In the biological aspect of the case, we must always remember that our bone is not only a living, but a highly plastic structure; the little trabeculae are constantly being formed and deformed, demolished and formed anew. Here, for once, it is safe to say that ãheredityã need not and cannot be invoked to account for the configuration and arrangement of the trabeculae: for we can see them, at any time of life, in the making, under the direct action and control of the forces to which the system is exposed. If a bone be broken and so repaired that its parts lie somewhat out of their former place, so that the pressure-and tension-lines have now a new distribution, before many weeks are over the trabecular system will be found to have been entirely remodelled, so as to fall into line with the new system of forces. And as Wolff pointed out, this process of reconstruction extends a long way off from the seat of injury, and so cannot be looked upon as a mere accident of the physiological process of healing and repair; for instance, it may happen that, after a fracture of the shaft of a long bone, the trabecular meshwork is wholly altered and reconstructed within the distant extremities of the bone. Moreover, in cases of transplantation of bone, for example when a diseased metacarpal is repaired by means of a portion taken from the lower end of the ulna, with astonishing quickness the plastic capabilities of the bony tissue are so manifested that neither in outward form nor inward structure can the old portion be distinguished from the new.
Herein then lies, so far as we can discern it, a great part at least of the physical causation of what at first sight strikes us as a purely functional adaptation: as a phenomenon, in other words, {688} whose physical cause is as obscure as its final cause or end is, apparently, manifest.
Partly associated with the same phenomenon, and partly to be looked upon (meanwhile at least) as a fact apart, is the very important physiological truth that a condition of strain, the result of a stress, is a direct stimulus to growth itself. This indeed is no less than one of the cardinal facts of theoretical biology. The soles of our boots wear thin, but the soles of our feet grow thick, the more we walk upon them: for it would seem that the living cells are ãstimulatedã by pressure, or by what we call ãexercise,ã to increase and multiply. The surgeon knows, when he bandages a broken limb, that his bandage is doing something more than merely keeping the parts together: and that the even, constant pressure which he skilfully applies is a direct encouragement of growth and an active agent in the process of repair. In the classical experiments of Sûˋdillot17, the greater part of the shaft of the tibia was excised in some young puppies, leaving the whole weight of the body to rest upon the fibula. The latter bone is normally about one-fifth or sixth of the diameter of the tibia; but under the new conditions, and under the ãstimulusã of the increased load, it grew till it was as thick or even thicker than the normal bulk of the larger bone. Among plant tissues this phenomenon is very apparent, and in a somewhat remarkable way; for a strain caused by a constant or increasing weight (such as that in the stalk of a pear while the pear is growing and ripening) produces a very marked increase of strength without any necessary increase of bulk, but rather by some histological, or molecular, alteration of the tissues. Hegler, and also Pfeffer, have investigated this subject, by loading the young shoot of a plant nearly to its breaking point, and then redetermining the breaking-strength after a few days. Some young shoots of the sunflower were found to break with a strain of 160 gms.; but when loaded with 150 gms., and retested after two days, they were able to support 250 gms.; and being again loaded with something short of this, by next day they sustained 300 gms., and a few days later even 400 gms. {689}
Such experiments have been amply confirmed, but so far as I am aware, we do not know much more about the matter: we do not know, for instance, how far the change is accompanied by increase in number of the bast-fibres, through transformation of other tissues; or how far it is due to increase in size of these fibres; or whether it be not simply due to strengthening of the original fibres by some molecular change. But I should