Commentary: The epidemiology of epigenetics: Figure 1

of the conventional kind; one can make operations on the developing wings, using fine glass needles and the other micro-surgical instruments of experimental embryology. A considerable series of such operations has been made by Lees 6 who was able to confirm, and in some cases to extend, many of the previous deductions. For this particular organ there remains very little of that gap between genetics and experimental embry-ology which has been so frequently lamented as one of the main flaws in the structure of biological theory. As would be expected, many of the general principles of experimental embryology reveal themselves again in the epigenetical analysis. For instance, we are familiar with the fact that there are critical periods in development, such as the time of gastrulation at which the primary organizer is active. Similarly we find that in the developmental of the wing there are certain periods at which many deviations of development , which had previously seemed of only minor importance, suddenly entail radical and far-reaching consequences. To give a concrete example: the wing is essentially a sac the two surfaces of which are, at one period, forced apart by a considerable pressure of the contained body-fluid, which is later withdrawn so that the two epithelia come together again. The process of contraction is a critical one. Slight irregularities in it are responsible for most of the abnormalities in the development of the wing-veins, and minor deviations in the relative positions of the wings and legs may, by impeding the flow of body-fluid, lead to crippling malformations of those organs. It would take us too far to attempt to discuss in detail the general characteristics of such epigenetic crises as these. We should find ourselves involved with the same highly complex and little-understood series of problems which confront the experimental embryologist; with the problem of structures of various ranges of size, with the differentiation of cells and of tissues, and with the question of whether differentiation is into sharply contrasted alternatives or into a continuously varying range of products. Without attempting to answer any of these questions here, we may be content to point out that the analysis of the effects of genes has now progressed far enough to become merged with experimental embryology. The two methods of analysis who rapprochement has for so long been no more than a pious hope can now actually and in practice come …