Page:EB1911 - Volume 13.djvu/367

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HEREDITY
351

generic, specific, individual—of a new organism, or rather the directing substance which in appropriate surroundings of food, &c., forms a new organism. Each id is a veritable microcosm, possessed of an historic architecture that has been elaborated slowly through the multitudinous series of generations that stretch backwards in time from every living individual. This microcosm, again, consists of a number of minor vital units called “determinants,” which cohere according to the architecture of the whole id. The determinants are hypothetical units corresponding to the number of parts of the organism independently variable. Lastly, each determinant consists of a number of small hypothetical units, the “biophores.” These are adaptations of a conception of H. de Vries, and are supposed to become active by leaving the nucleus of the cell in which they lie, passing out into the general protoplasm of the cell and ruling its activities. Each new individual begins life as a nucleated cell, the nucleus of which contains germ-plasm of this complex structure derived from the parent. The reproductive cell gives rise to the new individual by continued absorption of food, by growth, cell-divisions and cell-specializations. The theory supposes that the first divisions of the nucleus are “doubling,” or homogeneous divisions. The germ-plasm has grown in bulk without altering its character in any respect, and, when it divides, each resulting mass is precisely alike. From these first divisions a chain of similar doubling divisions stretches along the “germ-tracks,” so marshalling unaltered germ-plasm to the generative organs of the new individual, to be ready to form the germ-cells of the next generation. In this mode the continuity of the germ-plasm from individual to individual is maintained. This also is the immortality of the germ-cells, or rather of the germ-plasm, the part of the theory which has laid so large a hold on the popular imagination, although it is really no more than a reassertion in new terms of biogenesis. With this also is connected the celebrated denial of the inheritance of acquired characters. 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 on her body in subsequent development. In the later elaboration of his theory Weismann has admitted the possibility of some direct modification of the germ-plasm within the body of the individual acting as its host.

The mass of germ-plasm which is not retained in unaltered form to provide for the generative cells is supposed to be employed for the elaboration of the individual body. It grows, dividing and multiplying, and forms the nuclear matter of the tissues of the individual, but the theory supposes this process to occur in a peculiar fashion. The nuclear divisions are what Weismann calls “differentiating” or heterogeneous divisions. In them the microcosms of the germ-plasm are not doubled, but slowly disintegrated in accordance with the historical architecture of the plasm, each division differentiating among the determinants and marshalling one set into one portion, another into another portion. There are differences in the observed facts of nuclear division which tend to support the theoretical possibility of two sorts of division, but as yet these have not been correlated definitely with the divisions along the germ-tracks and the ordinary divisions of embryological organogeny. The theoretical conception is, that when the whole body is formed, the cells contain only their own kind of determinants, and it would follow from this that the cells of the tissues cannot give rise to structures containing germ-plasm less disintegrated than their own nuclear material, and least of all to reproductive cells which must contain the undisintegrated microcosms of the germ-plasm. Cases of bud-formation and of reconstructions of lost parts (see Regeneration of Lost Parts) are regarded as special adaptations made possible by the provision of latent groups of accessory determinants, to become active only on emergency.

It is to be noticed that Weismann’s conception of the processes of ontogeny is strictly evolutionary, and in so far is a reversion to the general opinion of biologists of the 17th and 18th centuries. These supposed that the germ-cell contained an image-in-little of the adult, and that the process of development was a mere unfolding or evolution of this, under the influence of favouring and nutrient forces. Hartsoeker, indeed, went so far as to figure the human spermatozoon with a mannikin seated within the “head,” and similar extremes of imagination were indulged in by other writers for the spermatozoon or ovum, according to the view they took of the relative importance of these two bodies. C. F. Wolff, in his Theoria generationis (1759), was the first distinguished anatomist to make assault on these evolutionary views, but his direct observations on the process of development were not sufficient in bulk nor in clarity of interpretation to convince his contemporaries. Naturally the improved methods and vastly greater knowledge of modern days have made evolution in the old sense an impossible conception; we know that the egg is morphologically unlike the adult, that various external conditions are necessary for its subsequent progress through a slow series of stages, each of which is unlike the adult, but gradually approaching it until the final condition is reached. None the less, Weismann’s theory supposes that the important determining factor in these gradual changes lies in the historical architecture of the germ-plasm, and from the theoretical point of view his theory remains strictly an unfolding, a becoming manifest of hidden complexity.

Hertwig’s View.—The chief modern holder of the rival view, and the writer who has put together in most cogent form the objections to Weismann’s theory, is Oscar Hertwig. He points out that there is no direct evidence for the existence of differentiating as opposed to doubling divisions of the nuclear matter, and, moreover, he thinks that there is very generally diffused evidence as to the universality of doubling division. In the first place, there is the fundamental fact that single-celled organisms exhibit only doubling division, as by that the persistence of species which actually occurs alone is possible. In the case of higher plants, the widespread occurrence of tissues with power of reproduction, the occurrence of budding in almost any part of the body in lower animals and in plants, and the widespread powers of regeneration of lost parts, are easily intelligible if every cell like the egg-cell has been formed by doubling division, and so contains the germinal material for every part of the organism, and thus, on the call of special conditions, can become a germ-cell again. He lays special stress on those experiments in which the process of development has been interfered with in various ways at various stages, as showing that the cells which arise from the division of the egg-cell were not predestined unalterably for a particular rôle, according to a predetermined plan. He dismisses Weismann’s suggestion of the presence of accessory determinants which remain latent unless they happen to be required, as being too complicated a supposition to be supported without exact evidence, a view in which he has received strong support from those who have worked most at the experimental side of the question. From consideration of a large number of physiological facts, such as the results of grafting, transplantations of tissues and transfusions of blood, he concludes that the cells of an organism possess, in addition to their patent microscopical characters, latent characters peculiar to the species, and pointing towards a fundamental identity of the germinal substance in every cell.

The Nuclear Matter.—Apart from these two characteristic protagonists of extreme and opposing views, the general consensus of biological opinion does not take us very far beyond the plainest facts of observation. The resemblances of heredity are due to the fact that the new organism takes its origin from a definite piece of the substance of its parent or parents. This piece always contains protoplasm, and as the protoplasm of every animal and plant appears to have its own specific reactions, we cannot exclude this factor; indeed many, following the views of M. Verworn, and seeing in the specific metabolisms of protoplasm a large part of the meaning of life, attach an increasing importance to the protoplasm in the hereditary mass. Next, it always contains nuclear matter, and, in view of the extreme