Rpg1-mediated durable stem rust resistance: mechanisms of action
Department of Crop and Soil Sciences, Washington State University, USA
View klienhofs_2011.pdf (221.38 KB)
Stem rust, caused by Puccinia graminis f. sp. tritici, is a devastating disease on wheat and barley. A single barley gene, Rpg1, has provided durable resistance since its commercial introduction in the 1940s. The cloned Rpg1 gene encodes a protein with two tandem protein kinase domains, one an active kinase (pK2) and one a pseudokinase (pK1). Function of both domains is required for resistance. The gene is constitutively expressed in all tissues with elevated levels in the epidermis. It is mostly cytoplasmic with small, but significant amount associated with the cell membrane. We have been studying this gene and protein to try to understand how it works and why it has been so durable. Here we report our most recent results showing that RPG1 is phosphorylated within 5 min after urediniospores from avirulent, but not virulent, races land on the leaf surface. Two effector proteins were isolated from the ungerminated spores and shown to work cooperatively to induce RPG1 phosphorylation and eventual degradation. The proteins were identified as a hypothetical protein (PGTG10537.2) with a fibronectin type III and BRCA1 C-terminal domains and vacuolar protein sortingassociated protein 9 (PGTG_16791). The rapidity of the effector function and the nature of the two protein effectors indicate that a unique mechanism for effector entry and signaling in the host cell is involved. This hypothetical mechanism may be similar to what is observed in animal cells where fibronectin proteins with an RGD-binding domain act to mediate communications between the extracellular matrix and plasma membrane.
Towards an understanding of the molecular mechanisms of durable and non-durable resistance to stripe rust in wheat
USDA-ARS Wheat Genetics, Quality, Physiology and Disease Research Unit, Pullman, WA
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Stripe rust of wheat, caused by Puccinia striiformis f. sp. tritici, continues to cause severe damage worldwide. Durable resistance is a key for sustainable control of the disease. High-temperature adult-plant (HTAP) resistance, which expresses when the weather becomes warm and plants grow old, has been demonstrated to be durable. We have conducted numerous of studies for understanding molecular mechanisms of different types of stripe rust resistance using a transcriptomics approach. Through comparing gene expression patterns with racespecific, all-stage resistance controlled by various genes, we found that a greater diversity of genes is involved in HTAP resistance. The genes involved in HTAP resistance are induced more slowly and their expression induction is less dramatic than genes involved in all-stage resistance. The high diversity of genes and less dramatic expression induction may explain the durability and incomplete level of HTAP resistance. Identification of transcripts may be helpful in identifying resistance controlled by different genes and in selecting better combinations of genes for pyramiding to achieve adequate and more durable resistance.
Unraveling the entry mechanism of oomycete and fungal effector proteins into host cells
Shiv D. Kale
Virginia Bioinformatics Institute, Virginia Tech University, USA
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Oomycetes and fungi facilitate pathogenesis via secretion of effector proteins that have apoplastic and intracellular localizations. These effector proteins have a diverse array of functions that aid in pathogenesis, including modification of defense responses. In the oomycetes, well characterized effector proteins that can translocate into the host cells share a pair of conserved N-terminal motifs known as RXLR and dEER. The RXLR motif has been shown to mediate translocation of the oomycete avirulence proteins Avr1b and Avr3a into host cells. Detailed mutagenesis of the RXLR motif of Avr1b revealed that the motif is tolerant to several amino acid substitutions while retaining functional translocation activity, resulting in the definition of a broadened RXLR-like motif, [R,K,H] X[L/M/I/F/Y/W]X. This motif has been used to identify functional translocation motifs in several fungal effector proteins, AvrL567, Avr2, and AvrLm6. Effectors with both RXLR and RXLR-like motifs bind phosphatidylinositol- 3-phosphate (PI-3-P) to mediate translocation via lipid raft mediated endocytosis. Mutations in RXLR or RXLRlike motifs result in loss of phospholipid binding and translocation by effectors. Effector entry into plant cells can be blocked by proteins and inositides that disrupt binding to PI-3-P, suggesting effector-blocking technologies that could be used in agriculturally important plant species.
Investigating rust resistance with the model grass Brachypodium
USDA-ARS Plant Science Research Unit and Department of Agronomy and Plant Genetics, University of Minnesota, USA
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The model plant Arabidopsis thaliana has provided unique opportunities to explore and unravel many key biological features of plant biology including disease resistance. However, the inability of rust fungi of the genus Puccinia to infect Arabidopsis has prevented its use in exploring grass-rust interactions. The model plant Brachypodium distachyon is a member of the same grass subfamily as the principal cool-season grain crops, and can be infected with various Puccinia species. We have focused our efforts on establishing Brachypodium as a model for exploring grass - Puccinia graminis interactions. Brachypodium can be successfully infected by different formae speciales of the stem rust pathogen, including P. graminis f. sp. tritici. A wide range of response to stem rust occurs in Brachypodium and efforts are underway to decipher the genetic basis for this variation using recombinant inbred populations from parents with differing levels of response. Similarly, induced mutants with compromised stem rust resistance have been identified and are now being employed within a program to understand the molecular biology of stem rust resistance and susceptibility. Our results to date suggest that Brachypodium holds promise as a model plant for advancing our understanding of stem rust resistance.
International surveillance of wheat rust pathogens - progress and challenges
The University of Sydney, Plant Breeding Institute, Australia
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Surveillance of wheat rust pathogens, including assessments of rust incidence and virulence characterization via either trap plots or race (pathotype) surveys, has provided information fundamental in formulating and adopting appropriate national and international policies, investments and strategies in plant protection, plant breeding, seed systems, and in rust pathogen research. Despite many successes from national and regional co-ordination of rust surveillance, few attempts were made to extend rust surveillance to international or even global levels. The Global Cereal Rust Monitoring System was established to address this deficiency. It is underpinned by an information platform that includes standardized protocols for methods and systems used in surveys, preliminary virulence testing, data, sample transmission and management at the field and national and global levels, and includes two web-based visualization tools. While considerable progress has been made towards a global system for monitoring variability in the wheat stem rust pathogen, and linking this to the threat posed by this pathogen to regional wheat production, some challenges remain, including ongoing commitment to support rust surveillance, and the ability to share and compare surveillance data.
Genetic protection of wheat from rusts and development of resistant varieties in Russia and Ukraine
View morgunov_2010.pdf (677.32 KB)
Leaf rust represents the major threat to wheat production in Russia and Ukraine. It has been present for many years and epidemics occur in different regions on both winter and spring wheat. In some regions there is evidence of more frequent epidemics, probably due to higher precipitation as a result of climate change. There is evidence that the virulence of the leaf rust population in Ukraine and European Russia and on winter wheat and spring wheat is similar. The pathogen population structure in Western Siberia is also similar to the European part, although there are some significant differences based on the genes employed in different regions. Ukrainian wheat breeders mostly rely on major resistance genes from wide crosses and have succeeded in developing resistant varieties. The North Caucasus winter wheat breeding programs apply the strategy of deploying varieties with different types of resistance and genes. This approach resulted in decreased leaf rust incidence in the region. Genes Lr23 and Lr19 deployed in spring wheat in the Volga region were rapidly overcome by the pathogen. There are continuing efforts to incorporate resistance from wild species. The first leaf rust resistant spring wheat varieties released in Western Siberia possessed gene LrTR which protected the crop for 10-15 years, but was eventually broken in 2007. Slow rusting is being utilized in several breeding programs in Russia and Ukraine, but has not become a major strategy.
Global status of stripe rust
The University of Sydney, Plant Breeding Institute, Australia
View wellings_2010.pdf (283.25 KB)
Stripe rust, caused by Puccinia striiformis, has been an important disease of wheat, barley, rye, triticale and certain graminaceous hosts for centuries. The significance of the disease on cultivated cereals has waxed and waned according to the vagaries of climate, inoculum levels and susceptible varieties. A progressive understanding of pathogen biology has revealed levels of specialisation between and within host groups, and these had varying impacts on the hosts concerned. The most economically important form is P. striiformis f. sp. tritici (Pst), the causal pathogen of stripe (yellow) rust of wheat, which is the major focus of this paper. The recent discovery of the perfect stage of Pst on Berberis spp. will encourage further work to uncover the potential importance of the sexual stage in pathogen biology in regions where Berberis spp. occur. A review of the evolution of pathotypes within Pst over the past 50 years reveals recurrent pandemics emanating from a combination of specific virulence in the pathogen population, wide scale cultivation of genetically similar varieties, and agronomic practices that led to high yield potential. When these factors operate in concert, regional stripe rust epidemics have proven to be dramatic, extensive and serious in terms of the magnitude of losses and the economic hardships endured. A review of these epidemics suggests that little progress has been made in containing the worst effects of epidemics. The current status of stripe rust was gauged from a survey of 25 pathologists and breeders directly associated with the disease. It was evident that Pst remains a significant threat in the majority of wheat growing regions of the world with potential to inflict regular regional crop losses ranging from 0.1 to 5%, with rare events giving losses of 5 to 25%. Regions with current vulnerability include the USA (particularly Pacific North West), East Asia (China north-west and south-west), South Asia (Nepal), Oceania (Australia) and East Africa (Kenya). The resources deployed to contain the worst effects of Pst will need to find a balance between training a new generation of breeders and pathologists in host-pathogen genetics, and an investment in infrastructure in IARCs and NARs.
Status of wheat rust research and control in China
College of Plant Protection, Northwest A&F University, P.R. China
View kang_2010.pdf (284.43 KB)
In China, wheat is grown on approximately 24 million hectares with an annual yield of 100 million tonnes. Stem rust, caused by Puccinia graminis f. sp. tritici, is a threat mainly to spring wheat in northeastern China. Leaf rust, caused by P. triticina, occurs on crops in the late growth stages in the Yellow-Huai-Hai River regions. Stripe rust, caused by P. striiformis f. sp. tritici (Pst), is destructive in all winter wheat regions and is considered the most important disease of wheat in China. During the last 20 years, widespread stripe rust epidemics occurred in 2002, 2003, and 2009, and localized epidemics occurred in many other years. In recent years, major yield losses were prevented by widespread and timely applications of fungicides based on accurate monitoring and prediction of disease epidemics. A total of 68 Pst races or pathotypes have been identified using a set of 19 differential wheat genotypes. At present, races CYR32 and CYR33 virulent to resistance genes Yr9, Yr3b, Yr4b, YrSu and some other resistance genes are predominant. Moreover, these races are virulent on many cultivars grown in recent years. Of 501 recent cultivars and breeding lines 71.9% were susceptible, 7.0% had effective all-stage resistance, mostly Yr26 (= Yr24), and 21.2% had adult-plant resistance. Several resistance genes, including Yr5, Yr10, Yr15, Yr24/Yr26, YrZH84 and some unnamed genes, are still effective against the current Pst population. All have been widely used in breeding programs. Lines with one or more of Yr1, Yr2, Yr3, Yr4, Yr6, Yr7, Yr8, Yr9 and other unnamed resistance genes are susceptible to currently predominant races. Durable adult plant resistance sources are being increasingly used as parents in breeding programs. Progress has been made in genomics and population genetics of Pst, molecular mapping of resistance genes, and cytological and molecular mechanisms of the host-pathogen interactions involved in stripe rust.
Implications of climate change for diseases, crop yields and food security
Scottish Crop Research Institute (SCRI), U.K.
View newton_2010.pdf (531.13 KB)
Accelerated climate change affects components of complex biological interactions differentially, often causing changes that are difficult to predict. Crop yield and quality are affected by climate change directly, and indirectly, through diseases that themselves will change but remain important. These effects are difficult to dissect and model as their mechanistic bases are generally poorly understood. Nevertheless, a combination of integrated modelling from different disciplines and multi-factorial experimentation will advance our understanding and prioritisation of the challenges. Food security brings in additional socio-economic, geographical and political factors. Enhancing resilience to the effects of climate change is important for all these systems and functional diversity is one of the most effective targets for improved sustainability.
Rust-proofing wheat for a changing climate
CSIRO Plant Industry, Australia
View chakaborty_2010.pdf (360.95 KB)
This paper offers projections of potential effects of climate change on rusts of wheat and how we should factor in a changing climate when planning for the future management of these diseases. Even though the rusts of wheat have been extensively studied internationally, there is a paucity of information on the likely effects of a changing climate on the rusts and hence on wheat production. Due to the lack of published empirical research we relied on the few published studies of other plant diseases, our own unpublished work and relevant information from the vast literature on rusts of wheat to prepare this overview. Potential risks from a changing climate were divided into three major groups: increased loss from wheat rusts, new rust races evolving faster and the reduced effectiveness of rust resistances. Increased biomass of wheat crops grown in the presence of elevated CO2 concentrations and higher temperatures will increase the leaf area available for attack by the pathogen. This combined with increased speed of the pathogen’s life cycle, may increase the rate of epidemic development in many environments. Likewise, should the effects of climate change result in more conducive conditions for rust development there will also be a corresponding increase in the rate of evolution of new and presumably virulent races. The effectiveness of some rust resistance genes are influenced by temperature, crop development stage and even nitrogen status of the host. It is likely that direct and indirect changes on the host from climate change may influence the effectiveness of some of these resistance genes. Currently the likely effects of climate change on the effectiveness of disease resistance is not known and since disease resistance breeding is a long term strategy it is important to determine if any of the important genes may become less effective due to climate change. Studies must be made to acquire new information on the rust disease triangle to increase the adaptive capacity of wheat under climate change. BGRI leadership is needed to broker research on rust evolution and the durability of resistance under climate change.