I. EVOLUTION IN BIOLOGY
Evolution in Biology - Introduction
In the former half of the eighteenth century, the term "evolution" was introduced into biological writings, in order to denote the mode in which some of the most eminent physiologists of that time conceived that the generations of living things took place; in opposition to the hypothesis advocated, in the preceding century, by Harvey in that remarkable work [Footnote 744-1] which would give him a claim to rank among the founders of biological science, even had he not been the discoverer of the circulation of the blood.
One of Harvey's prime objects is to defend and establish, on the basis of direct observation, the opinion already held by Aristotle; that, in the higher animals at any rate, the formation of the [188] new organism by the process of generation takes place, not suddenly, by simultaneous accretion of rudiments of all, or of the most important, of the organs of the adult; nor by sudden metamorphosis of a formative substance into a miniature of the whole, which subsequently grows; but by epigenesis, or successive differentiation of a relatively homogeneous rudiment into the parts and structures which are characteristic of the adult.
"Et primò, quidem, quoniam per epigenesin sive partium superexorientium additamentum pullum fabricari certum est: quænam pars ante alias omnes exstruatur, et quid de illa ejusque generandi modo observandum veniat, dispiciemus. Ratum sane est et in ovo manifestè apparet quod Aristoteles de perfectorum animalium generatione enuntiat: nimirum, non omnes partes simul fieri, sed ordine aliam post aliam; primùmque existere particulam genitalem, cujus virtute postea (tanquam ex principio quodam) reliquæ omnes partes prosiliant. Qualem in plantarum seminibus (fabis, putà, aut glandibus) gemmam sive apicem protuberantem cernimus, totius futuræ arboris principium. Estque hæc particula velut filius emancipatus seorsumque collocatus, et principium per se vivens; unde postea membrorum ordo describitur; et quæcunque ad absolvendum animal pertinent, disponuntur. [Footnote 744-2] Quoniam enim nulla pars se ipsam generat; sed postquam generata est, se ipsam jam auget; ideo eam primùm oriri necesse est, quæ principium augendi contineat (sive enim planta, sive animal est, æque omnibus inest quod vim habeat vegetandi, sive nutriendi), [Footnote 744-3] simulque reliquas omnes partes suo quamque ordine distinguat et formet; proindeque in eadem primogenita particula anima primario inest, sensus, motusque, et totius vitæ auctor et principium." (Exercitatio 51.)
Harvey proceeds to contrast this view with that of the "Medici," or followers of Hippocrates and Galen, who, "badly philosophising," imagined that the brain, the heart, and the liver were simultaneously first generated in the form of vesicles; and, at the same time, while expressing his agreement with Aristotle in the principle of epigenesis, he maintains that it is the blood which is the primal generative part, and not, as Aristotle thought, the heart.
In the latter part of the 17th century, the doctrine of epigenesis, thus advocated by Harvey, was controverted, on the ground of direct observation, by Malpighi, who affirmed that the body of the chick is to be seen in the egg, before the punctum sanguineum makes its appearance. But, from this perfectly correct observation a conclusion which is by no means warranted was drawn; namely, that the chick, as a whole, really exists in the egg antecedently to incubation; and that what happens in the course of the latter process is no addition of new parts, "alias post alias natas," as Harvey puts it, but a simple expansion, or unfolding, of the organs which already exist, though they are too small and inconspicuous to be discovered. The weight of Malpighi's observations therefore fell into the scale of that doctrine which Harvey terms metamorphosis, in contradistinction to epigenesis.
The views of Malphigi were warmly welcomed, on philosophical grounds, by Leibnitz, [Footnote 744-4] who found in them a support to his hypothesis of monads, and by Malebranche; [Footnote 744-5] while, in the middle of the 18th century, not only speculative considerations, but a great number of new and interesting observations on the phenomena of generation, led the ingenious Bonnet, and Haller, [Footnote 744-6] the first physiologist of the age, to adopt, advocate, and extend them.
Bonnet affirms that, before fecundation, the hen's egg contains an excessively minute but complete chick; and that fecundation and incubation simply cause this germ to absorb nutritious matters, which are deposited in the interstices of the elementary structures of which the miniature chick, or germ, is made up. The consequence of this intussusceptive growth is the "development" or "evolution" of the germ into the visible bird. Thus an organised individual (tout organisé) "is a composite body consisting of the original, or elementary, parts and of the matters which have been associated with them by the aid of nutrition;" so that, if these matters could be extracted from the individual (tout), it would, so to speak, become concentrated in a point, and would thus be restored to its primitive condition of a germ; "just as by extracting from a bone the calcareous substance which is the source of its hardness, it is reduced to its primitive state of gristle or membrane." [Footnote 745-1]
"Evolution" and "development" are, for Bonnet, synonymous terms; and since by "evolution" he means simply the expansion of that which was invisible into visibility, he was naturally led to the conclusion, at which Leibnitz had arrived by a different line of reasoning, that no such thing as generation, in the proper sense of the word, exists in Nature. The growth of an organic being is simply a process of enlargement as a particle of dry gelatine may be swelled up by the intussusception of water; its death is a shrinkage, such as the swelled jelly might undergo on desiccation. Nothing really new is produced in the living world, but the germs which develop have existed since the beginning of things; and nothing really dies, but, when what we call death takes place, the living thing shrinks back into its germ state. [Footnote 745-2]
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 745-3]
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 Eloge, 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 745-4]
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 cœur 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'œuf avant qu'il ait été couvé." [Footnote 745-5]
"J'ai ouvert une grande quantité d'œufs 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 745-6]
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 745-7] 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 "Evolution" 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 "Evolution" 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.
Footnotes
744-1 The Exercitationes de Generatione Animalium, which Dr. George Ent extracted from him and published in 1651.
744-2 De Generatione Animalium, lib.ii. cap. x.
744-3 De Generatione, lib. ii. cap. iv.
744-4 "Cependant, pour revenir aux formes ordinaires, ou aux âmes matérielles, cette durée qu'il leur faut attribuer à la place de celle qu'on avoit attribuée aux atomes pourroit faire douter si elles ne vont pas de corps en corps; ce qui seroit la métempsychose, à peu près comme quelques philosophes ont cru la transmission du mouvement et celle des espèces. Mais cette imagination est bien éloignée de la nature des choses. II n'y a point de tel passage, et c'est ici où les transformations de Messieurs Swammerdam, Malpighi, et Leewenhoek, qui sont des plus excellens observateurs de notre tems, sont venues à mon secours, et m'ont fait admettre plus aisément, que l'animal, et toute autre substance organisée ne commence point lorsque nous le croyons, et que sa generation apparente n'est qu'une développement et une espèce d'augmentation. Aussi ai je remarqué que l'auteur de la Recherche da la Verité, M. Regis, M. Hartsoeker, et d'autres habiles hommes n'ont pas été fort éloignés de ce sentiment." Leibnitz, Système Nouveau de la Nature, 1695. The doctrine of "Emboîtement" is contained in the Considérations sur le Principe de Vie, 1705, the preface to the Theodicée, 1710, and the Principes de la Nature et da la Grace (§ 6), 1718.
744-5 "II est vrai que la pensée la plus raisonnable et la plus conforme à l'experience sur cette question très difficile de la formation du fœtus; c'est que les enfans sont déja presque tout formés avant même l'action par laquelle ils sont conçus; et que leurs mères ne font que leur donner l'accroissement ordinaire dans le temps de la grossesse." De la Recherche de la Verité, livre ii. chap. vii. p. 334, 7th ed., 1721.
744-6 The writer is indebted to Dr. Allen Thomson for reference to the evidence contained in a note to Haller's edition of Boerhaave's Prælectiones Academicæ, vol. v. pt. ii. p. 497, published in 1744, that Haller originally advocated epigenesis.
745-1 Considérations sur les Corps organisés, chap. x.
745-2 Bonnet had the courage of his opinions, and in the Palingénésie Philosophique, part vi. chap. iv., he develops a hypothesis which he terms "évolution naturelle;" and which, making allowance for his peculiar vievs of the nature of generation, bears no small resemblance to what is understood by "evolution" at the present day:–
"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 organisés 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 l'etoient probablement alors bien différens de ce qu'ils sont aujourd'hui. Ils l'etoient autent 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."
745-3 "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.
745-4 "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, Eloge de Cuvier, note 12.
745-5 Histoire Naturelle, tom. ii. ed. ii. 1750, p. 350.
745-6 Ibid., p. 351.
745-7 See particularly Buffon, l.c. p. 41.
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