You will readily understand that the presentation here given of the phenomena is only the barest possible outline. Some of the details we may now fill in. For example, I have spoken of the characters of the organism, its colour, shape, and the like, as if they were due each to one ingredient or factor. Some of them are no doubt correctly so represented; but already we know numerous bodily features which need the concurrence of several factors to produce them. Nevertheless though the character only appears when all the complementary ingredients are together present, each of these severally and independently follows, as regards its transmission, the simple rules I have described.

This complementary action may be illustrated by some curious results that Mr Punnett and I have encountered when experimenting with the height of Sweet Peas. There are two dwarf varieties, one the prostrate "Cupid," the other the half-dwarf or "Bush" Sweet Peas. Crossed together they give a cross-bred of full height. There is thus some element in the Cupid which when it meets the complementary element from the Bush, produces the characteristic length of the ordinary Sweet Pea. We may note in passing that such a fact demonstrates at once the nature of Variation and Reversion. The Reversion occurs because the two factors that made the height of the old Sweet Pea again come together after being parted: and the Variations by which each of the dwarfs came into existence must have taken place by the dropping out of one of these elements or of the other.

Conversely there are factors which by their presence can prevent or inhibit the development and appearance of others present and unperceived.

For example, all the factors for pigmentation may be present in a plant or an animal; but in addition there may be another factor present which keeps the individual white, or nearly so.

There are cases in which the action of the factors is superposed one on top of the other, and not until each factor is removed in turn can the effects of the underlying factors be perceived. So in the mouse if no other colour-factor is present, the fur is chocolate. If the next factor in the series be there, it is black. If still another factor be added, it has the brownish grey of the common wild mouse. Conversely, by the variation which dropped out the top factor, a black mouse came into existence. By the loss of the black factor, the chocolate mouse was created, and for aught we can tell there may be still more possibilities hidden beneath.

In the disentanglement of the properties and interactions of these elementary factors, the science we must call to our aid is Physiological Chemistry. The relations of Genetics with the other branches of biology are close. Such work can only be conducted by those who have the good fortune to be able to count upon continual help and advice from specialists in the various branches of Zoology, Physiology, and Botany. Often we have questions with which only a cytologist can deal, and often it is the experience of a systematist we must invoke. The school of Genetics in Cambridge starts under happy auspices in that we are surrounded by colleagues qualified, and as we have often found, willing to give us such aid unstinted. But with chemical physiology, we stand in an even closer relation; and from the little I have dared to say respecting the action and interaction of factors, it is evident that for their disentanglement there must one day be an intimate and enduring partnership arranged with the physiological chemists.

Now, as the whole of the elaborate process by which the various elements are apportioned among the gametes must be got through in a few cell-divisions at most, and perhaps in one division only, it is not surprising that there is sometimes an interaction between factors that have quite distinct rôles to perform. These interactions are probably of several kinds. One, which I shall illustrate presently, is probably to be represented as a repulsion between two factors. As a consequence of its operations when the various factors are sorted out into the gametes, if the individual be cross-bred in respect of the two repelling factors, having received so to speak only a single dose of each, then the gametes are made up in such a way that each takes one or other of the two repelling factors, not both.

Mutual repulsions of this kind probably play a significant part in the phenomena of heredity. A single concrete case which Mr Punnett and I have been investigating for some years will illustrate several of these principles. We crossed together a pure white Sweet Pea having an erect standard, with another pure white Sweet Pea having a hooded standard. The result is, as you see, a purple flower with an erect standard. The colour comes from the concurrence of complementary elements. A dose of a certain ingredient from one parent meets a dose of another ingredient from the other parent and the two make pigment in the flower. From other experiments we know that the purple colour of the pigment is due to a dose of a third ingredient brought in from the hooded parent; and that in the absence of that blue factor, as we may call it, the flower would be red. The standard is erect because it contains a dose of the erectness-factor from the erect parent, and the hooded parent can readily be proved to owe its peculiar shape to the absence of that element.

Our purple plant is thus cross-bred for four factors, containing only one dose of each.

We let it fertilise itself, and its offspring show all the possible combinations of the four different factors and their absences which the genetic constitution of the plant can make.