Mutations that a population accumulates during evolution in one ("home") environment may cause fitness gains or losses in other conditions. Such pleiotropic fitness effects determine the evolutionary fate of the population in variable environments and can lead to ecological specialization. It is unclear how the pleiotropic outcomes of evolution are shaped by the intrinsic randomness of the evolutionary process and by the deterministic variation in selection pressures across environments. To address this question, we evolved 20 replicate populations of the yeast Saccharomyces cerevisiae in 11 laboratory environments and measured their fitness across multiple other conditions. We found that evolution in all home environments led to a diversity of patterns of pleiotropic fitness gains and losses, driven by multiple types of mutations. Approximately 60% percent of this variation are explained by clone's home environment and the most common parallel genetic changes, while about 40% are attributed to the stochastic accumulation of mutations whose pleiotropic effects are unpredictable. On average, populations specialized to their home environment, but generalists also evolved in almost all conditions. Our results suggest that the mutations accumulating in a home environment incur a variety of pleiotropic effects, from costs to benefits, with different probabilities. Therefore, whether a population evolves towards a specialist or a generalist phenotype is heavily influenced by chance.frequencies in a population adapting to one condition typically affect fitness in other 14 conditions?
15Historically, it has been assumed that pleiotropy is often antagonistic, i.e., fitness 16 benefits in one environment should often come at a fitness cost in other 17 conditions [16][17][18]. If antagonistic pleiotropy was common, it would explain why 18 ecological specialization and local adaptation are so widespread in nature. However, 19 more recent field studies have found that adaptive alleles confer fitness defects much less 20 frequently than anticipated [8,[13][14][15]19]. Although theory suggests that ecological 21 specialization and local adaptation can arise without trade-offs [20-22], it is also 22 possible that field studies provide a skewed view of the structure of pleiotropy because 23 of statistical complications and confounding factors, such as migration and unknown 24 environmental variation [23-25]. 25 Laboratory microbial and viral populations are powerful model systems where the 26 structure of pleiotropy can be probed under controlled conditions and with a degree of 27 replication seldom achievable in natural systems. Experimental populations can be 28 evolved in a laboratory environment, adaptive mutations can be identified, and the 29 fitness of evolved genotypes can be directly measured in other conditions. Several dozen 30 such studies in a variety of organisms have been carried out so far (e.g., Refs. [26-45] 31and more references in recent reviews by Elena [15] and Bono et al [14,22]). Their 32 outcomes generall...