XI.EVOLUTION IN BIOLOGY.

Strike out the propositions about which neither controversialist does or can know anything, and there is nothing left for them to quarrel about. Make a desert of the Unknowable, and the divine Astræa of philosophic peace will commence her blessed reign.

Inthe 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 generation of living things took place; in opposition to the hypothesis advocated, in the preceding century, by Harvey in that remarkable work63which 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 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 byepigenesis, or successive differentiation of arelatively homogeneous rudiment into the parts and structures which are characteristic of the adult.

“Et primò, quidem, quoniam perepigenesinsive 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 quodAristotelesde 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.64Quoniam enimnulla 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),65simulque 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 seventeenth century, thedoctrine 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 thepunctum sanguineummakes 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 termsmetamorphosis, in contradistinction to epigenesis.

The views of Malpighi were warmly welcomed, on philosophical grounds, by Leibnitz,66who found in them a support to his hypothesis of monads, and byMalebranche;67while, in the middle of the eighteenth 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,68the 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, orelementary, 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 berestored to its primitive condition of agerm; “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.”69

“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.70

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.71

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 Laurillards’ “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.”72

Bonnet’s eminent contemporary, Buffon, held nearly the same views with respect to the nature of the germ, and expresses them even more confidently.73

“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 qui’il ait été couvé.”“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.”74

“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.”74

The “moule intérieur” of Buffon is the aggregate of elementary parts which constitute the individual,and is thus the equivalent of Bonnet’s germ,75as 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 Caspar 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.

No exception is, at this time, known to the general law, established upon an immense multitudeof 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 agerm. Harvey76says—

“Omnibus viventibus primordium insit, ex quo et a quo proveniant. Liceat hoc nobisprimordium vegetalenominare; 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 insectorumvermisab 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 mayarise 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 andphosphorus 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.77

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 germplusthe nutriment which it has absorbed, it is probable that the adult contains neither in form, nor in substance, more than an inappreciable fraction of theconstituents 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 theMetazoaare divisible, the cells of the cell-aggregate which results from the process of yelk-division, and which is termed amorula, 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, theplanula. The wall of the planula is next pushed in on one side, or invaginated, whereby it is convertedinto a double-walled sac with an opening, theblastopore, which leads into the cavity lined by the inner wall. This cavity is the primitive alimentary cavity orarchenteron; the inner, or invaginated, layer is thehypoblast, the outer theepiblast; and the embryo, in this stage, is termed agastrula. In all the higher animals, a layer of cells makes its appearance between the hypoblast and the epiblast, and is termed themesoblast. 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, Cœlenterates, Worms, Echinoderms, Tunicates, Arthropods, Mollusks, and Vertebrates; and there are valid reasons for the belief, that all animals of higher organisation than theProtozoaagree 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 the 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 ... perepigenesinprocreatur, 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.”78

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 farther, 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 sortof 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 substance79of 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 be 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.

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.

3. The observation of the existence of an analogy between the series of gradations presented by the species which compose any great group of animals or plants, and the series of embryonic conditions of the highest members of that group.

4. The observation that large groups of species of widely different habits present the same fundamental plan of structure; and that parts of the same animal or plant, the functions of which are very different, likewise exhibit modifications of a common plan.

5. The observation of the existence of structures, in a rudimentary and apparently useless condition, in one species of a group, which are fully developed andhave definite functions in other species of the same group.

6. The observation of the effects of varying conditions in modifying living organisms.

7. The observation of the facts of geographical distribution.

8. The observation of the facts of the geological succession of the forms of life.

1. Notwithstanding the elaborate disguise which fear of the powers that were led Descartes to throw over his real opinions, it is impossible to read the “Principes de la Philosophie” without acquiring the conviction that this great philosopher held that the physical world and all things in it, whether living or not living, have originated by a process of evolution, due to the continuous operation of purely physical causes, out of a primitive relatively formless matter.80

The following passage is especially instructive:—

“Et tant s’en faut que je veuille que l’on croie toutes les choses que j’écrirai, que même je pretends en proposer ici quelques unes que je crois absolument être fausses; à savoir, je ne doute point que le monde n’ait été créé au commencement avec autant de perfection qu’il en a; en sorte que le soleil, la terre, la lune, et les étoiles ont été dès lors; et que la terre n’a pas eu seulement en soi les semences des plantes, mais que les plantes même en ont couvert une partie; et qu’Adam et Eve n’ont pas été créés enfans mais en âge d’hommes parfaits. La religion chrétienne veut que nous le croyons ainsi, et la raison naturelle nous persuade entièrement cette vérité; car si nous considérons la toute puissancede Dieu, nous devons juger que tout ce qu’il a fait a eu dès le commencement toute la perfection qu’il devoit avoir. Mais néanmoins, comme on connôitroit beaucoup mieux quelle a été la nature d’Adam et celle des arbres de Paradis si on avoit examiné comment les enfants se forment peu à peu dans le ventre de leurs mères et comment les plantes sortent de leurs semences, que si on avoit seulement considéré quels ils ont été quand Dieu les a créés: tout de même, nous ferons mieux entendre quelle est généralement la nature de toutes les choses qui sont au monde si nous pouvons imaginer quelques principes qui soient fort intelligibles et fort simples, desquels nous puissions voir clairement que les astres et la terre et enfin tout ce monde visible auroit pu être produit ainsi que de quelques semences (bien que nous sachions qu’il n’a pas été produit en cette façon) que si nous la decrivions seulement comme il est, ou bien comme nous croyons qu’il a été créé. Et parceque je pense avoir trouvé des principes qui sont tels, je tacherai ici de les expliquer.”81

If we read between the lines of this singular exhibition of force of one kind and weakness of another, it is clear that Descartes believed that he had divined the mode in which the physical universe had been evolved; and the “Traité de l’homme,” and the essay “Sur les Passions” afford abundant additional evidence that he sought for, and thought he had found, an explanation of the phenomena of physical life by deduction from purely physical laws.

Spinoza abounds in the same sense, and is as usual perfectly candid—

“Naturæ leges et regulæ, secundum quas omnia fiunt et ex unis formis in alias mutantur, sunt ubique et semper eadem.”82

Leibnitz’s doctrine of continuity necessarily led him in the same direction; and, of the infinite multitude of monads with which he peopled the world, each is supposed to be the focus of an endless process of evolution and involution. In the “Protogæa,” xxvi., Leibnitz distinctly suggests the mutability of species—

“Alii mirantur in saxis passim species videri quas vel in orbe cognito, vel saltem in vicinis locis frustra quæras. Ita “Cornua Ammonis,” quæ ex nautilorum numero habeantur, passim et forma et magnitudine (nam et pedali diametro aliquando reperiuntur) ab omnibus illis naturis discrepare dicunt, quas præbet mare. Sed quis absconditos ejus recessus aut subterraneas abyssos pervestigavit? quam multa nobis animalia antea ignota offert novus orbis? Et credibile est per magnas illas conversiones etiam animalium species plurimum immutatas.”

Thus, in the end of the seventeenth century, the seed was sown which has, at intervals, brought forth recurrent crops of evolutional hypotheses, based, more or less completely, on general reasonings.

Among the earliest of these speculations is that put forward by Benoit de Maillet in his “Telliamed,” which, though printed in 1735, was not published until twenty-three years later. Considering that this book was written before the time of Haller, or Bonnet, or Linnæus, or Hutton, it surely deserves more respectful consideration than it usually receives. For De Maillet not only has a definite conception of the plasticity of living things, and of the production of existing species by the modification of their predecessors; but he clearly apprehends the cardinal maxim of modern geological science, that the explanation of the structure of the globe is to be sought in the deductive application to geological phenomena of theprinciples established inductively by the study of the present course of nature. Somewhat later, Maupertuis83suggested a curious hypothesis as to the causes of variation, which he thinks may be sufficient to account for the origin of all animals from a single pair. Robinet84followed out much the same line of thought as De Maillet, but less soberly; and Bonnet’s speculations in the “Palingénésie,” which appeared in 1769, have already been mentioned. Buffon (1753-1778), at first a partisan of the absolute immutability of species, subsequently appears to have believed that larger or smaller groups of species have been produced by the modification of a primitive stock; but he contributed nothing to the general doctrine of evolution.

Erasmus Darwin (“Zoonomia,” 1794), though a zealous evolutionist, can hardly be said to have made any real advance on his predecessors; and, notwithstanding that Goethe (1791-4) had the advantage of a wide knowledge of morphological facts, and a true insight into their signification, while he threw all the power of a great poet into the expression of his conceptions, it may be questioned whether he supplied the doctrine of evolution with a firmer scientific basis than it already possessed. Moreover, whatever the value of Goethe’s labours in that field, they were not published before 1820, long after evolutionism hadtaken a new departure from the works of Treviranus and Lamarck—the first of its advocates who were equipped for their task with the needful large and accurate knowledge of the phenomena of life, as a whole. It is remarkable that each of these writers seems to have been led, independently and contemporaneously, to invent the same name of “Biology” for the science of the phenomena of life; and thus, following Buffon, to have recognised the essential unity of these phenomena, and their contradistinction from those of inanimate nature. And it is hard to say whether Lamarck or Treviranus has the priority in propounding the main thesis of the doctrine of evolution; for though the first volume of Treviranus’s “Biologie” appeared only in 1802, he says, in the preface to his later work, the “Erscheinungen und Gesetze des organischen Lebens,” dated 1831, that he wrote the first volume of the “Biologie” “nearly five-and-thirty years ago,” or about 1796.

Now, in 1794, there is evidence that Lamarck held doctrines which present a striking contrast to those which are to be found in the“Philosophie Zoologique,”as the following passages show:—

“685. Quoique mon unique objet dans cet article n’ait été que de traiter de la cause physique de l’entretien de la vie des êtres organiques, malgré cela j’ai osé avancer en débutant, que l’existence de ces êtres étonnants n’appartiennent nullement à la nature; que tout ce qu’on peut entendre par le motnature, ne pouvoit donner la vie, c’est-à-dire, que toutes les qualités de la matière, jointes à toutes les circonstances possibles, et même à l’activité répandue dans l’univers, ne pouvaient point produire un être muni du mouvement organique, capable de reproduire son semblable, et sujet à la mort.“686. Tous les individus de cette nature, qui existent, proviennent d’individus semblables qui tous ensemble constituent l’espèce entière. Or, je crois qu’il est aussi impossible à l’homme de connôitre la cause physique du premier individu de chaque espèce, que d’assigner aussi physiquement la cause de l’existence de la matière ou de l’univers entier. C’est au moins ce que le résultat de mes connaissances et de mes réflexions me portent à penser. S’il existe beaucoup de variétés produites par l’effet des circonstances, ces variétés ne dénaturent point les espèces; mais on se trompe, sans doute souvent, en indiquant comme espèce, ce qui n’est que variété; et alors je sens que cette erreur peut tirer à conséquence dans les raisonnements que l’on fait sur cette matière.”85

“686. Tous les individus de cette nature, qui existent, proviennent d’individus semblables qui tous ensemble constituent l’espèce entière. Or, je crois qu’il est aussi impossible à l’homme de connôitre la cause physique du premier individu de chaque espèce, que d’assigner aussi physiquement la cause de l’existence de la matière ou de l’univers entier. C’est au moins ce que le résultat de mes connaissances et de mes réflexions me portent à penser. S’il existe beaucoup de variétés produites par l’effet des circonstances, ces variétés ne dénaturent point les espèces; mais on se trompe, sans doute souvent, en indiquant comme espèce, ce qui n’est que variété; et alors je sens que cette erreur peut tirer à conséquence dans les raisonnements que l’on fait sur cette matière.”85

The first three volumes of Treviranus’s “Biologie,” which contain his general views of evolution, appeared between 1802 and 1805. The “Recherches sur l’organisation des corps vivants,” in which the outlines of Lamarck’s doctrines are given, was published in 1802; but the full development of his views, in the “Philosophie Zoologique,” did not take place until 1809.

The “Biologie” and the “Philosophie Zoologique” are both very remarkable productions, and are still worthy of attentive study, but they fell upon evil times. The vast authority of Cuvier was employed in support of the traditionally respectable hypotheses of special creation and of catastrophism; and the wild speculations of the “Discours sur les Révolutions de la Surfacedu Globe” were held to be models of sound scientific thinking, while the really much more sober and philosophical hypotheses of the “Hydrogeologie” were scouted. For many years it was the fashion to speak of Lamarck with ridicule, while Treviranus was altogether ignored.

Nevertheless, the work had been done. The conception of evolution was henceforward irrepressible, and it incessantly reappears, in one shape or another,86up to the year 1858, when Mr. Darwin and Mr. Wallace published their “Theory of Natural Selection.” The “Origin of Species” appeared in 1859; and it is within the knowledge of all whose memories go back to that time, that, henceforward, the doctrine of evolution has assumed a position and acquired an importance which it never before possessed. In the “Origin of Species,” and in his other numerous and important contributions to the solution of the problem of biological evolution, Mr. Darwin confines himself to the discussion of the causes which have brought about the present condition of living matter, assuming such matter to have once come into existence. On the other hand, Mr. Spencer87and Professor Haeckel88have dealt with the whole problem of evolution. The profound and vigorous writings of Mr. Spencer embody the spirit of Descartes in the knowledge of our own day, and may be regarded as the “Principes de laPhilosophie” of the nineteenth century; while, whatever hesitation may not unfrequently be felt by less daring minds, in following Haeckel in many of his speculations, his attempt to systematise the doctrine of evolution and to exhibit its influence as the central thought of modern biology, cannot fail to have a far-reaching influence on the progress of science.

If we seek for the reason of the difference between the scientific position of the doctrine of evolution a century ago, and that which it occupies now, we shall find it in the great accumulation of facts, the several classes of which have been enumerated above, under the second to the eighth heads. For those which are grouped under the second to the seventh of these classes, respectively, have a clear significance on the hypothesis of evolution, while they are unintelligible if that hypothesis be denied. And those of the eighth group are not only unintelligible without the assumption of evolution, but can be proved never to be discordant with that hypothesis, while, in some cases, they are exactly such as the hypothesis requires. The demonstration of these assertions would require a volume, but the general nature of the evidence on which they rest may be briefly indicated.

2. The accurate investigation of the lowest forms of animal life, commenced by Leeuwenhoek and Swammerdam, and continued by the remarkable labours of Reaumur, Trembley, Bonnet, and a host of other observers, in the latter part of the seventeenth and the first half of the eighteenth centuries, drew the attention of biologists to the gradation in the complexity oforganisation which is presented by living beings, and culminated in the doctrine of the “échelle des êtres,” so powerfully and clearly stated by Bonnet; and, before him, adumbrated by Locke and by Leibnitz. In the then state of knowledge, it appeared that all the species of animals and plants could be arranged in one series; in such a manner that, by insensible gradations, the mineral passed into the plant, the plant into the polype, the polype into the worm, and so, through gradually higher forms of life, to man, at the summit of the animated world.

But, as knowledge advanced, this conception ceased to be tenable in the crude form in which it was first put forward. Taking into account existing animals and plants alone, it became obvious that they fell into groups which were more or less sharply separated from one another; and, moreover, that even the species of a genus can hardly ever be arranged in linear series. Their natural resemblances and differences are only to be expressed by disposing them as if they were branches springing from a common hypothetical centre.

Lamarck, while affirming the verbal proposition that animals form a single series, was forced by his vast acquaintance with the details of zoology to limit the assertion to such a series as may be formed out of the abstractions constituted by the common characters of each group.89

Cuvier on anatomical, and Von Baer on embryological grounds, made the further step of proving that, even in this limited sense, animals cannot be arranged in a single series, but that there are several distinct plans of organisation to be observed among them, no one of which, in its highest and most complicated modification, leads to any of the others.

The conclusions enunciated by Cuvier and Von Baer have been confirmed, in principle, by all subsequent research into the structure of animals and plants. But the effect of the adoption of these conclusions has been rather to substitute a new metaphor for that of Bonnet than to abolish the conception expressed by it. Instead of regarding living things as capable of arrangement in one series like the steps of a ladder, the results of modern investigation compel us to dispose them as if they were the twigs and branches of a tree. The ends of the twigs represent individuals, the smallest groups of twigs species, larger groups genera, and so on, until we arrive at the source of all these ramifications of the main branch, which is represented by a common plan of structure. At the present moment, it is impossible to draw up any definition, based on broad anatomical or developmental characters, by which any one of Cuvier’s great groups shall be separated from all the rest. On the contrary, the lower members of each tend to converge towards the lower members of all the others. The same may be said of the vegetable world. The apparently clear distinction between flowering and flowerless plants has been broken down by the series of gradationsbetween the two exhibited by theLycopodiaceæ,Rhizocarpeæ, andGymnospermeæ. The groups ofFungi,Lichenes, andAlgæhave completely run into one another, and, when the lowest forms of each are alone considered, even the animal and vegetable kingdoms cease to have a definite frontier.

If it is permissible to speak of the relations of living forms to one another metaphorically, the similitude chosen must undoubtedly be that of a common root, whence two main trunks, one representing the vegetable and one the animal world, spring; and, each dividing into a few main branches, these subdivide into multitudes of branchlets and these into smaller groups of twigs.

As Lamarck has well said—90

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