Evolution of ageing since Darwin.

In the late 19th century, the evolutionary approach to the problem of ageing was initiated by August Weismann, who argued that natural selection was more important for ageing than any physiological mechanism. In the mid-twentieth century, J. B. S. Haldane, P. B. Medawar and G. C. Williams informally argued that the force of natural selection falls with adult age. In 1966, W. D. Hamilton published formal equations that showed mathematically that two 'forces of natural selection' do indeed decline with age, though his analysis was not genetically explicit. Brian Charlesworth then developed the required mathematical population genetics for the evolution of ageing in the 1970's. In the 1980's, experiments using Drosophila showed that the rate of ageing evolves as predicted by Hamilton's 'forces of natural selection'. The discovery of the cessation of ageing late in life in the 1990's was followed by its explanation in terms of evolutionary theory based on Hamilton's forces. Recently, it has been shown that the cessation of ageing can also be manipulated experimentally using Hamilton's 'forces of natural selection'. Despite the success of evolutionary research on ageing, mainstream gerontological research has largely ignored both this work and the opportunity that it provides for effective intervention in ageing.