The Biological Problem of To-day: Preformation Or Epigenesis?. Oscar Hertwig

The nucleus and the protoplasm increase in size, and then, first the nucleus and next the protoplasm divide, so that there are formed two cells, each with a nucleus. Each of these again divides, and the process goes on continuously, the new-formed cells gradually being marshalled into their places to form the adult tissues and organs, and they gradually assume the special characters of these tissues and organs. Now, Weismann's theory supposes that the first division of the germplasm is what is called in this translation a doubling division (Erbgleiche Theilung). The mass has grown in bulk, without altering its character, so that each resulting mass is precisely like the other. One of the two portions subsequently increases in bulk, and may again divide repeatedly, but always by doubling division. It therefore remains unaltered germplasm, and eventually is marshalled to the part of the adult from which new organisms are to arise, becoming, for instance, in the case of a woman, the nuclear matter of the ovary. Thus, the germplasm is handed on continuously from generation to generation, forming an unbroken chain, through each individual, from grandparent to grandchild. This is the immortality of the germ-cells, the part of the theory which has laid so strong a hold on the popular imagination. And with this also is connected the equally celebrated denial of the inheritance of acquired characters. For, at first, it seemed a clear inference that, if the hereditary mass for the daughters were separated off from the hereditary mass that was to form the mother, at the very first, before the body of the mother was formed, the daughters were in all essentials the sisters of their mother, and could take from her nothing of any characters that might be impressed upon her body in subsequent development. As this treatise touches only indirectly on the question of acquired characters, it is necessary only to mention that while his early sharp denial of the possibility of inheritance of acquired characters has led to a damaging criticism of supposed cases, Weismann, in the riper development of his theory, has found a possibility for the partial transference of influences that affect the mother to the germplasm contained within her.

      It is with the fate of the other portion coming from the first division of the germplasm that we are concerned here. It is set apart to form the nuclear matter, and so to control the building up of the actual individual. Weismann supposes that the subsequent divisions it undergoes are what I call in this translation differentiating divisions (Erbungleiche Theilung). According to his theory, in each of these divisions the microcosms of the germplasm are not doubled, but are slowly disintegrated, the division differentiating among the determinants, and marshalling one set into one portion, the other set into the other portion. The differentiating process occurs in an order determined by the historic architecture of the microcosms, so that the proper determinants are liberated at the proper time for the modelling of the tissues and organs. Ultimately, when the whole body is formed, the cells contain only their own kind of determinants. It follows, of course, from this that the cells of the tissues cannot give rise to structures containing less disintegrated nuclear material than their own nuclear material, and least of all to reproductive cells, which must contain the undisintegrated microcosms of the germplasm. As special adaptations for the formation of buds and for the reconstruction of lost parts, cells may be provided with latent groups of determinants to become active only on emergency. But with these exceptions, the nuclear matter of the cells of the body contains only what is called idioplasm, a differentiated portion of the germplasm peculiar to cells of their own order, and it can give rise only to idioplasm of the same or of a lower order. And here we come round again to the original observations from which Weismann set out. For he found that among the Hydromedusæ, although the sexual cells seemed to arise in very different topographical positions, there had always been a migration to these localities of a material which he would now call the germplasm. And here also, that the point may be made plain, there may be mentioned the observations of surgeons and physicians, who insist that the growths of disease always conform strictly, in their cellular nature, to the tissues from which they arose, and that in the healing of wounds like only grows from cellular like.

      Dr. Oscar Hertwig is a scientific naturalist of the very first rank, and his name is peculiarly associated with many of the most important advances in our knowledge of cells and of embryology. To him chiefly, for instance, is due the discovery of the intimate nature of fertilisation—that it consists in the union of the nuclear matter of a cell from the male with the nuclear matter of a cell from the female. With the exception of Francis Balfour, no man has laboured more patiently, or achieved more wonderful results, in the investigation of the origin and marshalling of cells by which the egg changes into the adult. From his own experience, and from his study of the observations made by others, he has been led to doubt the validity of apparently fundamental parts of Weismann's conception. In the first place, he thinks that there is no evidence for the existence of differentiating as opposed to doubling divisions, and that there is evidence that divisions always are doubling divisions. He thinks, in fact, that when a portion of germplasm divides, the daughter-cells receive portions of germplasm exactly alike and exactly like the original portion in the parent-cell. The cells, indeed, become different from each other as the organism grows, some becoming muscle-cells, others nerve-cells, others digestive-cells, and so forth. Weismann thinks that the differences occur because, in the disintegration of the germplasm-microcosms, according to a prearranged plan, only the determinants for nerve-cells are marshalled into nerve-cells, only those for muscle-cells into muscle-cells, and so forth. The development is an evolution, an unfolding or unwrapping of little rudiments that lie in the germplasm. Hertwig insists that every cell receives the same kind of germplasm, but that, according to the situations in which they come to lie, different characters are impressed upon them. The development is an epigenesis, or impressing on identical material of different characters by different surrounding forces. His second line of argument against Weismann leads to a similar conclusion. A large number of the characters that arise in an organism during its development are due to the combination of many cells. They cannot come into existence until the multiplication of cells has made their existence possible, and he thinks, therefore, that they cannot have rudiments inside a single cell as their determining cause.

      It is no part of my present purpose to insist, even to the extent that in this treatise Hertwig himself insists, upon the points of agreement between the two views. We are only at the beginning of inquiry into the problems of heredity, and the protagonists of the opposing views, like all those who care more for knowledge than for argument, are concerned more for truth than for the establishment of a modus vivendi. Reconciliation is the parent of slothful thinking and of glosses; it is by sharp contrasting of the opposing views that we are like to have new facts elicited, and new lines of inquiry suggested.

      As many are interested in the problems who have little acquaintance with the technical facts of embryology, a simple account of the early stages in the development of an animal may be useful for reference. I shall choose back-boned animals, as, from the inclusion of man among them, they are of more general interest. The process begins with the fertilisation of the egg-cell by the fusion with its nucleus of the nucleus or head of a male-cell or spermatozoon. At their first origin the nuclei of the sperm and of the egg may be of very different appearance, while that of the sperm is invariably smaller than that of the egg. But before or during the process of fertilisation, changes take place, the result of which is that the fusing nuclei are exactly alike in morphological character. The chromatin, or peculiar substance of the nuclei, is transformed into a number of bodies known as chromosomes, which are of the same number, form, and size, in the two sexes. Form, size, and number are different in different animals, but there is reason to believe that they are normally the same in all the individuals of a species. The fertilised nucleus, thus consisting of chromosomes from male and female, then divides by a complicated process known as karyokinesis, in which each chromosome splits longitudinally, one half passing to each daughter-nucleus. Throughout the whole process of embryonic and post-embryonic growth, the chromatin is gradually increasing in bulk, and being distributed by karyokinesis. The normal character of these divisions is as follows: A daughter-nucleus, after separation, passes through a resting phase, in which the chromosomes, as definite structures, disappear, and in which growth of the nuclear matter occurs. Then chromosomes of definite size and form, and corresponding in number to those present in the fertilised egg-cell, again appear. These split longitudinally, and a half of each passes to each daughter-nucleus. The similarity of these processes among all living creatures, vegetable and animal, and their extreme complication, suggests that karyokinesis is the chief factor in distributing the