We have seen that this idea was not first expressed by Weismann; it was a modification of the views of Nussbaum and Hertwig. While it was not his individually, his conclusions were apparently reached independently. This idea was in the intellectual atmosphere of the times. Several investigators reached their conclusions independently, although there is great similarity between them. Although the credit for the first formulation of the law of germinal continuity does not belong to Weismann, that of the greatest elaboration of it does. This doctrine of germinal continuity is now so firmly embedded in biological ideas of inheritance and the evolution of animal life that we may say it has become the corner-stone of modern biology.

The conclusion reached—that the hereditary substance is the germ-plasm—is merely preliminary; the question remains, Is the germ-plasm homogeneous and endowed equally in all parts with a mixture of hereditary qualities? This leads to the second step.

The More Precise Investigation of the Material Basis of Inheritance.—The application of the microscope to critical studies of the structure of the germ-plasm has brought important results which merge with the development of the idea of germinal continuity. Can we by actual observation determine the particular part of the protoplasmic substance that carries the hereditary qualities? The earliest answer to this question was that the protoplasm, being the living substance, was the bearer of heredity. But close analysis of the behavior of the nucleus during development led, about 1875, to the idea that the hereditary qualities are located within the nucleus of the cell.

This idea, promulgated by Fol, Koelliker, and Oskar Hertwig, narrowed the attention of students of heredity from the general protoplasmic contents of the cell to the nucleus. Later investigations show that this restriction was, in a measure, right. The nucleus takes an active part during cell-division, and it was very natural to reach the conclusion that it is the particular bearer of hereditary substance. But, in 1883, Van Beneden and Boveri made the discovery that within the nucleus are certain distinct little rod-like bodies which make their appearance during cell-division. These little bodies, inasmuch as they stain very deeply with the dyes used in microscopic research, are called chromosomes. And continued investigation brought out the astounding fact that, although the number of chromosomes vary in different animals (commonly from two to twenty-four), they are of the same number in all the cells of any particular animal or plant. These chromosomes are regarded as the bearers of heredity, and their behavior during fertilization and development has been followed with great care.

Brilliant studies of the formation of the egg have shown that the egg nucleus, in the process of becoming mature, surrenders one-half its number of chromosomes; it approaches the surface of the egg and undergoes division, squeezing out one-half of its substance in the form of a polar globule; and this process is once repeated.[8] The formation of polar globules is accompanied by a noteworthy process of reduction in the number of chromosomes, so that when the egg nucleus has reached its mature condition it contains only one-half the number of chromosomes characteristic of the species, and will not ordinarily undergo development without fertilization.

The precise steps in the formation of the sperm have also been studied, and it has been determined that a parallel series of changes occur. The sperm, when it is fully formed, contains also one-half the number of chromosomes characteristic of the species. Now, egg and sperm are the two germinal elements which unite in development. Fertilization takes place by the union of sperm and egg, and inasmuch as the nuclei of each of these structures contain one-half of the number of chromosomes characteristic of the species, their union in fertilization results in the restoration of the original number of chromosomes. The fertilized ovum is the starting-point of a new organism, and from the method of its fertilization it appears that the parental qualities are passed along to the cells of every tissue.

The complex mechanism exhibited in the nucleus during segmentation is very wonderful. The fertilized ovum begins to divide, the nucleus passing through a series of complicated changes whereby its chromosomes undergo a lengthwise division—a division that secures an equable partition of the substance of which they are composed. With each successive division, this complicated process is repeated, and the many cells, arising from continued segmentation of the original cell, contain nuclei in which are embedded descendants of the chromosomes in unbroken succession. Moreover, since these chromosomes are bi-parental, we can readily understand that every cell in the body carries both maternal and paternal qualities.

The careful analysis of the various changes within the nuclei of the egg proves to be the key to some of the central questions of heredity. We see the force of the point which was made in a previous chapter, that inheritance is in the long run a cellular study, and we see in a new light the importance of the doctrine of germinal continuity. This conception, in fact, elucidates the general problem of inheritance in a way in which it has never been elucidated by any other means.

For some time the attention of investigators was concentrated upon the nucleus and the chromosomes, but it is now necessary to admit that the basis of some structures is discoverable within the cytoplasm that surrounds the nucleus. Experimental observations (Conklin, Lillie, Wilson) have shown the existence of particular areas within the apparently simple substance of the egg, areas which are definitely related to the development of particular parts of the embryo. The removal of any one of these pre-localized areas prevents the development of the part with which it is genetically related. Researches of this kind, necessitating great ingenuity in method and great talents in the observers, are widening the field of observation upon the phenomena of heredity.

The Inheritance of Acquired Characteristics.—The belief in the inheritance of acquired characteristics was generally accepted up to the middle of the nineteenth century, but the reaction against it started by Galton and others has assumed great proportions. Discussions in this line have been carried on extensively, and frequently in the spirit of great partizanship. These discussions cluster very much about the name and the work of Weismann, the man who has consistently stood against the idea of acquired characteristics. More in reference to this phase of the question is given in the chapter dealing with Weismann's theory of evolution (see p. 398). Wherever the truth may lie, the discussions regarding the inheritance of acquired characteristics provoked by Weismann's theoretical considerations, have resulted in stimulating experiment and research, and have, therefore, been beneficial to the advance of science.