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 to 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 (chapter 5), 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 (chapter 4). 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 (morphae = 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 chapter 4 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.
CHAPTER 1.5. THE MODERN SCIENCE OF EVOLUTION.
We owe so much of the progress of scientific knowledge to Darwin's Origin of Species that its influence is almost without parallel in the history of science. The literature of Darwinism grows from day to day, not only on the side of academic zoology and botany, the sciences which were chiefly affected by Darwin's theory, but in a far wider circle, so that we find Darwinism discussed in popular literature with a vigour and zest that are given to no other scientific conception. This remarkable success is due chiefly to two circumstances. In the first place, all the sciences, and especially biology, have made astounding progress in the last half-century, and have furnished a very vast quantity of proofs of the theory of evolution. In striking contrast to the failure of Lamarck and the older scientists to attract attention to their effort to explain the origin of living things and of man, we have this second and successful effort of Darwin, which was able to gather to its support a large number of established facts. Availing himself of the progress already made, he had very different scientific proofs to allege than Lamarck, or St. Hilaire, or Goethe, or Treviranus had had. But, in the second place, we must acknowledge that Darwin had the special distinction of approaching the subject from an entirely new side, and of basing the theory of descent on a consistent system, which now goes by the name of Darwinism.
Lamarck had unsuccessfully attempted to explain the modification of organisms that descend from a common form chiefly by the action of habit and the use of organs, though with the aid of heredity. But Darwin's success was complete when he independently sought to give a mechanical explanation, on a quite new ground, of this modification of plant and animal structures by adaptation and heredity. He was impelled to his theory of selection on the following grounds. He compared the origin of the various kinds of animals and plants which we modify artificially—by the action of artificial selection in horticulture and among domestic animals—with the origin of the species of animals and plants in their natural state. He then found that the agencies which we employ in the modification of forms by artificial selection are also at work in Nature. The chief of these agencies he held to be "the struggle for life." The gist of this peculiarly Darwinian idea is given in this formula: The struggle for existence produces new species without premeditated design in the life of Nature, in the same way that the will of man consciously selects new races in artificial conditions. The gardener or the farmer selects new forms as he wills for his own profit, by ingeniously using the agency of heredity and adaptation for the modification of structures; so, in the natural state, the struggle for life is always unconsciously modifying the various species of living things. This struggle for life, or competition of organisms in securing the means of subsistence, acts without any conscious design, but it is none the less effective in modifying structures. As heredity and adaptation enter into the closest reciprocal action under its influence, new structures, or alterations of structure, are produced; and these are purposive in the sense that they serve the organism when formed, but they were produced without any pre-conceived aim.
This simple idea is the central thought of Darwinism, or the theory of selection. Darwin conceived this idea at an early date, and then, for more than twenty years, worked at the collection of empirical evidence in support of it before he published his theory. His grandfather, Erasmus Darwin, was an able scientist of the older school of natural philosophy, who published a number of natural-philosophic works about the end of the eighteenth century. The most important of them is his Zoonomia, published in 1794, in which he expounds views similar to those of Goethe and Lamarck, without really knowing anything of the work of these contemporaries. However, in the writings of the grandfather the plastic imagination rather outran the judgment, while in Charles Darwin the two were better balanced.
Darwin did not publish any account of his theory until 1858, when Alfred Russel Wallace, who had independently reached the same theory of selection, published his own work. In the following year appeared the Origin of Species, in which he develops it at length and supports it with a mass of proof. Wallace had reached the same conclusion, but he had not so clear a perception as Darwin of the effectiveness of natural selection in forming species, and did not develop the theory so fully. Nevertheless, Wallace's writings, especially those on mimicry, etc., and an admirable work on The Geographical Distribution of Animals, contain many fine original contributions to the theory of selection. Unfortunately, this gifted scientist has since devoted himself to spiritism.* (* Darwin and Wallace arrived at the theory quite independently. Vide Wallace's Contributions to the Theory of Natural Selection (1870) and Darwinism (1891).)
Darwin's Origin of Species had an extraordinary influence, though not at first on the experts of the science. It took zoologists and botanists several years to recover from the astonishment into which they had been thrown through the revolutionary idea of the work. But its influence on the special sciences with which we zoologists and botanists are concerned has increased from year to year; it has introduced a most healthy fermentation in every branch of biology, especially in comparative anatomy and ontogeny, and in zoological and botanical classification. In this way it has brought about almost a revolution in the prevailing views.
However, the point which chiefly concerns us here—the extension of the theory to man—was not touched at all in Darwin's first work in 1859. It was believed for several years that he had no thought of applying his principles to man, but that he shared the current idea of man holding a special position in the universe. Not only ignorant laymen (especially several theologians), but also a number of men of science, said very naively that Darwinism in itself was not to be opposed; that it was quite right to use it to explain the origin of the various species of plants and animals, but that it was totally inapplicable to man.
In the meantime, however, it seemed to a good many thoughtful people, laymen as well as scientists, that this was wrong; that the descent of man from some other animal species, and immediately from some ape-like mammal, followed logically and necessarily from Darwin's reformed theory of evolution. Many of the acuter opponents of the theory saw at once the justice of this position, and, as this consequence was intolerable, they wanted to get rid of the whole theory.
The first scientific application of the Darwinian theory to man was made by Huxley, the greatest zoologist in England. This able and learned scientist, to whom zoology owes much of its progress, published in 1863 a small work entitled Evidence as to Man's Place in Nature. In the extremely important and interesting lectures which made up this work he proved clearly that the descent of man from the ape followed necessarily from the theory of descent. If that theory is true, we are bound to conceive the animals which most closely resemble man as those from which humanity has been gradually evolved. About the same time Carl Vogt published a larger work on the same subject. We must also mention Gustav Jaeger and Friedrich Rolle among the zoologists who accepted and taught the theory of evolution immediately after the publication of Darwin's book, and maintained that the descent of man from the lower animals logically followed from it. The latter published, in 1866, a work on the origin and position of man.
About the same time I attempted, in the second volume of my General Morphology (1866), to apply the theory of evolution to the whole organic kingdom, including man.* (* Huxley spoke of this "as one of the greatest scientific works ever published."—Translator.) I endeavoured to sketch the probable ancestral trees of the various classes of the animal world, the protists, and the plants, as it seemed necessary to do on Darwinian principles, and as we can actually do now with a high degree of confidence. If the theory of descent, which Lamarck first clearly formulated and Darwin thoroughly established, is true, we should be able to draw up a natural classification of plants and animals in the light of their genealogy, and to conceive the large and small divisions of the system as the branches and twigs of an ancestral tree. The eight genealogical tables which I inserted in the second volume of the General Morphology are the first sketches of their kind. In Chapter 2.27, particularly, I trace the chief stages in man's ancestry, as far as it is possible to follow it through the vertebrate stem. I tried especially to determine, as well as one could at that time, the position of man in the classification of the mammals and its genealogical significance. I have greatly improved this attempt, and treated it in a more popular form, in chapters 26 to 28 of my History of Creation (1868).* (* Of which Darwin said that the Descent of Man would probably never have been written if he had seen it earlier.—Translator.)
It was not until 1871, twelve years after the appearance of The Origin of Species, that Darwin published the famous work which made the much-contested application of his theory to man, and crowned the splendid structure of his system. This important work was The Descent of Man, and Selection in Relation to Sex. In this Darwin expressly drew the conclusion, with rigorous logic, that man also must have been developed out of lower species, and described the important part played by sexual selection in the elevation of man and the other higher animals. He showed that the careful selection which the sexes exercise on each other in regard to sexual relations and procreation, and the aesthetic feeling which the higher animals develop through this, are of the utmost importance in the progressive development of forms and the differentiation of the sexes. The males choosing the handsomest females in one class of animals, and the females choosing only the finest-looking males in another, the special features and the sexual characteristics are increasingly accentuated. In fact, some of the higher animals develop in this connection a finer taste and judgment than man himself. But, even as regards man, it is to this sexual selection that we owe the family-life, which is the chief foundation of civilisation. The rise of the human race is due for the most part to the advanced sexual selection which our ancestors exercised in choosing their mates.
Darwin accepted in the main the general outlines of man's ancestral tree, as I gave it in the General Morphology and the History of Creation, and admitted that his studies led him to the same conclusion. That he did not at once apply the theory to man in his first work was a commendable piece of discretion; such a sequel was bound to excite the strongest opposition to the whole theory. The first thing to do was to establish it as regards the animal and plant worlds. The subsequent extension to man was bound to be made sooner or later.
It is important to understand this very clearly. If all living things come from a common root, man must be included in the general scheme of evolution. On the other hand, if the various species were separately created, man, too, must have been created, and not evolved. We have to choose between these two alternatives. This cannot be too frequently or too strongly emphasised. EITHER all the species of animals and plants are of supernatural origin—created, not evolved—and in that case man also is the outcome of a creative act, as religion teaches, OR the different species have been evolved from a few common, simple ancestral forms, and in that case man is the highest fruit of the tree of evolution.
We may state this briefly in the following principle—The descent of man from the lower animals is a special deduction which inevitably follows from the general inductive law of the whole theory of evolution. In this principle we have a clear and plain statement of the matter. Evolution is in reality nothing but a great induction, which we are compelled to make by the comparative study of the most important facts of morphology and physiology. But we must draw our conclusion according to the laws of induction, and not attempt to determine scientific truths by direct measurement and mathematical calculation. In the study of living things we can scarcely ever directly and fully, and with mathematical accuracy, determine the nature of phenomena, as is done in the simpler study of the inorganic world—in chemistry, physics, mineralogy, and astronomy. In the latter, especially, we can always use the simplest and absolutely safest method—that of mathematical determination. But in biology this is quite impossible for various reasons; one very obvious reason being that most of the facts of the science are very complicated and much too intricate to allow a direct mathematical analysis. The greater part of the phenomena that biology deals with are complicated HISTORICAL PROCESSES, which are related to a far-reaching past, and as a rule can only be approximately estimated. Hence we have to proceed by INDUCTION—that is to say, to draw general conclusions, stage by stage, and with proportionate confidence, from the accumulation of detailed observations. These inductive conclusions cannot command absolute confidence, like mathematical axioms; but they approach the truth, and gain increasing probability, in proportion as we extend the basis of observed facts on which we build. The importance of these inductive laws is not diminished from the circumstance that they are looked upon merely as temporary acquisitions of science, and may be improved to any extent in the progress of scientific knowledge. The same may be said of the attainments of many other sciences, such as geology or archeology. However much they may be altered and improved in detail in the course of time, these inductive truths may retain their substance unchanged.
Now, when we say that the theory of evolution in the sense of Lamarck and Darwin is an inductive law—in fact, the greatest of all biological inductions—we rely, in the first place, on the facts of paleontology. This science gives us some direct acquaintance with the historical phenomena of the changes of species. From the situations in which we find the fossils in the various strata of the earth we gather confidently, in the first place, that the living population of the earth has been gradually developed, as clearly as the earth's crust itself; and that, in the second place, several different populations have succeeded each other in the various geological periods. Modern geology teaches that the formation of the earth has been gradual, and unbroken by any violent revolutions. And when we compare together the various kinds of animals and plants which succeed each other in the history of our planet, we find, in the first place, a constant and gradual increase in the number of species from the earliest times until the present day; and, in the second place, we notice that the forms in each great group of animals and plants also constantly improve as the ages advance. Thus, of the vertebrates there are at first only the lower fishes; then come the higher fishes, and later the amphibia. Still later appear the three higher classes of vertebrates—the reptiles, birds, and mammals, for the first time; only the lowest and least perfect forms of the mammals are found at first; and it is only at a very late period that placental mammals appear, and man belongs to the latest and youngest branch of these. Thus perfection of form increases as well as variety from the earliest to the latest stage. That is a fact of the greatest importance. It can only be explained by the theory of evolution, with which it is in perfect harmony. If the different groups of plants and animals do really descend from each other, we must expect to find this increase in their number and perfection under the influence of natural selection, just as the succession of fossils actually discloses it to us.
Comparative anatomy furnishes a second series of facts which are of great importance for the forming of our inductive law. This branch of morphology compares the adult structures of living things, and seeks in the great variety of organic forms the stable and simple law of organisation, or the common type or structure. Since Cuvier founded this science at the beginning of the nineteenth century it has been a favourite study of the most distinguished scientists. Even before Cuvier's time Goethe had been greatly stimulated by it, and induced to take up the study of morphology. Comparative osteology, or the philosophic study and comparison of the bony skeleton of the vertebrates—one of its most interesting sections—especially fascinated him, and led him to form the theory of the skull which I mentioned before. Comparative anatomy shows that the internal structure of the animals of each stem and the plants of each class is the same in its essential features, however much they differ in external appearance. Thus man has so great a resemblance in the chief features of his internal organisation to the other mammals that no comparative anatomist has ever doubted that he belongs to this class. The whole internal structure of the human body, the arrangement of its various systems of organs, the distribution of the bones, muscles, blood-vessels, etc., and the whole structure of these organs in the larger and the finer scale, agree so closely with those of the other mammals (such as the apes, rodents, ungulates, cetacea, marsupials, etc.) that their external differences are of no account whatever. We learn further from comparative anatomy that the chief features of animal structure are so similar in the various classes (fifty to sixty in number altogether) that they may all be comprised in from eight to twelve great groups. But even in these groups, the stem-forms or animal types, certain organs (especially the alimentary canal) can be proved to have been originally the same for all. We can only explain by the theory of evolution this essential unity in internal structure of all these animal forms that differ so much in outward appearance. This wonderful fact can only be really understood and explained when we regard the internal resemblance as an inheritance from common-stem forms, and the external differences as the effect of adaptation to different environments.
In recognising this, comparative anatomy has itself advanced to a higher stage. Gegenbaur, the most distinguished of recent students of this science, says that with the theory of evolution a new period began in comparative anatomy, and that the theory in turn found a touch stone in the science. "Up to now there is no fact in comparative anatomy that is inconsistent with the theory of evolution; indeed, they all lead to it. In this way the theory receives back from the science all the service it rendered to its method." Until then students had marvelled at the wonderful resemblance of living things in their inner structure without being able to explain it. We are now in a position to explain the causes of this, by showing that this remarkable agreement is the necessary consequence of the inheriting of common stem-forms; while the striking difference in outward appearance is a result of adaptation to changes of environment. Heredity and adaptation alone furnish the true explanation.
But one special part of comparative anatomy is of supreme interest and of the utmost philosophic importance in this connection. This is the science of rudimentary or useless organs; I have given it the name of "dysteleology" in view of its philosophic consequences. Nearly every organism (apart from the very lowest), and especially every highly-developed animal or plant, including man, has one or more organs which are of no use to the body itself, and have no share in its functions or vital aims. Thus we all have, in various parts of our frame, muscles which we never use, as, for instance, in the shell of the ear and adjoining parts. In most of the mammals, especially those with pointed ears, these internal and external ear-muscles are of great service in altering the shell of the ear, so as to catch the waves of sound as much as possible. But in the case of man and other short-eared mammals these muscles are useless, though they are still present. Our ancestors having long abandoned the use of them, we cannot work them at all to-day. In the inner corner of the eye we have a small crescent-shaped fold of skin; this is the last relic of a third inner eye-lid, called the nictitating (winking) membrane. This membrane is highly developed and of great service in some of our distant relations, such as fishes of the shark type and several other vertebrates; in us it is shrunken and useless. In the intestines we have a process that is not only quite useless, but may be very harmful—the vermiform appendage. This small intestinal appendage is often the cause of a fatal illness. If a cherry-stone or other hard body is unfortunately squeezed through its narrow aperture during digestion, a violent inflammation is set up, and often proves fatal. This appendix has no use whatever now in our frame; it is a dangerous relic of an organ that was much larger and was of great service in our vegetarian ancestors. It is still large and important in many vegetarian animals, such as apes and rodents.
There are similar rudimentary organs in all parts of our body, and in all the higher animals. They are among the most interesting phenomena to which comparative anatomy introduces us; partly because they furnish one of the clearest proofs of evolution, and partly because they most strikingly refute the teleology of certain philosophers. The theory of evolution enables us to give a very simple explanation of these phenomena.
We have to look on them as organs which have fallen into disuse in the course of many generations. With the decrease in the use of its function, the organ itself shrivels up gradually, and finally disappears. There is no other way of explaining rudimentary organs. Hence they are also of great interest in philosophy; they show clearly that the monistic or mechanical view of the organism is the only correct one, and that the dualistic or teleological conception is wrong. The ancient legend of the direct creation of man according to a pre-conceived plan and the empty phrases about "design" in the organism are completely shattered by them. It would be difficult to conceive a more thorough refutation of teleology than is furnished by the fact that all the higher animals have these rudimentary organs.
The theory of evolution finds its broadest inductive foundation in the natural classification of living things, which arranges all the various forms in larger and smaller groups, according to their degree of affinity. These groupings or categories of classification—the varieties, species, genera, families, orders, classes, etc.—show such constant features of coordination and subordination that we are bound to look on them as genealogical, and represent the whole system in the form of a branching tree. This is the genealogical tree of the variously related groups; their likeness in form is the expression of a real affinity. As it is impossible to explain in any other way the natural tree-like form of the system of organisms, we must regard it at once as a weighty proof of the truth of evolution. The careful construction of these genealogical trees is, therefore, not an amusement, but the chief task of modern classification.
Among the chief phenomena that bear witness to the inductive law of evolution we have the geographical distribution of the various species of animals and plants over the surface of the earth, and their topographical distribution on the summits of mountains and in the depths of the ocean. The scientific study of these features—the "science of distribution," or chorology (chora = a place)—has been pursued with lively interest since the discoveries made by Alexander von Humboldt. Until Darwin's time the work was confined to the determination of the facts of the science, and chiefly aimed at settling the spheres of distribution of the existing large and small groups of living things. It was impossible at that time to explain the causes of this remarkable distribution, or the reasons why one group is found only in one locality and another in a different place, and why there is this manifold distribution at all. Here, again, the theory of evolution has given us the solution of the problem. It furnishes the only possible explanation when it teaches that the various species and groups of species descend from common stem-forms, whose ever-branching offspring have gradually spread themselves by migration over the earth. For each group of species we must admit a "centre of production," or common home; this is the original habitat in which the ancestral form was developed, and from which its descendants spread out in every direction. Several of these descendants became in their turn the stem-forms for new groups of species, and these also scattered themselves by active and passive migration, and so on. As each migrating organism found a different environment in its new home, and adapted itself to it, it was modified, and gave rise to new forms.
This very important branch of science that deals with active and passive migration was founded by Darwin, with the aid of the theory of evolution; and at the same time he advanced the true explanation of the remarkable relation or similarity of the living population in any locality to the fossil forms found in it. Moritz Wagner very ably developed his idea under the title of "the theory of migration." In my opinion, this famous traveller has rather over-estimated the value of his theory of migration when he takes it to be an indispensable condition of the formation of new species and opposes the theory of selection. The two theories are not opposed in their main features. Migration (by which the stem-form of a new species is isolated) is really only a special case of selection. The striking and interesting facts of chorology can be explained only by the theory of evolution, and therefore we must count them among the most important of its inductive bases.
The same must be said of all the remarkable phenomena which we perceive in the economy of the living organism. The many and various relations of plants and animals to each other and to their environment, which are treated in bionomy (from nomos, law or norm, and bios, life), the interesting facts of parasitism, domesticity, care of the young, social habits, etc., can only be explained by the action of heredity and adaptation. Formerly people saw only the guidance of a beneficent Providence in these phenomena; to-day we discover in them admirable proofs of the theory of evolution. It is impossible to understand them except in the light of this theory and the struggle for life.
Finally, we must, in my opinion, count among the chief inductive bases of the theory of evolution the foetal development of the individual organism, the whole science of embryology or ontogeny. But as the later chapters will deal with this in detail, I need say nothing further here. I shall endeavour in the following pages to show, step by step, how the whole of the embryonic phenomena form a massive chain of proof for the theory of evolution; for they can be explained in no other way. In thus appealing to the close causal connection between ontogenesis and phylogenesis, and taking our stand throughout on the biogenetic law, we shall be able to prove, stage by stage, from the facts of embryology, the evolution of man from the lower animals.
The general adoption of the theory of evolution has definitely closed the controversy as to the nature or definition of the species. The word has no ABSOLUTE meaning whatever, but is only a group-name, or category of classification, with a purely relative value. In 1857, it is true, a famous and gifted, but inaccurate and dogmatic, scientist, Louis Agassiz, attempted to give an absolute value to these "categories of classification." He did this in his Essay on Classification, in which he turns upside down the phenomena of organic nature, and, instead of tracing them to their natural causes, examines them through a theological prism. The true species (bona species) was, he said, an "incarnate idea of the Creator." Unfortunately, this pretty phrase has no more scientific value than all the other attempts to save the absolute or intrinsic value of the species.
The dogma of the fixity and creation of species lost its last great champion when Agassiz died in 1873. The opposite theory, that all the different species descend from common stem-forms, encounters no serious difficulty to-day. All the endless research into the nature of the species, and the possibility of several species descending from a common ancestor, has been closed to-day by the removal of the sharp limits that had been set up between species and varieties on the one hand, and species and genera on the other. I gave an analytic proof of this in my monograph on the sponges (1872), having made a very close study of variability in this small but highly instructive group, and shown the impossibility of making any dogmatic distinction of species. According as the classifier takes his ideas of genus, species, and variety in a broader or in a narrower sense, he will find in the small group of the sponges either one genus with three species, or three genera with 238 species, or 113 genera with 591 species. Moreover, all these forms are so connected by intermediate forms that we can convincingly prove the descent of all the sponges from a common stem-form, the olynthus.
Here, I think, I have given an analytic solution of the problem of the origin of species, and so met the demand of certain opponents of evolution for an actual instance of descent from a stem-form. Those who are not satisfied with the synthetic proofs of the theory of evolution which are provided by comparative anatomy, embryology, paleontology, dysteleology, chorology, and classification, may try to refute the analytic proof given in my treatise on the sponge, the outcome of five years of assiduous study. I repeat: It is now impossible to oppose evolution on the ground that we have no convincing example of the descent of all the species of a group from a common ancestor. The monograph on the sponges furnishes such a proof, and, in my opinion, an indisputable proof. Any man of science who will follow the protracted steps of my inquiry and test my assertions will find that in the case of the sponges we can follow the actual evolution of species in a concrete case. And if this is so, if we can show the origin of all the species from a common form in one single class, we have the solution of the problem of man's origin, because we are in a position to prove clearly his descent from the lower animals.
At the same time, we can now reply to the often-repeated assertion, even heard from scientists of our own day, that the descent of man from the lower animals, and proximately from the apes, still needs to be "proved with certainty." These "certain proofs" have been available for a long time; one has only to open one's eyes to see them. It is a mistake to seek them in the discovery of intermediate forms between man and the ape, or the conversion of an ape into a human being by skilful education. The proofs lie in the great mass of empirical material we have already collected. They are furnished in the strongest form by the data of comparative anatomy and embryology, completed by paleontology. It is not a question now of detecting new proofs of the evolution of man, but of examining and understanding the proofs we already have.
I was almost alone thirty-six years ago when I made the first attempt, in my General Morphology, to put organic science on a mechanical foundation through Darwin's theory of descent. The association of ontogeny and phylogeny and the proof of the intimate causal connection between these two sections of the science of evolution, which I expounded in my work, met with the most spirited opposition on nearly all sides. The next ten years were a terrible "struggle for life" for the new theory. But for the last twenty-five years the tables have been turned. The phylogenetic method has met with so general a reception, and found so prolific a use in every branch of biology, that it seems superfluous to treat any further here of its validity and results. The proof of it lies in the whole morphological literature of the last three decades. But no other science has been so profoundly modified in its leading thoughts by this adoption, and been forced to yield such far-reaching consequences, as that science which I am now seeking to establish—monistic anthropogeny.
This statement may seem to be rather audacious, since the very next branch of biology, anthropology in the stricter sense, makes very little use of these results of anthropogeny, and sometimes expressly opposes them.* (*This does not apply to English anthropologists, who are almost all evolutionists.) This applies especially to the attitude which has characterised the German Anthropological Society (the Deutsche Gesellschaft fur Anthropologie) for some thirty years. Its powerful president, the famous pathologist, Rudolph Virchow, is chiefly responsible for this. Until his death (September 5th, 1902) he never ceased to reject the theory of descent as unproven, and to ridicule its chief consequence—the descent of man from a series of mammal ancestors—as a fantastic dream. I need only recall his well-known expression at the Anthropological Congress at Vienna in 1894, that "it would be just as well to say man came from the sheep or the elephant as from the ape."
Virchow's assistant, the secretary of the German Anthropological Society, Professor Johannes Ranke of Munich, has also indefatigably opposed transformism: he has succeeded in writing a work in two volumes (Der Mensch), in which all the facts relating to his organisation are explained in a sense hostile to evolution. This work has had a wide circulation, owing to its admirable illustrations and its able treatment of the most interesting facts of anatomy and physiology—exclusive of the sexual organs! But, as it has done a great deal to spread erroneous views among the general public, I have included a criticism of it in my History of Creation, as well as met Virchow's attacks on anthropogeny.
Neither Virchow, nor Ranke, nor any other "exact" anthropologist, has attempted to give any other natural explanation of the origin of man. They have either set completely aside this "question of questions" as a transcendental problem, or they have appealed to religion for its solution. We have to show that this rejection of the rational explanation is totally without justification. The fund of knowledge which has accumulated in the progress of biology in the nineteenth century is quite adequate to furnish a rational explanation, and to establish the theory of the evolution of man on the solid facts of his embryology.
CHAPTER 1.6. THE OVUM AND THE AMOEBA.
In order to understand clearly the course of human embryology, we must select the more important of its wonderful and manifold processes for fuller explanation, and then proceed from these to the innumerable features of less importance. The most important feature in this sense, and the best starting-point for ontogenetic study, is the fact that man is developed from an ovum, and that this ovum is a simple cell. The human ovum does not materially differ in form and composition from that of the other mammals, whereas there is a distinct difference between the fertilised ovum of the mammal and that of any other animal.
(FIGURE 1.1. The human ovum, magnified 100 times. The globular mass of yelk (b) is enclosed by a transparent membrane (the ovolemma or zona pellucida [a]), and contains a noncentral nucleus (the germinal vesicle, c). Cf. Figure 1.14.)
This fact is so important that few should be unaware of its extreme significance; yet it was quite unknown in the first quarter of the nineteenth century. As we have seen, the human and mammal ovum was not discovered until 1827, when Carl Ernst von Baer detected it. Up to that time the larger vesicles, in which the real and much smaller ovum is contained, had been wrongly regarded as ova. The important circumstance that this mammal ovum is a simple cell, like the ovum of other animals, could not, of course, be recognised until the cell theory was established. This was not done, by Schleiden for the plant and Schwann for the animal, until 1838. As we have seen, this cell theory is of the greatest service in explaining the human frame and its embryonic development. Hence we must say a few words about the actual condition of the theory and the significance of the views it has suggested.
In order properly to appreciate the cellular theory, the most important element in our science, it is necessary to understand in the first place that the cell is a UNIFIED ORGANISM, a self-contained living being. When we anatomically dissect the fully-formed animal or plant into its various organs, and then examine the finer structure of these organs with the microscope, we are surprised to find that all these different parts are ultimately made up of the same structural element or unit. This common unit of structure is the cell. It does not matter whether we thus dissect a leaf, flower, or fruit, or a bone, muscle, gland, or bit of skin, etc.; we find in every case the same ultimate constituent, which has been called the cell since Schleiden's discovery. There are many opinions as to its real nature, but the essential point in our view of the cell is to look upon it as a self-contained or independent living unit. It is, in the words of Brucke, "an elementary organism." We may define it most precisely as the ultimate organic unit, and, as the cells are the sole active principles in every vital function, we may call them the "plastids," or "formative elements." This unity is found in both the anatomic structure and the physiological function. In the case of the protists, the entire organism usually consists of a single independent cell throughout life. But in the tissue-forming animals and plants, which are the great majority, the organism begins its career as a simple cell, and then grows into a cell-community, or, more correctly, an organised cell-state. Our own body is not really the simple unity that it is generally supposed to be. On the contrary, it is a very elaborate social system of countless microscopic organisms, a colony or commonwealth, made up of innumerable independent units, or very different tissue-cells.
In reality, the term "cell," which existed long before the cell theory was formulated, is not happily chosen. Schleiden, who first brought it into scientific use in the sense of the cell theory, gave this name to the elementary organisms because, when you find them in the dissected plant, they generally have the appearance of chambers, like the cells in a bee-hive, with firm walls and a fluid or pulpy content. But some cells, especially young ones, are entirely without the enveloping membrane, or stiff wall. Hence we now generally describe the cell as a living, viscous particle of protoplasm, enclosing a firmer nucleus in its albuminoid body. There may be an enclosing membrane, as there actually is in the case of most of the plants; but it may be wholly lacking, as is the case with most of the animals. There is no membrane at all in the first stage. The young cells are usually round, but they vary much in shape later on. Illustrations of this will be found in the cells of the various parts of the body shown in Figures 1.3 to 1.7.
Hence the essential point in the modern idea of the cell is that it is made up of two different active constituents—an inner and an outer part. The smaller and inner part is the nucleus (or caryon or cytoblastus, Figure 1.1 c and Figure 1.2 k). The outer and larger part, which encloses the other, is the body of the cell (celleus, cytos, or cytosoma). The soft living substance of which the two are composed has a peculiar chemical composition, and belongs to the group of the albuminoid plasma-substances ("formative matter"), or protoplasm. The essential and indispensable element of the nucleus is called nuclein (or caryoplasm); that of the cell body is called plastin (or cytoplasm). In the most rudimentary cases both substances seem to be quite simple and homogeneous, without any visible structure. But, as a rule, when we examine them under a high power of the microscope, we find a certain structure in the protoplasm. The chief and most common form of this is the fibrous or net-like "thready structure" (Frommann) and the frothy "honeycomb structure" (Butschli).
(FIGURE 1.2. Stem-cell of one of the echinoderms (cytula, or "first segmentation-cell" = fertilised ovum), after Hertwig. k is the nucleus or caryon.)
The shape or outer form of the cell is infinitely varied, in accordance with its endless power of adapting itself to the most diverse activities or environments. In its simplest form the cell is globular (Figure 1.2). This normal round form is especially found in cells of the simplest construction, and those that are developed in a free fluid without any external pressure. In such cases the nucleus also is not infrequently round, and located in the centre of the cell-body (Figure 1.2 k). In other cases, the cells have no definite shape; they are constantly changing their form owing to their automatic movements. This is the case with the amoebae (Figures 1.15 and 1.16) and the amoeboid travelling cells (Figure 1.11), and also with very young ova (Figure 1.13). However, as a rule, the cell assumes a definite form in the course of its career. In the tissues of the multicellular organism, in which a number of similar cells are bound together in virtue of certain laws of heredity, the shape is determined partly by the form of their connection and partly by their special functions. Thus, for instance, we find in the mucous lining of our tongue very thin and delicate flat cells of roundish shape (Figure 1.3). In the outer skin we find similar, but harder, covering cells, joined together by saw-like edges (Figure 1.4). In the liver and other glands there are thicker and softer cells, linked together in rows (Figure 1.5).
The last-named tissues (Figures 1.3 to 1.5) belong to the simplest and most primitive type, the group of the "covering-tissues," or epithelia. In these "primary tissues" (to which the germinal layers belong) simple cells of the same kind are arranged in layers. The arrangement and shape are more complicated in the "secondary tissues," which are gradually developed out of the primary, as in the tissues of the muscles, nerves, bones, etc. In the bones, for instance, which belong to the group of supporting or connecting organs, the cells (Figure 1.6) are star-shaped, and are joined together by numbers of net-like interlacing processes; so, also, in the tissues of the teeth (Figure 1.7), and in other forms of supporting-tissue, in which a soft or hard substance (intercellular matter, or base) is inserted between the cells.
(FIGURE 1.3. Three epithelial cells from the mucous lining of the tongue.
FIGURE 1.4. Five spiny or grooved cells, with edges joined, from the outer skin (epidermis): one of them (b) is isolated.
FIGURE 1.5. Ten liver-cells: one of them (b) has two nuclei.)
The cells also differ very much in size. The great majority of them are invisible to the naked eye, and can be seen only through the microscope (being as a rule between 1/2500 and 1/250 inch in diameter). There are many of the smaller plastids—such as the famous bacteria—which only come into view with a very high magnifying power. On the other hand, many cells attain a considerable size, and run occasionally to several inches in diameter, as do certain kinds of rhizopods among the unicellular protists (such as the radiolaria and thalamophora). Among the tissue-cells of the animal body many of the muscular fibres and nerve fibres are more than four inches, and sometimes more than a yard, in length. Among the largest cells are the yelk-filled ova; as, for instance, the yellow "yolk" in the hen's egg, which we shall describe later (Figure 1.15).
Cells also vary considerably in structure. In this connection we must first distinguish between the active and passive components of the cell. It is only the former, or active parts of the cell, that really live, and effect that marvellous world of phenomena to which we give the name of "organic life." The first of these is the inner nucleus (caryoplasm), and the second the body of the cell (cytoplasm). The passive portions come third; these are subsequently formed from the others, and I have given them the name of "plasma-products." They are partly external (cell-membranes and intercellular matter) and partly internal (cell-sap and cell-contents).
The nucleus (or caryon), which is usually of a simple roundish form, is quite structureless at first (especially in very young cells), and composed of homogeneous nuclear matter or caryoplasm (Figure 1.2 k). But, as a rule, it forms a sort of vesicle later on, in which we can distinguish a more solid nuclear base (caryobasis) and a softer or fluid nuclear sap (caryolymph). In a mesh of the nuclear network (or it may be on the inner side of the nuclear envelope) there is, as a rule, a dark, very opaque, solid body, called the nucleolus. Many of the nuclei contain several of these nucleoli (as, for instance, the germinal vesicle of the ova of fishes and amphibia). Recently a very small, but particularly important, part of the nucleus has been distinguished as the central body (centrosoma)—a tiny particle that is originally found in the nucleus itself, but is usually outside it, in the cytoplasm; as a rule, fine threads stream out from it in the cytoplasm. From the position of the central body with regard to the other parts it seems probable that it has a high physiological importance as a centre of movement; but it is lacking in many cells.
The cell-body also consists originally, and in its simplest form, of a homogeneous viscid plasmic matter. But, as a rule, only the smaller part of it is formed of the living active cell-substance (protoplasm); the greater part consists of dead, passive plasma-products (metaplasm). It is useful to distinguish between the inner and outer of these. External plasma-products (which are thrust out from the protoplasm as solid "structural matter") are the cell-membranes and the intercellular matter. The internal plasma-products are either the fluid cell-sap or hard structures. As a rule, in mature and differentiated cells these various parts are so arranged that the protoplasm (like the caryoplasm in the round nucleus) forms a sort of skeleton or framework. The spaces of this network are filled partly with the fluid cell-sap and partly by hard structural products.
(FIGURE 1.6. Nine star-shaped bone-cells, with interlaced branches.
FIGURE 1.7. Eleven star-shaped cells from the enamel of a tooth, joined together by their branchlets.)
The simple round ovum, which we take as the starting-point of our study (Figures 1.1 and 1.2), has in many cases the vague, indifferent features of the typical primitive cell. As a contrast to it, and as an instance of a very highly differentiated plastid, we may consider for a moment a large nerve-cell, or ganglionic cell, from the brain. The ovum stands potentially for the entire organism—in other words, it has the faculty of building up out of itself the whole multicellular body. It is the common parent of all the countless generations of cells which form the different tissues of the body; it unites all their powers in itself, though only potentially or in germ. In complete contrast to this, the neural cell in the brain (Figure 1.9) develops along one rigid line. It cannot, like the ovum, beget endless generations of cells, of which some will become skin-cells, others muscle-cells, and others again bone-cells. But, on the other hand, the nerve-cell has become fitted to discharge the highest functions of life; it has the powers of sensation, will, and thought. It is a real soul-cell, or an elementary organ of the psychic activity. It has, therefore, a most elaborate and delicate structure. Numbers of extremely fine threads, like the electric wires at a large telegraphic centre, cross and recross in the delicate protoplasm of the nerve cell, and pass out in the branching processes which proceed from it and put it in communication with other nerve-cells or nerve-fibres (a, b). We can only partly follow their intricate paths in the fine matter of the body of the cell.
Here we have a most elaborate apparatus, the delicate structure of which we are just beginning to appreciate through our most powerful microscopes, but whose significance is rather a matter of conjecture than knowledge. Its intricate structure corresponds to the very complicated functions of the mind. Nevertheless, this elementary organ of psychic activity—of which there are thousands in our brain—is nothing but a single cell. Our whole mental life is only the joint result of the combined activity of all these nerve-cells, or soul-cells. In the centre of each cell there is a large transparent nucleus, containing a small and dark nuclear body. Here, as elsewhere, it is the nucleus that determines the individuality of the cell; it proves that the whole structure, in spite of its intricate composition, amounts to only a single cell.
(FIGURE 1.8. Unfertilised ovum of an echinoderm (from Hertwig). The vesicular nucleus (or "germinal vesicle") is globular, half the size of the round ovum, and encloses a nuclear framework, in the central knot of which there is a dark nucleolus (the "germinal spot").
FIGURE 1.9. A large branching nerve-cell, or "soul-cell," from the brain of an electric fish (Torpedo), magnified 600 times. In the middle of the cell is the large transparent round nucleus, one nucleolus, and, within the latter again, a nucleolinus. The protoplasm of the cell is split into innumerable fine threads (or fibrils), which are embedded in intercellular matter, and are prolonged into the branching processes of the cell (b). One branch (a) passes into a nerve-fibre. (From Max Schultze.))
In contrast with this very elaborate and very strictly differentiated psychic cell (Figure 1.9), we have our ovum (Figures 1.1 and 1.2), which has hardly any structure at all. But even in the case of the ovum we must infer from its properties that its protoplasmic body has a very complicated chemical composition and a fine molecular structure which escapes our observation. This presumed molecular structure of the plasm is now generally admitted; but it has never been seen, and, indeed, lies far beyond the range of microscopic vision. It must not be confused—as is often done—with the structure of the plasm (the fibrous network, groups of granules, honey-comb, etc.) which does come within the range of the microscope.
But when we speak of the cells as the elementary organisms, or structural units, or "ultimate individualities," we must bear in mind a certain restriction of the phrases. I mean, that the cells are not, as is often supposed, the very lowest stage of organic individuality. There are yet more elementary organisms to which I must refer occasionally. These are what we call the "cytodes" (cytos = cell), certain living, independent beings, consisting only of a particle of plasson—an albuminoid substance, which is not yet differentiated into caryoplasm and cytoplasm, but combines the properties of both. Those remarkable beings called the monera—especially the chromacea and bacteria—are specimens of these simple cytodes. (Compare Chapter 2.19.) To be quite accurate, then, we must say: the elementary organism, or the ultimate individual, is found in two different stages. The first and lower stage is the cytode, which consists merely of a particle of plasson, or quite simple plasm. The second and higher stage is the cell, which is already divided or differentiated into nuclear matter and cellular matter. We comprise both kinds—the cytodes and the cells—under the name of plastids ("formative particles"), because they are the real builders of the organism. However, these cytodes are not found, as a rule, in the higher animals and plants; here we have only real cells with a nucleus. Hence, in these tissue-forming organisms (both plant and animal) the organic unit always consists of two chemically and anatomically different parts—the outer cell-body and the inner nucleus.
In order to convince oneself that this cell is really an independent organism, we have only to observe the development and vital phenomena of one of them. We see then that it performs all the essential functions of life—both vegetal and animal—which we find in the entire organism. Each of these tiny beings grows and nourishes itself independently. It takes its food from the surrounding fluid; sometimes, even, the naked cells take in solid particles at certain points of their surface—in other words, "eat" them—without needing any special mouth and stomach for the purpose (cf. Figure 1.19).
Further, each cell is able to reproduce itself. This multiplication, in most cases, takes the form of a simple cleavage, sometimes direct, sometimes indirect; the simple direct (or "amitotic") division is less common, and is found, for instance, in the blood cells (Figure 1.10). In these the nucleus first divides into two equal parts by constriction. The indirect (or "mitotic") cleavage is much more frequent; in this the caryoplasm of the nucleus and the cytoplasm of the cell-body act upon each other in a peculiar way, with a partial dissolution (caryolysis), the formation of knots and loops (mitosis), and a movement of the halved plasma-particles towards two mutually repulsive poles of attraction (caryokinesis, Figure 1.11.)
(FIGURE 1.10. Blood-cells, multiplying by direct division, from the blood of the embryo of a stag. Originally, each blood-cell has a nucleus and is round (a). When it is going to multiply, the nucleus divides into two (b, c, d). Then the protoplasmic body is constricted between the two nuclei, and these move away from each other (e). Finally, the constriction is complete, and the cell splits into two daughter-cells (f). (From Frey.))
FIGURE 1.11. Indirect or mitotic cell-division (with caryolysis and caryokinesis) from the skin of the larva of a salamander. (From Rabl.). A. Mother-cell (Knot, spirema), with Nuclear threads (chromosomata) (coloured nuclear matter, chromatin), Cytosoma, Nuclear membrane, Protoplasm of the cell-body and Nuclear sap. B. Mother-star, the loops beginning to split lengthways (nuclear membrane gone), with Star-like appearance in cytoplasm, Centrosoma (sphere of attraction), Nuclear spindle (achromin, colourless matter) and Nuclear loops (chromatin, coloured matter). C. The two daughter-stars, produced by the breaking of the loops of the mother-star (moving away), with Upper daughter-crown, Connecting threads of the two crowns (achromin), Lower daughter-crown and Double-star (amphiaster). D. The two daughter-cells, produced by the complete division of the two nuclear halves (cytosomata still connected at the equator) (Double-knot, Dispirema), with Upper daughter-nucleus, Equatorial constriction of the cell-body and Lower daughter-nucleus.)
The intricate physiological processes which accompany this "mitosis" have been very closely studied of late years. The inquiry has led to the detection of certain laws of evolution which are of extreme importance in connection with heredity. As a rule, two very different parts of the nucleus play an important part in these changes. They are: the chromatin, or coloured nuclear substance, which has a peculiar property of tingeing itself deeply with certain colouring matters (carmine, haematoxylin, etc.), and the achromin (or linin, or achromatin), a colourless nuclear substance that lacks this property. The latter generally forms in the dividing cell a sort of spindle, at the poles of which there is a very small particle, also colourless, called the "central body" (centrosoma). This acts as the centre or focus in a "sphere of attraction" for the granules of protoplasm in the surrounding cell-body, and assumes a star-like appearance (the cell-star, or monaster). The two central bodies, standing opposed to each other at the poles of the nuclear spindle, form "the double-star" (or amphiaster, Figure 1.11, BC). The chromatin often forms a long, irregularly-wound thread—"the coil" (spirema, Figure A). At the commencement of the cleavage it gathers at the equator of the cell, between the stellar poles, and forms a crown of U-shaped loops (generally four or eight, or some other definite number). The loops split lengthwise into two halves (B), and these back away from each other towards the poles of the spindle (C). Here each group forms a crown once more, and this, with the corresponding half of the divided spindle, forms a fresh nucleus (D). Then the protoplasm of the cell-body begins to contract in the middle, and gather about the new daughter-nuclei, and at last the two daughter-cells become independent beings.
Between this common mitosis, or indirect cell-division—which is the normal cleavage-process in most cells of the higher animals and plants—and the simple direct division (Figure 1.10) we find every grade of segmentation; in some circumstances even one kind of division may be converted into another.
The plastid is also endowed with the functions of movement and sensation. The single cell can move and creep about, when it has space for free movement and is not prevented by a hard envelope; it then thrusts out at its surface processes like fingers, and quickly withdraws them again, and thus changes its shape (Figure 1.12). Finally, the young cell is sensitive, or more or less responsive to stimuli; it makes certain movements on the application of chemical and mechanical irritation. Hence we can ascribe to the individual cell all the chief functions which we comprehend under the general heading of "life"—sensation, movement, nutrition, and reproduction. All these properties of the multicellular and highly developed animal are also found in the single animal-cell, at least in its younger stages. There is no longer any doubt about this, and so we may regard it as a solid and important base of our physiological conception of the elementary organism.
Without going any further here into these very interesting phenomena of the life of the cell, we will pass on to consider the application of the cell theory to the ovum. Here comparative research yields the important result that EVERY OVUM IS AT FIRST A SIMPLE CELL. I say this is very important, because our whole science of embryology now resolves itself into the problem: "How does the multicellular organism arise from the unicellular?" Every organic individual is at first a simple cell, and as such an elementary organism, or a unit of individuality. This cell produces a cluster of cells by segmentation, and from these develops the multicellular organism, or individual of higher rank.
When we examine a little closer the original features of the ovum, we notice the extremely significant fact that in its first stage the ovum is just the same simple and indefinite structure in the case of man and all the animals (Figure 1.13). We are unable to detect any material difference between them, either in outer shape or internal constitution. Later, though the ova remain unicellular, they differ in size and shape, enclose various kinds of yelk-particles, have different envelopes, and so on. But when we examine them at their birth, in the ovary of the female animal, we find them to be always of the same form in the first stages of their life. In the beginning each ovum is a very simple, roundish, naked, mobile cell, without a membrane; it consists merely of a particle of cytoplasm enclosing a nucleus (Figure 1.13). Special names have been given to these parts of the ovum; the cell-body is called the yelk (vitellus), and the cell-nucleus the germinal vesicle. As a rule, the nucleus of the ovum is soft, and looks like a small pimple or vesicle. Inside it, as in many other cells, there is a nuclear skeleton or frame and a third, hard nuclear body (the nucleolus). In the ovum this is called the germinal spot. Finally, we find in many ova (but not in all) a still further point within the germinal spot, a "nucleolin," which goes by the name of the germinal point. The latter parts (germinal spot and germinal point) have, apparently, a minor importance, in comparison with the other two (the yelk and germinal vesicle). In the yelk we must distinguish the active formative yelk (or protoplasm = first plasm) from the passive nutritive yelk (or deutoplasm = second plasm).
(FIGURE 1.12. Mobile cells from the inflamed eye of a frog (from the watery fluid of the eye, the humor aqueus). The naked cells creep freely about, by (like the amoeba or rhizopods) protruding fine processes from the uncovered protoplasmic body. These bodies vary continually in number, shape, and size. The nucleus of these amoeboid lymph-cells ("travelling cells," or planocytes) is invisible, because concealed by the numbers of fine granules which are scattered in the protoplasm. (From Frey.))
In many of the lower animals (such as sponges, polyps, and medusae) the naked ova retain their original simple appearance until impregnation. But in most animals they at once begin to change; the change consists partly in the formation of connections with the yelk, which serve to nourish the ovum, and partly of external membranes for their protection (the ovolemma, or prochorion). A membrane of this sort is formed in all the mammals in the course of the embryonic process. The little globule is surrounded by a thick capsule of glass-like transparency, the zona pellucida, or ovolemma pellucidum (Figure 1.14). When we examine it closely under the microscope, we see very fine radial streaks in it, piercing the zona, which are really very narrow canals. The human ovum, whether fertilised or not, cannot be distinguished from that of most of the other mammals. It is nearly the same everywhere in form, size, and composition. When it is fully formed, it has a diameter of (on an average) about 1/120 of an inch. When the mammal ovum has been carefully isolated, and held against the light on a glass-plate, it may be seen as a fine point even with the naked eye. The ova of most of the higher mammals are about the same size. The diameter of the ovum is almost always between 1/250 to 1/125 inch. It has always the same globular shape; the same characteristic membrane; the same transparent germinal vesicle with its dark germinal spot. Even when we use the most powerful microscope with its highest power, we can detect no material difference between the ova of man, the ape, the dog, and so on. I do not mean to say that there are no differences between the ova of these different mammals. On the contrary, we are bound to assume that there are such, at least as regards chemical composition. Even the ova of different men must differ from each other; otherwise we should not have a different individual from each ovum. It is true that our crude and imperfect apparatus cannot detect these subtle individual differences, which are probably in the molecular structure. However, such a striking resemblance of their ova in form, so great as to seem to be a complete similarity, is a strong proof of the common parentage of man and the other mammals. From the common germ-form we infer a common stem-form. On the other hand, there are striking peculiarities by which we can easily distinguish the fertilised ovum of the mammal from the fertilised ovum of the birds, amphibia, fishes, and other vertebrates (see the close of Chapter 2.29).