A new avirulence gene of Leptosphaeria maculans, AvrLm14, identifies a resistance source in American broccoli (Brassica oleracea) genotypes.

In many cultivated crops, sources of resistance to diseases are sparse and rely on introgression from wild relatives. Agricultural crops often are allopolyploids resulting from interspecific crosses between related species, which are sources of diversity for resistance genes. This is the case for Brassica napus (oilseed rape, canola), an interspecific hybrid between Brassica rapa (turnip) and Brassica oleracea (cabbage). B. napus has a narrow genetic basis and few effective resistance genes against stem canker (blackleg) disease, caused by the fungus Leptosphaeria maculans, are currently available. B. rapa diversity has proven to be a valuable source of resistance (Rlm, LepR) genes, while B. oleracea genotypes were mostly considered susceptible. Here we identified a new resistance source in B. oleracea genotypes from America, potentially effective against French L. maculans isolates under both controlled and field conditions. Genetic analysis of fungal avirulence and subsequent cloning and validation identified a new avirulence gene termed AvrLm14 and suggested a typical gene-for-gene interaction between AvrLm14 and the postulated Rlm14 gene. AvrLm14 shares all the usual characteristics of L. maculans avirulence genes: it is hosted in a genomic region enriched in transposable elements and heterochromatin marks H3K9me3, its expression is repressed during vegetative growth but shows a strong overexpression 5-9 days following cotyledon infection, and it encodes a small secreted protein enriched in cysteine residues with few matches in databases. Similar to the previously cloned AvrLm10-A, AvrLm14 contributes to reduce lesion size on susceptible cotyledons, pointing to a complex interplay between effectors promoting or reducing lesion development.

Early evolution of endoparasitic group in powdery mildews: molecular phylogeny suggests missing link between Phyllactinia and Leveillula.

Of the 17 genera of the Erysiphaceae, only four genera (viz. Leveillula, Phyllactinia, Pleochaeta and Queirozia) exhibit (partly) endoparasitism. To investigate early evolution of this endoparasitic nature, we performed molecular phylogenetic analyses of powdery mildews belonging to the tribe Phyllactinieae collected in North and South America. The most ancestral taxa in the tribe Phyllactinieae belong to the Pleochaeta/Queirozia group, from which the genus Phyllactinia was derived. Finally, the truly endoparasitic genus Leveillula emerged from a part of Phyllactinia The present study showed clear evolutional polarity in the powdery mildews concerned (that is, partly endoparasitic group evolved from ectoparasitic group) and then a truly endoparasitic group emerged from a partly endoparasitic group. In addition, a group with distinctly dimorphic conidia proved to be basal in the Phyllactinieae, and a group without distinctly dimorphic conidia was derived from that group. The present analyses clearly showed that Leveillula derived from a part of the "Basal Phyllactinia group". However, all sister taxa to Leveillula were distributed in North and South America. Because the putative geographic origin of Leveillula is assumed to be Central and Western Asia or the Mediterranean region, we postulate a missing link during the evolution of Leveillula from Phyllactinia Based on the present phylogenetic studies and the new rules of the International Code of Nomenclature for algae, fungi, and plants (McNeill et al. 2012), the following new species and taxonomic re-allocations are proposed: Phyllactinia bougainvilleae sp. nov., Ph. caricae comb. nov., Ph. caricicola comb. nov., Ph. durantae comb. nov., Ph. leveilluloides sp. nov., Ph. obclavata comb. nov., and Ph. papayae comb. nov.

Foliicolous fungi from Arctostaphylos pungens in Mexico.

Arctostaphylos pungens "Manzanita" is an important shrub in the southwestern USA, and northern and central Mexico. Manzanita bears apple-like fruit that is utilised for a range of edible products. Over the past two years, several foliar disease problems were noted on this host in the San José de Gracia region of Mexico. The aim of the present study was to elucidate their identity through the analysis of morphological characters and DNA phylogeny (based on the large subunit nuclear ribosomal RNA gene and the ITS spacers and the intervening 5.8S rRNA gene of the nrDNA operon) of the fungi associated with these disease symptoms. Three species are newly described: Phaeococcomyces mexicanus sp. nov., a presumed epiphyte, and two species associated with leaf spots and defoliation, namely Coccomyces arctostaphyloides sp. nov. and Passalora arctostaphyli sp. nov. A fourth species is also associated with leaf spots and tip dieback is Harknessia arctostaphyli, for which an epitype is designated. All species can co-occur on the same shrub, which adds to the stress experienced by the plant, leading to further defoliation and dieback.

Genome structure impacts molecular evolution at the AvrLm1 avirulence locus of the plant pathogen Leptosphaeria maculans.

Leptosphaeria maculans, a dothideomycete fungus causing stem canker on oilseed rape, develops gene-for-gene interactions with its host plants. It has the ability to rapidly adapt to selection pressure exerted by cultivars harbouring novel resistance genes as exemplified recently by the 3-year evolution towards virulence at the AvrLm1 locus in French populations. The AvrLm1 avirulence gene was recently cloned and shown to be a solo gene within a 269 kb non-coding, heterochromatin-like region. Here we describe the sequencing of the AvrLm1 genomic region in one avirulent and two virulent isolates to investigate the molecular basis of evolution towards virulence at the AvrLm1 locus. For these virulent isolates, the gain of virulence was linked to a 260 kb deletion of a chromosomal segment spanning AvrLm1 and deletion breakpoints were identical or similar. Among the 460 isolates analysed from France, Australia and Mexico, a similar large deletion was apparent in > 90% of the virulent isolates. Deletion breakpoints were also strongly conserved in most of the virulent isolates, which led to the hypothesis that a unique deletion event leading to the avrLm1 virulence has diffused in pathogen populations. These data finally suggest that retrotransposons are key drivers in genome evolution and adaptation to novel selection pressure in L. maculans.