Horizontal Transfer and Selection in the Evolution of P Elements

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Abstract
The roles of selection and horizontal transfer in the evolution of the canonical subfamily of P elements were studied in the saltans and willistoni species groups of the genus Drosophila (subgenus Sophophora). We estimate that the common ancestor of the canonical P subfamily dates back 2–3 Myr at the most, despite the much older age (more than 40 Myr) of the P family as a whole. The evolution of the canonical P subfamily is characterized by weak selection at nonsynonymous sites. These sites have evolved at three quarters the rate of synonymous sites, in which no selective constraints were detected. Their recent horizontal transfer best explains the high degree of similarity among canonical P elements from the saltans and willistoni species groups. These results are consistent with a model of P-element evolution in which selective constraints are imposed at the time of horizontal transfer. Furthermore, it is estimated that the spread and diversification of the canonical subfamily involved a minimum of 11 horizontal transfer events among the 18 species surveyed within the past 3 Myr. The presence of multiple P subfamilies in the saltans and willistoni species groups is likely to be the result of multiple invasions that have previously swept through these taxa in a succession of horizontal transfer events. These results suggest that horizontal transfer among eukaryotes might be more common than anticipated.