Chapter 3

So that the one end to which, in all living beings, the formative impulse is tending--the one scheme which the Archæus of the old speculators strives to carry out, seems to be to mould the offspring into the likeness of the parent. It is the first great law of reproduction, that the offspring tends to resemble its parent or parents, more closely than anything else.

Science will some day show us how this law is a necessary consequence of the more general laws which govern matter; but, for the present, more can hardly be said than that it appears to be in harmony with them. We know that the phænomena of vitality are not something apart from other physical phænomena, but one with them; and matter and force are the two names of the one artist who fashions the living as well as the lifeless. Hence living bodies should obey the same great laws as other matter--nor, throughout Nature, is there a law of wider application than this, that a body impelled by two forces takes the direction of their resultant. But living bodies may be regarded as nothing but extremely complex bundles of forces held in a mass of matter, as the complex forces of a magnet are held in the steel by its coercive force; and, since the differences of sex are comparatively slight, or, in other words, the sum of the forces in each has a very similar tendency, their resultant, the offspring, may reasonably be expected to deviate but little from a course parallel to either, or to both.

Represent the reason of the law to ourselves by what physical metaphor or analogy we will, however, the great matter is to apprehend its existence and the importance of the consequences deducible from it. For things which are like to the same are like to one another; and if, in a great series of generations, every offspring is like its parent, it follows that all the offspring and all the parents must be like one another; and that, given an original parental stock, with the opportunity of undisturbed multiplication, the law in question necessitates the production, in course of time, of an indefinitely large group, the whole of the members of which are at once very similar and are blood relations, having descended from the same parent, or pair of parents. The proof that all the members of any given group of animals, or plants, had thus descended, would be ordinarily considered sufficient to entitle them to the rank of physiological species, for most physiologists consider species to be definable as "the offspring of a single primitive stock."

But though it is quite true that all those groups we call species _may_, according to the known laws of reproduction, have descended from a single stock, and though it is very likely they really have done so, yet this conclusion rests on deduction and can hardly hope to establish itself upon a basis of observation. And the primitiveness of the supposed single stock, which, after all, is the essential part of the matter, is not only a hypothesis, but one which has not a shadow of foundation, if by "primitive" be meant "independent of any other living being." A scientific definition, of which an unwarrantable hypothesis forms an essential part, carries its condemnation within itself; but, even supposing such a definition were, in form, tenable, the physiologist who should attempt to apply it in Nature would soon find himself involved in great, if not inextricable, difficulties. As we have said, it is indubitable that offspring _tend_ to resemble the parental organism, but it is equally true that the similarity attained never amounts to identity either in form or in structure. There is always a certain amount of deviation, not only from the precise characters of a single parent, but when, as in most animals and many plants, the sexes are lodged in distinct individuals, from an exact mean between the two parents. And indeed, on general principles, this slight deviation seems as intelligible as the general similarity, if we reflect how complex the co-operating "bundles of forces" are, and how improbable it is that, in any case, their true resultant shall coincide with any mean between the more obvious characters of the two parents. Whatever be its cause, however, the co-existence of this tendency to minor variation with the tendency to general similarity, is of vast importance in its bearing on the question of the origin of species.

As a general rule, the extent to which an offspring differs from its parent is slight enough; but, occasionally, the amount of difference is much more strongly marked, and then the divergent offspring receives the name of a Variety. Multitudes, of what there is every reason to believe are such varieties, are known, but the origin of very few has been accurately recorded, and of these we will select two as more especially illustrative of the main features of variation. The first of them is that of the "Ancon" or "Otter" sheep, of which a careful account is given by Colonel David Humphreys, F.R.S., in a letter to Sir Joseph Banks, published in the "Philosophical Transactions" for 1813. It appears that one Seth Wright, the proprietor of a farm on the banks of the Charles River, in Massachusetts, possessed a flock of fifteen ewes and a ram of the ordinary kind. In the year 1791, one of the ewes presented her owner with a male lamb, differing, for no assignable reason, from its parents by a proportionally long body and short bandy legs, whence it was unable to emulate its relatives in those sportive leaps over the neighbours' fences, in which they were in the habit of indulging, much to the good farmer's vexation.

The second case is that detailed by a no less unexceptionable authority than Réaumur, in his "Art de faire éclore les Poulets." A Maltese couple, named Kelleia, whose hands and feet were constructed upon the ordinary human model, had born to them a son, Gratio, who possessed six perfectly movable fingers on each hand, and six toes, not quite so well formed, on each foot. No cause could be assigned for the appearance of this unusual variety of the human species.

Two circumstances are well worthy of remark in both these cases. In each, the variety appears to have arisen in full force, and, as it were, _per saltum_; a wide and definite difference appearing, at once, between the Ancon ram and the ordinary sheep; between the six-fingered and six-toed Gratio Kelleia and ordinary men. In neither case is it possible to point out any obvious reason for the appearance of the variety. Doubtless there were determining causes for these as for all other phenomena; but they do not appear, and we can be tolerably certain that what are ordinarily understood as changes in physical conditions, as in climate, in food, or the like, did not take place and had nothing to do with the matter. It was no case of what is commonly called adaptation to circumstances; but, to use a conveniently erroneous phrase, the variations arose spontaneously. The fruitless search after final causes leads their pursuers a long way; but even those hardy teleologists, who are ready to break through all the laws of physics in chase of their favourite will-o'-the-wisp, may be puzzled to discover what purpose could be attained by the stunted legs of Seth Wright's ram or the hexadactyle members of Gratio Kelleia.

Varieties then arise we know not why; and it is more than probable that the majority of varieties have arisen in this "spontaneous" manner, though we are, of course, far from denying that they may be traced, in some cases, to distinct external influences; which are assuredly competent to alter the character of the tegumentary covering, to change colour, to increase or diminish the size of muscles, to modify constitution, and, among plants, to give rise to the metamorphosis of stamens into petals, and so forth. But however they may have arisen, what especially interests us at present is, to remark that, once in existence, many varieties obey the fundamental law of reproduction that like tends to produce like; and their offspring exemplify it by tending to exhibit the same deviation from the parental stock as themselves. Indeed, there seems to be, in many instances, a prepotent influence about a newly-arisen variety which gives it what one may call an unfair advantage over the normal descendants from the same stock. This is strikingly exemplified by the case of Gratio Kelleia, who married a woman with the ordinary pentadactyle extremities, and had by her four children, Salvator, George, André, and Marie. Of these children Salvator, the eldest boy, had six fingers and six toes, like his father; the second and third, also boys, had five fingers and five toes, like their mother, though the hands and feet of George were slightly deformed. The last, a girl, had five fingers and five toes, but the thumbs were slightly deformed. The variety thus reproduced itself purely in the eldest, while the normal type reproduced itself purely in the third, and almost purely in the second and last: so that it would seem, at first, as if the normal type were more powerful than the variety. But all these children grew up and intermarried with normal wives and husband, and then, note what took place: Salvator had four children, three of whom exhibited the hexadactyle members of their grandfather and father, while the youngest had the pentadactyle limbs of the mother and grandmother; so that here, notwithstanding a double pentadactyle dilution of the blood, the hexadactyle variety had the best of it. The same pre-potency of the variety was still more markedly exemplified in the progeny of two of the other children, Marie and George. Marie (whose thumbs only were deformed) gave birth to a boy with six toes, and three other normally formed children; but George, who was not quite so pure a pentadactyle, begot, first, two girls, each of whom had six fingers and toes; then a girl with six fingers on each hand and six toes on the right foot, but only five toes on the left; and lastly, a boy with only five fingers and toes. In these instances, therefore, the variety, as it were, leaped over one generation to reproduce itself in full force in the next. Finally, the purely pentadactyle André was the father of many children, not one of whom departed from the normal parental type.

If a variation which approaches the nature of a monstrosity can strive thus forcibly to reproduce itself, it is not wonderful that less aberrant modifications should tend to be preserved even more strongly; and the history of the Ancon sheep is, in this respect, particularly instructive. With the "'cuteness" characteristic of their nation, the neighbours of the Massachusetts farmer imagined it would be an excellent thing if all his sheep were imbued with the stay-at-home tendencies enforced by Nature upon the newly-arrived ram; and they advised Wright to kill the old patriarch of his fold, and install the Ancon ram in his place. The result justified their sagacious anticipations, and coincided very nearly with what occurred to the progeny of Gratio Kelleia. The young lambs were almost always either pure Ancons, or pure ordinary sheep.[Footnote: Colonel Humphreys' statements are exceedingly explicit on this point:--. "When an Ancon ewe is impregnated by a common ram, the increase resembles wholly either the ewe or the ram. The increase of the common ewe impregnated by an Ancon ram follows entirely the one or the other, without blending any of the distinguishing and essential peculiarities of both. Frequent instances have happened where common ewes have had twins by Ancon rams, when one exhibited the complete marks and features of the ewe, the other of the ram. The contrast has been rendered singularly striking, when one short-legged and one long-legged lamb, produced at a birth, have been seen sucking the dam at the same time."--_Philosophical Transactions_, 1813, Ft. I. pp. 89, 90.] But when sufficient Ancon sheep were obtained to interbreed with one another, it was found that the offspring was always pure Ancon. Colonel Humphreys, in fact, states that he was acquainted with only "one questionable case of a contrary nature." Here, then, is a remarkable and well-established instance, not only of a very distinct race being established _per saltum_, but of that race breeding "true" at once, and showing no mixed forms, even when crossed with another breed.

By taking care to select Ancons of both sexes, for breeding from, it thus became easy to establish an extremely well-marked race; so peculiar that, even when herded with other sheep, it was noted that the Ancons kept together. And there is every reason to believe that the existence of this breed might have been indefinitely protracted; but the introduction of the Merino sheep, which were not only very superior to the Ancons in wool and meat, but quite as quiet and orderly, led to the complete neglect of the new breed, so that, in 1813, Colonel Humphreys found it difficult to obtain the specimen, the skeleton of which was presented to Sir Joseph Banks. We believe that, for many years, no remnant of it has existed in the United States.

Gratio Kelleia was not the progenitor of a race of six-fingered men, as Seth Wright's ram became a nation of Ancon sheep, though the tendency of the variety to perpetuate itself appears to have been fully as strong in the one case as in the other. And the reason of the difference is not far to seek. Seth Wright took care not to weaken the Ancon blood by matching his Ancon ewes with any but males of the same variety, while Gratio Kelleia's sons were too far removed from the patriarchal times to intermarry with their sisters; and his grand-children seem not to have been attracted by their six-fingered cousins. In other words, in the one example a race was produced, because, for several generations, care was taken to _select_ both parents of the breeding stock from animals exhibiting a tendency to vary in the same direction; while, in the other, no race was evolved, because no such selection was exercised. A race is a propagated variety; and as, by the laws of reproduction, offspring tend to assume the parental forms, they will be more likely to propagate a variation exhibited by both parents than that possessed by only one.

There is no organ of the body of an animal which may not, and does not, occasionally, vary more or less from the normal type; and there is no variation which may not be transmitted and which, if selectively transmitted, may not become the foundation of a race. This great truth, sometimes forgotten by philosophers, has long been familiar to practical agriculturists and breeders; and upon it rest all the methods of improving the breeds of domestic animals, which, for the last century, have been followed with so much success in England. Colour, form, size, texture of hair or wool, proportions of various parts, strength or weakness of constitution, tendency to fatten or to remain lean, to give much or little milk, speed, strength, temper, intelligence, special instincts; there is not one of these characters the transmission of which is not an every-day occurrence within the experience of cattle-breeders, stock-farmers, horse-dealers, and dog and poultry fanciers. Nay, it is only the other day that an eminent physiologist, Dr. Brown-Séquard, communicated to the Royal Society his discovery that epilepsy, artificially produced in guinea-pigs, by a means which he has discovered, is transmitted to their offspring. [Footnote: Compare Weismann's _Essays Upon Heredity_, p. 310, _et seq_. 1893.]

But a race, once produced, is no more a fixed and immutable entity than the stock whence it sprang; variations arise among its members, and as these variations are transmitted like any others, new races may be developed out of the pre-existing one _ad infinitum_, or, at least, within any limit at present determined. Given sufficient time and sufficiently careful selection, and the multitude of races which may arise from a common stock is as astonishing as are the extreme structural differences which they may present. A remarkable example of this is to be found in the rock-pigeon, which Mr. Darwin has, in our opinion, satisfactorily demonstrated to be the progenitor of all our domestic pigeons, of which there are certainly more than a hundred well-marked races. The most noteworthy of these races are, the four great stocks known to the "fancy" as tumblers, pouters, carriers, and fantails; birds which not only differ most singularly in size, colour, and habits, but in the form of the beak and of the skull; in the proportions of the beak to the skull; in the number of tail-feathers; in the absolute and relative size of the feet; in the presence or absence of the uropygial gland; in the number of vertebræ in the back; in short, in precisely those characters in which the genera and species of birds differ from one another.

And it is most remarkable and instructive to observe, that none of these races can be shown to have been originated by the action of changes in what are commonly called external circumstances, upon the wild rock-pigeon. On the contrary, from time immemorial pigeon-fanciers have had essentially similar methods of treating their pets, which have been housed, fed, protected and cared for in much the same way in all pigeonries. In fact, there is no case better adapted than that of the pigeons to refute the doctrine which one sees put forth on high authority, that "no other characters than those founded on the development of bone for the attachment of muscles" are capable of variation. In precise contradiction of this hasty assertion, Mr. Darwin's researches prove that the skeleton of the wings in domestic pigeons has hardly varied at all from that of the wild type; while, on the other hand, it is in exactly those respects, such as the relative length of the beak and skull, the number of the vertebrae, and the number of the tail-feathers, in which muscular exertion can have no important influence, that the utmost amount of variation has taken place.

We have said that the following out of the properties exhibited by physiological species would lead us into difficulties, and at this point they begin to be obvious; for if, as the result of spontaneous variation and of selective breeding, the progeny of a common stock may become separated into groups distinguished from one another by constant, not sexual, morphological characters, it is clear that the physiological definition of species is likely to clash with the morphological definition. No one would hesitate to describe the pouter and the tumbler as distinct species, if they were found fossil, or if their skins and skeletons were imported, as those of exotic wild birds commonly are--and without doubt, if considered alone, they are good and distinct morphological species. On the other hand, they are not physiological species, for they are descended from a common stock, the rock-pigeon.

Under these circumstances, as it is admitted on all sides that races occur in Nature, how are we to know whether any apparently distinct animals are really of different physiological species, or not, seeing that the amount of morphological difference is no safe guide? Is there any test of a physiological species? The usual answer of physiologists is in the affirmative. It is said that such a test is to be found in the phænomena of hybridisation--in the results of crossing races, as compared with the results of crossing species.

So far as the evidence goes at present, individuals, of what are certainly known to be mere races produced by selection, however distinct they may appear to be, not only breed freely together, but the offspring of such crossed races are perfectly fertile with one another. Thus, the spaniel and the greyhound, the dray-horse and the Arab, the pouter and the tumbler, breed together with perfect freedom, and their mongrels, if matched with other mongrels of the same kind, are equally fertile.

On the other hand, there can be no doubt that the individuals of many natural species are either absolutely infertile if crossed with individuals of other species, or, if they give rise to hybrid offspring, the hybrids so produced are infertile when paired together. The horse and the ass, for instance, if so crossed, give rise to the mule, and there is no certain evidence of offspring ever having been produced by a male and female mule. The unions of the rock-pigeon and the ring-pigeon appear to be equally barren of result. Here, then, says the physiologist, we have a means of distinguishing any two true species from any two varieties. If a male and a female, selected from each group, produce offspring, and that offspring is fertile with others produced in the same way, the groups are races and not species. If, on the other hand, no result ensues, or if the offspring are infertile with others produced in the same way, they are true physiological species. The test would be an admirable one, if, in the first place, it were always practicable to apply it, and if, in the second, it always yielded results susceptible of a definite interpretation. Unfortunately, in the great majority of cases, this touchstone for species is wholly inapplicable.

The constitution of many wild animals is so altered by confinement that they will not breed even with their own females, so that the negative results obtained from crosses are of no value; and the antipathy of wild animals of different species for one another, or even of wild and tame members of the same species, is ordinarily so great, that it is hopeless to look for such unions in Nature. The hermaphrodism of most plants, the difficulty in the way of insuring the absence of their own or the proper working of other pollen, are obstacles of no less magnitude in applying the test to them. And, in both animals and plants, is super-added the further difficulty, that experiments must be continued over a long time for the purpose of ascertaining the fertility of the mongrel or hybrid progeny, as well as of the first crosses from which they spring.

Not only do these great practical difficulties lie in the way of applying the hybridisation test, but even when this oracle can be questioned, its replies are sometimes as doubtful as those of Delphi. For example, cases are cited by Mr. Darwin, of plants which are more fertile with the pollen of another species than with their own; and there are others, such as certain _Fuci,_ the male element of which will fertilise the ovule of a plant of distinct species, while the males of the latter species are ineffective with the females of the first. So that, in the last-named instance, a physiologist, who should cross the two species in one way, would decide that they were true species; while another, who should cross them in the reverse way, would, with equal justice, according to the rule, pronounce them to be mere races. Several plants, which there is great reason to believe are mere varieties, are almost sterile when crossed; while both animals and plants, which have always been regarded by naturalists as of distinct species, turn out, when the test is applied, to be perfectly fertile. Again, the sterility or fertility of crosses seems to bear no relation to the structural resemblances or differences of the members of any two groups.

Mr. Darwin has discussed this question with singular ability and circumspection, and his conclusions are summed up as follows, at page 276 of his work:--