The embryology of man and the animals, the history of which we have reviewed in the last two chapters, was mainly a descriptive science forty years ago. The earlier investigations in this province were chiefly directed to the discovery, by careful observation, of the wonderful facts of the embryonic development of the animal body from the ovum. Forty years ago no one dared attack the question of the causes of these phenomena. For fully a century, from the year 1759, when Wolff’s solid Theoria generationis appeared, until 1859, when Darwin published his famous Origin of Species, the real causes of the embryonic processes were quite unknown. No one thought of seeking the agencies that effected this marvellous succession of structures. The task was thought to be so difficult as almost to pass beyond the limits of human thought. It was reserved for Charles Darwin to initiate us into the knowledge of these causes. This compels us to recognise in this great genius, who wrought a complete revolution in the whole field of biology, a founder at the same time of a new period in embryology. It is true that Darwin occupied himself very little with direct embryological research, and even in his chief work he only touches incidentally on the embryonic phenomena; but by his reform of the theory of descent and the founding of the theory of selection he has given us the means of attaining to a real knowledge of the causes of embryonic formation. That is, in my opinion, the chief feature in Darwin’s incalculable influence on the whole science of evolution.

When we turn our attention to this latest period of embryological research, we pass into the second division of organic evolution—stem-evolution, or phylogeny. I have already indicated in Chapter I the important and intimate causal connection between these two sections of the science of evolution—between the evolution of the individual and that of his ancestors. We have formulated this connection in the biogenetic law; the shorter evolution, that of the individual, or ontogenesis, is a rapid and summary repetition, a condensed recapitulation, of the larger evolution, or that of the species. In this principle we express all the essential points relating to the causes of evolution; and we shall seek throughout this work to confirm this principle and lend it the support of facts. When we look to its causal significance, perhaps it would be better to formulate the biogenetic law thus: “The evolution of the species and the stem ( phylon) shows us, in the physiological functions of heredity and adaptation, the conditioning causes on which the evolution of the individual depends”; or, more briefly: “Phylogenesis is the mechanical cause of ontogenesis.”

But before we examine the great achievement by which Darwin revealed the causes of evolution to us, we must glance at the efforts of earlier scientists to attain this object. Our historical inquiry into these will be even shorter than that into the work done in the field of ontogeny. We have very few names to consider here. At the head of them we find the great French naturalist, Jean Lamarck, who first established evolution as a scientific theory in 1809. Even before his time, however, the chief philosopher, Kant, and the chief poet, Goethe, of Germany had occupied themselves with the subject. But their efforts passed almost without recognition in the eighteenth century. A “philosophy of nature” did not arise until the beginning of the nineteenth century. In the whole of the time before this no one had ventured to raise seriously the question of the origin of species, which is the culminating point of phylogeny. On all sides it was regarded as an insoluble enigma.

The whole science of the evolution of man and the other animals is intimately connected with the question of the nature of species, or with the problem of the origin of the various animals which we group together under the name of species. Thus the definition of the species becomes important. It is well known that this definition was given by LinnÉ, who, in his famous Systema NaturÆ (1735), was the first to classify and name the various groups of animals and plants, and drew up an orderly scheme of the species then known. Since that time “species” has been the most important and indispensable idea in descriptive natural history, in zoological and botanical classification; although there have been endless controversies as to its real meaning.

What, then, is this “organic species”? LinnÉ himself appealed directly to the Mosaic narrative; he believed that, as it is stated in Genesis, one pair of each species of animals and plants was created in the beginning, and that all the individuals of each species are the descendants of these created couples. As for the hermaphrodites (organisms that have male and female organs in one being), he thought it sufficed to assume the creation of one sole individual, since this would be fully competent to propagate its species. Further developing these mystic ideas, LinnÉ went on to borrow from Genesis the account of the deluge and of Noah’s ark as a ground for a science of the geographical and topographical distribution of organisms. He accepted the story that all the plants, animals, and men on the earth were swept away in a universal deluge, except the couples preserved with Noah in the ark, and ultimately landed on Mount Ararat. This mountain seemed to LinnÉ particularly suitable for the landing, as it reaches a height of more than 16,000 feet, and thus provides in its higher zones the several climates demanded by the various species of animals and plants: the animals that were accustomed to a cold climate could remain at the summit; those used to a warm climate could descend to the foot; and those requiring a temperate climate could remain half-way down. From this point the re-population of the earth with animals and plants could proceed.

It was impossible to have any scientific notion of the method of evolution in LinnÉ’s time, as one of the chief sources of information, paleontology, was still wholly unknown. This science of the fossil remains of extinct animals and plants is very closely bound up with the whole question of evolution. It is impossible to explain the origin of living organisms without appealing to it. But this science did not rise until a much later date. The real founder of scientific paleontology was Georges Cuvier, the most distinguished zoologist who, after LinnÉ, worked at the classification of the animal world, and effected a complete revolution in systematic zoology at the beginning of the nineteenth century. In regard to the nature of the species he associated himself with LinnÉ and the Mosaic story of creation, though this was more difficult for him with his acquaintance with fossil remains. He clearly showed that a number of quite different animal populations have lived on the earth; and he claimed that we must distinguish a number of stages in the history of our planet, each of which was characterised by a special population of animals and plants. These successive populations were, he said, quite independent of each other, and therefore the supernatural creative act, which was demanded as the origin of the animals and plants by the dominant creed, must have been repeated several times. In this way a whole series of different creative periods must have succeeded each other; and in connection with these he had to assume that stupendous revolutions or cataclysms—something like the legendary deluge—must have taken place repeatedly. Cuvier was all the more interested in these catastrophes or cataclysms as geology was just beginning to assert itself, and great progress was being made in our knowledge of the structure and formation of the earth’s crust. The various strata of the crust were being carefully examined, especially by the famous geologist Werner and his school, and the fossils found in them were being classified; and these researches also seemed to point to a variety of creative periods. In each period the earth’s crust, composed of the various strata, seemed to be differently constituted, just like the population of animals and plants that then lived on it. Cuvier combined this notion with the results of his own paleontological and zoological research; and in his effort to get a consistent view of the whole process of the earth’s history he came to form the theory which is known as “the catastrophic theory,” or the theory of terrestrial revolutions. According to this theory, there have been a series of mighty cataclysms on the earth, and these have suddenly destroyed the whole animal and plant population then living on it; after each cataclysm there was a fresh creation of living things throughout the earth. As this creation could not be explained by natural laws, it was necessary to appeal to an intervention on the part of the Creator. This catastrophic theory, which Cuvier described in a special work, was soon generally accepted, and retained its position in biology for half a century.

However, Cuvier’s theory was completely overthrown sixty years ago by the geologists, led by Charles Lyell, the most distinguished worker in this field of science. Lyell proved in his famous Principles of Geology (1830) that the theory was false, in so far as it concerned the crust of the earth; that it was totally unnecessary to bring in supernatural agencies or general catastrophes in order to explain the structure and formation of the mountains; and that we can explain them by the familiar agencies which are at work to-day in altering and reconstructing the surface of the earth. These causes are—the action of the atmosphere and water in its various forms (snow, ice, fog, rain, the wear of the river, and the stormy ocean), and the volcanic action which is exerted by the molten central mass. Lyell convincingly proved that these natural causes are quite adequate to explain every feature in the build and formation of the crust. Hence Cuvier’s theory of cataclysms was very soon driven out of the province of geology, though it remained for another thirty years in undisputed authority in biology. All the zoologists and botanists who gave any thought to the question of the origin of organisms adhered to Cuvier’s erroneous idea of revolutions and new creations.

In order to illustrate the complete stagnancy of biology from 1830 to 1859 on the question of the origin of the various species of animals and plants, I may say, from my own experience, that during the whole of my university studies I never heard a single word said about this most important problem of the science. I was fortunate enough at that time (1852–1857) to have the most distinguished masters for every branch of biological science. Not one of them ever mentioned this question of the origin of species. Not a word was ever said about the earlier efforts to understand the formation of living things, nor about Lamarck’s Philosophie Zoologique which had made a fresh attack on the problem in 1809. Hence it is easy to understand the enormous opposition that Darwin encountered when he took up the question for the first time. His views seemed to float in the air, without a single previous effort to support them. The whole question of the formation of living things was considered by biologists, until 1859, as pertaining to the province of religion and transcendentalism; even in speculative philosophy, in which the question had been approached from various sides, no one had ventured to give it serious treatment. This was due to the dualistic system of Immanuel Kant, who taught a natural system of evolution as far as the inorganic world was concerned; but, on the whole, adopted a supernaturalist system as regards the origin of living things. He even went so far as to say: “It is quite certain that we cannot even satisfactorily understand, much less explain, the nature of an organism and its internal forces on purely mechanical principles; it is so certain, indeed, that we may confidently say: ‘It is absurd for a man to imagine even that some day a Newton will arise who will explain the origin of a single blade of grass by natural laws not controlled by design’—such a hope is entirely forbidden us.” In these words Kant definitely adopts the dualistic and teleological point of view for biological science.

Nevertheless, Kant deserted this point of view at times, particularly in several remarkable passages which I have dealt with at length in my Natural History of Creation (chap. v), where he expresses himself in the opposite, or monistic, sense. In fact, these passages would justify one, as I showed, in claiming his support for the theory of evolution. However, these monistic passages are only stray gleams of light; as a rule, Kant adheres in biology to the obscure dualistic ideas, according to which the forces at work in inorganic nature are quite different from those of the organic world. This dualistic system prevails in academic philosophy to-day—most of our philosophers still regarding these two provinces as totally distinct. They put, on the one side, the inorganic or “lifeless” world, in which there are at work only mechanical laws, acting necessarily and without design; and, on the other, the province of organic nature, in which none of the phenomena can be properly understood, either as regards their inner nature or their origin, except in the light of preconceived design, carried out by final or purposive causes.

The prevalence of this unfortunate dualistic prejudice prevented the problem of the origin of species, and the connected question of the origin of man, from being regarded by the bulk of people as a scientific question at all until 1859. Nevertheless, a few distinguished students, free from the current prejudice, began, at the commencement of the nineteenth century, to make a serious attack on the problem. The merit of this attaches particularly to what is known as “the older school of natural philosophy,” which has been so much misrepresented, and which included Jean Lamarck, Buffon, Geoffroy St. Hilaire, and Blainville in France; Wolfgang Goethe, Reinhold Treviranus, Schelling, and Lorentz Oken in Germany [and Erasmus Darwin in England].

The gifted natural philosopher who treated this difficult question with the greatest sagacity and comprehensiveness was Jean Lamarck. He was born at Bazentin, in Picardy, on August 1st, 1744; he was the son of a clergyman, and was destined for the Church. But he turned to seek glory in the army, and eventually devoted himself to science.

His Philosophie Zoologique was the first scientific attempt to sketch the real course of the origin of species, the first “natural history of creation” of plants, animals, and men. But, as in the case of Wolff’s book, this remarkably able work had no influence whatever; neither one nor the other could obtain any recognition from their prejudiced contemporaries. No man of science was stimulated to take an interest in the work, and to develop the germs it contained of the most important biological truths. The most distinguished botanists and zoologists entirely rejected it, and did not even deign to reply to it. Cuvier, who lived and worked in the same city, has not thought fit to devote a single syllable to this great achievement in his memoir on progress in the sciences, in which the pettiest observations found a place. In short, Lamarck’s Philosophie Zoologique shared the fate of Wolff’s theory of development, and was for half a century ignored and neglected. The German scientists, especially Oken and Goethe, who were occupied with similar speculations at the same time, seem to have known nothing about Lamarck’s work. If they had known it, they would have been greatly helped by it, and might have carried the theory of evolution much farther than they found it possible to do.

To give an idea of the great importance of the Philosophie Zoologique, I will briefly explain Lamarck’s leading thought. He held that there was no essential difference between living and lifeless beings. Nature is one united and connected system of phenomena; and the forces which fashion the lifeless bodies are the only ones at work in the kingdom of living things. We have, therefore, to use the same method of investigation and explanation in both provinces. Life is only a physical phenomenon. All the plants and animals, with man at their head, are to be explained, in structure and life, by mechanical or efficient causes, without any appeal to final causes, just as in the case of minerals and other inorganic bodies. This applies equally to the origin of the various species. We must not assume any original creation, or repeated creations (as in Cuvier’s theory), to explain this, but a natural, continuous, and necessary evolution. The whole evolutionary process has been uninterrupted. All the different kinds of animals and plants which we see to-day, or that have ever lived, have descended in a natural way from earlier and different species; all come from one common stock, or from a few common ancestors. These remote ancestors must have been quite simple organisms of the lowest type, arising by spontaneous generation from inorganic matter. The succeeding species have been constantly modified by adaptation to their varying environment (especially by use and habit), and have transmitted their modifications to their successors by heredity.

Lamarck was the first to formulate as a scientific theory the natural origin of living things, including man, and to push the theory to its extreme conclusions—the rise of the earliest organisms by spontaneous generation (or abiogenesis) and the descent of man from the nearest related mammal, the ape. He sought to explain this last point, which is of especial interest to us here, by the same agencies which he found at work in the natural origin of the plant and animal species. He considered use and habit (adaptation) on the one hand, and heredity on the other, to be the chief of these agencies. The most important modifications of the organs of plants and animals are due, in his opinion, to the function of these very organs, or to the use or disuse of them. To give a few examples, the woodpecker and the humming-bird have got their peculiarly long tongues from the habit of extracting their food with their tongues from deep and narrow folds or canals; the frog has developed the web between his toes by his own swimming; the giraffe has lengthened his neck by stretching up to the higher branches of trees, and so on. It is quite certain that this use or disuse of organs is a most important factor in organic development, but it is not sufficient to explain the origin of species.

To adaptation we must add heredity as the second and not less important agency, as Lamarck perfectly recognised. He said that the modification of the organs in any one individual by use or disuse was slight, but that it was increased by accumulation in passing by heredity from generation to generation. But he missed altogether the principle which Darwin afterwards found to be the chief factor in the theory of transformation—namely, the principle of natural selection in the struggle for existence. It was partly owing to his failure to detect this supremely important element, and partly to the poor condition of all biological science at the time, that Lamarck did not succeed in establishing more firmly his theory of the common descent of man and the other animals.

Independently of Lamarck, the older German school of natural philosophy, especially Reinhold Treviranus, in his Biologie (1802), and Lorentz Oken, in his Naturphilosophie (1809), turned its attention to the problem of evolution about the end of the eighteenth and beginning of the nineteenth century. I have described its work in my History of Creation (chap. iv). Here I can only deal with the brilliant genius whose evolutionary ideas are of special interest—the greatest of German poets, Wolfgang Goethe. With his keen eye for the beauties of nature, and his profound insight into its life, Goethe was early attracted to the study of various natural sciences. It was the favourite occupation of his leisure hours throughout life. He gave particular and protracted attention to the theory of colours. But the most valuable of his scientific studies are those which relate to that “living, glorious, precious thing,” the organism. He made profound research into the science of structures or morphology (morphÆ = forms). Here, with the aid of comparative anatomy, he obtained the most brilliant results, and went far in advance of his time. I may mention, in particular, his vertebral theory of the skull, his discovery of the pineal gland in man, his system of the metamorphosis of plants, etc. These morphological studies led Goethe on to research into the formation and modification of organic structures which we must count as the first germ of the science of evolution. He approaches so near to the theory of descent that we must regard him, after Lamarck, as one of its earliest founders. It is true that he never formulated a complete scientific theory of evolution, but we find a number of remarkable suggestions of it in his splendid miscellaneous essays on morphology. Some of them are really among the very basic ideas of the science of evolution. He says, for instance (1807): “When we compare plants and animals in their most rudimentary forms, it is almost impossible to distinguish between them. But we may say that the plants and animals, beginning with an almost inseparable closeness, gradually advance along two divergent lines, until the plant at last grows in the solid, enduring tree and the animal attains in man to the highest degree of mobility and freedom.” That Goethe was not merely speaking in a poetical, but in a literal genealogical, sense of this close affinity of organic forms is clear from other remarkable passages in which he treats of their variety in outward form and unity in internal structure. He believes that every living thing has arisen by the interaction of two opposing formative forces or impulses. The internal or “centripetal” force, the type or “impulse to specification,” seeks to maintain the constancy of the specific forms in the succession of generations: this is heredity. The external or “centrifugal” force, the element of variation or “impulse to metamorphosis,” is continually modifying the species by changing their environment: this is adaptation. In these significant conceptions Goethe approaches very close to a recognition of the two great mechanical factors which we now assign as the chief causes of the formation of species.

However, in order to appreciate Goethe’s views on morphology, one must associate his decidedly monistic conception of nature with his pantheistic philosophy. The warm and keen interest with which he followed, in his last years, the controversies of contemporary French scientists, and especially the struggle between Cuvier and Geoffroy St. Hilaire (see chap. iv of The History of Creation), is very characteristic. It is also necessary to be familiar with his style and general tenour of thought in order to appreciate rightly the many allusions to evolution found in his writings. Otherwise, one is apt to make serious errors.

He approached so close, at the end of the eighteenth century, to the principles of the science of evolution that he may well be described as the first forerunner of Darwin, although he did not go so far as to formulate evolution as a scientific system, as Lamarck did.