to their ancestral strain has changed. All populations showed a pattern of rapid increase in relative fitness during early generations, with this increase decelerating over time. By 20,000 generations the populations grew approximately 70% faster than the ancestral strain. This increase and deceleration in increase has continued in subsequent generations. A 2013 study by Wiser et al. reported ongoing improvement at 50,000 generations relative to samples isolated at 40,000 generations. They found that the fitness increase fit to a power law model much better than the hyperbolic models that had been used earlier. As a power law model describes an ever-slowing increase that has no upper limit, while a hyperbolic model implies a hard limit, the work suggested that the increase would continue without bound as progressively lower benefit mutations were fixed in the populations. Further work published in 2015 reported the results of over 1100 new fitness assays that examined fitness changes through 60,000 generations. The data once again fit the proposed power law model, and, indeed, fit within predictions of the model from earlier data. These results suggest that, contrary to previous thinking, adaptation and adaptive divergence can potentially increase indefinitely, even in a constant environment. Genome evolution Of the 12 populations, six have so far been reported to have developed defects in their ability to repair DNA, greatly increasing the rate of mutation in those strains. Although the bacteria in each population are thought to have generated hundreds of millions of mutations over the first 20,000 generations, Lenski has estimated that within this time frame, only 10 to 20 beneficial mutations achieved fixation in each population, with fewer than 100 total point mutations (including neutral mutations) reaching fixation in each population. In 2009, Barrick et al. reported the results of genome sequences from multiple time points in population Ara-1. They found that, unlike the declining rate of fitness improvement, mutation accumulation was linear and clock like, even though several lines of evidence suggested that much of the accumulation was beneficial, rather than neutral. Evolution of increased cell size in all twelve populations All twelve of the experimental populations show an increase in cell size concurrent with a decline in maximum population density, and in many of the populations, a more rounded cell shape. This change was partly the result of a mutation that changed the expression of a gene for a penicillin-binding protein, which allowed the mutant