Are such differentiations among the muscles functionally-produced? or are they produced by the natural selection of variations distinguished as spontaneous? We have, I think, good grounds for concluding that they are functionally-produced. We know that in individual men and animals, the power of sustained action in muscles is rapidly adaptable to the amount of sustained action required. We know that being “out of condition,” is usually less shown by the inability to put out a violent effort than by the inability to continue making violent efforts; and we know that the result of training for prize-fights and races, is more shown in the prolongation of energy than in the intensification of energy. At the same time, experience has taught us that the structural change which accompanies this functional change, is not so much a change in the bulk of the muscles as a change in their internal state: instead of being soft and flabby they become hard. We have inductive proof, then, that exercise of a muscle causes some interstitial growth along with the power of more sustained action; and there can be no doubt that the one is a condition to the other. What is this interstitial growth? There is reason to suspect that it is in part an increased deposit of the sarcous substance and in part a development of blood-vessels. Microscopic observation tends to confirm the conclusions before drawn, that repetition of contractions furthers the formation of the matter which contracts, and that greater draughts of blood determine greater vascularity. And if the contrasts of molecular structure and the contrasts of vascularity, directly caused in muscles by contrasts in their activities, are to any degree inheritable; there results an explanation of those constitutional differences in the colours and textures of muscles, which accompany constitutional differences in their degrees of activity.
It may be added that if we are warranted in so ascribing the differentiations of muscles from one another to direct equilibration, then we have the more reason for thinking that the differentiation of muscles in general from other structures is also due to direct equilibration. That unlikenesses between parts of the contractile tissues having unlike functions, are caused by the unlikenesses of their functions, renders it the more probable that the unlikenesses between contractile tissue and other tissues, have been caused by analogous unlikenesses.
§ 304. These interpretations, which have already occupied too large a space, must here be closed. Of course out of phenomena so multitudinous and varied, it has been impracticable to deal with any but the most important; and it has been practicable to deal with these only in a general way. Much, however, as remains to be explained, I think the possibility of tracing, in so many cases, the actions to which these internal differentiations may rationally be ascribed, makes it likely that the remaining internal differentiations are due to kindred actions. We find evidence that, in more cases than seemed probable, these actions produce their effects directly on the individual; and that the unlikenesses are produced by accumulation of such effects from generation to generation. While for all the other unlikenesses, we have, as an adequate cause, the indirect effects wrought by the survival, generation after generation, of the individuals in which favourable variations have occurred—variations such as those of which human anatomy furnishes endless instances. Thus accounting for so much, we may not unreasonably presume that these co-operative processes of direct and indirect equilibration will account for what remains.
[Note.—After having dismissed this revised chapter as done with, and sent it to the printer, further thought concerning those differentiations which produce bone, has reminded me of a fact of extreme and varied significance named in the first volume. I refer to the formation of adaptive structures round the ends of dislocated bones, and to the formation of “false joints.”
These are ontogenetic changes of which phylogeny yields no explanation. They do not repeat the traits of ancestral organisms, and they cannot be ascribed to either of the recognized evolutionary factors. If a humerus be broken across and, failing to set, presently comes to have its two loose ends so modified as in a measure to simulate the parts of a normal joint—the ends becoming smooth, covered with periosteum and supplied with fibrous tissue, and attached by ligaments in such ways as to allow of restrained movements—it is impossible to think that natural selection has had anything to do with the power of adjustment thus shown. No survival of individuals in which adaptations of this kind, now in one place and now in another, were better and better effected, could account for acquirement of the ability. Nor can it be supposed that the ability might result from a functionally-produced habit; since it is scarcely conceivable that the number of cases in which individuals profited by it (at first a little and gradually more) could be such (even did they survive) as to affect the constitution of the species. Both of the alleged causes of structural modifications are out of court. It is manifest, too, that the foregoing hypothesis respecting bone-formation yields us not the slightest help.
But on carefully considering the facts, certain phenomena of profound meaning may strike us. Here, in a part of the body where no such tissues ordinarily exist and to which no such structures are ordinarily appropriate, there arise tissues and structures adapted to the physical circumstances imposed on that part. Out of what do these abnormal but appropriate tissues arise? The substances around—osseous, cartilaginous, membranous—consist of differentiated elements too far specialized to allow of transformation. These new tissues, then, must originate from the undifferentiated protoplasm pervading the part. The units of this protoplasm, subject to the actions proper to an articulation, begin to assume the appropriate histological traits—are determined by local stimuli to form tissues ordinarily associated with such stimuli. What is the inevitable implication? These units—physiological or constitutional, as we may call them—must have possessed latent potentialities of falling into these special arrangements under stress of such conditions. At one point there arises periosteum and at another ligamentous tissue, while for the shaping of the ends of the bones—here into a rude hinged form and there into a rude ball-and-socket form, according to the habitual movements—there goes on some appropriate deposit of bone. Hence we must conclude that in the units of protoplasm which have not yet been organized into special tissues, there resides the ability to take on one or other type of histological structure according to circumstances; and, further, that there resides in each of them the still more marvellous ability to co-operate with kindred units dispersed around in developing that arrangement of the parts required to constitute a “false joint.” So that while these units have a general proclivity towards the structure of the organism as a whole, they have also proclivities towards structures proper to the local conditions into which they fall. There is latent in each unit the constitution of the entire organism and by implication the constitution of every organ; and each unit while co-operating with the aggregate is ready to take part in that particular arrangement proper to the position it has fallen into. If the reader will refer back to [§§ 97d, 97e], in which it is shown that each member of a human society possesses a combination of potentialities like these, he will be the better enabled to believe that this thing may be so while he is unable to conceive how it is so.
And here, indeed, let it be pointed out how completely irrelevant is the test of conceivableness as applied to these ultimate physiological actions. For as here, from the un-united ends of the broken bone, there presently arises a rude joint with fit membranes, ligaments, and even synovial fluid, though we are absolutely unable to imagine the process by which the adjacent tissues produce this structure; so there may be from an organ enlarged by function, such reactive effect upon the system at large as eventually to influence the reproductive cells, though we may be absolutely unable to imagine how this can be done.]
CHAPTER IX.
PHYSIOLOGICAL INTEGRATION IN ANIMALS.
§ 305. Physiological differentiation and physiological integration, are correlatives that vary together. We have but to recollect the familiar parallel between the division of labour in a society and the physiological division of labour, to see that as fast as the kinds of work performed by the component parts of an organism become more numerous, and as fast as each part becomes more restricted to its own work, so fast must the parts have their actions combined in such ways that no one can go on without the rest and the rest cannot go on without each one.