Read On The Origin Of Species Online
Authors: Charles Darwin
Finally, we have seen that natural selection, which results from the struggle for existence, and which almost inevitably induces extinction and divergence of character in the many descendants from one dominant parent-species, explains that great and universal feature in the affinities of all organic beings, namely, their subordination in group under group. We use the element of descent in classing the individuals of both sexes and of all ages, although having few characters in common, under one species; we use descent in classing acknowledged varieties, however different they may be from their parent; and I believe this element of descent is the hidden bond of connexion which naturalists have sought under the term of the Natural System. On this idea of the natural system being, in so far as it has been perfected, genealogical in its arrangement, with the grades of difference between the descendants from a common parent, expressed by the terms genera, families, orders, etc., we can understand the rules which we are compelled to follow in our classification. We can understand why we value certain resemblances far more than others; why we are permitted to use rudimentary and useless organs, or others of trifling physiological importance; why, in comparing one group with a distinct group, we summarily reject analogical or adaptive characters, and yet use these same characters within the limits of the same group. We can clearly see how it is that all living and extinct forms can be grouped together in one great system; and how the several members of each class are connected together by the most complex and radiating lines of affinities. We shall never, probably, disentangle the inextricable web of affinities between the members of any one class; but when we have a distinct object in view, and do not look to some unknown plan of creation, we may hope to make sure but slow progress.
MORPHOLOGY.
We have seen that the members of the same class, independently of their habits of life, resemble each other in the general plan of their organisation. This resemblance is often expressed by the term "unity of type;" or by saying that the several parts and organs in the different species of the class are homologous. The whole subject is included under the general name of Morphology. This is the most interesting department of natural history, and may be said to be its very soul. What can be more curious than that the hand of a man, formed for grasping, that of a mole for digging, the leg of the horse, the paddle of the porpoise, and the wing of the bat, should all be constructed on the same pattern, and should include the same bones, in the same relative positions? Geoffroy St. Hilaire has insisted strongly on the high importance of relative connexion in homologous organs: the parts may change to almost any extent in form and size, and yet they always remain connected together in the same order. We never find, for instance, the bones of the arm and forearm, or of the thigh and leg, transposed. Hence the same names can be given to the homologous bones in widely different animals. We see the same great law in the construction of the mouths of insects: what can be more different than the immensely long spiral proboscis of a sphinx-moth, the curious folded one of a bee or bug, and the great jaws of a beetle?--yet all these organs, serving for such different purposes, are formed by infinitely numerous modifications of an upper lip, mandibles, and two pairs of maxillae. Analogous laws govern the construction of the mouths and limbs of crustaceans. So it is with the flowers of plants.
Nothing can be more hopeless than to attempt to explain this similarity of pattern in members of the same class, by utility or by the doctrine of final causes. The hopelessness of the attempt has been expressly admitted by Owen in his most interesting work on the 'Nature of Limbs.' On the ordinary view of the independent creation of each being, we can only say that so it is;--that it has so pleased the Creator to construct each animal and plant.
The explanation is manifest on the theory of the natural selection of successive slight modifications,--each modification being profitable in some way to the modified form, but often affecting by correlation of growth other parts of the organisation. In changes of this nature, there will be little or no tendency to modify the original pattern, or to transpose parts. The bones of a limb might be shortened and widened to any extent, and become gradually enveloped in thick membrane, so as to serve as a fin; or a webbed foot might have all its bones, or certain bones, lengthened to any extent, and the membrane connecting them increased to any extent, so as to serve as a wing: yet in all this great amount of modification there will be no tendency to alter the framework of bones or the relative connexion of the several parts. If we suppose that the ancient progenitor, the archetype as it may be called, of all mammals, had its limbs constructed on the existing general pattern, for whatever purpose they served, we can at once perceive the plain signification of the homologous construction of the limbs throughout the whole class. So with the mouths of insects, we have only to suppose that their common progenitor had an upper lip, mandibles, and two pair of maxillae, these parts being perhaps very simple in form; and then natural selection will account for the infinite diversity in structure and function of the mouths of insects. Nevertheless, it is conceivable that the general pattern of an organ might become so much obscured as to be finally lost, by the atrophy and ultimately by the complete abortion of certain parts, by the soldering together of other parts, and by the doubling or multiplication of others,--variations which we know to be within the limits of possibility. In the paddles of the extinct gigantic sea-lizards, and in the mouths of certain suctorial crustaceans, the general pattern seems to have been thus to a certain extent obscured.
There is another and equally curious branch of the present subject; namely, the comparison not of the same part in different members of a class, but of the different parts or organs in the same individual. Most physiologists believe that the bones of the skull are homologous with--that is correspond in number and in relative connexion with--the elemental parts of a certain number of vertebrae. The anterior and posterior limbs in each member of the vertebrate and articulate classes are plainly homologous. We see the same law in comparing the wonderfully complex jaws and legs in crustaceans. It is familiar to almost every one, that in a flower the relative position of the sepals, petals, stamens, and pistils, as well as their intimate structure, are intelligible on the view that they consist of metamorphosed leaves, arranged in a spire. In monstrous plants, we often get direct evidence of the possibility of one organ being transformed into another; and we can actually see in embryonic crustaceans and in many other animals, and in flowers, that organs, which when mature become extremely different, are at an early stage of growth exactly alike.
How inexplicable are these facts on the ordinary view of creation! Why should the brain be enclosed in a box composed of such numerous and such extraordinarily shaped pieces of bone? As Owen has remarked, the benefit derived from the yielding of the separate pieces in the act of parturition of mammals, will by no means explain the same construction in the skulls of birds. Why should similar bones have been created in the formation of the wing and leg of a bat, used as they are for such totally different purposes? Why should one crustacean, which has an extremely complex mouth formed of many parts, consequently always have fewer legs; or conversely, those with many legs have simpler mouths? Why should the sepals, petals, stamens, and pistils in any individual flower, though fitted for such widely different purposes, be all constructed on the same pattern?
On the theory of natural selection, we can satisfactorily answer these questions. In the vertebrata, we see a series of internal vertebrae bearing certain processes and appendages; in the articulata, we see the body divided into a series of segments, bearing external appendages; and in flowering plants, we see a series of successive spiral whorls of leaves. An indefinite repetition of the same part or organ is the common characteristic (as Owen has observed) of all low or little-modified forms; therefore we may readily believe that the unknown progenitor of the vertebrata possessed many vertebrae; the unknown progenitor of the articulata, many segments; and the unknown progenitor of flowering plants, many spiral whorls of leaves. We have formerly seen that parts many times repeated are eminently liable to vary in number and structure; consequently it is quite probable that natural selection, during a long-continued course of modification, should have seized on a certain number of the primordially similar elements, many times repeated, and have adapted them to the most diverse purposes. And as the whole amount of modification will have been effected by slight successive steps, we need not wonder at discovering in such parts or organs, a certain degree of fundamental resemblance, retained by the strong principle of inheritance.
In the great class of molluscs, though we can homologise the parts of one species with those of another and distinct species, we can indicate but few serial homologies; that is, we are seldom enabled to say that one part or organ is homologous with another in the same individual. And we can understand this fact; for in molluscs, even in the lowest members of the class, we do not find nearly so much indefinite repetition of any one part, as we find in the other great classes of the animal and vegetable kingdoms.
Naturalists frequently speak of the skull as formed of metamorphosed vertebrae: the jaws of crabs as metamorphosed legs; the stamens and pistils of flowers as metamorphosed leaves; but it would in these cases probably be more correct, as Professor Huxley has remarked, to speak of both skull and vertebrae, both jaws and legs, etc.,--as having been metamorphosed, not one from the other, but from some common element. Naturalists, however, use such language only in a metaphorical sense: they are far from meaning that during a long course of descent, primordial organs of any kind--vertebrae in the one case and legs in the other--have actually been modified into skulls or jaws. Yet so strong is the appearance of a modification of this nature having occurred, that naturalists can hardly avoid employing language having this plain signification. On my view these terms may be used literally; and the wonderful fact of the jaws, for instance, of a crab retaining numerous characters, which they would probably have retained through inheritance, if they had really been metamorphosed during a long course of descent from true legs, or from some simple appendage, is explained.
EMBRYOLOGY.
It has already been casually remarked that certain organs in the individual, which when mature become widely different and serve for different purposes, are in the embryo exactly alike. The embryos, also, of distinct animals within the same class are often strikingly similar: a better proof of this cannot be given, than a circumstance mentioned by Agassiz, namely, that having forgotten to ticket the embryo of some vertebrate animal, he cannot now tell whether it be that of a mammal, bird, or reptile. The vermiform larvae of moths, flies, beetles, etc., resemble each other much more closely than do the mature insects; but in the case of larvae, the embryos are active, and have been adapted for special lines of life. A trace of the law of embryonic resemblance, sometimes lasts till a rather late age: thus birds of the same genus, and of closely allied genera, often resemble each other in their first and second plumage; as we see in the spotted feathers in the thrush group. In the cat tribe, most of the species are striped or spotted in lines; and stripes can be plainly distinguished in the whelp of the lion. We occasionally though rarely see something of this kind in plants: thus the embryonic leaves of the ulex or furze, and the first leaves of the phyllodineous acaceas, are pinnate or divided like the ordinary leaves of the leguminosae.
The points of structure, in which the embryos of widely different animals of the same class resemble each other, often have no direct relation to their conditions of existence. We cannot, for instance, suppose that in the embryos of the vertebrata the peculiar loop-like course of the arteries near the branchial slits are related to similar conditions,--in the young mammal which is nourished in the womb of its mother, in the egg of the bird which is hatched in a nest, and in the spawn of a frog under water. We have no more reason to believe in such a relation, than we have to believe that the same bones in the hand of a man, wing of a bat, and fin of a porpoise, are related to similar conditions of life. No one will suppose that the stripes on the whelp of a lion, or the spots on the young blackbird, are of any use to these animals, or are related to the conditions to which they are exposed.
The case, however, is different when an animal during any part of its embryonic career is active, and has to provide for itself. The period of activity may come on earlier or later in life; but whenever it comes on, the adaptation of the larva to its conditions of life is just as perfect and as beautiful as in the adult animal. From such special adaptations, the similarity of the larvae or active embryos of allied animals is sometimes much obscured; and cases could be given of the larvae of two species, or of two groups of species, differing quite as much, or even more, from each other than do their adult parents. In most cases, however, the larvae, though active, still obey more or less closely the law of common embryonic resemblance. Cirripedes afford a good instance of this: even the illustrious Cuvier did not perceive that a barnacle was, as it certainly is, a crustacean; but a glance at the larva shows this to be the case in an unmistakeable manner. So again the two main divisions of cirripedes, the pedunculated and sessile, which differ widely in external appearance, have larvae in all their several stages barely distinguishable.
The embryo in the course of development generally rises in organisation: I use this expression, though I am aware that it is hardly possible to define clearly what is meant by the organisation being higher or lower. But no one probably will dispute that the butterfly is higher than the caterpillar. In some cases, however, the mature animal is generally considered as lower in the scale than the larva, as with certain parasitic crustaceans. To refer once again to cirripedes: the larvae in the first stage have three pairs of legs, a very simple single eye, and a probosciformed mouth, with which they feed largely, for they increase much in size. In the second stage, answering to the chrysalis stage of butterflies, they have six pairs of beautifully constructed natatory legs, a pair of magnificent compound eyes, and extremely complex antennae; but they have a closed and imperfect mouth, and cannot feed: their function at this stage is, to search by their well-developed organs of sense, and to reach by their active powers of swimming, a proper place on which to become attached and to undergo their final metamorphosis. When this is completed they are fixed for life: their legs are now converted into prehensile organs; they again obtain a well-constructed mouth; but they have no antennae, and their two eyes are now reconverted into a minute, single, and very simple eye-spot. In this last and complete state, cirripedes may be considered as either more highly or more lowly organised than they were in the larval condition. But in some genera the larvae become developed either into hermaphrodites having the ordinary structure, or into what I have called complemental males: and in the latter, the development has assuredly been retrograde; for the male is a mere sack, which lives for a short time, and is destitute of mouth, stomach, or other organ of importance, excepting for reproduction.