 It is well documented that Thomas Robert Malthus’ An  Essay on the Principle of Population greatly influenced both  Charles Darwin and Alfred Russell Wallace’s independent conception of  their theory of natural selection. In it, Malthus puts forward his  observation that the finite nature of resources is in conflict with the  potentially exponential rate of reproduction, leading to an inevitable  struggle between individuals. Darwin took this basic premise and applied  it to nature, as he notes in his autobiography:
It is well documented that Thomas Robert Malthus’ An  Essay on the Principle of Population greatly influenced both  Charles Darwin and Alfred Russell Wallace’s independent conception of  their theory of natural selection. In it, Malthus puts forward his  observation that the finite nature of resources is in conflict with the  potentially exponential rate of reproduction, leading to an inevitable  struggle between individuals. Darwin took this basic premise and applied  it to nature, as he notes in his autobiography:
In October 1838, that is, fifteen months after I had  begun my systematic inquiry, I happened to read for amusement Malthus on  Population, and being well prepared to appreciate the struggle for  existence which everywhere goes on  from long-continued observation of  the habits of animals and plants, it at once struck me that under these  circumstances favourable variations would tend to be preserved, and  unfavourable ones to be destroyed. The results of this would be the  formation of a new species. Here, then I had at last got a theory by  which to work.
The interaction of demographic and evolutionary processes is thus  central in understanding Darwin’s big idea: that exponential growth will  eventually lead to a large population, and in turn will generate  competition for natural selection to act on any heritable variation  which conferred a greater fitness advantage. Under these assumptions we  are able to interpret the evolutionary record of most species by  appealing to two basic causal elements: genes and the environment. As we  all know, in most cases the environment generates selection pressures  to which genes operate and respond. For humans, however, the situation  becomes more complicated when we consider another basic causal element:  culture. The current  paper by Richerson, Boyd & Bettinger (2009) offers one way to  view this muddied situation by delineating the demographic and  evolutionary processes through the notion of time scales:
The idea of time scales is used in the physical   environmental sciences to simplify problems with complex interactions   between processes. If one process happens on a short time scale and the   other one on a long time scale, then one can often assume that the  short  time scale process is at an equilibrium (or in some more complex  state  that can be described statistically) with respect to factors  governed by  the long scale process. If the short time scale and long  time scale  interact, we can often imagine that at each time step in the  evolution  of the long time scale process, the short time scale process  is at  “equilibrium.” A separation of time scales, if justified, makes  thinking  about many problems of coupled dynamics much easier.
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