The moulding of senescence by natural selection

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Abstract

The consequences to fitness of several types of small age-specific effects on mortality are formulated mathematically. An effect of given form always has a larger consequence, or at least one as large, when it occurs earlier. By reference to a model in which mortality is constant it is shown that this implication cannot be avoided by any conceivable organism. A basis for the theory that senescence is an inevitable outcome of evolution is thus established.

The simple theory cannot explain specially high infant mortalities. Fisher's “reproductive value”, the form of which gave rise to an erroneous opinion on this point, is shown to be not directly relevant to the situation. Infant mortality may evolve when the early death of one infant makes more likely the creation or survival of a close relative. Similarly, post-reproductive life-spans may evolve when the old animal still benefits its younger relatives.

The model shows that higher fertility will be a primary factor leading to the evolution of higher rates of senescence unless the resulting extra mortality is confined to the immature period. Some more general analytical notes on the consequences of modifications to the reproductive schedule are given.

Applications to species with populations in continual fluctuation are briefly discussed. Such species apart, it is argued that general stationarity of population can be assumed, in which case the measurement of consequences to fitness in terms of consequences to numerical expectation of offspring is justified.

All the age-functions discussed are illustrated by graphs derived from the life-table of the Taiwanese about 1906, and the method of computation is shown.

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    The work presented in this paper was mainly carried out during tenure of a Medical Research Council Scholarship at the London School of Economics and at University College, London.

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