Environmental heterogeneity has often been suggested as a general explanation for patterns of diversity at scales ranging from individuals within populations to communities within landscapes. I evaluate this proposition using laboratory experiments with two microbial species, the unicellular chlorophyte Chlamydomonas reinhardtii and the common bacterium Pseudomonas fluorescens. These experiments contrast the fate of diversity following selection in heterogeneous and homogeneous environments. Specifically, I show that (1) an individual's breadth of adaptation evolves to match the amount of environmental variation, specialists evolving in environments that remain constant through time and generalists evolving in environments that vary through time irrespective of the scale at which environmental variation occurs relative to the lifetime of an individual; (2) the maintenance of diversity in a spatially heterogeneous environment is context-dependent, diversity being more readily maintained when environmental conditions are very different and genotypes are widely divergent; (3) selection in heterogeneous environments represents a plausible mechanism for two well-known patterns of diversity at large spatial scales, namely that between species diversity and both productivity and disturbance. This thesis thus demonstrates that environmental heterogeneity is a plausible, and perhaps very general, factor responsible for the diversity of natural communities.

Determining the factors responsible for the origin and maintenance of diversity remains a difficult problem in evolutionary biology. There is extensive theoretical work which suggests that environmental heterogeneity plays a major role. This theory argues that diversification is ultimately due to divergent natural selection for alternative resources. In this thesis I investigate adaptation and the evolution of diversity in experimental populations of the asexual bacterium Pseudomonas fluorescens. In all experiments I introduce clonal isolates of Pseudomonas to a novel environment and allow evolution to occur through the substitution of random mutations. Adaptation can then be quantified by comparing evolved genotypes to the ancestor. These experiments show that when Pseudomonas is selected in a complex environment containing several resources, sympatric genotypes adapt to use different resources, leading to the evolution of genetically diverse populations. In environments containing just a single resource, most genotypes adapt to use the same resource and no such diversity is observed. Adaptation in the experimental populations is caused by the fixation of beneficial mutations of intermediate fitness effect. My results highlight the value of microbial model systems for answering evolutionary questions and provide strong evidence for the role of ecological factors in the origin of diversity.

No description available.

No description available.