identified processes by which populations might move from one peak to another. One process that can produce a peak shift, without environmental change, is random genetic drift (including founder effects). Although this hypothesis has been highly influential [e.g., for models of speciation (Mayr, 1954; Templeton, 1980)], the mathematical conditions conducive to such peak shifts appear to be restrictive (Barton and Rouhani, 1987). In this respect, it is important that, in our experiment, populations were not on one adaptive peak and asked to "jump" to another peak; instead, they were thrown into an arbitrary environment and asked to climb any accessible peak.

Adaptation, Diversification, and Stasis. For about 2000 generations after their introduction into the experimental environment, all 12 populations underwent rapid changes in both morphology and fitness, whereas these properties were nearly static between generations 5000 and 10,000. The initially rapid evolution was presumably due to intense selection triggered by the sudden environmental changes imposed at the start of our experiment. Although the ancestors of the founding bacterium used in this study had been "in captivity" for several decades, they were not systematically propagated under the experimental conditions that we imposed (serial dilution in glucose-minimal medium). The experimental regime was thus a novel environment. Unfortunately, we cannot say anything quantitative about the evolutionary dynamics of the study organism before the start of our experiment. However, in two other studies, we have used derived bacteria from this study to found new populations, which were introduced into environments that differed either in temperature (Bennett et al., 1992) or in limiting nutrient (Travisano, 1993). In both cases, these environmental changes led to more rapid adaptive evolution.

Our results also reveal the quasi-punctuated dynamics expected when selection depends on new mutations. However, we saw no compelling evidence for any more radical punctuation, such as when one adaptive change sets off a cascade of further changes [cf. "genetic revolutions" (Mayr, 1954; Templeton, 1980) and "epochal mutations" (Kubitschek, 1974)]. Such an effect might have been manifest by a period of renewed, rapid evolutionary change in a population that had previously been at or near stasis. Perhaps 12 populations and 10,000 generations were too few to see such rare events.

If environmental change and the dynamics of selection caused the initially rapid changes in morphology and fitness, then the eventual stasis (or near stasis) presumably resulted from the constancy of the environment and a paucity of mutations that would produce further improvements comparable to those seen earlier in the experiment. The



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