Hamilton's forces of natural selection after forty years

In 1966, William D. Hamilton published a landmark paper in evolutionary biology: "The Moulding of Senescence by Natural Selection." It is now apparent that this article is as important as his better-known 1964 articles on kin selection. Not only did the 1966 article explain aging, it also supplied the basic scaling forces for natural selection over the entire life history. Like the Lorentz transformations of relativistic physics, Hamilton's Forces of Natural Selection provide an overarching framework for understanding the power of natural selection at early ages, the existence of aging, the timing of aging, the cessation of aging, and the timing of the cessation of aging. His twin Forces show that natural selection shapes survival and fecundity in different ways, so their evolution can be somewhat distinct. Hamilton's Forces also define the context in which genetic variation is shaped. The Forces of Natural Selection are readily manipulable using experimental evolution, allowing the deceleration or acceleration of aging, and the shifting of the transition ages between development, aging, and late life. For these reasons, evolutionary research on the demographic features of life history should be referred to as "Hamiltonian."