Toward a General Theory of Adaptive Radiation Insights from Microbial Experimental Evolution

The history of life has been punctuated by unusually spectacular periods of evolutionary diversification called adaptive radiation. Darwin's finches in the Galapagos, cichlid fishes in African Rift and Nicaraguan crater lakes, and the emergence of mammals at the end of the Cretaceous are hallmark examples. Although we have learned much from these and other case studies about the mechanisms thought to drive adaptive radiations, convincing experimental tests of theory are often lacking for the simple reason that it is usually impossible to "rewind the tape of life," as Stephen Jay Gould was fond of saying, and run it again. This situation has changed dramatically in recent years with the increasing emphasis on the use of microbial populations which, because of their small size and rapid generation times, make possible the construction of replicated, manipulative experiments to study evolution in the laboratory. Here I review the contributions that microbial experimental evolution has made to our understanding of the ecological and genetic mechanisms underlying adaptive radiation. I focus on three major gaps in the theory of adaptive radiation-the paucity of direct tests of mechanism, the genetics of diversification, and the limits and constraints on the progress of radiations-with the aim of pointing the way toward the development of a more general theory of adaptive radiation.