The Origin of Species by means of Natural Selection (6th ed), Charles Darwin [the beginning after the end read novel .TXT] 📗

In the great class of molluscs, though the parts in distinct species can be shown to be homologous, only a few serial homologies; such as the valves of Chitons, can be indicated; that is, we are seldom enabled to say that one part is homologous with another part 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.

But morphology is a much more complex subject than it at first appears, as has lately been well shown in a remarkable paper by Mr. E. Ray Lankester, who has drawn an important distinction between certain classes of cases which have all been equally ranked by naturalists as homologous. He proposes to call the structures which resemble each other in distinct animals, owing to their descent from a common progenitor with subsequent modification, “homogenous”; and the resemblances which cannot thus be accounted for, he proposes to call “homoplastic”. For instance, he believes that the hearts of birds and mammals are as a whole homogenous—

that is, have been derived from a common progenitor; but that the four cavities of the heart in the two classes are homoplastic—that is, have been independently developed. Mr. Lankester also adduces the close resemblance of the parts on the right and left sides of the body, and in the successive segments of the same individual animal; and here we have parts commonly called homologous which bear no relation to the descent of distinct species from a common progenitor. Homoplastic structures are the same with those which I have classed, though in a very imperfect manner, as analogous modifications or resemblances. Their formation may be attributed in part to distinct organisms, or to distinct parts of the same organism, having varied in an analogous manner; and in part to similar modifications, having been preserved for the same general purpose or function, of which many instances have been given.

Naturalists frequently speak of the skull as formed of metamorphosed vertebrae; the jaws of crabs as metamorphosed legs; the stamens and pistils in flowers as metamorphosed leaves; but it would in most cases be more correct, as Professor Huxley has remarked, to speak of both skull and vertebrae, jaws and legs, etc., as having been metamorphosed, not one from the other, as they now exist, but from some common and simpler element.

Most 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 converted into skulls or jaws. Yet so strong is the appearance of this having occurred that naturalists can hardly avoid employing language having this plain signification. According to the views here maintained, such language may be used literally; and the wonderful fact of the jaws, for instance, of a crab retaining numerous characters, which they probably would have retained through inheritance, if they had really been metamorphosed from true though extremely simple legs, is in part explained.

DEVELOPMENT AND EMBRYOLOGY.

This is one of the most important subjects in the whole round of natural history. The metamorphoses of insects, with which every one is familiar, are generally effected abruptly by a few stages; but the transformations are in reality numerous and gradual, though concealed. A certain ephemerous insect (Chloeon) during its development, moults, as shown by Sir J. Lubbock, above twenty times, and each time undergoes a certain amount of change; and in this case we see the act of metamorphosis performed in a primary and gradual manner. Many insects, and especially certain crustaceans, show us what wonderful changes of structure can be effected during development. Such changes, however, reach their acme in the so-called alternate generations of some of the lower animals. It is, for instance, an astonishing fact that a delicate branching coralline, studded with polypi, and attached to a submarine rock, should produce, first by budding and then by transverse division, a host of huge floating jelly-fishes; and that these should produce eggs, from which are hatched swimming animalcules, which attach themselves to rocks and become developed into branching corallines; and so on in an endless cycle. The belief in the essential identity of the process of alternate generation and of ordinary metamorphosis has been greatly strengthened by Wagner’s discovery of the larva or maggot of a fly, namely the Cecidomyia, producing asexually other larvae, and these others, which finally are developed into mature males and females, propagating their kind in the ordinary manner by eggs.

It may be worth notice that when Wagner’s remarkable discovery was first announced, I was asked how was it possible to account for the larvae of this fly having acquired the power of a sexual reproduction. As long as the case remained unique no answer could be given. But already Grimm has shown that another fly, a Chironomus, reproduces itself in nearly the same manner, and he believes that this occurs frequently in the order. It is the pupa, and not the larva, of the Chironomus which has this power; and Grimm further shows that this case, to a certain extent, “unites that of the Cecidomyia with the parthenogenesis of the Coccidae;” the term parthenogenesis implying that the mature females of the Coccidae are capable of producing fertile eggs without the concourse of the male.

Certain animals belonging to several classes are now known to have the power of ordinary reproduction at an unusually early age; and we have only to accelerate parthenogenetic reproduction by gradual steps to an earlier and earlier age—Chironomus showing us an almost exactly intermediate stage, viz., that of the pupa—and we can perhaps account for the marvellous case of the Cecidomyia.

It has already been stated that various parts in the same individual, which are exactly alike during an early embryonic period, become widely different and serve for widely different purposes in the adult state. So again it has been shown that generally the embryos of the most distinct species belonging to the same class are closely similar, but become, when fully developed, widely dissimilar. A better proof of this latter fact cannot be given than the statement by Von Baer that “the embryos of mammalia, of birds, lizards and snakes, probably also of chelonia, are in the earliest states exceedingly like one another, both as a whole and in the mode of development of their parts; so much so, in fact, that we can often distinguish the embryos only by their size. In my possession are two little embryos in spirit, whose names I have omitted to attach, and at present I am quite unable to say to what class they belong. They may be lizards or small birds, or very young mammalia, so complete is the similarity in the mode of formation of the head and trunk in these animals.

The extremities, however, are still absent in these embryos. But even if they had existed in the earliest stage of their development we should learn nothing, for the feet of lizards and mammals, the wings and feet of birds, no less than the hands and feet of man, all arise from the same fundamental form.” The larvae of most crustaceans, at corresponding stages of development, closely resemble each other, however different the adults may become; and so it is with very many other animals. A trace of the law of embryonic resemblance occasionally lasts till a rather late age: thus birds of the same genus, and of allied genera, often resemble each other in their immature plumage; as we see in the spotted feathers in the young of the thrush group. In the cat tribe, most of the species when adult are striped or spotted in lines; and stripes or spots can be plainly distinguished in the whelp of the lion and the puma. We occasionally, though rarely, see something of the same kind in plants; thus the first leaves of the ulex or furze, and the first leaves of the phyllodineous acacias, are pinnate or divided like the ordinary leaves of the leguminosae.

The points of structure, in which the embryos of widely different animals within 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 courses 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 similar 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 supposes that the stripes on the whelp of a lion, or the spots on the young blackbird, are of any use to these animals.

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. In how important a manner this has acted, has recently been well shown by Sir J. Lubbock in his remarks on the close similarity of the larvae of some insects belonging to very different orders, and on the dissimilarity of the larvae of other insects within the same order, according to their habits of life. Owing to such adaptations the similarity of the larvae of allied animals is sometimes greatly obscured; especially when there is a division of labour during the different stages of development, as when the same larva has during one stage to search for food, and during another stage has to search for a place of attachment. Cases can even be given of the larvae of allied species, or groups of species, differing more from each other than do the adults. 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 a crustacean: but a glance at the larva shows this in an unmistakable manner. So again the two main divisions of cirripedes, the pedunculated and sessile, though differing widely in external appearance, have larvae in all their 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 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 must be 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 locomotive organs, a 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 out 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