discusses the very question at issue in a passage which is well worth reading (vol. i. p. 87), and also comes to a conclusion opposed to Mr. Mivart’s axiom. A proposition which has been so much disputed and repudiated, should, under no circumstances, have been thus confidently assumed to be true. For myself, I utterly reject it, inasmuch as the logical consequence of the adoption of any such principle is the denial of all moral value to sympathy and affection. According to Mr. Mivart’s axiom, the man who, seeing another struggling in the water, leaps in at the risk of his own life to save him, does that which is “destitute of the most incipient degree of real goodness,” unless, as he strips off his coat, he says to himself, “Now, mind, I am going to do this because it is my duty and for no other reason;” and the most beautiful character to which humanity can attain, that of the man who does good without thinking about it, because he loves justice and mercy and is repelled by evil, has no claim on our moral approbation. The denial that a man acts morally because he does not think whether he does so or not, may be put upon the same footing as the denial of the title of an arithmetician to the calculating boy, because he did not know how he worked his sums. If mankind ever generally accept and act upon Mr. Mivart’s axiom, they will simply become a set of most unendurable prigs; but they never have accepted it, and I venture to hope that evolution has nothing so terrible in store for the human race.
But if an action, the motive of which is nothing but affection or sympathy, may be deserving of moral approbation and really good, who that has ever had a dog of his own will deny that animals are capable of such actions? Mr. Mivart indeed says:–“It may be safely affirmed, however, that there is no trace in brutes of any actions simulating morality which are not explicable by the fear of punishment, by the hope of pleasure, or by personal affection” (p. 221). But it may be affirmed, with equal truth, that there is no trace in men of any actions which are not traceable to the same motives. If a man does anything, he does it either because he fears to be punished if he does not do it, or because he hopes to obtain pleasure by doing it, or because he gratifies his affections [Footnote: In separating pleasure and the gratification of affection, I simply follow Mr. Mivart without admitting the justice of the separation.] by doing it.
Assuming the position of the absolute moralists, let it be granted that there is a perception of right and wrong innate in every man. This means, simply, that when certain ideas are presented to his mind, the feeling of approbation arises; and when certain others, the feeling of disapprobation. To do your duty is to earn the approbation of your conscience, or moral sense; to fail in your duty is to feel its disapprobation, as we all say. Now, is approbation a pleasure or a pain? Surely a pleasure. And is disapprobation a pleasure or a pain? Surely a pain. Consequently, all that is really meant by the absolute moralists is that there is, in the very nature of man, something which enables him to be conscious of these particular pleasures and pains. And when they talk of immutable and eternal principles of morality, the only intelligible sense which I can put upon the words, is that the nature of man being what it is, he always has been, and always will be, capable of feeling these particular pleasures and pains. _À priori,_ I have nothing to say against this proposition. Admitting its truth, I do not see how the moral faculty is on a different footing from any of the other faculties of man. If I choose to say that it is an immutable and eternal law of human nature that “ginger is hot in the mouth,” the assertion has as much foundation of truth as the other, though I think it would be expressed in needlessly pompous language. I must confess that I have never been able to understand why there should be such a bitter quarrel between the intuitionists and the utilitarians. The intuitionist is, after all, only a utilitarian who believes that a particular class of pleasures and pains has an especial importance, by reason of its foundation in the nature of man, and its inseparable connection with his very existence as a thinking being. And as regards the motive of personal affection: Love, as Spinoza profoundly says, is the association of pleasure with that which is loved. [Footnote: “Nempe, Amor nihil aliud est, quam Lætitia, concomitante idea causæ externæ.”–_Ethices_, III. xiii.] Or, to put it to the common sense of mankind, is the gratification of affection a pleasure or a pain? Surely a pleasure. So that whether the motive which leads us to perform an action is the love of our neighbour, or the love of God, it is undeniable that pleasure enters into that motive.
Thus much in reply to Mr. Mivart’s arguments. I cannot but think that it is to be regretted that he ekes them out by ascribing to the doctrines of the philosophers with whom he does not agree, logical consequences which have been over and over again proved not to flow from them: and when reason fails him, tries the effect of an injurious nickname. According to the views of Mr. Spencer, Mr. Mill, and Mr. Darwin, Mr. Mivart tells us, “_virtue is a mere kind of retrieving:_” and, that we may not miss the point of the joke, he puts it in italics. But what if it is? Does that make it less virtue? Suppose I say that sculpture is a “mere way” of stone-cutting, and painting a “mere way” of daubing canvas, and music a “mere way” of making a noise, the statements are quite true; but they only show that I see no other method of depreciating some of the noblest aspects of humanity than that of using language in an inadequate and misleading sense about them. And the peculiar inappropriateness of this particular nickname to the views in question, arises from the circumstance which Mr. Mivart would doubtless have recollected, if his wish to ridicule had not for the moment obscured his judgment–that whether the law of evolution applies to man or not, that of hereditary transmission certainly does. Mr. Mivart will hardly deny that a man owes a large share of the moral tendencies which he exhibits to his ancestors; and the man who inherits a desire to steal from a kleptomaniac, or a tendency to benevolence from a Howard, is, so far as he illustrates hereditary transmission, comparable to the dog who inherits the desire to fetch a duck out of the water from his retrieving sire. So that, evolution, or no evolution, moral qualities are comparable to a “kind of retrieving;” though the comparison, if meant for the purposes of casting obloquy on evolution, does not say much for the fairness of those who make it.
The Quarterly Reviewer and Mr. Mivart base their objections to the evolution of the mental faculties of man from those of some lower animal form upon what they maintain to be a difference in kind between the mental and moral faculties of men and brutes; and I have endeavoured to show, by exposing the utter unsoundness of their philosophical basis, that these objections are devoid of importance.
The objections which Mr. Wallace brings forward to the doctrine of the evolution of the mental faculties of man from those of brutes by natural causes, are of a different order, and require separate consideration.
If I understand him rightly, he by no means doubts that both the bodily and the mental faculties of man have been evolved from those of some lower animal; but he is of opinion that some agency beyond that which has been concerned in the evolution of ordinary animals has been operative in the case of man. “A superior intelligence has guided the development of man in a definite direction and for a special purpose, just as man guides the development of many animal and vegetable forms.” [Footnote: “The Limits of Natural Selection as applied to Man” (_loc. cit._ p. 359).] I understand this to mean that, just as the rock-pigeon has been produced by natural causes, while the evolution of the tumbler from the blue rock has required the special intervention of the intelligence of man, so some anthropoid form may have been evolved by variation and natural selection; but it could never have given rise to man, unless some superior intelligence had played the part of the pigeon-fancier.
According to Mr. Wallace, “whether we compare the savage with the higher developments of man, or with the brutes around him, we are alike driven to the conclusion, that, in his large and well-developed brain, he possesses an organ quite disproportioned to his requirements” (p. 343); and he asks, “What is there in the life of the savage but the satisfying of the cravings of appetite in the simplest and easiest way? What thoughts, idea, or actions are there that raise him many grades above the elephant or the ape?” (p. 342.) I answer Mr. Wallace by citing a remarkable passage which occurs in his instructive paper on “Instinct in Man and Animals.”
“Savages make long journeys in many directions, and, their whole faculties being directed to the subject, they gain a wide and accurate knowledge of the topography, not only of their own district, but of all the regions round about. Every one who has travelled in a new direction communicates his knowledge to those who have travelled less, and descriptions of routes and localities, and minute incidents of travel, form one of the main staples of conversation around the evening fire. Every wanderer or captive from another tribe adds to the store of information, and, as the very existence of individuals and of whole families and tribes depends upon the completeness of this knowledge, all the acute perceptive faculties of the adult savage are directed to acquiring and perfecting it. The good hunter or warrior thus comes to know the bearing of every hill and mountain range, the directions and junctions of all the streams, the situation of each tract characterised by peculiar vegetation, not only within the area he has himself traversed, but perhaps for a hundred miles around it. His acute observation enables him to detect the slightest undulations of the surface, the various changes of subsoil and alterations in the character of the vegetation that would be quite imperceptible to a stranger. His eye is always open to the direction in which he is going; the mossy side of trees, the presence of certain plants under the shade of rocks, the morning and evening flight of birds, are to him indications of direction almost as sure as the sun in the heavens” (pp. 207, 208).
I have seen enough of savages to be able to declare that nothing can be more admirable than this description of what a savage has to learn. But it is incomplete. Add to all this the knowledge which a savage is obliged to gain of the properties of plants, of the characters and habits of animals, and of the minute indications by which their course is discoverable: consider that even an Australian can make excellent baskets and nets, and neatly fitted and beautifully balanced spears; that he learns to use these so as to be able to transfix a quartern loaf at sixty yards; and that very often, as in the case of the American Indians, the language of a savage exhibits complexities which a well-trained European finds it difficult to master: consider that every time a savage tracks his game he employs a minuteness of observation, and an accuracy of inductive and deductive reasoning which, applied to other matters, would assure some reputation to a man of science, and I think we need ask no further why he possesses such a fair supply of brains. In complexity and difficulty, I should say that the intellectual labour of a “good hunter or warrior” considerably exceeds that of an ordinary Englishman. The Civil Service Examiners are held in great terror by young Englishmen; but even their ferocity never tempted them to require a candidate to possess such a knowledge of a parish as Mr. Wallace justly points out savages may possess of an area a hundred miles or more in diameter.
But suppose, for the sake of argument, that a savage has more brains than seems proportioned to his wants, all that can be said is that the objection to natural selection, if it be one, applies quite as strongly to the lower animals. The brain of a porpoise is quite wonderful for its mass, and for the development of the cerebral convolutions. And yet since we have ceased to credit the story of Arion, it is hard to believe that porpoises are much troubled with intellect: and still more difficult is it to imagine that their big brains are only a preparation for the advent of some accomplished cetacean of the future. Surely, again, a wolf must have too much brains, or else how is it that a dog with only the same quantity and form of brain is able to develop such singular intelligence? The wolf stands to the dog in the same relation as the savage to the man; and, therefore, if Mr. Wallace’s doctrine holds good, a higher power must have superintended the breeding up of wolves from some inferior stock, in order to prepare them to become dogs.
Mr. Wallace further maintains that the origin of some of man’s mental faculties by the preservation of useful variations is not possible. Such, for example, are “the capacity to form ideal conceptions of space and time, of eternity and infinity; the capacity for intense artistic feelings of pleasure in form, colour, and composition; and for those abstract notions of form and number which render geometry and arithmetic possible.” “How,” he asks, “were all or any of these faculties first developed, when they could have been of no possible use to man in his early stages of barbarism?”
Surely the answer is not far to seek. The lowest savages are as devoid of any such conceptions as the brutes themselves. What sort of conceptions of space and time, of form and number, can be possessed by a savage who has not got so far as to be able to count beyond five or six, who does not know how to draw a triangle or a circle, and has not the remotest notion of separating the particular quality we call form, from the other qualities of bodies? None of these capacities are exhibited by men, unless they form part of a tolerably advanced society. And, in such a society, there are abundant conditions by which a selective influence is exerted in favour of those persons who exhibit an approximation towards the possession of these capacities.
The savage who can amuse his fellows by telling a good story over the nightly fire, is held by them in esteem and rewarded, in one way or another, for so doing–in other words, it is an advantage to him to possess this power. He who can carve a paddle, or the figure-head of a canoe better, similarly profits beyond his duller neighbour. He who counts a little better than others, gets most yams when barter is going on, and forms the shrewdest estimate of the numbers of an opposing tribe. The experience of daily life shows that the conditions of our present social existence exercise the most extraordinarily powerful selective influence in favour of novelists, artists, and strong intellects of all kinds; and it seems unquestionable that all forms of social existence must have had the same tendency, if we consider the indisputable facts that even animals possess the power of distinguishing form and number, and that they are capable of deriving pleasure from particular forms and sounds. If we admit, as Mr. Wallace does, that the lowest savages are not raised “many grades above the elephant and the ape;” and if we further admit, as I contend must be admitted, that the conditions of social life tend, powerfully, to give an advantage to those individuals who vary in the direction of intellectual or æsthetic excellence, what is there to interfere with the belief that these higher faculties, like the rest, owe their development to natural selection?
Finally, with respect to the development of the moral sense out of the simple feelings of pleasure and pain, liking and disliking, with which the lower animals are provided, I can find nothing in Mr. Wallace’s reasonings which has not already been met by Mr. Mill, Mr. Spencer, or Mr. Darwin.
I do not propose to follow the Quarterly Reviewer and Mr. Mivart through the long string of objections in matters of detail which they bring against Mr. Darwin’s views. Every one who has considered the matter carefully will be able to ferret out as many more “difficulties”; but he will also, I believe, fail as completely as they appear to me to have done, in bringing forward any fact which is really contradictory of Mr. Darwin’s views. Occasionally, too, their objections and criticisms are based upon errors of their own. As, for example, when Mr. Mivart and the Quarterly Reviewer insist upon the resemblances between the eyes of _Cephalopoda_ and _Vertebrata_, quite forgetting that there are striking and altogether fundamental differences between them; or when the Quarterly Reviewer corrects Mr. Darwin for saying that the gibbons, “without having been taught, can walk or run upright with tolerable quickness, though they move awkwardly, and much less securely than man.” The Quarterly Reviewer says, “This is a little misleading, inasmuch as it is not stated that this upright progression is effected by placing the enormously long arms behind the head, or holding them out backwards as a balance in progression.”
Now, before carping at a small statement like this, the Quarterly Reviewer should have made sure that he was quite right. But he happens to be quite wrong. I suspect he got his notion of the manner in which a gibbon walks from a citation in “Man’s Place in Nature.” But at that time I had not seen a gibbon walk. Since then I have, and I can testify that nothing can be more precise than Mr. Darwin’s statement. The gibbon I saw walked without either putting his arms behind his head or holding them out backwards. All he did was to touch the ground with the outstretched fingers of his long arms now and then, just as one sees a man who carries a stick, but does not need one, touch the ground with it as he walks along.
Again, a large number of the objections brought forward by Mr. Mivart and the Quarterly Reviewer apply to evolution in general, quite as much as to the particular form of that doctrine advocated by Mr. Darwin; or, to their notions of Mr. Darwin’s views and not to what they really are. An excellent example of this class of difficulties is to be found in Mr. Mivart’s chapter on “Independent Similarities of Structure.” Mr. Mivart says that these cannot be explained by an “absolute and pure Darwinian,” but “that an innate power and evolutionary law, aided by the corrective action of natural selection, should have furnished like needs with like aids, is not at all improbable” (p. 82).
I do not exactly know what Mr. Mivart means by an “absolute and pure Darwinian;” indeed Mr. Mivart makes that creature hold so many singular opinions that I doubt if I can ever have seen one alive. But I find nothing in his statement of the view which he imagines to be originated by himself, which is really inconsistent with what I understand to be Mr. Darwin’s views.
I apprehend that the foundation of the theory of natural selection is the fact that living bodies tend incessantly to vary. This variation is neither indefinite, nor fortuitous, nor does it take place in all directions, in the strict sense of these words.
Accurately speaking, it is not indefinite, nor does it take place in all directions, because it is limited by the general characters of the type to which the organism exhibiting the variation belongs. A whale does not tend to vary in the direction of producing feathers, nor a bird in the direction of developing whalebone. In popular language there is no harm in saying that the waves which break upon the sea-shore are indefinite, fortuitous, and break in all directions. In scientific language, on the contrary, such a statement would be a gross error, inasmuch as every particle of foam is the result of perfectly definite forces, operating according to no less definite laws. In like manner, every variation of a living form, however minute, however apparently accidental, is inconceivable except as the expression of the operation of molecular forces or “powers” resident within the organism. And, as these forces certainly operate according to definite laws, their general result is, doubtless, in accordance with some general law which subsumes them all. And there appears to be no objection to call this an “evolutionary law.” But nobody is the wiser for doing so, or has thereby contributed, in the least degree, to the advance of the doctrine of evolution, the great need of which is a theory of variation.
When Mr. Mivart tells us that his “aim has been to support the doctrine that these species have been evolved by ordinary _natural laws_ (for the most part unknown), aided by the _subordinate_ action of ‘natural selection'” (pp. 332-3), he seems to be of opinion that his enterprise has the merit of novelty. All I can say is that I have never had the slightest notion that Mr. Darwin’s aim is in any way different from this. If I affirm that “species have been evolved by variation [Footnote: Including under this head hereditary transmission.] (a natural process, the laws of which are for the most part unknown), aided by the subordinate action of natural selection,” it seems to me that I enunciate a proposition which constitutes the very pith and marrow of the first edition of the “Origin of Species.” And what the evolutionist stands in need of just now, is not an iteration of the fundamental principle of Darwinism, but some light upon the questions, What are the limits of variation? and, If a variety has arisen, can that variety be perpetuated, or even intensified, when selective conditions are indifferent, or perhaps unfavourable to its existence? I cannot find that Mr. Darwin has ever been very dogmatic in answering these questions. Formerly, he seems to have inclined to reply to them in the negative, while now his inclination is the other way. Leaving aside those broad questions of theology, philosophy, and ethics, by the discussion of which neither the Quarterly Reviewer nor Mr. Mivart can be said to have damaged Darwinism–whatever else they have injured–this is what their criticisms come to. They confound a struggle for some rifle-pits with an assault on the fortress.
In some respects, finally, I can only characterise the Quarterly Reviewer’s treatment of Mr. Darwin as alike unjust and unbecoming. Language of this strength requires justification, and on that ground I add the remarks which follow.
The Quarterly Reviewer opens his essay by a careful enumeration of all those points upon which, during the course of thirteen years of incessant labour, Mr. Darwin has modified his opinions. It has often and justly been remarked, that what strikes a candid student of Mr. Darwin’s works is not so much his industry, his knowledge, or even the surprising fertility of his inventive genius; but that unswerving truthfulness and honesty which never permit him to hide a weak place, or gloss over a difficulty, but lead him, on all occasions, to point out the weak places in his own armour, and even sometimes, it appears to me, to make admissions against himself which are quite unnecessary. A critic who desires to attack Mr. Darwin has only to read his works with a desire to observe, not their merits, but their defects, and he will find, ready to hand, more adverse suggestions than are likely ever to have suggested themselves to his own sharpness, without Mr. Darwin’s self-denying aid.
Now this quality of scientific candour is not so common that it needs to be discouraged; and it appears to me to deserve other treatment than that adopted by the Quarterly Reviewer, who deals with Mr. Darwin as an Old Bailey barrister deals with a man against whom he wishes to obtain a conviction, _per fas aut nefas_, and opens his case by endeavouring to create a prejudice against the prisoner in the minds of the jury. In his eagerness to carry out this laudable design, the Quarterly Reviewer cannot even state the history of the doctrine of natural selection without an oblique and entirely unjustifiable attempt to depreciate Mr. Darwin. “To Mr. Darwin,” says he, “and (through Mr. Wallace’s reticence) to Mr. Darwin alone, is due the credit of having first brought it prominently forward and demonstrated its truth.” No one can less desire than I do, to throw a doubt upon Mr. Wallace’s originality, or to question his claim to the honour of being one of the originators of the doctrine of natural selection; but the statement that Mr. Darwin has the sole credit of originating the doctrine because of Mr. Wallace’s reticence is simply ridiculous. The proof of this is, in the first place, afforded by Mr. Wallace himself, whose noble freedom from petty jealousy in this matter smaller folk would do well to imitate, and who writes thus:–“I have felt all my life, and I still feel, the most sincere satisfaction that Mr. Darwin had been at work long before me and that it was not left for me to attempt to write the ‘Origin of Species.’ I have long since measured my own strength, and know well that it would be quite unequal to that task.” So that if there was any reticence at all in the matter, it was Mr. Darwin’s reticence during the long twenty years of study which intervened between the conception and the publication of his theory, which gave Mr. Wallace the chance of being an independent discoverer of the importance of natural selection. And, finally, if it be recollected that Mr. Darwin’s and Mr. Wallace’s essays were published simultaneously in the “Journal of the Linnæan Society” for 1858, it follows that the Reviewer, while obliquely depreciating Mr. Darwin’s deserts, has in reality awarded to him a priority which, in legal strictness, does not exist.
Mr. Mivart, whose opinions so often concur with those of the Quarterly Reviewer, puts the case in a way, which I much regret to be obliged to say, is, in my judgment, quite as incorrect; though the injustice may be less glaring. He says that the theory of natural selection is, in general, exclusively associated with the name of Mr. Darwin, “on account of the noble self-abnegation of Mr. Wallace.” As I have said, no one can honour Mr. Wallace more than I do, both for what he has done and for what he has not done, in his relation to Mr. Darwin. And perhaps nothing is more creditable to him than his frank declaration that he could not have written such a work as the “Origin of Species.” But, by this declaration, the person most directly interested in the matter repudiates, by anticipation, Mr. Mivart’s suggestion that Mr. Darwin’s eminence is more or less due to Mr. Wallace’s modesty.
VI
EVOLUTION IN BIOLOGY
[1878]
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: The _Exercitationes de Generatione Animalium_, which Dr. George Ent extracted from him and published in 1651.] 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 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 seorsumquc collocatus, et principium per se vivens; unde postea, membrorum ordo describitur; et quæcunque ad absolvendum animal pertinent, disponuntur._ [Footnote: _De Generatione Animalium_, lib. ii. cap. x.] 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: _De Generatione_, lib. ii. cap. iv.] 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 seventeenth 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 it 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 Malpighi were warmly welcomed, on philosophical grounds, by Leibnitz, [Footnote: “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. Il 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 de 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 “Embôitement” is contained in the _Considérations sur le Principe de Vie_, 1705; the preface to the _Theodicée_, 1710; and the _Principes de la Nature et de la Grace_ (§ 6), 1718.] who found in them a support to his hypothesis of monads, and by Malebranche; [Footnote: “Il est vrai que la pensée la plus raisonnable et la plus conforme à l’experience sur cette question très difficile de la formation du foetus; 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.] while, 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, [Footnote: 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.] 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: _Considérations sur les Corps organisés, chap. x.] “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: 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 views 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 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.
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 and have 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. [Footnote: As Buffon has well said:–“L’idée de ramener l’explication de tous les phénomènes à des principes mecaniques est assurement grande et belle, ce pas est le plus hardi qu’on peut faire en philosophie, et c’est Descartes qui l’a fait.”–_l. c._ p. 50.]
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 quo le monde n’ait été créé au commencement avec autant de perfection qu’il eu 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 puissance de 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.” [Footnote: _Principes de la Philosophie_, Troisième partie, § 45.]
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.” [Footnote: _Ethices_, Pars tertia, Præfatio.] 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 the principles established inductively by the study of the present course of nature. Somewhat later, Maupertuis [Footnote: _Système de la Nature_. “Essai sur la Formation des Corps Organisés,” 1751, xiv.] suggested 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. Robinet [Footnote: _Considérations Philosophiques sur la gradation naturelle des formes de l’être; ou les essais de la nature qui apprend a faire l’homme,_ 1768.] followed 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 had taken 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 mot _nature_, 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 denaturent 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.” [Footnote: _Recherches sur les causes des principaux faits physiques_, par J.B. Lamarck. Paris. Seconde année de la République. In the preface, Lamarck says that the work was written in 1776, and presented to the Academy in 1780; but it was not published before 17994, and, at that time, it presumably expressed Lamarck’s mature views. It would be interesting to know what brought about the change of opinion manifested in the _Recherches sur l’organisation des corps vivants_, published only seven years later.]
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 Surface du 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, [Footnote: See the “Historical Sketch” prefixed to the last edition of the _Origin of Species_.] up 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. Spencer [Footnote: _First Principles_. and _Principles of Biology_, 1860-1864.] and Professor Haeckel [Footnote: _Generelle Marphologie_, 1866.] have 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 la Philosophie” 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 of organisation 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. [Footnote: “Il s’agit donc de prouver que la série qui constitue l’échelle animale réside essentiellement dans la distribution des masses principales qui la composent et non dans celle des espèces ni même toujours dans celle des genres.”–_Philosophie Zoologique_. chap. v.]
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 gradations between the two exhibited by the _Lycopodiaceæ, Rhizocarpeæ_, and _Gymnospermeæ_. The groups of _Fungi_, _Lichenes_, and _Algæ_ have 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–[Footnote: _Philosophie Zoologique_, première partie, chap. iii.] “Il n’y a que ceux qui se sont longtemps et fortement occupés de la détermination des espèces, et qui ont consulté de riches collections, qui peuvent savoir jusqu’à quel point les _espèces_, parmi les corps vivants se fondent les unes dans les autres, et qui ont pu se convaincre que, dans les parties où nous voyons des _espèces_ isolès, cela n’est ainsi que parcequ’il nous en manque d’autres qui en sont plus voisines et que nous n’avons pas encore recueillies.
“Je ne veux pas dire pour cela que les animaux qui existent forment une série très-simple et partout également nuancée; mais je dis qu’ils forment une série ramense, irréguliérement graduée et qui n’a point de discontinuité dans ses parties, ou qui, du moins, n’en a toujours pas eu, s’il est vrai que, par suite de quelques espèces perdues, il s’en trouve quelque part. Il en resulte que les _espèces_ qui terminent chaque rameau de la série générale tiennent, au moins d’un côté, à d’autres espèces voisines qui se nuancent avec elles. Voilà ce que l’état bien connu des choses me met maintenant à portée de demontrer. Je n’ai besoin d’aucune hypothèse ni d’aucune supposition pour cela: j’en atteste tous les naturalistes observateurs.”
3. In a remarkable essay [Footnote: “Entwurf einer Darstellung der zwischen dem Embryozustände der höheren Thiere und dem permanenten der niederen stattfindenden Parallele,” _Beyträge zur Vergleichenden Anatomie_, Bd. ii. 1811.] Meckel remarks–
“There is no good physiologist who has not been struck by the observation that the original form of all organisms is one and the same, and that out of this one form, all, the lowest as well as the highest, are developed in such a manner that the latter pass through the permanent forms of the former as transitory stages. Aristotle, Haller, Harvey, Kielmeyer, Autenrieth, and many others, have either made this observation incidentally, or, especially the latter, have drawn particular attention to it, and deduced therefrom results of permanent importance for physiology.”
Meckel proceeds to exemplify the thesis, that the lower forms of animals represent stages in the course of the development of the higher, with a large series of illustrations.
After comparing the Salamanders and the perennibranchiate _Urodela_ with the Tadpoles and the Frogs, and enunciating the law that the more highly any animal is organised the more quickly does it pass through the lower stages, Meckel goes on to say–
“From these lowest Vertebrata to the highest, and to the highest forms among these, the comparison between the embryonic conditions of the higher animals and the adult states of the lower can be more completely and thoroughly instituted than if the survey is extended to the Invertebrata, inasmuch as the latter are in many respects constructed upon an altogether too dissimilar type; indeed they often differ from one another far more than the lowest vertebrate does from the highest mammal; yet the following pages will show that the comparison may also be extended to them with interest. In fact, there is a period when, as Aristotle long ago said, the embryo of the highest animal has the form of a mere worm; and, devoid of internal and external organisation, is merely an almost structureless lump of polype substance. Notwithstanding the origin of organs, it still for a certain time, by reason of its want of an internal bony skeleton, remains worm and mollusk, and only later enters into the series of the Vertebrata, although traces of the vertebral column even in the earliest periods testify its claim to a place in that series.”–_Op, cit_ pp. 4, 5.
If Meckel’s proposition is so far qualified, that the comparison of adult with embryonic forms is restricted within the limits of one type of organisation; and, if it is further recollected that the resemblance between the permanent lower form and the embryonic stage of a higher form is not special but general, it is in entire accordance with modern embryology; although there is no branch of biology which has grown so largely, and improved its methods so much, since Meckel’s time, as this. In its original form, the doctrine of “arrest of development,” as advocated by Geoffroy Saint-Hilaire and Serres, was no doubt an overstatement of the case. It is not true, for example, that a fish is a reptile arrested in its development, or that a reptile was ever a fish: but it is true that the reptile embryo, at one stage of its development, is an organism which, if it had an independent existence, must be classified among fishes; and all the organs of the reptile pass, in the course of their development, through conditions which are closely analogous to those which are permanent in some fishes.
4. That branch of biology which is termed Morphology is a commentary upon, and expansion of, the proposition that widely different animals or plants, and widely different parts of animals or plants, are constructed upon the same plan. From the rough comparison of the skeleton of a bird with that of a man by Belon, in the sixteenth century (to go no farther back), down to the theory of the limbs and the theory of the skull at the present day; or, from the first demonstration of the homologies of the parts of a flower by C. F. Wolff, to the present elaborate analysis of the floral organs, morphology exhibits a continual advance towards the demonstration of a fundamental unity among the seeming diversities of living structures. And this demonstration has been completed by the final establishment of the cell theory, which involves the admission of a primitive conformity, not only of all the elementary structures in animals and plants respectively, but of those in the one of these great divisions of living things with those in the other. No _à priori_ difficulty can be said to stand in the way of evolution, when it can be shown that all animals and all plants proceed by modes of development, which are similar in principle, from a fundamental protoplasmic material.
5. The innumerable cases of structures, which are rudimentary and apparently useless, in species, the close allies of which possess well-developed and functionally important homologous structures, are readily intelligible on the theory of evolution, while it is hard to conceive their _raison d’être_ on any other hypothesis. However, a cautious reasoner will probably rather explain such cases deductively from the doctrine of evolution than endeavour to support the doctrine of evolution by them. For it is almost impossible to prove that any structure, however rudimentary, is useless–that is to say, that it plays no part whatever in the economy; and, if it is in the slightest degree useful, there is no reason why, on the hypothesis of direct creation, it should not have been created. Nevertheless, double-edged as is the argument from rudimentary organs, there is probably none which has produced a greater effect in promoting the general acceptance of the theory of evolution.
6. The older advocates of evolution sought for the causes of the process exclusively in the influence of varying conditions, such as climate and station, or hybridisation, upon living forms. Even Treviranus has got no farther than this point. Lamarck introduced the conception of the action of an animal on itself as a factor in producing modification. Starting from the well-known fact that the habitual use of a limb tends to develop the muscles of the limb, and to produce a greater and greater facility in using it, he made the general assumption that the effort of an animal to exert an organ in a given direction tends to develop the organ in that direction. But a little consideration showed that, though Lamarck had seized what, as far it goes, is a true cause of modification, it is a cause the actual effects of which are wholly inadequate to account for any considerable modification in animals, and which can have no influence at all in the vegetable world; and probably nothing contributed so much to discredit evolution, in the early part of this century, as the floods of easy ridicule which were poured upon this part of Lamarck’s speculation. The theory of natural selection, or survival of the fittest, was suggested by Wells in 1813, and further elaborated by Matthew in 1831. But the pregnant suggestions of these writers remained practically unnoticed and forgotten, until the theory was independently devised and promulgated by Darwin and Wallace in 1858, and the effect of its publication was immediate and profound.
Those who were unwilling to accept evolution, without better grounds than such as are offered by Lamarck, or the author of that particularly unsatisfactory book, the “Vestiges of the Natural History of the Creation,” and who therefore preferred to suspend their judgment on the question, found in the principle of selective breeding, pursued in all its applications with marvellous knowledge and skill by Mr. Darwin, a valid explanation of the occurrence of varieties and races; and they saw clearly that, if the explanation would apply to species, it would not only solve the problem of their evolution, but that it would account for the facts of teleology, as well as for those of morphology; and for the persistence of some forms of life unchanged through long epochs of time, while others undergo comparatively rapid metamorphosis.
How far “natural selection” suffices for the production of species remains to be seen. Few can doubt that, if not the whole cause, it is a very important factor in that operation; and that it must play a great part in the sorting out of varieties into those which are transitory and those which are permanent.
But the causes and conditions of variation have yet to be thoroughly explored; and the importance of natural selection will not be impaired, even if further inquiries should prove that variability is definite, and is determined in certain directions rather than in others, by conditions inherent in that which varies. It is quite conceivable that every species tends to produce varieties of a limited number and kind, and that the effect of natural selection is to favour the development of some of these, while it opposes the development of others along their predetermined lines of modification.
7. No truths brought to light by biological investigation were better calculated to inspire distrust of the dogmas intruded upon science in the name of theology, than those which relate to the distribution of animals and plants on the surface of the earth. Very skilful accommodation was needful, if the limitation of sloths to South America, and of the ornithorhynchus to Australia, was to be reconciled with the literal interpretation of the history of the deluge; and with the establishment of the existence of distinct provinces of distribution, any serious belief in the peopling of the world by migration from Mount Ararat came to an end.
Under these circumstances, only one alternative was left for those who denied the occurrence of evolution–namely, the supposition that the characteristic animals and plants of each great province were created as such, within the limits in which we find them. And as the hypothesis of “specific centres,” thus formulated, was heterodox from the theological point of view, and unintelligible under its scientific aspect, it may be passed over without further notice, as a phase of transition from the creational to the evolutional hypothesis.
8. In fact, the strongest and most conclusive arguments in favour of evolution are those which are based upon the facts of geographical, taken in conjunction with those of geological, distribution.
Both Mr. Darwin and Mr. Wallace lay great stress on the close relation which obtains between the existing fauna of any region and that of the immediately antecedent geological epoch in the same region; and rightly, for it is in truth inconceivable that there should be no genetic connection between the two. It is possible to put into words the proposition that all the animals and plants of each geological epoch were annihilated and that a new set of very similar forms was created for the next epoch; but it may be doubted if any one who ever tried to form a distinct mental image of this process of spontaneous generation on the grandest scale, ever really succeeded in realising it.
Within the last twenty years, the attention of the best palæontologists has been withdrawn from the hodman’s work of making “new species” of fossils, to the scientific task of completing our knowledge of individual species, and tracing out the succession of the forms presented by any given type in time.
Those who desire to inform themselves of the nature and extent of the evidence bearing on these questions may consult the works of Rütimeyer, Gaudry, Kowalewsky, Marsh, and the writer of the present article. It must suffice, in this place, to say that the successive forms of the Equine type have been fully worked out; while those of nearly all the other existing types of Ungulate mammals and of the _Carnivora_ have been almost as closely followed through the Tertiary deposits; the gradations between birds and reptiles have been traced; and the modifications undergone by the _Crocodilia_, from the Triassic epoch to the present day, have been demonstrated. On the evidence of palæontology, the evolution of many existing forms of animal life from their predecessors is no longer an hypothesis, but an historical fact; it is only the nature of the physiological factors to which that evolution is due which is still open to discussion.
[At page 209, the reference to Erasmus Darwin does not do justice to that ingenious writer, who, in the 39th section of the _Zoonomia_, clearly and repeatedly enunciates the theory of the inheritance of acquired modifications. For example “From their first rudiment, or primordium, to the termination of their lives, all animals undergo perpetual transformations; which are in part produced by their own exertions in consequence of their desires and aversions, of their pleasures and their pains, or of irritation, or of associations; and many of these acquired forms or propensities are transmitted to their posterity.” _Zoonomia_ I., p. 506. 1893.]
VII
THE COMING OF AGE OF “THE ORIGIN OF SPECIES”
[1880]
Many of you will be familiar with the aspect of this small green-covered book. It is a copy of the first edition of the “Origin of Species,” and bears the date of its production–the 1st of October 1859. Only a few months, therefore, are needed to complete the full tale of twenty-one years since its birthday.
Those whose memories carry them back to this time will remember that the infant was remarkably lively, and that a great number of excellent persons mistook its manifestations of a vigorous individuality for mere naughtiness; in fact there was a very pretty turmoil about its cradle. My recollections of the period are particularly vivid, for, having conceived a tender affection for a child of what appeared to me to be such remarkable promise, I acted for some time in the capacity of a sort of under-nurse, and thus came in for my share of the storms which threatened the very life of the young creature. For some years it was undoubtedly warm work; but considering how exceedingly unpleasant the apparition of the newcomer must have been to those who did not fall in love with him at first sight, I think it is to the credit of our age that the war was not fiercer, and that the more bitter and unscrupulous forms of opposition died away as soon as they did.
I speak of this period as of something past and gone, possessing merely an historical, I had almost said an antiquarian interest. For, during the second decade of the existence of the “Origin of Species,” opposition, though by no means dead, assumed a different aspect. On the part of all those who had any reason to respect themselves, it assumed a thoroughly respectful character. By this time, the dullest began to perceive that the child was not likely to perish of any congenital weakness or infantile disorder, but was growing into a stalwart personage, upon whom mere goody scoldings and threatenings with the birch-rod were quite thrown away.
In fact, those who have watched the progress of science within the last ten years will bear me out to the full, when I assert that there is no field of biological inquiry in which the influence of the “Origin of Species” is not traceable; the foremost men of science in every country are either avowed champions of its leading doctrines, or at any rate abstain from opposing them; a host of young and ardent investigators seek for and find inspiration and guidance in Mr. Darwin’s great work; and the general doctrine of evolution, to one side of which it gives expression, obtains, in the phenomena of biology, a firm base of operations whence it may conduct its conquest of the whole realm of Nature.
History warns us, however, that it is the customary fate of new truths to begin as heresies and to end as superstitions; and, as matters now stand, it is hardly rash to anticipate that, in another twenty years, the new generation, educated under the influences of the present day, will be in danger of accepting the main doctrines of the “Origin of Species,” with as little reflection, and it may be with as little justification, as so many of our contemporaries, twenty years ago, rejected them.
Against any such a consummation let us all devoutly pray; for the scientific spirit is of more value than its products, and irrationally held truths may be more harmful than reasoned errors. Now the essence of the scientific spirit is criticism. It tells us that whenever a doctrine claims our assent we should reply, Take it if you can compel it. The struggle for existence holds as much in the intellectual as in the physical world. A theory is a species of thinking, and its right to exist is coextensive with its power of resisting extinction by its rivals.
From this point of view, it appears to me that it would be but a poor way of celebrating the Coming of Age of the “Origin of Species,” were I merely to dwell upon the facts, undoubted and remarkable as they are, of its far-reaching influence and of the great following of ardent disciples who are occupied in spreading and developing its doctrines. Mere insanities and inanities have before now swollen to portentous size in the course of twenty years. Let us rather ask this prodigious change in opinion to justify itself: let us inquire whether anything has happened since 1859, which will explain, on rational grounds, why so many are worshipping that which they burned, and burning that which they worshipped. It is only in this way that we shall acquire the means of judging whether the movement we have witnessed is a mere eddy of fashion, or truly one with the irreversible current of intellectual progress, and, like it, safe from retrogressive reaction.
Every belief is the product of two factors: the first is the state of the mind to which the evidence in favour of that belief is presented; and the second is the logical cogency of the evidence itself. In both these respects, the history of biological science during the last twenty years appears to me to afford an ample explanation of the change which has taken place; and a brief consideration of the salient events of that history will enable us to understand why, if the “Origin of Species” appeared now, it would meet with a very different reception from that which greeted it in 1859.
One-and-twenty years ago, in spite of the work commenced by Hutton and continued with rare skill and patience by Lyell, the dominant view of the past history of the earth was catastrophic. Great and sudden physical revolutions, wholesale creations and extinctions of living beings, were the ordinary machinery of the geological epic brought into fashion by the misapplied genius of Cuvier. It was gravely maintained and taught that the end of every geological epoch was signalised by a cataclysm, by which every living being on the globe was swept away, to be replaced by a brand-new creation when the world returned to quiescence. A scheme of nature which appeared to be modelled on the likeness of a succession of rubbers of whist, at the end of each of which the players upset the table and called for a new pack, did not seem to shock anybody.
I may be wrong, but I doubt if, at the present time, there is a single responsible representative of these opinions left. The progress of scientific geology has elevated the fundamental principle of uniformitarianism, that the explanation of the past is to be sought in the study of the present, into the position of an axiom; and the wild speculations of the catastrophists, to which we all listened with respect a quarter of a century ago, would hardly find a single patient hearer at the present day. No physical geologist now dreams of seeking, outside the range of known natural causes, for the explanation of anything that happened millions of years ago, any more than he would be guilty of the like absurdity in regard to current events.
The effect of this change of opinion upon biological speculation is obvious. For, if there have been no periodical general physical catastrophes, what brought about the assumed general extinctions and re-creations of life which are the corresponding biological catastrophes? And, if no such interruptions of the ordinary course of nature have taken place in the organic, any more than in the inorganic, world, what alternative is there to the admission of evolution?
The doctrine of evolution in biology is the necessary result of the logical application of the principles of uniformitarianism to the phenomena of life. Darwin is the natural successor of Hutton and Lyell, and the “Origin of Species” the logical sequence of the “Principles of Geology.”
The fundamental doctrine of the “Origin of Species,” as of all forms of the theory of evolution applied to biology, is “that the innumerable species, genera, and families of organic beings with which the world is peopled have all descended, each within its own class or group, from common parents, and have all been modified in the course of descent.” [Footnote: _Origin of Species_, ed. I, p. 457.]
And, in view of the facts of geology, it follows that all living animals and plants “are the lineal descendants of those which lived long before the Silurian epoch.” [Footnote: _Origin of Species_, p. 458.]
It is an obvious consequence of this theory of descent with modification, as it is sometimes called, that all plants and animals, however different they may now be, must, at one time or other, have been connected by direct or indirect intermediate gradations, and that the appearance of isolation presented by various groups of organic beings must be unreal.
No part of Mr. Darwin’s work ran more directly counter to the prepossessions of naturalists twenty years ago than this. And such prepossessions were very excusable, for there was undoubtedly a great deal to be said, at that time, in favour of the fixity of species and of the existence of great breaks, which there was no obvious or probable means of filling up, between various groups of organic beings.
For various reasons, scientific and unscientific, much had been made of the hiatus between man and the rest of the higher mammalia, and it is no wonder that issue was first joined on this part of the controversy. I have no wish to revive past and happily forgotten controversies; but I must state the simple fact that the distinctions in the cerebral and other characters, which were so hotly affirmed to separate man from all other animals in 1860, have all been demonstrated to be non-existent, and that the contrary doctrine is now universally accepted and taught.
But there were other cases in which the wide structural gaps asserted to exist between one group of animals and another were by no means fictitious; and, when such structural breaks were real, Mr. Darwin could account for them only by supposing that the intermediate forms which once existed had become extinct. In a remarkable passage he says–
“We may thus account even for the distinctness of whole classes from each other–for instance, of birds from all other vertebrate animals–by the belief that many animal forms of life have been utterly lost, through which the early progenitors of birds were formerly connected with the early progenitors of the other vertebrate classes.” [Footnote: _Origin of Species_, p. 431.] Adverse criticism made merry over such suggestions as these. Of course it was easy to get out of the difficulty by supposing extinction; but where was the slightest evidence that such intermediate forms between birds and reptiles as the hypothesis required ever existed? And then probably followed a tirade upon this terrible forsaking of the paths of “Baconian induction.”
But the progress of knowledge has justified Mr. Darwin to an extent which could hardly have been anticipated. In 1862, the specimen of _Archæopteryx_, which, until the last two or three years, has remained unique, was discovered; and it is an animal which, in its feathers and the greater part of its organisation, is a veritable bird, while, in other parts, it is as distinctly reptilian.
In 1868, I had the honour of bringing under your notice, in this theatre, the results of investigations made, up to that time, into the anatomical characters of certain ancient reptiles, which showed the nature of the modifications in virtue of which the type of the quadrupedal reptile passed into that of a bipedal bird; and abundant confirmatory evidence of the justice of the conclusions which I then laid before you has since come to light.
In 1875, the discovery of the toothed birds of the cretaceous formation in North America by Professor Marsh completed the series of transitional forms between birds and reptiles, and removed Mr. Darwin’s proposition that “many animal forms of life have been utterly lost, through which the early progenitors of birds were formerly connected with the early progenitors of the other vertebrate classes,” from the region of hypothesis to that of demonstrable fact.
In 1859, there appeared to be a very sharp and clear hiatus between vertebrated and invertebrated animals, not only in their structure, but, what was more important, in their development. I do not think that we even yet know the precise links of connection between the two; but the investigations of Kowalewsky and others upon the development of _Amphioxus_ and of the _Tunicata_ prove, beyond a doubt, that the differences which were supposed to constitute a barrier between the two are non-existent. There is no longer any difficulty in understanding how the vertebrate type may have arisen from the invertebrate, though the full proof of the manner in which the transition was actually effected may still be lacking.
Again, in 1859, there appeared to be a no less sharp separation between the two great groups of flowering and flowerless plants. It is only subsequently that the series of remarkable investigations inaugurated by Hofmeister has brought to light the extraordinary and altogether unexpected modifications of the reproductive apparatus in the _Lycopodiaceæ_, the _Rhizocarpeæ_, and the _Gymnospermeæ_, by which the ferns and the mosses are gradually connected with the Phanerogamic division of the vegetable world.
So, again, it is only since 1859 that we have acquired that wealth of knowledge of the lowest forms of life which demonstrates the futility of any attempt to separate the lowest plants from the lowest animals, and shows that the two kingdoms of living nature have a common borderland which belongs to both, or to neither.
Thus it will be observed that the whole tendency of biological investigation, since 1859, has been in the direction of removing the difficulties which the apparent breaks in the series created at that time; and the recognition of gradation is the first step towards the acceptance of evolution.
As another great factor in bringing about the change of opinion which has taken place among naturalists, I count the astonishing progress which has been made in the study of embryology. Twenty years ago, not only were we devoid of any accurate knowledge of the mode of development of many groups of animals and plants, but the methods of investigation were rude and imperfect. At the present time, there is no important group of organic beings the development of which has not been carefully studied; and the modern methods of hardening and section-making enable the embryologist to determine the nature of the process, in each case, with a degree of minuteness and accuracy which is truly astonishing to those whose memories carry them back to the beginnings of modern histology. And the results of these embryological investigations are in complete harmony with the requirements of the doctrine of evolution. The first beginnings of all the higher forms of animal life are similar, and however diverse their adult conditions, they start from a common foundation. Moreover, the process of development of the animal or the plant from its primary egg, or germ, is a true process of evolution–a progress from almost formless to more or less highly organised matter, in virtue of the properties inherent in that matter.
To those who are familiar with the process of development, all _a priori_ objections to the doctrine of biological evolution appear childish. Any one who has watched the gradual formation of a complicated animal from the protoplasmic mass, which constitutes the essential element of a frog’s or a hen’s egg, has had under his eyes sufficient evidence that a similar evolution of the whole animal world from the like foundation is, at any rate, possible.
Yet another product of investigation has largely contributed to the removal of the objections to the doctrine of evolution current in 1859. It is the proof afforded by successive discoveries that Mr. Darwin did not over-estimate the imperfection of the geological record. No more striking illustration of this is needed than a comparison of our knowledge of the mammalian fauna of the Tertiary epoch in 1859 with its present condition. M. Gaudry’s researches on the fossils of Pikermi were published in 1868, those of Messrs. Leidy, Marsh, and Cope, on the fossils of the Western Territories of America, have appeared almost wholly since 1870, those of M. Filhol on the phosphorites of Quercy in 1878. The general effect of these investigations has been to introduce to us a multitude of extinct animals, the existence of which was previously hardly suspected; just as if zoologists were to become acquainted with a country, hitherto unknown, as rich in novel forms of life as Brazil or South Africa once were to Europeans. Indeed, the fossil fauna of the Western Territories of America bid fair to exceed in interest and importance all other known Tertiary deposits put together; and yet, with the exception of the case of the American tertiaries, these investigations have extended over very limited areas; and, at Pikermi, were confined to an extremely small space.
Such appear to me to be the chief events in the history of the progress of knowledge during the last twenty years, which account for the changed feeling with which the doctrine of evolution is at present regarded by those who have followed the advance of biological science, in respect of those problems which bear indirectly upon that doctrine.
But all this remains mere secondary evidence. It may remove dissent, but it does not compel assent. Primary and direct evidence in favour of evolution can be furnished only by palæontology. The geological record, so soon as it approaches completeness, must, when properly questioned, yield either an affirmative or a negative answer: if evolution has taken place, there will its mark be left; if it has not taken place, there will lie its refutation.
What was the state of matters in 1859? Let us hear Mr. Darwin, who may be trusted always to state the case against himself as strongly as possible.
“On this doctrine of the extermination of an infinitude of connecting links between the living and extinct inhabitants of the world, and at each successive period between the extinct and still older species, why is not every geological formation charged with such links? Why does not every collection of fossil remains afford plain evidence of the gradation and mutation of the forms of life? We meet with no such evidence, and this is the most obvious and plausible of the many objections which may be urged against my theory.” [Footnote: _Origin of Species_, ed. 1, p. 463.]
Nothing could have been more useful to the opposition than this characteristically candid avowal, twisted as it immediately was into an admission that the writer’s views were contradicted by the facts of palæontology. But, in fact, Mr. Darwin made no such admission. What he says in effect is, not that palæontological evidence is against him, but that it is not distinctly in his favour; and, without attempting to attenuate the fact, he accounts for it by the scantiness and the imperfection of that evidence.
What is the state of the case now, when, as we have seen, the amount of our knowledge respecting the mammalia of the Tertiary epoch is increased fifty-fold, and in some directions even approaches completeness?
Simply this, that, if the doctrine of evolution had not existed, palaeontologists must have invented it, so irresistibly is it forced upon the mind by the study of the remains of the Tertiary mammalia which have been brought to light since 1859.
Among the fossils of Pikermi, Gaudry found the successive stages by which the ancient civets passed into the more modern hyænas; through the Tertiary deposits of Western America, Marsh tracked the successive forms by which the ancient stock of the horse has passed into its present form; and innumerable less complete indications of the mode of evolution of other groups of the higher mammalia have been obtained. In the remarkable memoir on the phosphorites of Quercy, to which I have referred, M. Filhol describes no fewer than seventeen varieties of the genus _Cynodictis_, which fill up all the interval between the viverine animals and the bear-like dog _Amphicyon_; nor do I know any solid ground of objection to the supposition that, in this _Cynodictis-Amphicyon_ group, we have the stock whence all the Viveridæ, Felidæ, Hyænidæ, Canidæ, and perhaps the Procyonidæ and Ursidæ, of the present fauna have been evolved. On the contrary, there is a great deal to be said in favour.
In the course of summing up his results, M. Filhol observes:–
“During the epoch of the phosphorites, great changes took place in animal forms, and almost the same types as those which now exist became defined from one another.
“Under the influence of natural conditions of which we have no exact knowledge, though traces of them are discoverable, species have been modified in a thousand ways: races have arisen which, becoming fixed, have thus produced a corresponding number of secondary species.”
In 1859, language of which this is an unintentional paraphrase, occurring in the “Origin of Species,” was scouted as wild speculation; at present, it is a sober statement of the conclusions to which an acute and critically-minded investigator is led by large and patient study of the facts of palæontology. I venture to repeat what I have said before, that so far as the animal world is concerned, evolution is no longer a speculation, but a statement of historical fact. It takes its place alongside of those accepted truths which must be reckoned with by philosophers of all schools.
Thus when, on the first day of October next, “The Origin of Species” comes of age, the promise of its youth will be amply fulfilled; and we shall be prepared to congratulate the venerated author of the book, not only that the greatness of his achievement and its enduring influence upon the progress of knowledge have won him a place beside our Harvey; but, still more, that, like Harvey, he has lived long enough to outlast detraction and opposition, and to see the stone that the builders rejected become the head-stone of the corner.
VIII
CHARLES DARWIN
[_Nature_, April 27th, 1882]
Very few, even among those who have taken the keenest interest in the progress of the revolution in natural knowledge set afoot by the publication of “The Origin of Species,” and who have watched, not without astonishment, the rapid and complete change which has been effected both inside and outside the boundaries of the scientific world in the attitude of men’s minds towards the doctrines which are expounded in that great work, can have been prepared for the extraordinary manifestation of affectionate regard for the man, and of profound reverence for the philosopher, which followed the announcement, on Thursday last, of the death of Mr. Darwin.
Not only in these islands, where so many have felt the fascination of personal contact with an intellect which had no superior, and with a character which was even nobler than the intellect; but, in all parts of the civilised world, it would seem that those whose business it is to feel the pulse of nations and to know what interests the masses of mankind, were well aware that thousands of their readers would think the world the poorer for Darwin’s death, and would dwell with eager interest upon every incident of his history. In France, in Germany, in Austro-Hungary, in Italy, in the United States, writers of all shades of opinion, for once unanimous, have paid a willing tribute to the worth of our great countryman, ignored in life by the official representatives of the kingdom, but laid in death among his peers in Westminster Abbey by the will of the intelligence of the nation.
It is not for us to allude to the sacred sorrows of the bereaved home at Down; but it is no secret that, outside that domestic group, there are many to whom Mr. Darwin’s death is a wholly irreparable loss. And this not merely because of his wonderfully genial, simple, and generous nature; his cheerful and animated conversation, and the infinite variety and accuracy of his information; but because the more one knew of him, the more he seemed the incorporated ideal of a man of science. Acute as were his reasoning powers, vast as was his knowledge, marvellous as was his tenacious industry, under physical difficulties which would have converted nine men out of ten into aimless invalids; it was not these qualities, great as they were, which impressed those who were admitted to his intimacy with involuntary veneration, but a certain intense and almost passionate honesty by which all his thoughts and actions were irradiated, as by a central fire.
It was this rarest and greatest of endowments which kept his vivid imagination and great speculative powers within due bounds; which compelled him to undertake the prodigious labours of original investigation and of reading, upon which his published works are based; which made him accept criticisms and suggestions from anybody and everybody, not only without impatience, but with expressions of gratitude sometimes almost comically in excess of their value; which led him to allow neither himself nor others to be deceived by phrases, and to spare neither time nor pains in order to obtain clear and distinct ideas upon every topic with which he occupied himself.
One could not converse with Darwin without being reminded of Socrates. There was the same desire to find some one wiser than himself; the same belief in the sovereignty of reason; the same ready humour; the same sympathetic interest in all the ways and works of men. But instead of turning away from the problems of Nature as hopelessly insoluble, our modern philosopher devoted his whole life to attacking them in the spirit of Heraclitus and of Democritus, with results which are the substance of which their speculations were anticipatory shadows.
The due appreciation, or even enumeration, of these results is neither practicable nor desirable at this moment. There is a time for all things–a time for glorying in our ever-extending conquests over the realm of Nature, and a time for mourning over the heroes who have led us to victory.
None have fought better, and none have been more fortunate, than Charles Darwin. He found a great truth trodden underfoot, reviled by bigots, and ridiculed by all the world; he lived long enough to see it, chiefly by his own efforts, irrefragably established in science, inseparably incorporated with the common thoughts of men, and only hated and feared by those who would revile, but dare not. What shall a man desire more than this? Once more the image of Socrates rises unbidden, and the noble peroration of the “Apology” rings in our ears as if it were Charles Darwin’s farewell:–
“The hour of departure has arrived, and we go our ways–I to die and you to live. Which is the better, God only knows.”
IX
THE DARWIN MEMORIAL
[June 9th, 1885]
_Address by the President of the Royal Society, in the name of the Memorial Committee, on handing over the statue of Darwin to H.R.H. the Prince of Wales, as representative of the Trustees of the British Museum_.
YOUR ROYAL HIGHNESS,–It is now three years since the announcement of the death of our famous countryman, Charles Darwin, gave rise to a manifestation of public feeling, not only in these realms, but throughout the civilised world, which, if I mistake not, is without precedent in the modest annals of scientific biography.
The causes of this deep and wide outburst of emotion are not far to seek. We had lost one of these rare ministers and interpreters of Nature whose names mark epochs in the advance of natural knowledge. For, whatever be the ultimate verdict of posterity upon this or that opinion which Mr. Darwin has propounded; whatever adumbrations or anticipations of his doctrines may be found in the writings of his predecessors; the broad fact remains that, since the publication and by reason of the publication, of “The Origin of Species” the fundamental conceptions and the aims of the students of living Nature have been completely changed. From that work has sprung a great renewal, a true “instauratio magna” of the zoological and botanical sciences.
But the impulse thus given to scientific thought rapidly spread beyond the ordinarily recognised limits of biology. Psychology, Ethics, Cosmology were stirred to their foundations, and the “Origin of Species” proved itself to be the fixed point which the general doctrine of evolution needed in order to move the world. “Darwinism,” in one form or another, sometimes strangely distorted and mutilated, became an everyday topic of men’s speech, the object of an abundance both of vituperation and of praise, more often than of serious study.
It is curious now to remember how largely, at first, the objectors predominated; but considering the usual fate of new views, it is still more curious to consider for how short a time the phase of vehement opposition lasted. Before twenty years had passed, not only had the importance of Mr. Darwin’s work been fully recognised, but the world had discerned the simple, earnest, generous character of the man, that shone through every page of his writings.
I imagine that reflections such as these swept through the minds alike of loving friends and of honourable antagonists when Mr. Darwin died; and that they were at one in the desire to honour the memory of the man who, without fear and without reproach, had successfully fought the hardest intellectual battle of these days.
It was in satisfaction of these just and generous impulses that our great naturalist’s remains were deposited in Westminster Abbey; and that, immediately afterwards, a public meeting, presided over by my lamented predecessor, Mr. Spottiswoode, was held in the rooms of the Royal Society, for the purpose of considering what further step should be taken towards the same end.
It was resolved to invite subscriptions, with the view of erecting a statue of Mr. Darwin in some suitable locality; and to devote any surplus to the advancement of the biological sciences.
Contributions at once flowed in from Austria, Belgium, Brazil, Denmark, France, Germany, Holland, Italy, Norway, Portugal, Russia, Spain, Sweden, Switzerland, the United States, and the British Colonies, no less than from all parts of the three kingdoms; and they came from all classes of the community. To mention one interesting case, Sweden sent in 2296 subscriptions “from all sorts of people,” as the distinguished man of science who transmitted them wrote, “from the bishop to the seamstress, and in sums from five pounds to two pence.”
The Executive Committee has thus been enabled to carry out the objects proposed. A “Darwin Fund” has been created, which is to be held in trust by the Royal Society, and is to be employed in the promotion of biological research.
The execution of the statue was entrusted to Mr. Boehm; and I think that those who had the good fortune to know Mr. Darwin personally will admire the power of artistic divination which has enabled the sculptor to place before us so very characteristic a likeness of one whom he had not seen.
It appeared to the Committee that, whether they regarded Mr. Darwin’s career or the requirements of a work of art, no site could be so appropriate as this great hall, and they applied to the Trustees of the British Museum for permission to erect it in its present position.
That permission was most cordially granted, and I am desired to tender the best thanks of the Committee to the Trustees for their willingness to accede to our wishes.
I also beg leave to offer the expression of our gratitude to your Royal Highness for kindly consenting to represent the Trustees to-day. It only remains for me, your Royal Highness, my Lords and Gentlemen, Trustees of the British Museum, in the name of the Darwin Memorial Committee, to request you to accept this statue of Charles Darwin.
We do not make this request for the mere sake of perpetuating a memory; for so long as men occupy themselves with the pursuit of truth, the name of Darwin runs no more risk of oblivion than does that of Copernicus, or that of Harvey.
Nor, most assuredly, do we ask you to preserve the statue in its cynosural position in this entrance-hall of our National Museum of Natural History as evidence that Mr. Darwin’s views have received your official sanction; for science does not recognise such sanctions, and commits suicide when it adopts a creed.
No; we beg you to cherish this Memorial as a symbol by which, as generation after generation of students of Nature enter yonder door, they shall be reminded of the ideal according to which they must shape their lives, if they would turn to the best account the opportunities offered by the great institution under your charge.
X
OBITUARY [Footnote: From the Obituary Notices of the _Proceedings of the Royal Society_, vol. 44.]
[1888]
Charles Robert Darwin was the fifth child and second son of Robert Waring Darwin and Susannah Wedgwood, and was born on the 12th February, 1809, at Shrewsbury, where his father was a physician in large practice.
Mrs. Robert Darwin died when her son Charles was only eight years old, and he hardly remembered her. A daughter of the famous Josiah Wedgwood, who created a new branch of the potter’s art, and established the great works of Etruria, could hardly fail to transmit important mental and moral qualities to her children; and there is a solitary record of her direct influence in the story told by a schoolfellow, who remembers Charles Darwin “bringing a flower to school, and saying that his mother had taught him how, by looking at the inside of the blossom, the name of the plant could be discovered.” (I., p. 28. [Footnote: The references throughout this notice are to the _Life and Letters_, unless the contrary is expressly stated.])
The theory that men of genius derive their qualities from their mothers, however, can hardly derive support from Charles Darwin’s case, in the face of the patent influence of his paternal forefathers. Dr. Darwin, indeed, though a man of marked individuality of character, a quick and acute observer, with much practical sagacity, is said not to have had a scientific mind. But when his son adds that his father “formed a theory for almost everything that occurred” (I., p. 20), he indicates a highly