variation in even greater degree to some of their offspring. A competitive struggle for existence working in combination with certain principles of variation and heredity results in a slow and continuous transformation of species through the operation of a process which Darwin termed natural selection.

The coherence and simplicity of the theory, supported as it was by the great array of facts which Darwin had patiently marshalled together, rapidly gained the enthusiastic support of the great majority of biologists. The problem of the relation of species at last appeared to be solved, and for the next forty years zoologists and botanists were busily engaged in classifying by the light of Darwin's theory the great masses of anatomical facts which had already accumulated and in adding and classifying fresh ones. The study of comparative anatomy and embryology received a new stimulus, for with the acceptance of the theory of descent with modification it became incumbent upon the biologist to demonstrate the manner in which animals and plants differing widely in structure and appearance could be conceivably related to one another. Thenceforward the energies of both

botanists and zoologists have been devoted to the construction of hypothetical pedigrees suggesting the various tracks of evolution by which one group of animals or plants may have arisen from another through a long continued process of natural selection. The result of such work on the whole may be said to have shown that the diverse forms under which living things exist to-day, and have existed in the past so far as palaeontology can tell us, are consistent with the view that they are all related by the community of descent which the accepted theory of evolution demands, though as to the exact course of descent for any particular group of animals there is often considerable diversity of opinion. It is obvious that all this work has little or nothing to do with the manner in which species are formed. Indeed, the effect of Darwin's Origin of Species was to divert attention from the way in which species originate. At the time that it was put forward his explanation appeared so satisfying that biologists accepted the notions of variation and heredity there set forth and ceased to take any further interest in the work of the hybridisers. Had Mendel's paper appeared a dozen years earlier it is difficult to believe that it could have failed to attract the attention it deserved. Coming as it did a few years after the publication of Darwin's great work, it found men's minds set at rest on the problems that he raised and their thoughts and energies directed to other matters.

Nevertheless one interesting and noteworthy attempt to give greater precision to the term heredity was made about this time. Francis Galton, a cousin of Darwin, working upon data relating to the breeding of Basset hounds, found that he could express on a definite statistical scheme the proportion in which the different colours appeared in successive generations. Every individual was conceived of as possessing a definite heritage which might be expressed as unity. Of this, ½ was on the average derived from the two parents (i.e. ¼ from each parent), ¼ from the four grandparents, ⅛ from the eight great-grandparents, and so on. The Law of Ancestral Heredity, as it was termed, expresses with fair accuracy some of the statistical phenomena relating to the transmission of characters in a mixed population. But the problem of the way in which characters are distributed from gamete to zygote and from zygote to gamete remained as before. Heredity is essentially a physiological problem, and though statistics may be suggestive in the initiation of experiment, it is upon the basis of experimental fact that progress must ultimately rest. For this reason, in spite of its ingenuity and originality, Galton's theory and the subsequent statistical work that has been founded upon it failed to give us any deeper insight into the nature of the hereditary process.

While Galton was working in England the German zoologist August Weismann was elaborating the complicated

theory of heredity which eventually appeared in his work on The Germplasm (1885), a book which will be remembered for one notable contribution to the subject. Until the publication of Weismann's work it had been generally accepted that the modifications brought about in the individual during its lifetime, through the varying conditions of nutrition and environment, could be transmitted to the offspring. In this biologists were but following Darwin, who held that the changes in the parent resulting from increased use or disuse of any part or organ were passed on to the children. Weismann's theory involved the conception of a sharp cleavage between the general body tissues or somatoplasm and the reproductive glands or germplasm. The individual was merely a carrier for the essential germplasm whose properties had been determined long before he was capable of leading a separate existence. As this conception ran counter to the possibility of the inheritance of "acquired characters," Weismann challenged the evidence upon which it rested and showed that it broke down wherever it was critically examined. By thus compelling biologists to revise their ideas as to the inherited effects of use and disuse, Weismann rendered a valuable service to the study of genetics and did much to clear the way for subsequent research.

A further important step was taken in 1895, when Bateson once more drew attention to the problem of the origin

of species, and questioned whether the accepted ideas of variation and heredity were after all in consonance with the facts. Speaking generally, species do not grade gradually from one to the other, but the differences between them are sharp and specific. Whence comes this prevalence of discontinuity if the process by which they have arisen is one of accumulation of minute and almost imperceptible differences? Why are not intermediates of all sorts more abundantly produced in nature than is actually known to be the case? Bateson saw that if we are ever to answer this question we must have more definite knowledge of the nature of variation and of the nature of the hereditary process by which these variations are transmitted. And the best way to obtain that knowledge was to let the dead alone and to return to the study of the living. It was true that the past record of experimental breeding had been mainly one of disappointment. It was true also that there was no tangible clue by which experiments might be directed in the present. Nevertheless in this kind of work alone there seemed any promise of ultimate success.

A few years later appeared the first volume of de Vries' remarkable book on The Mutation Theory. From a prolonged study of the evening primrose (Oenothera) de Vries concluded that new varieties suddenly arose from older ones by sudden sharp steps or mutations, and not by any process involving the gradual accumulation of minute

differences. The number of striking cases from among widely different plants which he was able to bring forward went far to convincing biologists that discontinuity in variation was a more widespread phenomenon than had hitherto been suspected, and not a few began to question whether the account of the mode of evolution so generally accepted for forty years was after all the true account. Such in brief was the outlook in the central problem of biology at the time of the rediscovery of Mendel's work.

CHAPTER III

MENDEL'S WORK

The task that