Chapter 15
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, Pt. 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, whose skeleton 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 grandchildren 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 condition; 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 whose transmission 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-Sequard, 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.
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 Dr. 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 vertebrae 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 phenomena of hybridization--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 only 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 the same 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 superadded 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 hybridization 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, whose male element will fertilize 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:--
"First crosses between forms sufficiently distinct to be ranked as species, and their hybrids, are very generally, but not universally, sterile. The sterility is of all degrees, and is often so slight that the two most careful experimentalists who have ever lived have come to diametrically opposite conclusions in ranking forms by this test. The sterility is innately variable in individuals of the same species, and is eminently susceptible of favourable and unfavourable conditions. The degree of sterility does not strictly follow systematic affinity, but is governed by several curious and complex laws. It is generally different and sometimes widely different, in reciprocal crosses between the same two species. It is not always equal in degree in a first cross, and in the hybrid produced from this cross.
"In the same manner as in grafting trees, the capacity of one species or variety to take on another is incidental on generally unknown differences in their vegetative systems; so in crossing, the greater or less facility of one species to unite with another is incidental on unknown differences in their reproductive systems. There is no more reason to think that species have been specially endowed with various degrees of sterility to prevent them crossing and breeding in Nature, than to think that trees have been specially endowed with various and somewhat analogous degrees of difficulty in being grafted together, in order to prevent them becoming inarched in our forests.
"The sterility of first crosses between pure species, which have their reproductive systems perfect, seems to depend on several circumstances; in some cases largely on the early death of the embryo. The sterility of hybrids which have their reproductive systems imperfect, and which have had this system and their whole organization disturbed by being compounded of two distinct species, seems closely allied to that sterility which so frequently affects pure species when their natural conditions of life have been disturbed. This view is supported by a parallelism of another kind: namely, that the crossing of forms, only slightly different, is favourable to the vigour and fertility of the offspring; and that slight changes in the conditions of life are apparently favourable to the vigour and fertility of all organic beings. It is not surprising that the degree of difficulty in uniting two species, and the degree of sterility of their hybrid offspring, should generally correspond, though due to distinct causes; for both depend on the amount of difference of some kind between the species which are crossed. Nor is it surprising that the facility of effecting a first cross, the fertility of hybrids produced from it, and the capacity of being grafted together--though this latter capacity evidently depends on widely different circumstances--should all run to a certain extent parallel with the systematic affinity of the forms which are subjected to experiment; for systematic affinity attempts to express all kinds of resemblance between all species.
"First crosses between forms known to be varieties, or sufficiently alike to be considered as varieties, and their mongrel offspring, are very generally, but not quite universally, fertile. Nor is this nearly general and perfect fertility surprising, when we remember how liable we are to argue in a circle with respect to varieties in a state of Nature; and when we remember that the greater number of varieties have been produced under domestication by the selection of mere external differences, and not of differences in the reproductive system. In all other respects, excluding fertility, there is a close general resemblance between hybrids and mongrels."--Pp. 276-8.
We fully agree with the general tenor of this weighty passage; but forcible as are these arguments, and little as the value of fertility or infertility as a test of species may be, it must not be forgotten that the really important fact, so far as the inquiry into the origin of species goes, is, that there are such things in Nature as groups of animals and of plants, whose members are incapable of fertile union with those of other groups; and that there are such things as hybrids, which are absolutely sterile when crossed with other hybrids. For, if such phenomena as these were exhibited by only two of those assemblages of living objects, to which the name of species (whether it be used in its physiological or in its morphological sense) is given, it would have to be accounted for by any theory of the origin of species, and every theory which could not account for it would be, so far, imperfect.
Up to this point, we have been dealing with matters of fact, and the statements which we have laid before the reader would, to the best of our knowledge, be admitted to contain a fair exposition of what is at present known respecting the essential properties of species, by all who have studied the question. And whatever may be his theoretical views, no naturalist will probably be disposed to demur to the following summary of that exposition:--
Living beings, whether animals or plants, are divisible into multitudes of distinctly definable kinds, which are morphological species. They are also divisible into groups of individuals, which breed freely together, tending to reproduce their like, and are physiological species. Normally resembling their parents, the offspring of members of these species are still liable to vary; and the variation may be perpetuated by selection, as a race, which race, in many cases, presents all the characteristics of a morphological species. But it is not as yet proved that a race ever exhibits, when crossed with another race of the same species, those phenomena of hybridization which are exhibited by many species when crossed with other species. On the other hand, not only is it not proved that all species give rise to hybrids infertile inter se, but there is much reason to believe that, in crossing, species exhibit every gradation from perfect sterility to perfect fertility.
Such are the most essential characteristics of species. Even were man not one of them--a member of the same system and subject to the same laws--the question of their origin, their causal connexion, that is, with the other phenomena of the universe, must have attracted his attention, as soon as his intelligence had raised itself above the level of his daily wants.
Indeed history relates that such was the case, and has embalmed for us the speculations upon the origin of living beings, which were among the earliest products of the dawning intellectual activity of man. In those early days positive knowledge was not to be had, but the craving after it needed, at all hazards, to be satisfied, and according to the country, or the turn of thought, of the speculator, the suggestion that all living things arose from the mud of the Nile, from a primeval egg, or from some more anthropomorphic agency, afforded a sufficient resting-place for his curiosity. The myths of Paganism are as dead as Osiris or Zeus, and the man who should revive them, in opposition to the knowledge of our time, would be justly laughed to scorn; but the coeval imaginations current among the rude inhabitants of Palestine, recorded by writers whose very name and age are admitted by every scholar to be unknown, have unfortunately not yet shared their fate, but, even at this day, are regarded by nine-tenths of the civilized world as the authoritative standard of fact and the criterion of the justice of scientific conclusions, in all that relates to the origin of things, and, among them, of species. In this nineteenth century, as at the dawn of modern physical science, the cosmogony of the semi-barbarous Hebrew is the incubus of the philosopher and the opprobrium of the orthodox. Who shall number the patient and earnest seekers after truth, from the days of Galileo until now, whose lives have been embittered and their good name blasted by the mistaken zeal of Bibliolaters? Who shall count the host of weaker men whose sense of truth has been destroyed in the effort to harmonize impossibilities--whose life has been wasted in the attempt to force the generous new wine of Science into the old bottles of Judaism, compelled by the outcry of the same strong party?
It is true that if philosophers have suffered, their cause has been amply avenged. Extinguished theologians lie about the cradle of every science as the strangled snakes beside that of Hercules; and history records that whenever science and orthodoxy have been fairly opposed, the latter has been forced to retire from the lists, bleeding and crushed if not annihilated; scotched, if not slain. But orthodoxy is the Bourbon of the world of thought. It learns not, neither can it forget; and though, at present, bewildered and afraid to move, it is as willing as ever to insist that the first chapter of Genesis contains the beginning and the end of sound science; and to visit, with such petty thunderbolts as its half-paralysed hands can hurl, those who refuse to degrade Nature to the level of primitive Judaism.
Philosophers, on the other hand, have no such aggressive tendencies. With eyes fixed on the noble goal to which "per aspera et ardua" they tend, they may, now and then, be stirred to momentary wrath by the unnecessary obstacles with which the ignorant, or the malicious, encumber, if they cannot bar, the difficult path; but why should their souls be deeply vexed? The majesty of Fact is on their side, and the elemental forces of Nature are working for them. Not a star comes to the meridian at its calculated time but testifies to the justice of their methods--their beliefs are "one with falling rain and with the growing corn." By doubt they are established, and open inquiry is their bosom friend. Such men have no fear of traditions however venerable, and no respect for them when they become mischievous and obstructive; but they have better than mere antiquarian business in hand, and if dogmas, which ought to be fossil but are not, are not forced upon their notice, they are too happy to treat them as non-existent.
The hypotheses respecting the origin of species which profess to stand upon a scientific basis, and, as such, alone demand serious attention, are of two kinds. The one, the "special creation" hypothesis, presumes every species to have originated from one or more stocks, these not being the result of the modification of any other form of living matter--or arising by natural agencies--but being produced, as such, by a supernatural creative act.