title: The R1 resistance gene cluster contains three groups of independently evolving, type I R1 homologues and shows substantial structural variation among haplotypes of Solanum demissum.
abstract: Cultivated and wild potatoes contain a major disease-resistance cluster on the short arm of chromosome V, including the R1 resistance (R) gene against potato late blight. To explore the functional and evolutionary significance of clustering in the generation of novel disease-resistance genes, we constructed three approximately 1 Mb physical maps in the R1 gene region, one for each of the three genomes (haplotypes) of allohexaploid Solanum demissum, the wild potato progenitor of the R1 locus. Totals of 691, 919 and 559 kb were sequenced for each haplotype, and three distinct resistance-gene families were identified, one homologous to the potato R1 gene and two others homologous to either the Prf or the Bs4 R-gene of tomato. The regions with R1 homologues are highly divergent among the three haplotypes, in contrast to the conserved flanking non-resistance gene regions. The R1 locus shows dramatic variation in overall length and R1 homologue number among the three haplotypes. Sequence comparisons of the R1 homologues show that they form three distinct clades in a distance tree. Frequent sequence exchanges were detected among R1 homologues within each clade, but not among those in different clades. These frequent sequence exchanges homogenized the intron sequences of homologues within each clade, but did not homogenize the coding sequences. Our results suggest that the R1 homologues represent three independent groups of fast-evolving type I resistance genes, characterized by chimeric structures resulting from frequent sequence exchanges among group members. Such genes were first identified among clustered RGC2 genes in lettuce, where they were distinguished from slow-evolving type II R-genes. Our findings at the R1 locus in S. demissum may indicate that a common or similar mechanism underlies the previously reported differentiation of type I and type II R-genes and the differentiation of type I R-genes into distinct groups, identified here.
authors.affiliation: Plant Gene Expression Center, USDA-ARS and Department of Plant and Microbial Biology, University of California, Berkeley, 800 Buchanan Street, Albany, CA 94710, USA.
authors.forename: Hanhui
authors.initials: H
authors.lastname: Kuang
authors.forename: Fusheng
authors.initials: F
authors.lastname: Wei
authors.forename: María Rosa
authors.initials: MR
authors.lastname: Marano
authors.forename: Uwe
authors.initials: U
authors.lastname: Wirtz
authors.forename: Xiaoxue
authors.initials: X
authors.lastname: Wang
authors.forename: Jia
authors.initials: J
authors.lastname: Liu
authors.forename: Wai Pun
authors.initials: WP
authors.lastname: Shum
authors.forename: Jennifer
authors.initials: J
authors.lastname: Zaborsky
authors.forename: Luke J
authors.initials: LJ
authors.lastname: Tallon
authors.forename: Willem
authors.initials: W
authors.lastname: Rensink
authors.forename: Stacey
authors.initials: S
authors.lastname: Lobst
authors.forename: Peifen
authors.initials: P
authors.lastname: Zhang
authors.forename: Carl-Erik
authors.initials: CE
authors.lastname: Tornqvist
authors.forename: Ahmet
authors.initials: A
authors.lastname: Tek
authors.forename: John
authors.initials: J
authors.lastname: Bamberg
authors.forename: John
authors.initials: J
authors.lastname: Helgeson
authors.forename: William
authors.initials: W
authors.lastname: Fry
authors.forename: Frank
authors.initials: F
authors.lastname: You
authors.forename: Ming-Cheng
authors.initials: MC
authors.lastname: Luo
authors.forename: Jiming
authors.initials: J
authors.lastname: Jiang
authors.forename: C
authors.initials: C
authors.lastname: Robin Buell
authors.forename: Barbara
authors.initials: B
authors.lastname: Baker
mesh_terms: D022202:Chromosomes, Artificial, Bacterial / Q000235:genetics; D003001:Cloning, Molecular; D017124:Conserved Sequence / Q000235:genetics; D019143:Evolution, Molecular; D017343:Genes, Plant / Q000235:genetics*; D014644:Genetic Variation / Q000235:genetics*; D006239:Haplotypes; D007438:Introns / Q000235:genetics; D008969:Molecular Sequence Data; D005810:Multigene Family / Q000235:genetics*; D010802:Phylogeny; D010935:Plant Diseases / Q000235:genetics*; D010940:Plant Proteins / Q000235:genetics; D032322:Solanum / Q000235:genetics* / Q000378:metabolism; D012997:Solvents
chemical_list: D010940:Plant Proteins; D012997:Solvents