The Origin of Species.
by Thomas H. Huxley.
MR. DARWIN"S long-standing and well-earned scientific eminence probably renders him indifferent to that social notoriety which pa.s.ses by the name of success; but if the calm spirit of the philosopher have not yet wholly superseded the ambition and the vanity of the carnal man within him, he must be well satisfied with the results of his venture in publishing the "Origin of Species". Overflowing the narrow bounds of purely scientific circles, the "species question" divides with Italy and the Volunteers the attention of general society. Everybody has read Mr.
Darwin"s book, or, at least, has given an opinion upon its merits or demerits; pietists, whether lay or ecclesiastic, decry it with the mild railing which sounds so charitable; bigots denounce it with ignorant invective; old ladies of both s.e.xes consider it a decidedly dangerous book, and even savants, who have no better mud to throw, quote antiquated writers to show that its author is no better than an ape himself; while every philosophical thinker hails it as a veritable Whitworth gun in the armoury of liberalism; and all competent naturalists and physiologists, whatever their opinions as to the ultimate fate of the doctrines put forth, acknowledge that the work in which they are embodied is a solid contribution to knowledge and inaugurates a new epoch in natural history.
Nor has the discussion of the subject been restrained within the limits of conversation. When the public is eager and interested, reviewers must minister to its wants; and the genuine "litterateur" is too much in the habit of acquiring his knowledge from the book he judges--as the Abyssinian is said to provide himself with steaks from the ox which carries him--to be withheld from criticism of a profound scientific work by the mere want of the requisite preliminary scientific acquirement; while, on the other hand, the men of science who wish well to the new views, no less than those who dispute their validity, have naturally sought opportunities of expressing their opinions. Hence it is not surprising that almost all the critical journals have noticed Mr.
Darwin"s work at greater or less length; and so many disquisitions, of every degree of excellence, from the poor product of ignorance, too often stimulated by prejudice, to the fair and thoughtful essay of the candid student of Nature, have appeared, that it seems an almost hopeless task to attempt to say anything new upon the question.
But it may be doubted if the knowledge and ac.u.men of prejudged scientific opponents, or the subtlety of orthodox special pleaders, have yet exerted their full force in mystifying the real issues of the great controversy which has been set afoot, and whose end is hardly likely to be seen by this generation; so that, at this eleventh hour, and even failing anything new, it may be useful to state afresh that which is true, and to put the fundamental positions advocated by Mr. Darwin in such a form that they may be grasped by those whose special studies lie in other directions. And the adoption of this course may be the more advisable, because, notwithstanding its great deserts, and indeed partly on account of them, the "Origin of Species" is by no means an easy book to read--if by reading is implied the full comprehension of an author"s meaning.
We do not speak jestingly in saying that it is Mr. Darwin"s misfortune to know more about the question he has taken up than any man living.
Personally and practically exercised in zoology, in minute anatomy, in geology; a student of geographical distribution, not on maps and in museums only, but by long voyages and laborious collection; having largely advanced each of these branches of science, and having spent many years in gathering and sifting materials for his present work, the store of accurately registered facts upon which the author of the "Origin of Species" is able to draw at will is prodigious.
But this very superabundance of matter must have been embarra.s.sing to a writer who, for the present, can only put forward an abstract of his views; and thence it arises, perhaps, that notwithstanding the clearness of the style, those who attempt fairly to digest the book find much of it a sort of intellectual pemmican--a ma.s.s of facts crushed and pounded into shape, rather than held together by the ordinary medium of an obvious logical bond; due attention will, without doubt, discover this bond, but it is often hard to find.
Again, from sheer want of room, much has to be taken for granted which might readily enough be proved; and hence, while the adept, who can supply the missing links in the evidence from his own knowledge, discovers fresh proof of the singular thoroughness with which all difficulties have been considered and all unjustifiable suppositions avoided, at every reperusal of Mr. Darwin"s pregnant paragraphs, the novice in biology is apt to complain of the frequency of what he fancies is gratuitous a.s.sumption.
Thus while it may be doubted if, for some years, any one is likely to be competent to p.r.o.nounce judgment on all the issues raised by Mr. Darwin, there is a.s.suredly abundant room for him, who, a.s.suming the humbler, though perhaps as useful, office of an interpreter between the "Origin of Species" and the public, contents himself with endeavouring to point out the nature of the problems which it discusses; to distinguish between the ascertained facts and the theoretical views which it contains; and finally, to show the extent to which the explanation it offers satisfies the requirements of scientific logic. At any rate, it is this office which we purpose to undertake in the following pages.
It may be safely a.s.sumed that our readers have a general conception of the nature of the objects to which the word "species" is applied; but it has, perhaps, occurred to a few, even to those who are naturalists "ex professo", to reflect, that, as commonly employed, the term has a double sense and denotes two very different orders of relations. When we call a group of animals, or of plants, a species, we may imply thereby, either that all these animals or plants have some common peculiarity of form or structure; or, we may mean that they possess some common functional character. That part of biological science which deals with form and structure is called Morphology--that which concerns itself with function, Physiology--so that we may conveniently speak of these two senses, or aspects, of "species"--the one as morphological, the other as physiological. Regarded from the former point of view, a species is nothing more than a kind of animal or plant, which is distinctly definable from all others, by certain constant, and not merely s.e.xual, morphological peculiarities. Thus horses form a species, because the group of animals to which that name is applied is distinguished from all others in the world by the following constantly a.s.sociated characters.
They have--1, A vertebral column; 2, Mammae; 3, A placental embryo; 4, Four legs; 5, A single well-developed toe in each foot provided with a hoof; 6, A bushy tail; and 7, Callosities on the inner sides of both the fore and the hind legs. The a.s.ses, again, form a distinct species, because, with the same characters, as far as the fifth in the above list, all a.s.ses have tufted tails, and have callosities only on the inner side of the fore-legs. If animals were discovered having the general characters of the horse, but sometimes with callosities only on the fore-legs, and more or less tufted tails; or animals having the general characters of the a.s.s, but with more or less bushy tails, and sometimes with callosities on both pairs of legs, besides being intermediate in other respects--the two species would have to be merged into one. They could no longer be regarded as morphologically distinct species, for they would not be distinctly definable one from the other.
However bare and simple this definition of species may appear to be, we confidently appeal to all practical naturalists, whether zoologists, botanists, or palaeontologists, to say if, in the vast majority of cases, they know, or mean to affirm anything more of the group of animals or plants they so denominate than what has just been stated.
Even the most decided advocates of the received doctrines respecting species admit this.
"I apprehend," says Professor Owen [2], "that few naturalists nowadays, in describing and proposing a name for what they call "a new species,"
use that term to signify what was meant by it twenty or thirty years ago; that is, an originally distinct creation, maintaining its primitive distinction by obstructive generative peculiarities. The proposer of the new species now intends to state no more than he actually knows; as, for example, that the differences on which he founds the specific character are constant in individuals of both s.e.xes, so far as observation has reached; and that they are not due to domestication or to artificially superinduced external circ.u.mstances, or to any outward influence within his cognizance; that the species is wild, or is such as it appears by Nature."
If we consider, in fact, that by far the largest proportion of recorded existing species are known only by the study of their skins, or bones, or other lifeless exuvia; that we are acquainted with none, or next to none, of their physiological peculiarities, beyond those which can be deduced from their structure, or are open to cursory observation; and that we cannot hope to learn more of any of those extinct forms of life which now const.i.tute no inconsiderable proportion of the known Flora and Fauna of the world: it is obvious that the definitions of these species can be only of a purely structural, or morphological, character. It is probable that naturalists would have avoided much confusion of ideas if they had more frequently borne the necessary limitations of our knowledge in mind. But while it may safely be admitted that we are acquainted with only the morphological characters of the vast majority of species--the functional or physiological, peculiarities of a few have been carefully investigated, and the result of that study forms a large and most interesting portion of the physiology of reproduction.
The student of Nature wonders the more and is astonished the less, the more conversant he becomes with her operations; but of all the perennial miracles she offers to his inspection, perhaps the most worthy of admiration is the development of a plant or of an animal from its embryo. Examine the recently laid egg of some common animal, such as a salamander or newt. It is a minute spheroid in which the best microscope will reveal nothing but a structureless sac, enclosing a glairy fluid, holding granules in suspension. But strange possibilities lie dormant in that semi-fluid globule. Let a moderate supply of warmth reach its watery cradle, and the plastic matter undergoes changes so rapid, yet so steady and purposelike in their succession, that one can only compare them to those operated by a skilled modeller upon a formless lump of clay. As with an invisible trowel, the ma.s.s is divided and subdivided into smaller and smaller portions, until it is reduced to an aggregation of granules not too large to build withal the finest fabrics of the nascent organism. And, then, it is as if a delicate finger traced out the line to be occupied by the spinal column, and moulded the contour of the body; pinching up the head at one end, the tail at the other, and fashioning flank and limb into due salamandrine proportions, in so artistic a way, that, after watching the process hour by hour, one is almost involuntarily possessed by the notion, that some more subtle aid to vision than an achromatic, would show the hidden artist, with his plan before him, striving with skilful manipulation to perfect his work.
As life advances, and the young amphibian ranges the waters, the terror of his insect contemporaries, not only are the nutritious particles supplied by its prey, by the addition of which to its frame, growth takes place, laid down, each in its proper spot, and in such due proportion to the rest, as to reproduce the form, the colour, and the size, characteristic of the parental stock; but even the wonderful powers of reproducing lost parts possessed by these animals are controlled by the same governing tendency. Cut off the legs, the tail, the jaws, separately or all together, and, as Spallanzani showed long ago, these parts not only grow again, but the redintegrated limb is formed on the same type as those which were lost. The new jaw, or leg, is a newt"s, and never by any accident more like that of a frog. What is true of the newt is true of every animal and of every plant; the acorn tends to build itself up again into a woodland giant such as that from whose twig it fell; the spore of the humblest lichen reproduces the green or brown incrustation which gave it birth; and at the other end of the scale of life, the child that resembled neither the paternal nor the maternal side of the house would be regarded as a kind of monster.
So that the one end to which, in all living beings, the formative impulse is tending--the one scheme which the Archaeus 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 phenomena of vitality are not something apart from other physical phenomena, 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 ma.s.s of matter, as the complex forces of a magnet are held in the steel by its coercive force; and, since the differences of s.e.x 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 a.n.a.logy 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 whose members 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 ent.i.tle 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" he 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 ident.i.ty, 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 s.e.xes 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. Mult.i.tudes, 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 ill.u.s.trative 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 Ma.s.sachusetts, 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 a.s.signable 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 Reaumur, in his "Art de faire eclore 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 a.s.signed for the appearance of this unusual variety of the human species.
Two circ.u.mstances 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 circ.u.mstances; 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 a.s.suredly competent to alter the character of the tegumentary covering, to change colour, to increase or diminish the size of muscles, to modify const.i.tution, 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, 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 pre-potent 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, Andre, 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 Andre 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 Ma.s.sachusetts 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 antic.i.p.ations, 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. [3] 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 s.e.xes, 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 a.s.sume 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 const.i.tution, 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 ent.i.ty 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 mult.i.tude 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 circ.u.mstances, 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 a.s.sertion, 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 s.e.xual, 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 circ.u.mstances, 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 a.s.s, 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 const.i.tution 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, p.r.o.nounce 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 circ.u.mspection, 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 a.n.a.logous 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 circ.u.mstances; 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 circ.u.mstances--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 pa.s.sage; 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 a.s.semblages 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.