Much more important than these theories is that of Wilhelm Roux[132] of "the struggle of parts within the organism, a supplementation of the theory of mechanical adaptation." He explains the functional autoformation of the purposive structure by a combination of Darwin's principle of selection with Lamarck's idea of transformative heredity, and applies the two in conjunction to the facts of histology. He lays stress on the significance of functional adaptation, which I had described in 1866, under the head of cumulative adaptation, as the most important factor in evolution. Pointing out its influence in the cell-life of the tissues, he puts "cellular selection" above "personal selection," and shows how the finest conceivable adaptations in the structure of the tissue may be brought about quite mechanically, without preconceived plan. This "mechanical teleology" is a valuable extension of Darwin's monistic principle of selection to the whole field of cellular physiology and histology, and is wholly destructive of dualistic vitalism.
The most important advance that evolution has made since Darwin and the most valuable amplification of his theory of selection is, in my opinion, the work of Richard Semon: Die Mneme als erhaltendes Prinzip im Wechsel des organischen Geschehens.[133] He offers a psychological explanation of the facts of heredity by reducing them to a process of (unconscious) memory. The physiologist Ewald Hering had shown in 1870 that memory must be regarded as a general function of organic matter, and that we are quite unable to explain the chief vital phenomena, especially those of reproduction and inheritance, unless we admit this unconscious memory. In my essay Die Perigenesis der Plastidule[134] I elaborated this far-reaching idea, and applied the physical principle of transmitted motion to the plastidules, or active molecules of plasm. I concluded that "heredity is the memory of the plastidules, and variability their power of comprehension." This "provisional attempt to give a mechanical explanation of the elementary processes of evolution" I afterwards extended by showing that sensitiveness is (as Carl Nägeli, Ernst Mach, and Albrecht Rau express it) a general quality of matter. This form of panpsychism finds its simplest expression in the "trinity of substance."
To the two fundamental attributes that Spinoza ascribed to substance—Extension (matter as occupying space) and Cogitation (energy, force)—we now add the third fundamental quality of Psychoma (sensitiveness, soul). I further elaborated this trinitarian conception of substance in the nineteenth chapter of my Die Lebenswunder (1904),[135] and it seems to me well calculated to afford a monistic solution of many of the antitheses of philosophy.
This important Mneme-theory of Semon and the luminous physiological experiments and observations associated with it not only throw considerable light on transformative inheritance, but provide a sound physiological foundation for the biogenetic law. I had endeavoured to show in 1874, in the first chapter of my Anthropogenie,[136] that this fundamental law of organic evolution holds good generally, and that there is everywhere a direct causal connection between ontogeny and phylogeny. "Phylogenesis is the mechanical cause of ontogenesis;" in other words, "The evolution of the stem or race is—in accordance with the laws of heredity and adaptation—the real cause of all the changes that appear, in a condensed form, in the development of the individual organism from the ovum, in either the embryo or the larva."
It is now fifty years since Charles Darwin pointed out, in the thirteenth chapter of his epoch-making Origin of Species, the fundamental importance of embryology in connection with his theory of descent:
"The leading facts in embryology, which are second to none in importance, are explained on the principle of variations in the many descendants from some one ancient progenitor, having appeared at a not very early period of life, and having been inherited at a corresponding period."[137]
He then shows that the striking resemblance of the embryos and larvae of closely related animals, which in the mature stage belong to widely different species and genera, can only be explained by their descent from a common progenitor. Fritz Müller made a closer study of these important phenomena in the instructive instance of the Crustacean larva, as given in his able work Für Darwin[138] (1864). I then, in 1872, extended the range so as to include all animals (with the exception of the unicellular Protozoa) and showed, by means of the theory of the Gastraea, that all multicellular, tissue-forming animals—all the Metazoa—develop in essentially the same way from the primary germ-layers.
I conceived the embryonic form, in which the whole structure consists of only two layers of cells, and is known as the gastrula, to be the ontogenetic recapitulation, maintained by tenacious heredity, of a primitive common progenitor of all the Metazoa, the Gastraea. At a later date (1895) Monticelli discovered that this conjectural ancestral form is still preserved in certain primitive Coelenterata—Pemmatodiscus, Kunstleria, and the nearly-related Orthonectida.
The general application of the biogenetic law to all classes of animals and plants has been proved in my Systematische Phylogenie.[139] It has, however, been frequently challenged, both by botanists and zoologists, chiefly owing to the fact that many have failed to distinguish its two essential elements, palingenesis and cenogenesis. As early as 1874 I had emphasised, in the first chapter of my Evolution of Man, the importance of discriminating carefully between these two sets of phenomena: