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adds, “That certain correspondences should occur between the embryonic states of some animals and the adult states of others seems inevitable and of no significance(?). They could not fail, since the embryos lie within the animal sphere, and the variations of which the animal body is capable are determined for each type by the internal connection and mutual reaction of its organs, so that particular repetitions are inevitable.” A profound remark, to which I shall hereafter have occasion to return, but its bearing on the present question is inconclusive. The fact that the embryonic stages of the higher animals resemble in general characters the permanent stages of the lower animals, and very closely resemble the embryonic stages of those animals, is all that the Development Hypothesis requires. Nor is its value lessened by the fact that many of the details and intermediate stages seem passed over in the development of the higher forms, for the recapitulation can only be of outlines, not of details; since there are differences in the forms, there must be differences in their histories.

107. In the preceding observations the object has simply been to show that the phenomena to be explained can be rationally conceived as resulting from gradual Evolution, whereas they cannot be so rationally interpreted on any other hypothesis. And here it may be needful to say a word respecting Epigenesis.

The Preformation hypothesis, which regarded every organism as a simple educt and not the product of a germ, was called by its advocates an evolution hypothesis—meaning that the adult form was an outgrowth of the germ, the miniature magnified. Wolff, who replaced that conception by a truer one, called his, by contrast, Epigenesis, meaning that there was not simply out-growth but new growth. “The various parts,” he says, “arise one after the other, so that always one is secreted from (excernirt), or deposited (deponirt) on the other; and then it is either a free and independent part, or is only fixed to that which gave it existence, or else is contained within it. So that every part is the effect of a pre-existing part, and in turn the cause of a succeeding part.”51 The last sentence expresses the conception of Epigenesis which embryologists now adopt; and having said this, we may admit that Wolff, in combating the error of preformation, replacing it with the truer notion of gradual and successive formation, was occasionally open to the criticism made by Von Baer, that he missed the true sense of Evolution, since the new parts are not added on to the old parts as new formations, but evolved from them as transformations. “The word Evolution, therefore, seems to me more descriptive of the process than Epigenesis. It is true that the organism is not preformed, but the course of its development is precisely the course which its parents formerly passed through. Thus it is the Invisible—the course of development—which is predetermined.”52 When the word Epigenesis is used, therefore, the reader will understand it to signify that necessary succession which determines the existence of new forms. Just as the formation of chalk is not the indifferent product of any combination of its elements, carbon, oxygen, and calcium, but is the product of only one series of combinations, an evolution through necessary successions, the carbon uniting with oxygen to form carbonic acid, and this combining with the oxide of calcium to form chalk, so likewise the formation of a muscle, a bone, a limb, or a joint has its successive stages, each of which is necessary, none of which can be transposed. The formation of bone is peculiarly instructive, because the large proportion of inorganic matter in its substance, and seemingly deposited in the organic tissue, would lead one to suppose that it was almost an accidental formation, which might take place anywhere; yet although what is called connective tissue will ossify under certain conditions, true bone is the product of a very peculiar modification, which almost always needs to be preceded by cartilage. That the formation of bone has its special history may be seen in the fact that it is the last to appear in the animal series, many highly organized fishes being without it, and all the other systems appearing before it in the development of the embryo. Thus although the mother’s blood furnishes all the requisite material, the fœtus is incapable of assimilating this material and of forming bone, until its own development has reached a certain stage. Moreover, when ossification does begin, it generally begins in the skull (in man in the clavicle); and the only approach to an internal skeleton in the Invertebrates is the so-called skull of the Cephalopoda. Not only is bone a late development, but cartilage is also; and although it is an error to maintain that the Invertebrates are wholly destitute of cartilage, its occasional presence having been fully proved by Claparède and Gegenbaur, the rarity of its presence is very significant. The animals which can form shells of chalk and chitine are yet incapable of forming even an approach to bone.

108. Epigenesis depends on the laws of succession, which may be likened to the laws of crystallization, if we bear in mind the essential differences between a crystal and an organism, the latter retaining its individuality through an incessant molecular change, the former only by the exclusion of all change. When a crystalline solution takes shape, it will always take a definite shape, which represents what may be called the direction of its forces, the polarity of its constituent molecules. In like manner, when an organic plasmode takes shape—crystallizes, so to speak—it always assumes a specific shape dependent on the polarity of its molecules. Crystallographers have determined the several forms possible to crystals; histologists have recorded the several forms of Organites, Tissues, and Organs. Owing to the greater variety in elementary composition, there is in organic substance a more various polar distribution than in crystals; nevertheless, there are sharply defined limits never overstepped, and these constitute what may be called the specific forms of Organites, Tissues, Organs, Organisms. An epithelial cell, for example, may be ciliated or columnar, a muscle-fibre striated or non-striated, a nerve-fibre naked or enveloped in a sheath, but the kind is always sharply defined. An intestinal tube may be a uniform canal, or a canal differentiated into several unlike compartments, with several unlike glandular appendages. A spinal column may be a uniform solid axis, or a highly diversified segmented axis. A limb may be an arm, or a leg, a wing, or a paddle. In every case the anatomist recognizes a specific type. He assigns the uniformities to the uniformity of the substance thus variously shaped, under a history which has been similar; the diversities he assigns to the various conditions under which the processes of growth have been determined. He never expects a muscular tissue to develop into a skeleton, a nervous tissue into a gland, an osseous tissue into a sensory organ. He never expects a tail to become a hand or a foot, though he sees it in monkeys and marsupials serving the offices of prehension and locomotion. He never expects to find fingers growing anywhere except from metacarpal bones, or an arm developed from a skull. The well-known generalization of Geoffroy St. Hilaire that an organ is more easily annihilated than transposed, points to the fundamental law of Epigenesis. In the same direction point all the facts of growth. Out of a formless germinal membrane we see an immense variety of forms evolved; and out of a common nutritive fluid this variety of organs is sustained, repaired, replaced; and this not indifferently, not casually, but according to rigorous laws of succession; that which precedes determining that which succeeds as inevitably as youth precedes maturity, and maturity decay. The nourishment of various organs from plasmodes derived from a common fluid, each selecting from that fluid only those molecules that are like its own, rejecting all the rest, is very similar to the formation of various crystals in a solution of different salts, each salt separating from the solution only those molecules that are like itself. Reil long ago called attention to this analogy. He observed that if in a solution of nitre and sulphate of soda a crystal of nitre be dropped, all the dissolved nitre crystallizes, the sulphate remaining in solution; whereas on reversing the experiment, a crystal of sulphate of soda is found to crystallize all the dissolved sulphate, leaving the nitre undisturbed. In like manner muscle selects from the blood its own materials which are there in solution, rejecting those which the nerve will select.

109. Nay, so definite is the course of growth, that when a limb or part of a limb is cut off from a crab or salamander, a new limb or new part is reproduced in the old spot, exactly like the one removed. Bonnet startled the world by the announcement that the Naïs, a worm common in ponds, spontaneously divided itself into two worms; and that when he cut it into several pieces, each piece reproduced head and tail, and grew into a perfect worm. This had been accepted by all naturalists without demur, until Dr. Williams, in his “Report on British Annelida, 1851,” declared it to be a fable. In 1858, under the impulse of Dr. Williams’s very emphatic denial, I repeated experiments similar to those of Bonnet, with similar results. I cut two worms in half, and threw away the head-bearing segments, placing the others in two separate vessels, with nothing but water and a little mud, which was first carefully inspected to see that no worm lay concealed therein. In a few days the heads were completely reformed, and I had the pleasure of watching them during their reconstruction. When the worms were quite perfect, I again cut away their heads, and again saw these reformed. This was repeated, till I had seen four heads reproduced; after which the worms succumbed.

110. The question naturally arises, Why does the nutritive fluid furnish only material which is formed into a part like the old one, instead of reproducing another part, or one having a somewhat different structure? The answer to this question is the key to the chief problem of organic life. That a limb in situ should replace its molecular waste by molecules derived from the blood, seems intelligible enough (because we are familiar with it), and may be likened to the formation of crystals in a solution; but how is it that the limb which is not in existence can assimilate materials from the blood? How is it that the blood, which elsewhere in the organism will form other parts, here will only form this particular part? There is, probably, no one who has turned his attention to these subjects who has not paused to consider this mystery. The most accredited answer at present before the world is one so metaphysiological that I should pass it by, were it not intimately allied with that conception of Species, which it is the object of these pages to root out. It is this:

111. The organism is determined by its Type, or, as the Germans say, its Idea. All its parts take shape according to this ruling plan; consequently, when any part is removed, it is reproduced according to the Idea of the whole of which it forms a part. Milne Edwards, in a very interesting and suggestive work, concludes his survey of organic phenomena in these words: “Dans l’organisme tout semble calculé en vue d’un résultat déterminé, et l’harmonie des parties ne résulte pas de l’influence qu’elles peuvent exercer les unes sur les autres, mais de leur co-ordination sous l’empire d’une puissance commune, d’un plan préconçu, d’une force pré-existante.”53 This is eminently metaphysiological. It

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