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, "there is no metamorphosis; the cephalopodic character is manifested long before the parts of the embryo are completed;" and again in spiders, "there is nothing worthy to be called a metamorphosis." The larvæ 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, namely the very general, but not universal difference in structure between the embryo and the adult;

of parts in the same individual embryo, which ultimately become very unlike and serve for diverse purposes, being at this early period of growth alike;-of embryos of different species within the same class, generally, but not universally, resembling each other;-of 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;—of 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, that slight variations necessarily appear at an equally early period. But we have little evidence on this headindeed the evidence rather points the other way; for it is notorious that breeders of cattle, horses, and various fancy animals, cannot positively tell, until some time after the animal has been born, what its merits or form will ultimately turn out. We see this plainly in our own children; we cannot always tell whether the child will be tall or short, or what its precise features will be. The question is not, at what period of life any variation has been caused, but at what period it is fully displayed.

The cause may have acted, and I believe generally has acted, even before the embryo is formed; and the variation may be due to the male and female sexual elements having been affected by the conditions to which either parent, or their ancestors, have been exposed. Nevertheless an effect thus caused at a very early period, even before the formation of the embryo, may appear late in life; as when an hereditary disease, which appears in old age alone, has been communicated to the offspring from the reproductive element of one parent. Or again, as when the horns of cross-bred cattle have been affected by the shape of the horns of either parent. For the welfare of a very young animal, as long as it remains in its mother's womb, or in the egg, or as long as it is nourished and protected by its parent, it must be quite unimportant whether most of its characters are fully acquired a little earlier or later in life. It would not signify, for instance, to a bird which obtained its food best by having a long beak, whether or not it assumed a beak of this particular length, as long as it was fed by its parents. Hence, I conclude, that it is quite possible, that each of the many successive modifications, by which each species has acquired its present structure, may have supervened at a not very early period of life; and some direct evidence from our domestic animals supports this view. But in other cases it is quite possibly that each successive modification, or most of them, may have appeared at an extremely early period.

I have stated in the first chapter, that there is some evidence to render it probable, that at whatever age any variation first appears in the parent, it tends to reappear at a corresponding age in the offspring. Certain variations can only appear at corresponding ages, for instance, peculiarities in the caterpillar, cocoon, or imago states of the silk-moth; or, again, in the horns of almost fullgrown cattle. But further than this, variations which, for all that we can see, might have appeared earlier or later in life, tend to appear at a corresponding age in the offspring and parent. I am far from meaning that this is invariably the case; and I could give a good many cases of variations (taking the word in the largest sense) which have supervened at an earlier age in the child than in the parent.

These two principles, if their truth be admitted, will, I believe, explain all the above specified leading facts

in embryology. But first let us look at a few analogous cases in domestic varieties. Some authors who have written on Dogs, maintain that the greyhound and bulldog, though appearing so "different, are really varieties closely allied, and have probably descended from the same wild stock; hence I was curious to see how far their puppies differed from each other: I was told by breeders that they differed just as much as their parents, and this, judging by the eye, seemed almost to be the case; but on actually measuring the old dogs and their six-days old puppies, I found that the puppies had not nearly acquired their full amount of proportional difference. So, again, I was told that the foals of cart and race-horses differed as much as the full-grown animals ; and this surprised me greatly, as I think it probable that the difference between these two breeds has been wholly caused by selection under domestication; but having had careful measurements made of the dam and of a three-days old colt of a race and heavy cart-horse, I find that the colts have by no means acquired their full amount of proportional difference.

As the evidence appears to me conclusive, that the several domestic breeds of Pigeon have descended from one wild species, I compared young pigeons of various breeds, within twelve hours after being hatched; I carefully measured the proportions (but will not here give details) of the beak, width of mouth, length of nostril and of eyelid, size of feet and length of leg, in the wild stock, in pouters, fantails, runts, barbs, dragons, carriers, and tumblers. Now some of these birds, when mature, differ so extraordinarily in length and form of beak, that they would, I cannot doubt, be ranked in distinct genera, had they been natural productions. But when the nestling birds of these several breeds were placed in a row, though most of them could be distinguished from each other, yet their proportional differences in the above specified several points were incomparably less than in the full-grown birds. Some characteristic points of difference-for instance, that of the width of mouth— could hardly be detected in the young. But there was one remarkable exception to this rule, for the young of the short-faced tumbler differed from the young of the wild rock-pigeon and of the other breeds, in all its proportions, almost exactly as much as in the adult state.

The two principles above given seem to me to explain these facts in regard to the later embryonic stages of our domestic varieties. Fanciers select their horses, dogs, and pigeons, for breeding, when they are nearly grown up: they are indifferent whether the desired qualities and structures have been acquired earlier or later in life, if the full-grown animal possesses them. And the cases just given, more especially that of pigeons, seem to show that the characteristic differences which give value to each breed, and which have been accumulated by man's selection, have not generally first appeared at an early period of life, and have been inherited by the offspring at a corresponding not early period. But the case of the short-faced tumbler, which when twelve hours old had acquired its proper proportions, proves that this is not the universal rule; for here characteristic differences must either have appeared at an earlier period than usual, or, if not so, the differences must have been inherited, not at the corresponding, but at an earlier age.

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Now let us apply these facts and the above two principles-which latter, though not proved true, can be shown to be in some degree probable-to species in a state of nature. Let us take a genus of birds, descended on my theory from some one parent-species, and of which the several new species have become modified through natural selection in accordance with their diverse habits. Then, from the many slight successive steps of variation having supervened at a rather late age, and having been inherited at a corresponding age, the young of the new species of our supposed genus will manifestly tend to resemble each other much more closely than do the adults, just as we have seen in the case of pigeons. We may extend this view to whole families or even classes. The fore-limbs, for instance, which served as legs in the parent-species, may have become, by a long course of modification, adapted in one descendant to act as hands, in another as paddles, in another as wings; and on the above two principles-namely of each successive modification supervening at a rather late age, and being inherited at a corresponding late age—the fore-limbs in the embryos of the several descendants of the parent species will still resemble each other closely, for they I will not have been modified. But in each of our new species, the embryonic fore-limbs will differ greatly from

the fore-limbs in the mature animal; the limbs in the latter having undergone much modification at a rather late period of life, and having thus been converted into hands, or paddles, or wings. Whatever influence longcontinued exercise or use on the one hand, and disuse on the other, may have in modifying an organ, such influence will mainly affect the mature animal, which has come to its full powers of activity and has to gain its own living; and the effects thus produced will be inherited at a corresponding mature age. Whereas the young will remain unmodified, or be modified in a lesser degree, by the effects of use and disuse.

In certain cases the successive steps of variation might supervene, from causes of which we are wholly ignorant, at a very early period of life, or each step might be inherited at an earlier period than that at which it first appeared. In either case (as with the short-faced tumbler) the young or embryo would closely resemble the mature parent-form. We have seen that this is the rule of development in certain whole groups of animals, as with cuttle-fish and spiders, and with a few members of the great class of insects, as with Aphis. With respect to the final cause of the young in these cases not undergoing any metamorphosis, or closely resembling their parents from their earliest age, we can see that this would result from the two following contingencies: firstly, from the young, during a course of modification carried on for many generations, having to provide for their own wants at a very early stage of development, and secondly, from their following exactly the same habits of life with their parents; for in this case, it would be indispensable for the existence of the species, that the child should be modified at a very early age in the same manner with its parents, in accordance with their similar habits. Some further explanation, however, of the embryo not undergoing any metamorphosis is perhaps requisite. If, on the other hand, it profited the young to follow habits of life in any degree different from those of their parent, and consequently to be constructed in a slightly different manner, then, on the principle of inheritance at corresponding ages, the active young or larvæ might be rendered by natural selection different to any conceivable extent from their parents. Such differences might, also, become correlated with successive stages of development; so that the larvæ in the first