Thus the first physiology, borrowing its ideas and methods from the first physics, was, like the latter, a mechanical science. After Galileo and Torricelli came Borelli with his purely mechanical conceptions of animal movement, and of the blood circulation, introducing even then mathematics into biology. There was no chemistry in these speculations, though Basil Valentine and Paracelsus and Van Helmont had preceded Descartes and Borelli. This chemistry was mystical, and though chemical reactions had been studied in the organism, they were supposed to be controlled by spiritual agencies, the “archei” of the first bio-chemists. But that notion was to disappear, and with Sylvius the conception of the animal body as a chemical mechanism arose. All that was valuable in Van Helmont’s chemistry was taken up by Sylvius, but in his mind the fermentations of the older chemists were sufficient in themselves without the mystical “sensitive soul” and “archei.” With Sylvius and Mayow physiology became based upon chemical discovery and again became mechanistic, and remained so until the time of Stahl, when chemical discovery attained for the time its greatest development.

The seventeenth century ended with the work of Stahl. It is well known to students of science how the views of this great chemist sterilised chemical investigation almost until the time of Lavoisier. The notion of phlogiston as an active constituent of material bodies entering and leaving them in their reactions with each other was a clear and simple one, and it served as a working hypothesis for the chemists who immediately followed Stahl. It was, of course, a false hypothesis, and retarded discovery to the extent that the greater part of the eighteenth century is a blank for chemistry, when compared with the seventeenth and nineteenth centuries. Deprived therefore of the stimulus afforded by new physico-chemical methods of investigation, physiology ceased to maintain the progress it had made during the previous century, and the only great name of this period is that of von Haller. Comparative anatomy, and zoological exploration, on the other hand, made enormous advances, and for these branches of biology the eighteenth century was the great period. It was the period of the historic vitalistic views—vital principles, and vital and formative forces. Stahl’s teaching dominated physiology just as it did chemistry. Chemical and physical reactions occurred in the living body just as they did in non-living matter, but they were controlled and modified by the soul, or vital principle. It has been said that Stahl’s vitalistic teaching retarded the progress of physiology, but it does not seem clear that this was the case. What did retard physiological discovery was the lack of progress made by chemistry and physics, and this may have been the result of the Stahlian phlogistic hypothesis.

However this may be, it seems clear that it was the discoveries of the great chemists of the close of the eighteenth century that again introduced mechanistic views into physiology. With the discoveries of Lavoisier and his successors the latter science acquired new methods of research and the older working hypotheses were re-introduced. There has been no recession from this position during the nineteenth century. Mechanistic biology culminated in the writings of Huxley and Max Verworn and received a new accession of strength almost in our own day in the modern discoveries of physical chemistry; and when physiology became truly a comparative science, and embraced the lower invertebrates, it became perhaps most mechanistic—witness the writings of Jacques Loeb.

Of far greater philosophical importance than the physico-chemical investigation of the functioning of individual organisms has been the essentially modern experimental study of embryological processes. The former deals essentially with the means of growth, reproduction, and so on. We can no longer doubt that the changes which we can observe taking place in the organism, either the developing embryo or the fully formed animal, are, in the long run, physico-chemical changes; and in ultimate analysis we cannot expect to find anything else than processes of this nature.

But physiological investigation has failed to provide anything more than this analysis. If we watch the development of the egg of an animal into a larval form, and continue to trace the metamorphosis of the larva into the perfect animal, we cannot fail to conclude that, beside the individual physico-chemical reactions which proceed, there is also organisation. The elementary processes must be integrated. There must be a due order and succession in them. In studying developmental processes, in considering the developing organism as a whole, we are impressed above all else with the notion that not only do physico-chemical reactions occur, but that these are marshalled into place, so to speak. When we attempt to make a description of this integration of those ultimate processes which we can describe in terms of physical chemistry, physiology fails us. “At present,” says Morgan, “we cannot see how any known principles of chemistry or of physics can explain the development of a definite form by the organism or by a piece of the organism.” It is true that we can attempt to imagine a physico-chemical mechanism which is the organisation of the developing embryo; but this must be a logically constructed mechanism, not only incapable of experimental verification, but which can also be demonstrated, purely by physical arguments, to be false. This conclusion may, without exaggeration, be said to be that of modern experimental embryology.

There have always been (in modern times) two views as to the nature of the embryological process: (1) that the egg contained the fully formed organism in a kind of rolled-up condition, and that the process of development consisted merely in the unfolding (evolution) of this embryonic organism, and in the increase in volume of its parts. This was the hypothesis of preformation held in the beginning of embryological science. It involved various consequences: the limitation, for instance, of the duration of a species, since each generation of female organisms contained in their ovaries all the future generations; with other consequences which the preformationists did not hesitate to accept. (2) The other view was the later one of epigenesis: the egg was truly homogeneous and the embryo grew from it. Obviously the acceptance of this hypothesis led to vitalism, and we find that it was abandoned just as soon as the embryologists recognised that physics provided a corpuscular theory of matter, when a return was made to the preformation views of earlier times; views which lent themselves to the construction of a mechanistic hypothesis of development.

We may state very briefly the main facts of the development of a typical animal ovum, such as that of the sea-urchin.

Fig. 12.

The fertilised ovum divides into two (2), and then each of these blastomeres divides again in a plane perpendicular to the first division plane (3). The third division plane is at right angles to the first two, and it cuts off a tier of smaller blastomeres from the tops of the first four. There are now (4) two tiers of blastomeres, a lower tier of large blastomeres and an upper tier of smaller ones. This is the 8-cell stage. Next, each of these blastomeres divides in two simultaneously so that the embryo now consists of sixteen cells. After this the divisions proceed with less regularity, but after about ten divisions the embryo consists of about 1000 cells (210), and these are arranged to form a hollow sphere consisting of a single layer of cells. The latter are furnished with cilia, and the whole embryo, now known as the blastula, can swim about by the movements of these cilia. Further development results in another larval form—the gastrula, and yet another, the pluteus larva. After this the transformation into the fully formed sea-urchin occurs.