Evolution in space and timethe second synthesis between ecology, evolutionary biology, and the philosophy of biology

Résumé

It is a surprising fact that for the majority of evolutionary biology, we have rarely studied how evolution typically unfolds in nature, in changing ecological environments, over space and time. While ecology played a major role in the eventual acceptance of the population genetic viewpoint of evolution in the Modern Synthesis (circa 1918-1950), it held a lesser role in the development of evolutionary theory until the 1980s, when we began to systematically study the evolutionary dynamics of natural populations in space and time. As a result, evolutionary theory was initially constructed in an abstract vacuum that was unrepresentative of evolution in nature.Evolutionary biology has since undergone a profound shift in thinking about evolution spurred by its recent synthesis with ecology. Ecological insight has revealed how natural selection varies in strength, direction, form, and more surprisingly level of biological organization, all dependent on ecological conditions. The novel concept of evolvability plays an organizing role throughout this thesis, since its recent rise to popularity provides some of the best evidence of how biologists have persistently neglected evolution in space and time. Evolvability, as an emergent adaptation of populations whose manifestation is prompted by ecological changes, thus finally became revealed within an evolutionary biology that embraced evolution in space and time. How such a central process as evolvability can go relatively unnoticed in theory until recent times thus highlights the areas of biology that warrant urgent progress, as well as science more generally.In this thesis, I offer an historical reconstruction of the philosophical, technological, and natural forces that led to the Second Synthesis of biology, in hopes of recognizing the significant advancements that have overtaken biology in the past generation. I then offer my normative recommendations, prescribing a pluralistic theory of natural selection that can explain complex emergent phenomena (like evolvability) to finally resolve the paradox of adaptive variation. I do so by building a bridge between greater biology and the history/philosophy of biology, bringing into focus the primary achievements made by historians and philosophers over the past generation and how these advancements can modernize biological thought