Contrasting evolutionary patterns of the Rp1 resistance gene family in different species of Poaceae.

Disease-resistance genes (R-genes) in plants show complex evolutionary patterns. We investigated the evolution of the Rp1 R-gene family in Poaceae, and 409 Rp1 fragments were sequenced from 21 species. Our data showed that the common ancestor of Poaceae had two Rp1 loci, but the number of Rp1 locus in extant species varies from one to five. Some wheat and Zea genotypes have dozens of Rp1 homologues in striking contrast to one or two copies in Brachypodium distachyon. The large number of diverse Rp1 homologues in Zea was the result of duplications followed by extensive sequence exchanges among paralogues, and all genes in maize have evolved in a pattern of Type I R-genes. The high frequency of sequence exchanges did not cause concerted evolution in Zea species, but concerted evolution was obvious between Rp1 homologues from genera Zea and Sorghum. Differentiation of Type I and Type II Rp1 homologues was observed in Oryza species, likely occurred in their common ancestor. One member (Type II R-gene) in the Oryza Rp1 cluster did not change sequences with its paralogues, whereas the other paralogues (Type I R-genes) had frequent sequence exchanges. The functional Pi37 resistance gene in rice was generated through an unequal crossover between two neighboring paralogues followed by four point mutations. The Rp1 homologues in wheat and barley were most divergent, probably due to lack of sequence exchanges among them. Our results shed more light on R-gene evolution, particularly on the differentiation of Type I and Type II R-genes.