No abstract

Tobacco was transformed with three different alleles (L2, L6, and L10) of the flax rust resistance gene L, a member of the toll interleukin-1 receptor, nucleotide-binding site, leucine-rich repeat (TIR-NBS-LRR) class of plant disease resistance genes. L6 transgenics had a stunted phenotype, expressed several defense response genes constitutively, and had increased resistance to the fungus Cercospora nicotianae and the oomycete Phytophthora parasitica pv. nicotianae. L2 and L10 transgenics, with one exception for L10, did not express these phenotypes, indicating that the activation of tobacco defense responses is L6 allele-specific. The phenotype of the exceptional L10 transgenic plant was associated with the presence of a truncated L10 gene resulting from an aberrant T-DNA integration. The truncated gene consisted of the promoter, the complete TIR region, and 39 codons of the NBS domain fused inframe to a tobacco retrotransposon-like sequence. A similar truncated L10 gene, constructed in vitro, was transiently expressed in tobacco leaves and gave rise to a strong localized necrotic reaction. Together, these results suggest that defense signaling properties of resistance genes can be expressed in an allele-specific and pathogen-independent manner when transferred between plant genera.

The L6 and M rust-resistance genes, representing two of the five rust-resistance gene loci in flax (Linum usitatissimum), have been cloned. The molecular data are fully consistent with earlier genetic data: the L locus is a single gene with multiple alleles expressing different rust resistance specificities, and the M locus is complex, containing an array of about 15 similar genes. Thus, while L6 and M resistance genes have 86% nucleotide identity, their locus structure is very different. These genes encode products belonging to the nucleotide binding site-leucine-rich repeat class of disease-resistance proteins. Analysis of alleles from the L locus and chimeric genes is providing evidence suggesting that important specificity determinants occur in the C-terminal half of the proteins, the region containing the leucine-rich repeats. The isolation and characterization of the rust (Melampsora lini) avirulence genes that correspond to the cloned rust-resistance genes is one of the major challenges remaining to the understanding of this system.