine and well-flavoured, his brother will not be amiss for breeding purposes for the table."
The great principle of the division of labour has been brought to bear on selection. In certain districts (20/12. Wilson in 'Transact. Highland Agricult. Soc.' quoted in 'Gardener's Chronicle' 1844 page 29.) "the breeding of bulls is confined to a very limited number of persons, who by devoting their whole attention to this department, are able from year to year to furnish a class of bulls which are steadily improving the general breed of the district." The rearing and letting of choice rams has long been, as is well known, a chief source of profit to several eminent breeders. In parts of Germany this principle is carried with merino sheep to an extreme point. (20/13. Simmonds quoted in 'Gardener's Chronicle' 1855 page 637. And for the second quotation see 'Youatt on Sheep' page 171.) So "important is the proper selection of breeding animals considered, that the best flock-masters do not trust to their own judgment or to that of their shepherds, but employ persons called 'sheep-classifiers' who make it their special business to attend to this part of the management of several flocks, and thus to preserve, or if possible to improve, the best qualities of both parents in the lambs." In Saxony, "when the lambs are weaned, each in his turn is placed upon a table that his wool and form may be minutely observed. The finest are selected for breeding and receive a first mark. When they are one year old, and prior to shearing them, another close examination of those previously marked takes place: those in which no defect can be found receive a second mark, and the rest are condemned. A few months afterwards a third and last scrutiny is made; the prime rams and ewes receive a third and final mark, but the slightest blemish is sufficient to cause the rejection of the animal." These sheep are bred and valued almost exclusively for the fineness of their wool; and the result corresponds with the labour bestowed on their selection. Instruments have been invented to measure accurately the thickness of the fibres; and "an Austrian fleece has been produced of which twelve hairs equalled in thickness one from a Leicester sheep."
