1. In some cases of sexless multiplication the germ is a cell-aggregate--if we call germ only that which is already detached from the parent organism.

In all cases the process of evolution consists in a succession of changes of the form, structure, and functions of the germ, by which it passes, step by step, from an extreme simplicity, or relative homogeneity, of visible structure, to a greater or less degree of complexity or heterogeneity; and the course of progressive differentiation is usually accompanied by growth, which is effected by intussusception. This intussusception, however, is a very different process from that imagined either by Buffon or by Bonnet. The substance by the addition of which the germ is enlarged is in no case simply absorbed, ready-made, from the not-living world and packed between the elementary constituents of the germ, as Bonnet imagined; still less does it consist of the "molecules organiques" of Buffon. The new material is, in great measure, not only absorbed but assimilated, so that it becomes part and parcel of the molecular structure of the living body into which it enters. And, so far from the fully developed organism being simply the germ plus the nutriment which it has absorbed, it is probable that the adult contains neither in form, nor in substance, more than an inappreciable fraction of the constituents of the germ, and that it is almost, if not wholly, made up of assimilated and metamorphosed nutriment. In the great majority of cases, at any rate, the full-grown organism becomes what it is by the absorption of not-living matter, and its conversion into living matter of a specific type. As Harvey says (Ex. 45), all parts of the body are nourished "ab eodem succo alibili, aliter aliterque cambiato," "ut plantæ omnes ex eodem communi nutrimento (sive rore seu terræ humore)."

In all animals and plants above the lowest the germ is a nucleated cell, using that term in its broadest sense; and the first step in the process of the evolution of the individual is the division of this cell into two or more portions. The process of division is repeated, until the organism, from being unicellular, becomes multicellular. The single cell becomes a cell-aggregate; and it is to the growth and metamorphosis of the cells of the cell-aggregate thus produced, that all the organs and tissues of the adult owe their origin.

In certain animals belonging to every one of the chief groups into which the Metazoa are divisible, the cells of the cell-aggregate which results from the process of yelk-division, and which is termed a morula, diverge from one another in such a manner as to give rise to a central space, around which they dispose themselves as a coat or envelope; and thus the morula becomes a vesicle filled with fluid, the planula. The wall of the planula is next pushed in on one side, or invaginated, whereby it is converted into a double-walled sac with an opening, the blastopore, which leads into the cavity lined by the inner wall. This cavity is the primitive alimentary cavity or archenteron; the inner or invaginated layer is the hypoblast; the outer the epiblast; and the embryo, in this stage, is termed a gastrula. In all the higher animals a layer of cells makes its appearance between the hypoblast and the epiblast, and is termed the mesoblast. In the further course of development the epiblast becomes the ectoderm or epidermic layer of the body; the hypoblast becomes the epithelium of the middle portion of the alimentary canal; and the mesoblast gives rise to all the other tissues, except the central nervous system, which originates from an ingrowth of the epiblast.

With more or less modification in detail, the embryo has been observed to pass through these successive evolutional stages in sundry Sponges, Coelenterates, Worms, Echinoderms, Tunicates, Arthropods, Mollusks, and Vertebrates; and there are valid reasons for the belief that all animals of higher organisation than the Protozoa, agree in the general character of the early stages of their individual evolution. Each, starting from the condition of a simple nucleated cell, becomes a cell-aggregate; and this passes through a condition which represents the gastrula stage, before taking on the features distinctive of the group to which it belongs. Stated in this form, the "gastræa theory" of Haeckel appears to the present writer to be one of most important and best founded of recent generalisations. So far as individual plants and animals are concerned, therefore, evolution is not a speculation but a fact; and it takes place by epigenesis.

"Animal...per epigenesin procreatur, materiam simul attrahit, parat, concoquit, et eâdem utitur; formatur simul et augetur ... primum futuri corporis concrementum ... prout augetur, dividitur sensim et distinguitur in partes, non simul omnes, sed alias post alias natas, et ordine quasque suo emergentes." [*] In these words, by the divination of genius, Harvey, in the seventeenth century, summed up the outcome of the work of all those who, with appliances he could not dream of, are continuing his labours in the nineteenth century.

  1. Harvey, Exercitationes de Generatione. Ex. 45, "Quænam sit pulli materia et quomodo fiat in Ovo."

Nevertheless, though the doctrine of epigenesis, as understood by Harvey, has definitively triumphed over the doctrine of evolution, as understood by his opponents of the eighteenth century, it is not impossible that, when the analysis of the process of development is carried still further, and the origin of the molecular components of the physically gross, though sensibly minute, bodies which we term germs is traced, the theory of development will approach more nearly to metamorphosis than to epigenesis. Harvey thought that impregnation influenced the female organism as a contagion; and that the blood, which he conceived to be the first rudiment of the germ, arose in the clear fluid of the "colliquamentum" of the ovum by a process of concrescence, as a sort of living precipitate. We now know, on the contrary, that the female germ or ovum, in all the higher animals and plants, is a body which possesses the structure of a nucleated cell; that impregnation consists in the fusion of the substance [*] of another more or less modified nucleated cell, the male germ, with the ovum; and that the structural components of the body of the embryo are all derived, by a process of division, from the coalesced male and female germs. Hence it is conceivable, and indeed probable, that every part of the adult contains molecules, derived both from the male and from the female parent; and that, regarded as a mass of molecules, the entire organism may he compared to a web of which the warp is derived from the female and the woof from the male. And each of these may constitute one individuality, in the same sense as the whole organism is one individual, although the matter of the organism has been constantly changing. The primitive male and female molecules may play the part of Buffon's "moules organiques," and mould the assimilated nutriment, each according to its own type, into innumerable new molecules. From this point of view the process, which, in its superficial aspect, is epigenesis, appears in essence, to be evolution, in the modified sense adopted in Bonnet's later writings; and development is merely the expansion of a potential organism or "original preformation" according to fixed laws.

  1. [At any rate of the nuclei of the two germ-cells. 1893]

II. The Evolution of the Sum of Living Beings.