On the Origin of Species by Charles Darwin (grave mercy .txt) π
From these considerations, I shall devote the first chapter of this Abstract to Variation under Domestication. We shall thus see that a large amount of hereditary modification is at least possible, and, what is equally or more important, we shall see how great is the power of man in accumulating by his Selection successive slight variations. I will then pass on to the variability of species in a state of nature; but I shall, unfortunately, be compelled to treat this subject far too briefly, as it can be treated properly only by giving long catalogues of facts. We shall, however, be enabled to discuss what circumstances are most favourable to variation. In the
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- Author: Charles Darwin
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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.,βas 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 kindβvertebrae in the one case and legs in the otherβhave 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,βin 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.
We are so much accustomed to see differences in structure between the embryo and the adult, and likewise a close similarity in the embryos of widely different animals within the same class, that we might be led to look at these facts as necessarily contingent in some manner on growth. But there is no obvious reason why, for instance, the wing of a bat, or the fin of a porpoise, should not have been sketched out with all the parts in proper proportion, as soon as any structure became visible in the embryo. And in some whole groups of animals and in certain members of other groups, the embryo does not at any period differ widely from the adult: thus Owen has remarked in regard to cuttle-fish, βthere is no metamorphosis; the cephalopodic character is manifested long before the parts of the embryo are completed;β and again in spiders, βthere is nothing worthy to be called a metamorphosis.β The larvae of insects, whether adapted to the most diverse and active habits, or quite inactive, being fed by their parents or placed in the midst of proper nutriment, yet nearly all pass through a similar worm-like stage of development; but in some few cases, as in that of Aphis, if we look to the admirable drawings by Professor Huxley of the development of this insect, we see no trace of the vermiform stage.
How, then, can we explain these several facts in embryology,βnamely the very general, but not universal difference in structure between the embryo and the adult;βof parts in the same individual embryo, which ultimately become very unlike and serve for diverse purposes, being at this early period of growth alike;βof embryos of different species within the same class, generally, but not universally, resembling each other;βof the structure of the embryo not being closely related to its conditions of existence, except when the embryo becomes at any period of life active and has to provide for itself;βof the embryo apparently having sometimes a higher organisation than the mature animal, into which it is developed. I believe that all these facts can be explained, as follows, on the view of descent with modification.
It is commonly assumed, perhaps from monstrosities often affecting the embryo at a very early period,
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