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effects of disuse, rudimentary and would at last be wholly suppressed; for the variations towards diminished size would no longer be checked by natural selection. The principle of the economy of growth, explained in a former chapter, by which the materials forming any part, if not useful to the possessor, are saved as far as is possible, will perhaps come into play in rendering a useless part rudimentary. But this principle will almost necessarily be confined to the earlier stages of the process of reduction; for we cannot suppose that a minute papilla, for instance, representing in a male flower the pistil of the female flower, and formed merely of cellular tissue, could be further reduced or absorbed for the sake of economising nutriment.
Finally, as rudimentary organs, by whatever steps they may have been degraded into their present useless condition, are the record of a former state of things, and have been retained solely through the power of inheritance,—we can understand, on the genealogical view of classification, how it is that systematists, in placing organisms in their proper places in the natural system, have often found rudimentary parts as useful as, or even sometimes more useful than, parts of high physiological importance. Rudimentary organs may be compared with the letters in a word, still retained in the spelling, but become useless in the pronunciation, but which serve as a clue for its derivation. On the view of descent with modification, we may conclude that the existence of organs in a rudimentary, imperfect, and useless condition, or quite aborted, far from presenting a strange difficulty, as they assuredly do on the old doctrine of creation, might even have been anticipated in accordance with the views here explained.
In this chapter I have attempted to show, that the arrangement of all organic beings throughout all time in groups under groups— that the nature of the relationships by which all living and extinct organisms are united by complex, radiating, and circuitous lines of affinities into a few grand classes,—the rules followed and the difficulties encountered by naturalists in their classifications,— the value set upon characters, if constant and prevalent, whether of high or of the most trifling importance, or, as with rudimentary organs, of no importance,—the wide opposition in value between analogical or adaptive characters, and characters of true affinity; and other such rules ;—all naturally follow if we admit the common parentage of allied forms, together with their modification through variation and natural selection, with the contingencies of extinction and divergence of character. In considering this view of classification, it should be borne in mind that the element of descent has been universally used in ranking together the sexes, ages, dimorphic forms, and acknowledged varieties of the same species, however much they may differ from each other in structure. If we extend the use of this element of descent,—the one certainly known cause of similarity in organic beings,— we shall understand what is meant by the Natural System: it is genealogical in its attempted arrangement, with the grades of acquired difference marked by the terms, varieties, species, genera, families, orders, and classes.
On this same view of descent with modification, most of the great facts in Morphology become intelligible,—whether we look to the same pattern displayed by the different species of the same class in their homologous organs, to whatever purpose applied; or to the serial and lateral homologies in each individual animal and plant.
On the principle of successive slight variations, not necessarily or generally supervening at a very early period of life, and being inherited at a corresponding period, we can understand the leading facts in Embryology ; namely, the close resemblance in the individual embryo of the parts which are homologous, and which when matured become widely different in structure and function; and the resemblance of the homologous parts or organs in allied though distinct species, though fitted in the adult state for habits as different as is possible. Larva, are active embryos, which have been specially modified in a greater or less degree in relation to their habits of life, with their modifications inherited at a corresponding early age. On these same principles,—and bearing in mind, that when organs are reduced in size, either from disuse or through natural selection, it will generally be at that period of life when the being has to provide for its own wants, and bearing in mind how strong is the force of inheritance—the occurrence of rudimentary organs might even have been anticipated. The importance of embryological characters and of rudimentary organs in classification is intelligible, on the view that a natural arrangement must be genealogical.
Finally, the several classes of facts which have been considered in this chapter, seem to me to proclaim so plainly, that the innumerable species, genera and families, with which this world is peopled, are all descended, each within its own class or group, from common parents, and have all been modified in the course of descent, that I should without hesitation adopt this view, even if it were unsupported by other facts or arguments.
Eecapitulation And Conclusion.
Recapitulation of the objections to the theory of Natural Selection — Recapitulation of the general and special circumstances in its favour — Causes of the general belief in the immutability of species — How far the theory of Natural Selection may be extended — Effects of its adoption on the study of Natural History — Concluding remarks.
As this whole volume is one long argument, it may be convenient to the reader to have the leading facts and inferences briefly recapitulated.
That many and serious objections may be advanced against the theory of descent with modification through variation and natural selection, I do not deny. I have endeavoured to give to them their full force. Nothing at first can appear more difficult to believe than that the more complex organs and instincts have been perfected, not by means superior to, though analogous with, human reason, but by the accumulation of innumerable slight variations, each good for the individual possessor. Nevertheless, this difficulty, though appearing to our imagination insuperably great, cannot be considered real if we admit the following propositions, namely, that all parts of the organisation and instincts offer, at least, individual differences—that there is a struggle for existence leading to the preservation of profitable deviations of structure or instinct—and, lastly, that gradations in the state of perfection of each organ may have existed, each good of its kind. The truth of these proposition* cannot, I think, be disputed.
It is, no doubt, extremely difficult even to conjecture by what gradations many structures have been perfected, more especially amongst broken and failing groups of organic beings, which have suffered much extinction; but we see so many strange gradations in nature, that we ought to be extremely cautious in saying that any organ or instinct, or any whole structure, could not have arrived at its present state by many graduated steps. There are, *\ it must be admitted, cases of special difficulty opposed to the 1 theory of natural selection; and one of the most curious of these . is the existence in the same community of two or three defined castes of workers or sterile female ants; but I have attempted to show how these difficulties can be mastered.
With respect to the almost universal sterility of species when first crossed, which forms so remarkable a contrast with the almost universal fertility of varieties when crossed, I must refer the reader to the recapitulation of the facts given at the end of the ninth chapter, which seem to me conclusively to show that this sterility is no more a special endowment than is the incapacity of two distinct kinds of trees to be grafted together; but that it is incidental on differences confined to the reproductive systems of the intercrossed species. We see the truth of this conclusion in the vast difference in the results of crossing the same two species reciprocally,—that is, when one species is first used as the father and then as the mother. Analogy from the consideration of dimorphic and trimorphic plants clearly leads to the same conclusion, for when the forms are illegitimately united, they yield few or no seed, and their offspring are more or less sterile; and these forms belong to the same undoubted species, and differ from each other in no respect except in their reproductive organs and functions.
Although the fertility of varieties when intercrossed and of their mongrel offspring has been asserted by so many authors to be universal, this cannot be considered as quite correct after the facts given on the high authority of Gartner and Kb'lreuter. Most of the varieties which have been experimented on have been produced under domestication; and as domestication (I do not mean mere confinement) almost certainly tends to eliminate that sterility which, judging from analogy, would have affected the parent-species if intercrossed, we ought not to expect that domestication would likewise induce sterility in their modified descendants when crossed. This elimination of sterility apparently follows from the same cause which allows our domestic animals to breed freely under diversified circumstances; and this again apparently follows from their having been gradually accustomed to frequent changes in their conditions of life.
A double and parallel series of facts seems to throw much light on the sterility of species, when first crossed, and of their hybrid offspring. On the one side, there is good reason to believe that slight changes in the conditions of life give vigour and fertility to all organic beings. We know also that a cross between the distinct individuals of the same variety, and between distinct varieties, increases the number of their offspring, and certainly gives to them increased size and vigour. This is chiefly owing to the forms which are crossed having been exposed to somewhat different conditions of life; for I have ascertained by a laborious series of experiments that if all the individuals of the same variety be subjected during several generations to the same conditions, the good derived from crossing is often much diminished or wholly disappears. This is one side of the case. On the other side, we know that species which have long been exposed to nearly uniform conditions, when they are subjected under confinement to new and greatly changed conditions, either perish, or if they survive, are rendered sterile, though retaining perfect health. This does not occur, or only in a very slight degree, with our domesticated productions, which have long been exposed to fluctuating conditions. Hence, when we find that hybrids produced by a cross between two distinct species are few in number, owing to their perishing soon alter conception or at a very early age, or if surviving that they are rendered more or less sterile, it seems highly probable that this result is due to their having been in fact subjected to a great change in their conditions of life, from being compounded of two distinct organisations. He who will explain in a definite manner why, for instance, an elephant or a fox will not breed under confinement in its native country, whilst the domestic pig or dog will breed freely under the most diversified conditions, will at the same time be able to give a definite answer to the question why two distinct species, when crossed, as well as their hybrid offspring, are generally rendered more or less sterile, whilst two domesticated varieties when crossed and their mongrel offspring are perfectly fertile.
Turning to geographical distribution, the difficulties encountered on the theory of descent with modification are serious enough. All the individuals of the same species, and all the species of the same genus, or even higher group, are descended from common parents; and therefore, in however distant and isolated parts of the world they may now be found, they must in the course of successive generations have travelled from some one point to all the others. We are often wholly unable even to conjecture how this could have been effected. Yet, as we have reason to believe that some species have retained the same specific form for very long periods of time, immensely long as measured by years, too much stress ought not to be laid on the occasional wide diffusion of the same species; for during very long periods there will always have been a good chance for wide migration by many means. A broken or interrupted range may often be accounted for by the extinction of the species in the intermediate regions. It cannot be denied