It should be kept in mind that, while all direct proof of independent origination is attainable from the nature of the case, the overthrow of particular schemes of derivation has not established the opposite proposition. The futility of each hypothesis thus far proposed to account for derivation may be made apparent, or unanswerable objections may be urged against it; and each victory of the kind may render derivation more improbable, and therefore specific creation more probable, without settling the question either way. New facts, or new arguments and a new mode of viewing the question, may some day change the whole aspect of the case. It is with the latter that Mr. Darwin now reopens the discussion.
Having conceived the idea that varieties are incipient species, he is led to study variation in the field where it shows itself most strikingly, and affords the greatest facilities to investigation. Thoughtful naturalists have had increasing grounds to suspect that a reexamination of the question of species in zoology and botany, commencing with those races which man knows most about, viz., the domesticated and cultivated races, would be likely somewhat to modify the received idea of the entire fixity of species. This field, rich with various but unsystematized stores of knowledge accumulated by cultivators and breeders, has been generally neglected by naturalists, because these races are not in a state of nature; whereas they deserve particular attention on this very account, as experiments, or the materials for experiments, ready to our hand. In domestication we vary some of the natural conditions of a species, and thus learn experimentally what changes are within the reach of varying conditions in Nature. We separate and protect a favorite race against its foes or its competitors, and thus learn what it might become if Nature ever afforded it equal opportunities. Even when, to subserve human uses, we modify a domesticated race to the detriment of its native vigor, or to the extent of practical monstrosity, although we secure forms which would not be originated and could not be perpetuated in free Nature, yet we attain wider and juster views of the possible degree of variation. We perceive that some species are more variable than others, but that no species subjected to the experiment persistently refuses to vary; and that, when it has once begun to vary, its varieties are not the less but the more subject to variation. "No case is on record of a variable being ceasing to be variable under cultivation." It is fair to conclude, from the observation of plants and animals in a wild as well as domesticated state, that the tendency to vary is general, and even universal. Mr. Darwin does "not believe that variability is an inherent and necessary contingency, under all circumstances, with all organic beings, as some authors have thought." No one supposes variation could occur under all circumstances; but the facts on the whole imply a universal tendency, ready to be manifested under favorable circumstances. In reply to the assumption that man has chosen for domestication animals and plants having an extraordinary inherent tendency to vary, and likewise to withstand diverse climates, it is asked:
"How could a savage possibly know, when he first tamed an animal, whether it would vary in succeeding generations and whether it would endure other climates? Has the little variability of the ass or Guinea-fowl, or the small power of endurance of warmth by the reindeer, or of cold by the common camel, prevented their domestication? I cannot doubt that if other animals and plants, equal in number to our domesticated productions, and belonging to equally diverse classes and countries, were taken from a state of nature, and could be made to breed for an equal number of generations under domestication, they would vary on an average as largely as the parent species of our existing domesticated productions have varied."
As to amount of variation, there is the common remark of naturalists that the varieties of domesticated plants or animals often differ more widely than do the individuals of distinct species in a wild state: and even in Nature the individuals of some species are known to vary to a degree sensibly wider than that which separates related species. In his instructive section on the breeds of the domestic pigeon, our author remarks that "at least a score of pigeons might be chosen which if shown to an ornithologist, and he were told that they were wild birds, would certainly be ranked by him as well-defined species. Moreover, I do not believe that any ornithologist would place the English carrier, the short-faced tumbler, the runt, the barb, pouter, and fantail, in the same genus; more especially as in each of these breeds several truly-inherited sub-breeds, or species, as he might have called them, could be shown him." That this is not a case like that of dogs, in which probably the blood of more than one species is mingled, Mr. Darwin proceeds to show, adducing cogent reasons for the common opinion that all have descended from the wild rock-pigeon. Then follow some suggestive remarks:
"I have discussed the probable origin of domestic pigeons at some, yet quite insufficient, length; because when I first kept pigeons and watched the several kinds, knowing well how true they bred, I felt fully as much difficulty in believing that they could ever have descended from a common parent as any naturalist could in coming to a similar conclusion in regard to many species of finches, or other large groups of birds, in Nature. One circumstance has struck me much; namely, that all the breeders of the various domestic animals and the cultivators of plants, with whom I have ever conversed, or whose treatises I have read, are firmly convinced that the several breeds to which each has attended are descended from so many aboriginally distinct species. Ask, as I have asked, a celebrated raiser of Hereford cattle, whether his cattle might not have descended from long-horns, and he will laugh you to scorn. I have never met a pigeon, or poultry, or duck, or rabbit fancier, who was not fully convinced that each main breed was descended from a distinct species. Van Mons, in his treatise on pears and apples, shows how utterly he disbelieves that the several sorts, for instance a Ribston-pippin or Codlin-apple, could ever have proceeded from the seeds of the same tree. Innumerable other examples could be given. The explanation, I think, is simple: from long-continued study they arc strongly impressed with the differences between the several races; and though they well know that each race varies slightly, for they win their prizes by selecting such slight differences, yet they ignore all general arguments, and refuse to sum up in their minds slight differences accumulated during many successive generations. May not those naturalists who, knowing far less of the laws of inheritance than does the breeder, and knowing no more than he does of the intermediate links in the long lines of descent, yet admit that many of our domestic races have descended from the same parents—may they not learn a lesson of caution, when they deride the idea of species in a state of nature being lineal descendants of other species?"
The actual causes of variation are unknown. Mr. Darwin favors the opinion of the late Mr. Knight, the great philosopher of horticulture, that variability tinder domestication is somehow connected with excess of food. He regards the unknown cause as acting chiefly upon the reproductive system of the parents, which system, judging from the effect of confinement or cultivation upon its functions, he concludes to be more susceptible than any other to the action of changed conditions of life. The tendency to vary certainly appears to be much stronger under domestication than in free Nature. But we are not sure that the greater variableness of cultivated races is not mainly owing to the far greater opportunities for manifestation and accumulation—a view seemingly all the more favorable to Mr. Darwin's theory. The actual amount of certain changes, such as size or abundance of fruit, size of udder, stands of course in obvious relation to supply of food. Really, we no more know the reason why the progeny occasionally deviates from the parent than we do why it usually resembles it. Though the laws and conditions governing variation are known to a certain extent, those governing inheritance are apparently inscrutable. "Perhaps," Darwin remarks, "the correct way of viewing the whole subject would be, to look at the inheritance of every character whatever as the rule, and non-inheritance as the anomaly." This, from general and obvious considerations, we have long been accustomed to do. Now, as exceptional instances are expected to be capable of explanation, while ultimate laws are not, it is quite possible that variation may be accounted for, while the great primary law of inheritance remains a mysterious fact.
The common proposition is, that species reproduce their like; this is a sort of general inference, only a degree closer to fact than the statement that genera reproduce their like. The true proposition, the fact incapable of further analysis, is, that individuals reproduce their like—that characteristics are inheritable. So varieties, or deviations, once originated, are perpetuable, like species. Not so likely to be perpetuated, at the outset; for the new form tends to resemble a grandparent and a long line of similar ancestors, as well as to resemble its immediate progenitors. Two forces which coincide in the ordinary case, where the offspring resembles its parent, act in different directions when it does not and it is uncertain which will prevail. If the remoter but very potent ancestral influence predominates, the variation disappears with the life of the individual. If that of the immediate parent—feebler no doubt, but closer—the variety survives in the offspring; whose progeny now has a redoubled tendency to produce its own like; whose progeny again is almost sure to produce its like, since it is much the same whether it takes after its mother or its grandmother.
In this way races arise, which under favorable conditions may be as hereditary as species. In following these indications, watching opportunities, and breeding only from those individuals which vary most in a desirable direction, man leads the course of variation as he leads a streamlet—apparently at will, but never against the force of gravitation—to a long distance from its source, and makes it more subservient to his use or fancy. He unconsciously strengthens those variations which he prizes when he plants the seed of a favorite fruit, preserves a favorite domestic animal, drowns the uglier kittens of a litter, and allows only the handsomest or the best mousers to propagate. Still more, by methodical selection, in recent times almost marvelous results have been produced in new breeds of cattle, sheep, and poultry, and new varieties of fruit of greater and greater size or excellence.
It is said that all domestic varieties, if left to run wild, would revert to their aboriginal stocks. Probably they would wherever various races of one species were left to commingle. At least the abnormal or exaggerated characteristics induced by high feeding, or high cultivation and prolonged close breeding, would promptly disappear; and the surviving stock would soon blend into a homogeneous result (in a way presently explained), which would naturally be taken for the original form; but we could seldom know if it were so. It is by no means certain that the result would be the same if the races ran wild each in a separate region. Dr. Hooker doubts if there is a true reversion in the case of plants. Mr. Darwin's observations rather favor it in the animal kingdom. With mingled races reversion seems well made out in the case of pigeons. The common opinion upon this subject therefore probably has some foundation, But even if we regard varieties as oscillations around a primitive centre or type, still it appears from the readiness with which such varieties originate that a certain amount of disturbance would carry them beyond the influence of the primordial attraction, where they may become new centres of variation.
Some suppose that races cannot be perpetuated indefinitely even by keeping up the conditions under which they were fixed; but the high antiquity of several, and the actual fixity of many of them, negative this assumption. "To assert that we could not breed our cart and race horses, long and short horned cattle, and poultry of various breeds, for almost an infinite number of generations, would be opposed to all experience."