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 Linné—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 III.
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 on extensive observations of fact—mentioned here and there as erroneous; their opponents, who adhered to the dominant theory of preformation, did not even deign to reply to them. This unjust treatment was chiefly due to the extraordinary authority of Albrecht von Haller; it is one of the most astonishing instances of a great authority, as such, preventing for a long time the recognition of established facts.

The general ignorance of Wolff’s work was so great that at the beginning of the nineteenth century two scientists of Jena, Oken (1806) and Kieser (1810), began independent research into the development of the alimentary canal of the chick, and hit upon the right clue to the embryonic puzzle, without knowing a word about Wolff’s important treatise on the same subject. They were treading in his very footsteps without suspecting it. This can be easily proved from the fact that they did not travel as far as Wolff. It was not until Meckel translated into German Wolff’s book on the alimentary system, and pointed out its great importance, that the eyes of anatomists and physiologists were suddenly opened. At once a number of biologists instituted fresh embryological inquiries, and began to confirm Wolff’s theory of epigenesis.

This resuscitation of embryology and development of the epigenesis-theory was chiefly connected with the university of Würtzburg. One of the professors there at that time was Döllinger, an eminent biologist, and father of the famous Catholic historian who later distinguished himself by his opposition to the new dogma of papal infallibility. Döllinger was both a profound thinker and an accurate observer. He took the keenest interest in embryology, and worked at it a good deal. However, he is not himself responsible for any important result in this field. In 1816 a young medical doctor, whom we may at once designate as Wolff’s chief successor, Karl Ernst von Baer, came to Würtzburg. Baer’s conversations with Döllinger on embryology led to a fresh series of most extensive investigations. Döllinger had expressed a wish that some young scientist should begin again under his guidance an independent inquiry into the development of the chick during the hatching of the egg. As neither he nor Baer had money enough to pay for an incubator and the proper control of the experiments, and for a competent artist to illustrate the various stages observed, the lead of the enterprise was given to Christian Pander, a wealthy friend of Baer’s who had been induced by Baer to come to Würtzburg. An able engraver, Dalton, was engaged to do the copper-plates. In a short time the embryology of the chick, in which Baer was taking the greatest indirect interest, was so far advanced that Pander was able to sketch the main features of it on the ground of Wolff’s theory in the dissertation he published in 1817. He clearly enunciated the theory of germinal layers which Wolff had anticipated, and established the truth of Wolff’s idea of a development of the complicated systems of organs out of simple leaf-shaped primitive structures. According to Pander, the leaf-shaped object in the hen’s egg divides, before the incubation has proceeded twelve hours, into two different layers, an external serous layer and an internal mucous layer; between the two there develops later a third layer, the vascular (blood-vessel) layer.[[8]]