Biology and Its Makers / With Portraits and Other Illustrations

by William A. Locy

Darwin's Theory of Pangenesis.—In attempting to account for heredity, Darwin saw clearly the necessity of providing some means of getting all hereditary qualities combined within the egg and the sperm. Accordingly he originated his provisional theory of pangenesis. Keeping in mind the fact that all organisms begin their lives in the condition of single cells, the idea of inheritance through these microscopic particles becomes difficult to understand. How is it possible to conceive of all the hereditary qualities being contained within the microscopic germ of the future being? Darwin supposed that very minute particles, which he called gemmules, were set free from all the cells in the body, those of the muscular system, of the nervous system, of the bony tissues, and of all other tissues contributing their part. These liberated gemmules were supposed to be carried by the circulation and ultimately to be aggregated within the germinal elements (ovum and sperm). Thus the germinal elements would be a composite of substances derived from all organs and all tissues.

With this conception of the blending of the parental qualities within the germinal elements we can conceive how inheritance would be possible and how there might be included in the egg and the sperm a representative in material substance of all the qualities of the parents. Since development begins in a fertilized ovum, this complex would contain minute particles derived from every part of the bodies of both parents, which by growth would give rise to new tissues, all of them containing representatives of the tissues of the parent form.

Theory of Pangenesis Replaced by that of Germinal Continuity.—This theory of Darwin served as the basis for other theories founded upon the conception of the existence of pangens; and although the modifications of Spencer, Brooks, and others were important, it is not necessary to indicate them in detail in order to understand what is to follow. The various theories founded upon the idea of pangens were destined to be replaced by others founded on the conception of germinal continuity—the central idea in nineteenth-century biology.

The four chief steps which have led to the advancement of the knowledge of heredity, as suggested by Thomson, are as follows: "(a) The exposition of the doctrine of germinal continuity, (b) More precise investigation of the material basis of inheritance, (c) Suspicions regarding the inheritance of acquired characteristics, (d) Application of statistical methods which have led to the formulation of the law of ancestral heredity." We shall take these up in order.

Exposition of the Doctrine of Germinal Continuity.—From parent to offspring there passes some hereditary substance; although small in amount, it is the only living thread that connects one generation with another. It thus appears that there enters into the building of the body of a new organism some of the actual substance of both parents, and that this transmitted substance must be the bearer of hereditary qualities. Does it also contain some characteristics inherited from grandparents and previous generations? If so, how far back in the history of the race does unbroken continuity extend?

Briefly stated, genetic continuity means that the ovum and its fertilizing agent are derived by continuous cell-lineage from the fertilized ovum of previous generations, extending back to the beginning of life. The first clear exposition of this theory occurs in the classical work of Virchow on Cellular Pathology, published in 1858. Virchow (1821-1902), the distinguished professor of the University of Berlin, has already been spoken of in connection with the development of histology. He took the step of overthrowing the theory of free cell-formation, and replacing it by the doctrine of cell-succession. According to the theory of Schleiden and Schwann, cells arose from a blastema by a condensation of matter around a nucleus, and the medical men prior to 1858 believed in free cell-formation within a matrix of secreted or excreted substance. This doctrine was held with tenacity especially for pathological growths. Virchow demonstrated, however, that there is a continuity of living substance in all growths—that cells, both in health and in disease, arise only by the growth and division of previously existing living cells; and to express this truth he coined the formula "omnis cellula e cellula." Manifestly it was necessary to establish this law of cell-succession before any idea of germinal continuity could prevail. Virchow's work in this connection is of undying value.

When applied to inheritance the idea of the continuity of living substance leads to making a distinction between germ-cells and body-cells. This had been done before the observations of Virchow made their separation of great theoretical value. Richard Owen, in 1849, pointed out certain differences between the body-cells and the germinal elements, but he did not follow up the distinction which he made. Haeckel's General Morphology, published in 1866, forecasts the idea also, and in 1878 Jaeger made use of the phrase "continuity of the germ protoplasm." Other suggestions and modifications led to the clear expression by Nussbaum, about 1875, that the germinal substance was continued by unbroken generations from the past, and is the particular substance in which all hereditary qualities are included. But the conception finds its fullest expression in the work of Weismann.

Weismann's explanation of heredity is at first sight relatively simple. In reply to the question, "Why is the offspring like the parent?" he says, "Because it is composed of some of the same stuff." In other words, there has been unbroken germinal continuity between generations. His idea of germinal continuity, i.e., unbroken continuity, through all time, of the germinal substance, is a conception of very great extent, and now underlies all discussion of heredity.

In order to comprehend it, we must first distinguish between the germ-cells and the body-cells. Weismann regards the body, composed of its many cells, as a derivative that becomes simply a vehicle for the germ-cells. Owen's distinction between germ-cells and body-cells, made in 1849, was not of much importance, but in the theory of Weismann it is of vital significance. The germ-cells are the particular ones which carry forward from generation to generation the life of the individual. The body-cells are not inherited directly, but in the transmission of life the germ-cells pass to the succeeding generation, and they in turn have been inherited from the previous generation, and, therefore, we have the phenomenon of an unbroken connection with all previous generations.

When the full significance of this conception comes to us, we see why the germ-cells have an inherited organization of remarkable complexity. This germinal substance embodies all the past history of the living, impressionable protoplasm, which has had an unbroken series of generations. During all time it has been subjected to the molding influence of external circumstances to which it has responded, so that the summation of its experiences becomes in some way embedded within its material substance. Thus we have the germinal elements possessing an inherited organization made up of all the previous experiences of the protoplasm, some of which naturally are much more dominant than the others.