Part 16

"Si la volonté divine a créé par un seul Acte l'Universalité des êtres, d'où venoient ces plantes et ces animaux dont Moyse nous decrit la Production au troisieme et au cinquieme jour du renouvellement de notre monde?

"Abuserois-je de la liberté de conjectures si je disois, que les Plantes et les Animaux qui existent aujourd'hui sont parvenus par une sorte d'evolution naturelle des Etres organises qui peuplaient ce premier Monde, sorti immédiatement des MAINS du CREATEUR?...

"Ne supposons que trois révolutions. La Terre vient de sortir des MAINS du CREATEUR. Des causes preparées par sa SAGESSE font développer de toutes parts les Germes. Les Etres organisés commencent à jouir de l'existence. Ils étoient probablement alors bien différens de ce qu'ils sont aujourd'hui. Ils l'etoient autant que ce premier Monde différoit de celui que nous habitons. Nous manquons de moyens pour juger de ces dissemblances, et peut-être que le plus habile Naturaliste qui auroit été placé dans ce premier Monde y auroit entièrement méconnu nos Plantes et nos Animaux."]

The two parts of Bonnet's hypothesis, namely, the doctrine that all living things proceed from pre-existing germs, and that these contain, one inclosed within the other, the germs of all future living things, which is the hypothesis of "emboîtement;" and the doctrine that every germ contains in miniature all the organs of the adult, which is the hypothesis of evolution or development, in the primary senses of these words, must be carefully distinguished. In fact, while holding firmly by the former, Bonnet more or less modified the latter in his later writings, and, at length, he admits that a "germ" need not be an actual miniature of the organism; but that it may be merely an "original preformation" capable of producing the latter. [Footnote: "Ce mot (germe) ne désignera pas seulement un corps organisé réduit en petit; il désignera encore toute espèce de préformation originelle dont un Tout organique peut résulter comme de son principe immédiat."--Palingénésie Philosophique, part X. chap. II.]

But, thus defined, the germ is neither more nor less than the "particula genitalis" of Aristotle, or the "primordium vegetale" or "ovum" of Harvey; and the "evolution" of such a germ would not be distinguishable from "epigenesis."

Supported by the great authority of Haller, the doctrine of evolution, or development, prevailed throughout the whole of the eighteenth century, and Cuvier appears to have substantially adopted Bonnet's later views, though probably he would not have gone all lengths in the direction of "emboîtement." In a well-known note to Laurillard's "Éloge," prefixed to the last edition of the "Ossemens fossiles," the "radical de l'être" is much the same thing as Aristotle's "particula genitalis" and Harvey's "ovum." [Footnote: "M. Cuvier considérant que tous les êtres organisés sont dérivés de parens, et ne voyant dans la nature aucune force capable de produire l'organisation, croyait à la pré-existence des germes; non pas à la pré-existence d'un être tout formé, puisqu'il est bien évident que ce n'est que par des développemens successifs que l'être acquiert sa forme; mais, si l'on peut s'exprimer ainsi, à la pré-existence du radical de l'être, radical qui existe avant que la série des évolutions ne commence, et qui remonte certainement, suivant la belle observation de Bonnet, à plusieurs generations."--Laurillard, Éloge de Cuvier, note 12.]

Bonnet's eminent contemporary, Buffon, held nearly the same views with respect to the nature of the germ, and expresses them even more confidently.

"Ceux qui ont cru que le coeur étoit le premier formé, se sont trompés; ceux qui disent que c'est le sang se trompent aussi: tout est formé en même temps. Si l'on ne consulte que l'observation, le poulet se voit dans l'oeuf avant qu'il ait été couvé." [Footnote: Histoire Naturelle, tom. ii. ed. ii. 1750, p. 350.]

"J'ai ouvert une grande quantité d'oeufs à differens temps avant et après l'incubation, et je me suis convaincu par mes yeux que le poulet existe en entier dans le milieu de la cicatricule au moment qu'il sort du corps de la poule." [Footnote: Ibid., p. 351.]

The "moule intérieur" of Buffon is the aggregate of elementary parts which constitute the individual, and is thus the equivalent of Bonnet's germ, [Footnote: See particularly Buffon, l. c. p. 41.] as defined in the passage cited above. But Buffon further imagined that innumerable "molecules organiques" are dispersed throughout the world, and that alimentation consists in the appropriation by the parts of an organism of those molecules which are analogous to them. Growth, therefore, was, on this hypothesis, a process partly of simple evolution, and partly of what has been termed "syngenesis." Buffon's opinion is, in fact, a sort of combination of views, essentially similar to those of Bonnet, with others, somewhat similar to those of the "Medici" whom Harvey condemns. The "molecules organiques" are physical equivalents of Leibnitz's "monads."

It is a striking example of the difficulty of getting people to use their own powers of investigation accurately, that this form of the doctrine of evolution should have held its ground so long; for it was thoroughly and completely exploded, not long after its enunciation, by Casper Friederich Wolff, who in his "Theoria Generationis," published in 1759, placed the opposite theory of epigenesis upon the secure foundation of fact, from which it has never been displaced. But Wolff had no immediate successors. The school of Cuvier was lamentably deficient in embryologists; and it was only in the course of the first thirty years of the present century, that Prévost and Dumas in France, and, later on, Döllinger, Pander, Von Bär, Rathke, and Remak in Germany, founded modern embryology; while, at the same time, they proved the utter incompatibility of the hypothesis of evolution, as formulated by Bonnet and Haller, with easily demonstrable facts.

Nevertheless, though the conceptions originally denoted by "evolution" and "development" were shown to be untenable, the words retained their application to the process by which the embryos of living beings gradually make their appearance; and the terms "Development," "Entwickelung," and "Evolutio," are now indiscriminately used for the series of genetic changes exhibited by living beings, by writers who would emphatically deny that "Development" or "Entwickelung" or "Evolutio," in the sense in which these words were usually employed by Bonnet or by Haller, ever occurs.

Evolution, or development, is, in fact, at present employed in biology as a general name for the history of the steps by which any living being has acquired the morphological and the physiological characters which distinguish it. As civil history may be divided into biography, which is the history of individuals, and universal history, which is the history of the human race, so evolution falls naturally into two categories--the evolution of the individual, and the evolution of the sum of living beings. It will be convenient to deal with the modern doctrine of evolution under these two heads.

I. The Evolution of the Individual.

No exception is at this time, known to the general law, established upon an immense multitude of direct observations, that every living thing is evolved from a particle of matter in which no trace of the distinctive characters of the adult form of that living thing is discernible. This particle is termed a germ. Harvey [Footnote: Execitationes de Generatione. Ex. 62, "Ovum esse primordium commune omnibus animalibus."] says--

"Omnibus viventibus primordium insit, ex quo et a quo proveniant. Liceat hoc nobis primordium vegetale nominare; nempe substantiam quandam corpoream vitam habentem potentiâ; vel quoddam per se existens, quod aptum sit, in vegetativam formam, ab interno principio operante, mutari. Quale nempe primordium, ovum est et plantarum semen; tale etiam viviparorum conceptus, et insectorum vermis ab Aristotele dictus: diversa scilicet diversorum viventium primordia."

The definition of a germ as "matter potentially alive, and having within itself the tendency to assume a definite living form," appears to meet all the requirements of modern science. For, notwithstanding it might be justly questioned whether a germ is not merely potentially, but rather actually, alive, though its vital manifestations are reduced to a minimum, the term "potential" may fairly be used in a sense broad enough to escape the objection. And the qualification of "potential" has the advantage of reminding us that the great characteristic of the germ is not so much what it is, but what it may, under suitable conditions, become. Harvey shared the belief of Aristotle--whose writings he so often quotes and of whom he speaks as his precursor and model, with the generous respect with which one genuine worker should regard another--that such germs may arise by a process of "equivocal generation" out of not-living matter; and the aphorism so commonly ascribed to him, "omne vivum ex ovo" and which is indeed a fair summary of his reiterated assertions, though incessantly employed against the modern advocates of spontaneous generation, can be honestly so used only by those who have never read a score of pages of the "Exercitationes." Harvey, in fact, believed as implicitly as Aristotle did in the equivocal generation of the lower animals. But, while the course of modern investigation has only brought out into greater prominence the accuracy of Harvey's conception of the nature and mode of development of germs, it has as distinctly tended to disprove the occurrence of equivocal generation, or abiogenesis, in the present course of nature. In the immense majority of both plants and animals, it is certain that the germ is not merely a body in which life is dormant or potential, but that it is itself simply a detached portion of the substance of a pre-existing living body; and the evidence has yet to be adduced which will satisfy any cautious reasoner that "omne vivum ex vivo" is not as well-established a law of the existing course of nature as "omne vivum ex ovo."

In all instances which have yet been investigated, the substance of this germ has a peculiar chemical composition, consisting of at fewest four elementary bodies, viz., carbon, hydrogen, oxygen, and nitrogen, united into the ill-defined compound known as protein, and associated with much water, and very generally, if not always, with sulphur and phosphorus in minute proportions. Moreover, up to the present time, protein is known only as a product and constituent of living matter. Again, a true germ is either devoid of any structure discernible by optical means, or, at most, it is a simple nucleated cell. [Footnote: 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." [Footnote: Harvey, Exercitationes de Generatione. Ex. 45, "Quænam sit pulli materia et quomodo fiat in Ovo."] 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.

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

II. The Evolution of the Sum of Living Beings.

The notion that all the kinds of animals and plants may have come into existence by the growth and modification of primordial germs is as old as speculative thought; but the modern scientific form of the doctrine can be traced historically to the influence of several converging lines of philosophical speculation and of physical observation, none of which go farther back than the seventeenth century. These are:--

1. The enunciation by Descartes of the conception that the physical universe, whether living or not living, is a mechanism, and that, as such, it is explicable on physical principles.

2. The observation of the gradations of structure, from extreme simplicity to very great complexity, presented by living things, and of the relation of these graduated forms to one another.