We thus know now what 'fertilization' is. Through the labours of the last decade the veil has been torn from a mystery of nature which for thousands of years confronted humanity as unapproachable; a riddle has been solved for the solution of which a few centuries ago men did not even dare to hope. Not a few have taken part in these labours; some I have already named, but it is impossible that I should here mention all who have shared in the achievement by observation or reflection. Whoever has helped it on even a single step may say to himself that he has taken an active part in bringing about what must be called essential progress in human knowledge.

But in the science of nature every new solution implies the cropping up of a new riddle, and we are immediately confronted with the problem, Why should nature, in the course of evolution, have interpolated this process of the mingling of different hereditary substances almost everywhere in the organic world? This, however, is a problem which we cannot attack until we have first made ourselves more fully acquainted with the phenomena of inheritance, and have attempted to reason back from these to the nature of the hereditary substance. We must, in short, think out a theory of heredity.


LECTURE XVII

THE GERM-PLASM THEORY

Conception of the 'id' deduced from the process of fertilization—Hereditary substance, 'idioplasm' and 'germ-plasm'—'Idants'—Evolution or Epigenesis—Herbert Spencer's uniform germinal substance—Determinants—Illustrations: Lycæna agestis—The leaf-butterflies—Insect metamorphosis, limbs of segmented animals—Heterotopia—The ultimate living units or biophors—Number of determinants—Stridulating organ of the grasshopper.

In proceeding to expound the theory of heredity which has shaped itself in my mind in the course of my own scientific development, I should like to begin by pointing out that the hereditary substance of the germ-cell of an animal or of a plant contains not only the primary constituents (Anlagen) of a single individual of the species, but rather those of several, often even of many individuals. That this is so can be proved in several ways.

I start from what I hold to be the proved proposition, that the chromatin substance of the nucleus is the hereditary substance. We have seen that this is present in the germ-cells of every species in the form of a definite number of chromosomes, and that in germ-cells destined for fertilization, that is, in sex-cells, this number is first reduced to half, the reduction being effected, as is now proved in regard to a whole series of animals, by the two last cell-divisions, the so-called maturation divisions.

We know that the full number is only reached again through amphimixis, by which process the half number of chromosomes in the male and female germ-cells are united in a single cell, the 'fertilized ovum,' and in a single nucleus, the so-called segmentation nucleus. Thus the hereditary substance of the child is formed half from the paternal, half from the maternal hereditary substance, and we have seen that this remains so during the whole development of the child, since, at every succeeding cell-division each of the paternal and each of the maternal chromosomes doubles by dividing, and the resulting halves are distributed between the two daughter-nuclei.