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Endeavour 134-139 (July 1953)
The evolution of adaptations
C. H. WADDINGTON

Current biological belief regards evolution as being primarily the result of the natural selection of random mutations, useful adaptations gradually spreading throughout a race. Professor Waddington regards this as an 'extreme view, and in this article puts forward a hypothesis to explain how acquired characteristics may become hereditarily fixed by a process of genetic assimilation not invoking the generally discredited theory of direct inheritance.

It is abundantly found in the living world that the structure of an animal or plant is very precisely adapted to the functions which it has to perform. The nature of the processes by which this situation has been brought about during evolution provides one of the major problems for biological theory. The hypothesis of the inheritance of acquired characters suggested that in some way or other the effects of functioning become themselves inherited. It has usually been interpreted to mean that the reaction between the organism and its surroundings has, as one of its results, an effect on the germ-plasm such that new hereditary changes occur, of a kind which determines the development in later generations of individuals suited to these particular conditions of life. Although this idea has recently been revived in a rather nebulous form in the Soviet Union, it has been so completely rejected by the rest of the scientific world that it is hardly considered to be worthy of discussion in most of the important recent works on evolution. The reigning modern view is that, in nature, the direction of mutational change is entirely at random, and that adaptation results solely from the natural selection of mutations which happen to give rise to individuals with suitable characteristics. I want to argue that this theory is an extremist one, and that, in essaying to account for adaptation, it neglects to call to its aid the doctrines emerging in other fields of modern biology which can quite properly be combined with the conclusions of genetics in the strict sense. In the discussion which follows, attention will be confined to animals, but there is no reason to doubt that similar arguments could be advanced in the botanical field.

It will be advisable first to glance briefly at the phenomena which are usually referred to under the heading of adaptation, since they are of several different kinds which must be distinguished from one another.

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