Phoma stem canker is an internationally important disease of oilseed rape (Brassica napus, canola, rapeseed), causing serious losses in Europe, Australia and North America. UK losses of e56M per season are estimated using national disease survey data and a yield loss formula. Phoma stem canker pathogen populations comprise two main species, Leptosphaeria maculans, associated with damaging stem base cankers, and Leptosphaeria biglobosa, often associated with less damaging upper stem lesions. Both major gene and quantitative trait loci mediated resistance to L. maculans have been identified in B. napus, but little is known about resistance to L. biglobosa. Leptosphaeria maculans, which has spread into areas in North America and eastern Europe where only L. biglobosa was previously identified, now poses a threat to large areas of oilseed rape production in Asia. Epidemics are initiated by air-borne ascospores; major gene resistance to initial infection by L. maculans operates in the leaf lamina of B. napus. It is not clear whether the quantitative trait loci involved in the resistance to the pathogen that can be assessed only at the end of the season operate in the leaf petioles or stems. In countries where serious phoma stem canker epidemics occur, a minimum standard for resistance to L. maculans is included in national systems for registration of cultivars. This review provides a background to a series of papers on improving strategies for managing B. napus resistance to L. maculans, which is a model system for studying genetic interactions between hemibiotrophic pathogens and their hosts.
Summary
LepR3, found in the Brassica napus cv ‘Surpass 400’, provides race‐specific resistance to the fungal pathogen Leptosphaeria maculans, which was overcome after great devastation in Australia in 2004. We investigated the LepR3 locus to identify the genetic basis of this resistance interaction.
We employed a map‐based cloning strategy, exploiting collinearity with the Arabidopsis thaliana and Brassica rapa genomes to enrich the map and locate a candidate gene. We also investigated the interaction of LepR3 with the L. maculans avirulence gene AvrLm1 using transgenics.
LepR3 was found to encode a receptor‐like protein (RLP). We also demonstrated that avirulence towards LepR3 is conferred by AvrLm1, which is responsible for both the Rlm1 and LepR3‐dependent resistance responses in B. napus.
LepR3 is the first functional B. napus disease resistance gene to be cloned. AvrLm1's interaction with two independent resistance loci, Rlm1 and LepR3, highlights the need to consider redundant phenotypes in ‘gene‐for‐gene’ interactions and offers an explanation as to why LepR3 was overcome so rapidly in parts of Australia.
The most common and effective way to control phoma stem canker (blackleg) caused by Leptosphaeria maculans in oilseed rape (Brassica napus) is through the breeding of resistant cultivars. Race specific major genes that mediate resistance from the seedling stage have been identified in B. napus or have been introgressed from related species. Many race specific major genes have been described and some of them are probably identical in B. napus (allotetraploid AACC) and the parental species B. rapa (diploid AA). More work is needed using a set of well-characterised isolates to determine the number of different major resistance genes available. In some B. napus cultivars, there is resistance which is polygenic (mediated by Quantitative Trait Loci) and postulated to be race non-specific. Many of these major genes and Quantitative Trait Loci for resistance to L. maculans have been located on B. napus genetic maps. Genes involved in race specific and polygenic resistance are generally distinct.
SUMMARY The fungal pathogen Sclerotinia sclerotiorum infects a broad range of dicotyledonous plant species and causes stem rot in Brassica napus. To elucidate the mechanisms underlying the defence response, the patterns of gene expression in the partially resistant B. napus cultivar ZhongYou 821 (ZY821) and the susceptible cultivar Westar were studied using a B. napus oligonucleotide microarray. Although maximum differential gene expression was observed at 48 h post-inoculation (hpi) in both cultivars, increased transcript levels were detected in cv. ZY821 at the earlier stages of infection (6-12 hpi) for many genes, including those encoding defence-associated proteins, such as chitinases, glucanases, osmotins and lectins, as well as genes encoding transcription factors belonging to the zinc finger, WRKY, APETALA2 (AP2) and MYB classes. In both cultivars, genes encoding enzymes involved in jasmonic acid, ethylene and auxin synthesis were induced, as were those for gibberellin degradation. In addition, changes in the expression of genes encoding enzymes involved in carbohydrate and energy metabolism appeared to be directed towards shuttling carbon reserves to the tricarboxylic acid cycle and generating reactive oxygen species. Transcripts from genes encoding enzymes involved in glucosinolate and phenylpropanoid biosynthesis were highly elevated in both cultivars, suggesting that secondary metabolites are also components of the response to S. sclerotiorum in B. napus.
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