NC Article 3 - O RIGINAL ARTICLE...

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ORIGINAL ARTICLE doi:10.1111/j.1558-5646.2007.00291.x THE ADAPTIVE DYNAMICS OF NICHE CONSTRUCTING TRAITS IN SPATIALLY SUBDIVIDED POPULATIONS: EVOLVING POSTHUMOUS EXTENDED PHENOTYPES Laurent Lehmann Stanford University, California 94305 E-mail: [email protected] Received March 7, 2007 Accepted October 18, 2007 Niche construction, by which organisms modify the environment in which they live, has been proposed to affect the evolution of many phenotypic traits. But what about the evolution of a niche constructing trait itself, whose expression changes the pattern of natural selection to which the trait is exposed in subsequent generations? This article provides an inclusive fitness analysis of selection on niche constructing phenotypes, which can affect their environment from local to global scales in arbitrarily spatially subdivided populations. The model shows that phenotypic effects of genes extending far beyond the life span of the actor can be affected by natural selection, provided they modify the fitness of those individuals living in the future that are likely to have inherited the niche construction lineage of the actor. Present benefits of behaviors are thus traded off against future indirect costs. The future costs will generally result from a complicated interplay of phenotypic effects, population demography and environmental dynamics. To illustrate these points, I derive the adaptive dynamics of a trait involved in the consumption of an abiotic resource, where resource abundance in future generations feeds back to the evolutionary dynamics of the trait. KEY WORDS: Consumer-resource system, extended phenotype, inclusive fitness, niche construction, social evolution, spatial population subdivision, spite. Adaptations, by which organisms appear to fit their particular en- vironment in form and function, have captured the imagination of humans for millennia (Orzack and Sober 2001; Vincent and Brown 2005). But organisms not only adapt to the demands of their environment, but they also modify it and construct it from the elements of the world in which they reside (Lewontin 2000). Although adaptations are often regarded as the phenotypic vari- ants reaching the highest fitness among a set of alternatives in a given environment (Reeve and Sherman 1993), it has also been stressed that adaptations, by modifying the environment, partici- pate in a feedback between the selective pressures and the adap- tations themselves (Odling-Smee et al. 1996, 2003). The view that organisms construct their niches leads to a picture in which organisms and their environment are tightly intertwined, both de- pending on the histories of the organism and the environment (Lewontin 2000). A classical example of this process is the case of the earthworms described by Darwin (1883), which through their burrowing change the structure and chemistry of the soil, with the consequence that some earthworm adaptations, such as epidermis structure or the amount of mucus secreted are likely to have coevolved with earthworm niche construction.
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