105. Humanly appreciated, not only is it difficult to justify the successive stages of development, the incessant building up of structures immediately to be taken down, but also to explain why development was necessary at all. Why are not plants and animals formed at once, as Eve was mythically affirmed to be taken from Adam’s rib, and Minerva from Jupiter’s head? The theory of Evolution answers this question very simply; the theory of Creation can only answer it by affirming that such was the ordained plan. But the theory of Evolution not only gives the simpler and more intelligible answer to this question, it gives an answer to the further question which leaves the theory of Creation no loophole except a sophism—namely, why the formation of organisms is constantly being frustrated or perverted? And, further, it gives an explanation of the law noticed by Milne Edwards, that Nature is as economical in her means as she is prodigal in her variation of them: “On dirait qu’avant de recourir à des ressources nouvelles elle a voulu épuiser, en quelque sorte, chacun des procédés qu’elle avait mis en jeu.”[49] The applause bestowed on Nature for being economical is a curious transference to Nature of human necessities. Why, with a whole universe at her disposal, should Nature be economical? Why must she always be working in the same groove, and using but a few out of the many substances at her command? Economy is a virtue only in the poor. If Nature, in organic evolutions, is restricted to a very few substances, and a very few modes of combination, always creating new forms by modification of the old, and apparently incapable of creating an organism at once, this must imply an inherent necessity which is very unlike the free choice that can render economy a merit.
106. There may indeed be raised an objection to the Development Hypothesis on the ground that if the complex forms were all developed from the simpler forms, we ought to trace the identities through all their stages. If the fish developed into the reptile, the reptile into the bird, and the bird into the mammal (which I, for one, think questionable), we ought to find, it is urged, evidence of this passage. And at one time it was asserted that the evidence existed; but this has been disproved, and on the disproof the opponents of Evolution take their stand. Although I cannot feel much confidence in the idea of such a passage from Type to Type, and although the passage, if ever it occurred, must have occurred at so remote a period as to leave no evidence more positive than inference, I cannot but think the teaching of Embryology far more favorable to it than to our opponents. Supposing, for the sake of argument, that the passage did take place, ought we to find the embryonic stages accurately reproducing the permanent forms of lower types? Von Baer thinks we ought; and lesser men may follow him without reproach. But it seems to me that he starts from an inadmissible assumption, namely, that the development must necessarily be in a straight line rather than in a multiplicity of divergent lines. “When we find the embryonic condition,” he says, “differing from the adult, we ought to find a corresponding condition somewhere in the lower animals.”[50] Not necessarily. We know that the mental development of a civilized man passes through the stages which the race passed through in the course of its long history, and the psychology of the child reproduces the psychology of the savage. But as this development takes place under conditions in many respects different, and as certain phases are hurried over, we do not expect to find a complete parallel. It is enough if we can trace general resemblances. Von Baer 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 refuses to acknowledge the operation of immanent properties, refuses to admit that the harmony of a complex structure results from the mutual relations of its parts, and seeks outside the organism for some mysterious force, some plan, not otherwise specified, which regulates and shapes the parts. Von Baer, in his great work, has a section entitled, “The nature of the animal determines its development”; and he thus explains himself: “Although every stage in development is only made possible by the pre-existing condition [which is another mode of expressing Epigenesis], nevertheless the entire development is ruled and guided by the Nature of the animal which is about to be (von der gesammten Wesenheit des Thieres welches werden soll), and it is not the momentary condition which alone and absolutely determines the future, but more general and higher relations.”[54] One must always be slow in rejecting the thoughts of a master, and feel sure that one sees the source of the error before regarding it as an error; but in the present case I think the positive biologist will be at no loss to assign Von Baer’s error to its metaphysical origin. Without pausing here to accumulate examples both of anomalies and slighter deviations which are demonstrably due to the “momentary conditions” that preceded them, let us simply note the logical inconsistency of a position which, while assuming that every separate stage in development is the necessary sequence of its predecessor, declares the whole of the stages independent of such relations! Such a position is indeed reconcilable on the assumption that animal forms are moulded “like clay in the hands of the potter.” But this is a theological dogma, which leads to very preposterous and impious conclusions; and whether it leads to these conclusions or to others, positive Biology declines theological explanations altogether. Von Baer, although he held the doctrine of Epigenesis, coupled it, as many others have done, with metaphysical doctrines to which it is radically opposed. He believed in Types as realities; he was therefore consistent in saying, “It is not the Matter and its arrangements which determine the product, but the nature of the parent form—the Idea, according to the new school.” How are we to understand this Idea? If it mean an independent Entity, an agency external to the organism, we refuse to acknowledge its existence. If it mean only an a posteriori abstraction expressing the totality of the conditions, then, indeed, we acknowledge that it determines the animal form; but this is only an abbreviated way of expressing the law of Evolution, by which each stage determines its successor. The Type does not dominate the conditions, it emerges from them; the animal organism is not cast in a mould, but the imaginary mould is the form which the polarities of the organic substance assume. It would seem very absurd to suppose that crystals assumed their definite shapes (when the liquid which held their molecules in solution is evaporated) under the determining impulse of phantom-crystals, or Ideas; yet it has not been thought absurd to assume phantom forms of organisms.
112. The conception of Type as a determining influence arises from that fallacy of taking a resultant for a principle, which has played so conspicuous a part in the history of philosophy. Like many others of its class it exhibits an interesting evolution from the crude metaphysical to the subtle metaphysical point of view, which at last insensibly blends into the positive point of view. At first the Type or Idea was regarded as an objective reality, external to the organism it was supposed to rule. Then this notion was replaced by an approach to the more rational interpretation, the idea was made an internal not an external force, and was incorporated with the material elements of the organism, which were said to “endeavor” to arrange themselves according to the Type. Thus Treviranus declares that the seed “dreams of the future flower”; and “Henle, when he affirms that hair and nails grow in virtue of the Idea, is forced to add that the parts endeavor to arrange themselves according to this Idea.”[55] Even Lotze, who has argued so victoriously against the vitalists, and has made it clear that an organism is a vital mechanism, cannot relinquish this conception of legislative Ideas, though he significantly adds, “these have no power in themselves, but only in as far as they are grounded in mechanical conditions.” Why then superfluously add them to the conditions? If every part of a watch, in virtue of the properties inherent in its substance, and of the mutual reactions of these and other parts, has a mechanical value, and if the sum of all these parts is the time-indicating mechanism, do we add to our knowledge of the watch, and our means of repairing or improving it, by assuming that the parts have over and above their physical properties the metaphysical “tendency” or “desire” to arrange themselves into this specific form? When we see that an organism is constructed of various parts, each of which has its own properties inalienable from its structure, and its uses dependent on its relation to other parts, do we gain any larger insight by crediting these parts with desires or “dreams” of a future result which their union will effect? That which is true in this conception of legislative Ideas is that when the parts come together there is mutual reaction, and the resultant of the whole is something very unlike the mere addition of the items, just as water is very unlike oxygen or hydrogen; further, the connexus of the whole impresses a peculiar direction on the development of the parts, and the law of Epigenesis necessitates a serial development, which may easily be interpreted as due to a preordained plan.
113. In a word, this conception of Type only adds a new name to the old difficulty, adding mist to darkness. The law of Epigenesis, which is simply the expression of the material process determined by the polarity of molecules, explains as much of the phenomena as is explicable. A lost limb is replaced by the very processes, and through the same progressive stages as those which originally produced it. We have a demonstration of its not being reformed according to any Idea or Type which exists apart from the immanent properties of the organic molecules, in the fact that it is not reformed at once, but by gradual evolution; the mass of cells at the stump are cells of embryonic character, cells such as those which originally “crystallized” into muscles, nerves, vessels, and integument, and each cell passes through all its ordinary stages of development. It is to be remembered that so intimately dependent is the result on the determining conditions, that any external influence which disturbs the normal course of development will either produce an anomaly, or frustrate the formation of a new limb altogether. One of my tritons bit off the leg of his female;[56] the leg which replaced it was much malformed, and curled over the back so as to be useless; was this according to the Idea? I cut it off, and examined it; all the bones were present, but the humerus was twisted, and of small size. In a few weeks a new leg was developed, and this leg was normal. If the Idea, as a ruling power, determined the growth of this third leg, what determined the second, which was malformed? Are we to suppose that in normal growth the Idea prevails, in abnormal the conditions? That it is the polarity of the molecules which at each moment determines the group those molecules will assume, is well seen in the experiment of Lavalle mentioned by Bronn.[57] He showed that if when an octohedral crystal is forming, an angle be cut away, so as to produce an artificial surface, a similar surface is produced spontaneously on the corresponding angle, whereas all the other angles are sharply defined. “Valentin,” says Mr. Darwin, “injured the caudal extremity of an embryo, and three days afterwards it produced rudiments of a double pelvis, and of double hind limbs. Hunter and others have observed lizards with their tails reproduced and doubled. When Bonnet divided longitudinally the foot of the salamander, several additional digits were occasionally formed.”[58] Where is the evidence of the Idea in these cases?