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The Factors Of Organic Evolution
by
As with plants so with animals, we find good reason for inferring that while the specialities of the tegumentary parts must be ascribed to the natural selection of favourable variations, their most general traits are due to the direct action of surrounding agencies. Here we come upon the border of those changes which are ascribable to use and disuse. But from this class of changes we may fitly exclude those in which the parts concerned are wholly or mainly passive. A corn and a blister will conveniently serve to illustrate the way in which certain outer actions initiate in the superficial tissues, effects of very marked kinds, which are related neither to the needs of the organism nor to its normal structure. They are neither adaptive changes nor changes towards completion of the type. After noting them we may pass to allied, but still more instructive, changes. Continuous pressure on any portion of the surface causes absorption, while intermittent pressure causes growth: the one impeding circulation and the passage of plasma from the capillaries into the tissues, and the other aiding both. There are yet further mechanically-produced effects. That the general character of the ribbed skin on the under surfaces of the feet and insides of the hands is directly due to friction and intermittent pressure, we have the proofs:–first, that the tracts most exposed to rough usage are the most ribbed; second, that the insides of hands subject to unusual amounts of rough usage, as those of sailors, are strongly ribbed all over; and third, that in hands which are very little used, the parts commonly ribbed become quite smooth. These several kinds of evidence, however, full of meaning as they are, I give simply to prepare the way for evidence of a much more conclusive kind.
Where a wide ulcer has eaten away the deep-seated layer out of which the epidermis grows, or where this layer has been destroyed by an extensive burn, the process of healing is very significant. From the subjacent tissues, which in the normal order have no concern with outward growth, there is produced a new skin, or rather a pro-skin; for this substituted outward-growing layer contains no hair-follicles or other specialities of the original one. Nevertheless, it is like the original one in so far that it is a continually renewed protective covering. Doubtless it may be contended that this make-shift skin results from the inherited proclivity of the type–the tendency to complete afresh the structure of the species when injured. We cannot, however, ignore the immediate influence of the medium, on recalling the facts above named, or on remembering the further fact that an inflamed surface of skin, when not sheltered from the air, will throw out a film of coagulable lymph. But that the direct action of the medium is a chief factor we are clearly shown by another case. Accident or disease occasionally causes permanent eversion, or protrusion, of mucous membrane. After a period of irritability, great at first but decreasing as the change advances, this membrane assumes the general character of ordinary skin. Nor is this all: its microscopic structure changes. Where it is a mucous membrane of the kind covered by cylinder-epithelium, the cylinders gradually shorten, becoming finally flat, and there results a squamous epithelium: there is a near approach in minute composition to epidermis. Here a tendency towards completion of the type cannot be alleged; for there is, contrariwise, divergence from the type. The effect of the medium is so great that, in a short time, it overcomes the inherited proclivity and produces a structure of opposite kind to the normal one.
With but little break we come here upon a significant analogy, parallel to an analogy already described. As was pointed out, an inorganic body that is modifiable by its medium, acquires, after a time, an outer coat which has already undergone such change as surrounding agencies can effect; has a contained mass which is as yet unchanged, because unreached; and has a surface between the two where change is going on–a region of activity. And we saw that alike in the vegetal cell and the animal cell there exist analogous distributions: of course with the difference that the innermost part is not inert. Now we have to note that in those aggregates of cells constituting the Metaphyta and Metazoa, analogous distributions also exist. In plants they are of course not to be looked for in leaves and other deciduous portions, but only in portions of long duration–stems and branches. Naturally, too, we need not expect them in plants having modes of growth which early produce an outer practically dead part, that effectually shields the inner actively living part of the stem from the influence of the medium–long-lived acrogens such as tree-ferns and long-lived endogens such as palms. But in the highest plants, exogens, which have the actively living part of their stems within reach of environing agencies, we find this part,–the cambium layer,–is one from which there is a growth inwards forming wood, and a growth outwards forming bark: there is an increasingly thick covering (where it does not scale off) of tissue changed by the medium, and inside this a film of highest vitality. In so far as concerns the present argument, it is the same with the Metazoa, or at least all of them which have developed organizations. The outer skin grows up from a limiting plane, or layer, a little distance below the surface–a place of predominant vital activity. Here perpetually arise new cells, which, as they develop, are thrust outwards and form the epidermis: flattening and drying up as they approach the surface, whence, having for a time served to shield the parts below, they finally scale off and leave younger ones to take their places. This still undifferentiated tissue forming the base of the epidermis, and existing also as a source of renewal in internal organs, is the essentially living substance; and facts above given imply that it was the action of the medium on this essentially living substance, which, during early stages in the organization of the Metazoa, initiated that protective envelope which presently became an inherited structure–a structure which, though now mainly inherited, still continues to be modifiable by its initiator.