Resistance May Not Be Futile: Recently, Ug99, a particularly devastating strain of wheat stem rust fungus, has emerged, which could potentially threaten food security. Now, two genes have been cloned that offer resistance to Ug99. Saintenac et al. (p. 783 , published online 27 June) cloned Sr35 from Triticum monococcum , a diploid wheat species not often cultivated. Periyannan et al. (p. 786 , published online 27 June) cloned Sr33 from Aegilops tauschii , a diploid wild grass that contributed to the hexaploid genome of cultivated wheat. The genes both encode proteins that show features typical of other disease resistance proteins and offer opportunities to slow the pace of Ug99 progression.

Background: Bread wheat is an allopolyploid species with a large, highly repetitive genome. To investigate the impact of selection on variants distributed among homoeologous wheat genomes and to build a foundation for understanding genotype-phenotype relationships, we performed population-scale re-sequencing of a diverse panel of wheat lines. Results: A sample of 62 diverse lines was re-sequenced using the whole exome capture and genotyping-by-sequencing approaches. We describe the allele frequency, functional significance, and chromosomal distribution of 1.57 million single nucleotide polymorphisms and 161,719 small indels. Our results suggest that duplicated homoeologous genes are under purifying selection. We find contrasting patterns of variation and inter-variant associations among wheat genomes; this, in addition to demographic factors, could be explained by differences in the effect of directional selection on duplicated homoeologs. Only a small fraction of the homoeologous regions harboring selected variants overlapped among the wheat genomes in any given wheat line. These selected regions are enriched for loci associated with agronomic traits detected in genome-wide association studies. Conclusions: Evidence suggests that directional selection in allopolyploids rarely acted on multiple parallel advantageous mutations across homoeologous regions, likely indicating that a fitness benefit could be obtained by a mutation at any one of the homoeologs. Additional advantageous variants in other homoelogs probably either contributed little benefit, or were unavailable in populations subjected to directional selection. We hypothesize that allopolyploidy may have increased the likelihood of beneficial allele recovery by broadening the set of possible selection targets.

Stem rust susceptibility of European wheats under Australian conditions posed a significant threat to wheat production for the early British settlers in Australia. The famous Australian wheat breeder, William Farrer, tackled the problem of stem rust susceptibility through breeding fast-maturing wheat cultivars. South-eastern Australia suffered a severe stem rust epidemic in 1973, which gave rise to a national approach to breeding for rust resistance. The National Wheat Rust Control Program was set up in 1975, modelled on the University of Sydney’s own rust resistance breeding program, at the University of Sydney Plant Breeding Institute, Castle Hill (now Cobbitty). Back-crossing of a range of sources of resistance provided genetically diverse germplasm for evaluation in various breeding programs. Current efforts are directed to building gene combinations through marker-assisted selection. Major genes for resistance to stem rust and leaf rust are being used in the back-crossing program of the ACRCP to create genetic diversity among Australian germplasm. Stripe rust and to a lesser extent leaf rust resistance in the Australian germplasm is largely based on combinations of adult plant resistance genes and our knowledge of their genomic locations has increased. Additional genes, other than Yr18/Lr34 and Yr29/Lr46, appeared to control adult plant resistance to both leaf rust and stripe rust. Two adult-plant stem rust resistance genes have also been identified. The development of selection technologies to achieve genotype-based selection of resistance gene combinations in the absence of bioassays has evolved in the last 5 years. Robust molecular markers are now available for several commercially important rust resistance genes. Marker-assisted selection for rust resistance is performed routinely in many wheat-breeding programs. Modified pedigree and limited back-cross methods have been used for breeding rust-resistant wheat cultivars in the University of Sydney wheat-breeding program. The single back-cross methodology has proved more successful in producing cultivars with combinations of adult plant resistance genes.