As the first step towards the evolution of selfing from obligate outcrossing, identifying the key mutations underlying the loss of self-incompatibility is of particular interest. However, our current knowledge is primarily based on sequence-based comparisons between selfing species and their self-incompatible relatives, which makes it hard to distinguish causal from secondary mutations. To by-pass this problem, we inferred the genetic basis of the loss of self-incompatibility by intercrossing plants from twelve geographically interspersed outcrossing and selfing populations of North-American Arabidopsis lyrata and determining the breeding system of 1,580 progeny. Self-incompatibility was not restored after crosses between different self-compatible populations. Equal frequencies of self-compatible and self-incompatible progeny emerged from crosses between parents with different breeding systems. We propose a two-locus genetic model for the loss of self-incompatibility in which specific S- locus haplotypes ( S1 and S19 ) are associated with loss of self-incompatibility through their interaction with an unlinked modifier.
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