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- The Evolution of Man, V.1. - 10/54 -


there, as the plant seed does in the fruitful earth. Hence, every spermatozoon was regarded as a homunculus, a tiny complete man; all the parts were believed to be pre-formed in it, and merely grew larger when it reached its proper medium in the female ovum. This theory, also, was consistently developed in the sense that in each of these thread-like bodies the whole of its posterity was supposed to be present in the minutest form. Adam's sexual glands were thought to have contained the germs of the whole of humanity.

This "theory of male scatulation" found itself at once in keen opposition to the prevailing "female" theory. The two rival theories at once opened a very lively campaign, and the physiologists of the eighteenth century were divided into two great camps--the Animalculists and the Ovulists--which fought vigorously. The animalculists held that the spermatozoa were the true germs, and appealed to the lively movements and the structure of these bodies. The opposing party of the Ovulists, who clung to the older "evolution theory," affirmed that the ovum is the real germ, and that the spermatozoa merely stimulate it at conception to begin its growth; all the future generations are stored in the ovum. This view was held by the great majority of the biologists of the eighteenth century, in spite of the fact that Wolff proved it in 1759 to be without foundation. It owed its prestige chiefly to the circumstance that the most weighty authorities in the biology and philosophy of the day decided in favour of it, especially Haller, Bonnet, and Leibnitz.

Albrecht Haller, professor at Gottingen, who is often called the father of physiology, was a man of wide and varied learning, but he does not occupy a very high position in regard to insight into natural phenomena. He made a vigorous defence of the "evolutionary theory" in his famous work, Elementa physiologiae, affirming: "There is no such thing as formation (nulla est epigenesis). No part of the animal frame is made before another; all were made together." He thus denied that there was any evolution in the proper sense of the word, and even went so far as to say that the beard existed in the new-born child and the antlers in the hornless fawn; all the parts were there in advance, and were merely hidden from the eye of man for the time being. Haller even calculated the number of human beings that God must have created on the sixth day and stored away in Eve's ovary. He put the number at 200,000 millions, assuming the age of the world to be 6000 years, the average age of a human being to be thirty years, and the population of the world at that time to be 1000 millions. And the famous Haller maintained all this nonsense, in spite of its ridiculous consequences, even after Wolff had discovered the real course of embryonic development and established it by direct observation!

Among the philosophers of the time the distinguished Leibnitz was the chief defender of the "preformation theory," and by his authority and literary prestige won many adherents to it. Supported by his system of monads, according to which body and soul are united in inseparable association and by their union form the individual, or the "monad," Leibnitz consistently extended the "scatulation theory" to the soul, and held that this was no more evolved than the body. He says, for instance, in his Theodicee: "I mean that these souls, which one day are to be the souls of men, are present in the seed, like those of other species; in such wise that they existed in our ancestors as far back as Adam, or from the beginning of the world, in the forms of organised bodies."

The theory seemed to receive considerable support from the observations of one of its most zealous supporters, Bonnet. In 1745 he discovered, in the plant-louse, a case of parthenogenesis, or virgin-birth, an interesting form of reproduction that has lately been found by Siebold and others among various classes of the articulata, especially crustacea and insects. Among these and other animals of certain lower species the female may reproduce for several generations without having been fertilised by the male. These ova that do not need fertilisation are called "false ova," pseudova or spores. Bonnet saw that a female plant-louse, which he had kept in cloistral isolation, and rigidly removed from contact with males, had on the eleventh day (after forming a new skin for the fourth time) a living daughter, and during the next twenty days ninety-four other daughters; and that all of them went on to reproduce in the same way without any contact with males. It seemed as if this furnished an irrefutable proof of the truth of the scatulation theory, as it was held by the Ovulists; it is not surprising to find that the theory then secured general acceptance.

This was the condition of things when suddenly, in 1759, Caspar Friedrich Wolff appeared, and dealt a fatal blow at the whole preformation theory with his new theory of epigenesis. Wolff, the son of a Berlin tailor, was born in 1733, and went through his scientific and medical studies, first at Berlin under the famous anatomist Meckel, and afterwards at Halle. Here he secured his doctorate in his twenty-sixth year, and in his academic dissertation (November 28th, 1759), the Theoria generationis, expounded the new theory of a real development on a basis of epigenesis. This treatise is, in spite of its smallness and its obscure phraseology, one of the most valuable in the whole range of biological literature. It is equally distinguished for the mass of new and careful observations it contains, and the far-reaching and pregnant ideas which the author everywhere extracts from his observations and builds into a luminous and accurate theory of generation. Nevertheless, it met with no success at the time. Although scientific studies were then assiduously cultivated owing to the impulse given by Linne--although botanists and zoologists were no longer counted by dozens, but by hundreds, hardly any notice was taken of Wolff's theory. Even when he established the truth of epigenesis by the most rigorous observations, and demolished the airy structure of the preformation theory, the "exact" scientist Haller proved one of the most strenuous supporters of the old theory, and rejected Wolff's correct view with a dictatorial "There is no such thing as evolution." He even went on to say that religion was menaced by the new theory! It is not surprising that the whole of the physiologists of the second half of the eighteenth century submitted to the ruling of this physiological pontiff, and attacked the theory of epigenesis as a dangerous innovation. It was not until more than fifty years afterwards that Wolff's work was appreciated. Only when Meckel translated into German in 1812 another valuable work of Wolff's on The Formation of the Alimentary Canal (written in 1768), and called attention to its great importance, did people begin to think of him once more; yet this obscure writer had evinced a profounder insight into the nature of the living organism than any other scientist of the eighteenth century.

Wolff's idea led to an appreciable advance over the whole field of biology. There is such a vast number of new and important observations and pregnant thoughts in his writings that we have only gradually learned to appreciate them rightly in the course of the nineteenth century. He opened up the true path for research in many directions. In the first place, his theory of epigenesis gave us our first real insight into the nature of embryonic development. He showed convincingly that the development of every organism consists of a series of NEW FORMATIONS, and that there is no trace whatever of the complete form either in the ovum or the spermatozoon. On the contrary, these are quite simple bodies, with a very different purport. The embryo which is developed from them is also quite different, in its internal arrangement and outer configuration, from the complete organism. There is no trace whatever of preformation or in-folding of organs. To-day we can scarcely call epigenesis a THEORY, because we are convinced it is a fact, and can demonstrate it at any moment with the aid of the microscope.

Wolff furnished the conclusive empirical proof of his theory in his classic dissertation on The Formation of the Alimentary Canal (1768). In its complete state the alimentary canal of the hen is a long and complex tube, with which the lungs, liver, salivary glands, and many other small glands, are connected. Wolff showed that in the early stages of the embryonic chick there is no trace whatever of this complicated tube with all its dependencies, but instead of it only a flat, leaf-shaped body; that, in fact, the whole embryo has at first the appearance of a flat, oval-shaped leaf. When we remember how difficult the exact observation of so fine and delicate a structure as the early leaf-shaped body of the chick must have been with the poor microscopes then in use, we must admire the rare faculty for observation which enabled Wolff to make the most important discoveries in this most difficult part of embryology. By this laborious research he reached the correct opinion that the embryonic body of all the higher animals, such as the birds, is for some time merely a flat, thin, leaf-shaped disk--consisting at first of one layer, but afterwards of several. The lowest of these layers is the alimentary canal, and Wolff followed its development from its commencement to its completion. He showed how this leaf-shaped structure first turns into a groove, then the margins of this groove fold together and form a closed canal, and at length the two external openings of the tube (the mouth and anus) appear.

Moreover, the important fact that the other systems of organs are developed in the same way, from tubes formed out of simple layers, did not escape Wolff. The nerveless system, muscular system, and vascular (blood-vessel) system, with all the organs appertaining thereto, are, like the alimentary system, developed out of simple leaf-shaped structures. Hence, Wolff came to the view by 1768 which Pander developed in the Theory of Germinal Layers fifty years afterwards. His principles are not literally correct; but he comes as near to the truth in them as was possible at that time, and could be expected of him.

Our admiration of this gifted genius increases when we find that he was also the precursor of Goethe in regard to the metamorphosis of plants and of the famous cellular theory. Wolff had, as Huxley showed, a clear presentiment of this cardinal theory, since he recognised small microscopic globules as the elementary parts out of which the germinal layers arose.

Finally, I must invite special attention to the MECHANICAL character of the profound philosophic reflections which Wolff always added to his remarkable observations. He was a great monistic philosopher, in the best meaning of the word. It is unfortunate that his philosophic discoveries were ignored as completely as his observations for more than half a century. We must be all the more careful to emphasise the fact of their clear monistic tendency.

CHAPTER 1.3. MODERN EMBRYOLOGY.

We may distinguish three chief periods in the growth of our science of human embryology. The first has been considered in the preceding chapter; it embraces the whole of the preparatory period of research, and extends from Aristotle to Caspar Friedrich Wolff, or to the year 1759, in which the epoch-making Theoria generationis was published. The second period, with which we have now to deal, lasts about a century--that is to say, until the appearance of Darwin's Origin of Species, which brought about a change in the very foundations of biology, and, in particular, of embryology. The third period begins with Darwin. When we say that the second period lasted a full century, we must remember that Wolff's work had remained almost unnoticed during half the time--namely, until the year 1812. During the whole of these fifty-three years not a single book that appeared followed up the path that Wolff had opened, or extended his theory of embryonic development. We merely find his views--perfectly correct views, based


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