"The specific meaning of the term 'evolvability' is heavily debated, but most definitions can be summarized as: the potential of populations and genomes to produce adaptive variation and complex structures in response to mutation and selection. Changing environments are thought to play a significant role in shaping and promoting evolvability through alternating selective pressures. In this dissertation, I will discuss my recent research on the interplay between changing environments, evolvability, genetic architecture, and the evolution of horizontal gene transfer (HGT), an information-rich mutagenic function that is ubiquitous in nature. Before delving into my own research, however, I begin in the first chapter by providing a survey of current literature on each of these topics, with emphases on how they are believed to arise, how they affect subsequent evolution, and how they relate to each other. Genetic architecture and population dynamics clearly have a complex interplay in ongoing evolutionary dynamics. Evolutionary history, population diversity, modularity, and task size all play a role in determining the location and characteristics of populations in genotype space, and alter the genotype to phenotype map that permits neutral genetic variation. All of these features contribute to evolvability. In Chapter 2, I demonstrate how changing environments provided a sufficient selective pressure to produce quasi-modular genetic architectures that allow for rapid adaptation to the meta-environment of environmental change. Horizontal gene transfer is a highly regulated, ubiquitous, and ancient mechanism for exchanging genetic material between unrelated organisms. In the third chapter, I explore conditions which may have led to the evolution of horizontal gene transfer through transformation, and identify mechanisms that might support its continued performance. In Chapter 4, I compare the fitness and phenotypic effects of the HGT process against other types of increasingly less information rich mutational operators. I demonstrate that not only is HGT selected for in harsh changing environments, but that other mutagenic instructions that contain less information, or provide lesser fitness benefits are not similarly selected for. In the fifth chapter, I explore the long-term evolutionary potential of populations evolved in changing environments by evolving two different populations, one evolved in a minimal changing environment, and the other in a rich changing environment, and exposing them to a brand new environment. I demonstrate that while populations adapted to harsh changing environments are indeed able to adapt quickly to previously seen environmental changes, that these populations do not fare as well in brand new environments. Rather, benign changing environments perform best in measures of task discovery and exploration. In the final chapter, I conclude with a synthesis of my results, along with implications for the field, as well as identification of some new directions for pursuing my research into changing environments."--Pages ii-iii.
Read
