The Significance of the Material Continuity in Inheritance

But what about the material continuity appearing in inheritance, which we have said to be almost self-evident, as life is only known to exist on material bodies? Is there not, in fact, a serious contradiction in admitting at the same time entelechy on the one side and a sort of material condition on the other as the basis of all that leads to and from inheritance? Next summer the relation between matter and our autonomous agent of life will be studied more fully; at present it must be enough to state in a more simple and realistic way, what we hold this relation to be. There is no contradiction at all in stating that material continuity is the basis of inheritance on the one side, and entelechy on the other. It would be very inconvenient for us if there were any: for the material continuity is a mere fact and our entelechy we hope we have proved to exist also; if now there were any sort of contradiction in assuming the existence of both of them, of course it would be fatal to our proof.

Let us try to comprehend what is meant by the statement that entelechy and something material are at work in inheritance at the same time. Entelechy has ruled the individual morphogenesis of the generation which is regarded as being the starting-point for inheritance, and will rule also the morphogenesis of the generation which is to follow; entelechy determines the egg to be what it is, and the morphogenesis starting from this egg to be what it is also. Entelechy, at present, is not much more for us than a mere word, to signify the autonomous, the irreducible of all that happens in morphogenesis with respect to order, in the one generation and in the next. But may not the material continuity which exists in inheritance account perhaps for the material elements which are to be ordered? In such a way, indeed, I hope we shall be able to reconcile entelechy and the material basis of heredity. May it not be that there exist some “means” for morphogenesis, which are handed down from generation to generation, always controlled by entelechy, and which constitute the real significance of the continuity of matter during inheritance?

The Experimental Facts about Inheritance

Discoveries of the last few years do seem to show that such means of a material character, though not the foundation of that order of processes which is inherited, are nevertheless among the most necessary conditions for the accomplishment of inheritance in general. It is scarcely necessary to remind you that for very many years all concrete research on heredity proper—that is, the actual comparison of the various specific characters in the generations of the grandfather, the father, and the child—was due to Galton. You may also be aware that in spite of Galton’s inestimable services it was not till 1900 that one of the active principles concerned in inheritance was found independently by de Vries, Correns, and Tschermak, and that this principle happened to be one that had been discovered already, stated with the utmost clearness and precision by the Augustinian monk, Gregor Mendel,[129] as early as 1865, though it had been completely forgotten ever since.

The so-called “rule of Mendel” is based upon experiments with hybrids, that is, with the offspring of parents belonging to different species, or, at least, varieties, but it relates not to the characters of the generation resulting immediately from hybridisation, the “first” generation of hybrids, as we shall call it, but to the characters of that generation which is the result of crossing the hybrids with each other, provided that this leads to any offspring at all. There are many cases indeed, both amongst animals and plants, where the offspring of the hybrids, or in other terms the “second” generation, is found to consist of individuals of three different types—the mixed[130] type of the hybrids themselves, and the two pure types of the grandparents. Whenever the individuals of the “second” generation are separated into these three different types, hybrids are said to “split.” It is the fact of this splitting on the one hand, and on the other hand a certain statement about the numbers of individuals in the three different types of the “second” generation, that gives its real importance to Mendel’s rule.

Before discussing what may follow from Mendel’s discovery for the theory of heredity, we must lay stress on the fact that there are many exceptions to his rule. In quite a number of cases the hybrids are of one or more types, which remain constant: there is no splitting at all in the second generation. But that does not affect the rule of Mendel in those cases where it is true. Where there is a “splitting” in the second generation, there also are the numerical proportions stated by Mendel; there never are other relations among the numbers of individuals of the mixed and of the two pure types than those given by his rule. I regard it as very important that this real meaning of Mendel’s principle should be most clearly understood.

From the fact of the splitting of hybrids in the second generation most important consequences may be drawn for the theory of inheritance; the split individuals, if crossed with each other, always give an offspring which remains pure; there is no further splitting and no other change whatever. The germ-cells produced by the split individuals of the second generation may therefore be said to be “pure,” as pure as were those of the grandparents. But that is as much as to say that the pureness of the germ-cells has been preserved in spite of their passing through the “impure” generation of the hybrids, and from this fact it follows again that the union of characters in the hybrids must have been such as to permit pure separation: in fact, the germ-cells produced by Mendelian hybrids may hypothetically be regarded as being pure themselves.[131]

We have not yet considered one feature of all experiments in hybridisation, which indeed seems to be the most important of all for the theory of inheritance, if taken together with the fact of the pureness of the germs. The rule of Mendel always relates to one single character of the species or varieties concerned in hybridisation, and if it deals with more than one character, it regards every one of them separately; indeed, the rule holds for every one of them irrespective of the others. We cannot study here how this most important fact of the independence of the single characters of a species with regard to inheritance leads to the production of new races, by an abnormal mixture of those characters. We only take advantage of the fact theoretically, and in doing so, I believe, we can hardly escape the conclusion that the independence of the single characters in inheritance, taken together with the pureness of the germ-cells in the most simple form of hybrids, proves that there occurs in inheritance a sort of handing over of single and separate morphogenetic agents which relate to the single morphogenetic characters of the adult. We may use Bateson’s word “allelomorphs” for these agents, or units, as they may be called, thereby giving expression to the fact that the single and separate units, which are handed over in inheritance, correspond to each other in nearly related species without being the same.

And so we have at least an inkling of what the material continuity of inheritance is to mean, though, of course, our “single and separate morphogenetic agents,” or “units” or “allelomorphs” are in themselves not much more than unknown somethings described by a word; but even then they are “somethings.”