Figure 4 Trajectory for mean fitness relative to the ancestor in one population of E. coli during 10,000 generations of experimental evolution. Each point is the mean of three assays. Curve is the best fit of a hyperbolic model.

proved, thus demonstrating adaptation by natural selection? Is the evolutionary trajectory for fitness similar to that for cell size, or did fitness improve at a constant rate throughout the experimental evolution? Have the replicate populations also diverged from one another in mean fitness, suggesting that they scaled different adaptive peaks? Or have the populations converged on similar fitnesses, implying that their differences in cell size are inconsequential for fitness? How tightly correlated are morphology and fitness?

Figure 4 shows the dynamics of mean fitness relative to the ancestor for the same population whose evolutionary trajectory for average cell size is shown in Figure 1. Mean fitness evolved rapidly for 2000 generations in the experimental environment but was nearly constant for the final several thousand generations. Although the fitness data are subject to more "noise" than the data on cell size, the hyperbolic model explains 70% of the variation in mean fitness (r = 0.843, n = 21, P < 0.001; relative to linear model, partial F = 17.9, 1 and 18 df, P < 0.001). The fact that fitness shows the same decelerating trajectory belies the hypothesis that adaptation continued apace for the 10,000 generations but was not reflected in further changes in cell size.

Figure 5 provides finer resolution of the fitness trajectory for the first 2000 generations in the same population; fitness was estimated every 100 (rather than 500) generations and with 10-fold (rather than 3-fold) replication. A step model (Lenski et al., 1991) [y = c0 + c1 (if t > T1) +



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