Downy mildew of Arabidopsis thaliana caused by Hyaloperonospora parasitica (formerly Peronospora parasitica).

UNLABELLED: SUMMARY Downy mildew of Arabidopsis is not a hugely destructive disease of an important crop plant, neither is it of any economic importance. The most obvious symptom, the aerial conidiophores, might, at a glance to the casual observer, be mistaken for the trichomes normally present on the leaves. However, a huge research effort is being devoted to this humble pathosystem which became established as a laboratory model in the 1990s. Since then, enormous progress has been made in cloning and characterizing major genes for resistance (RPP genes) and in defining many of their downstream signalling components, some of them RPP-gene specific. Resistance is generally associated with an oxidative burst and a salicylic acid dependent hypersensitive reaction type of programmed cell death. Biological and chemical induction of systemic acquired resistance (SAR) in Arabidopsis protecting against downy mildew were demonstrated early on, and investigations of mutants have contributed fundamentally to our understanding of host-pathogen interactions and the mechanisms of plant defence. This review will attempt to collate the wealth of information which has accrued with this pathosystem in the last decade and will attempt to predict future research directions by drawing attention to some still unanswered questions.

TAXONOMY: Hyaloperonospora Constant. parasitica (Pers.:Fr) Fr. (formerly Peronospora parasitica), Kingdom Chromista, Phylum Oomycota, Order Peronosporales, Family Peronosporaceae, Genus Hyaloperonospora, of which it is the type species. The taxonomy of the group of organisms causing downy mildew of brassicas has undergone a number of revisions since Corda (1837) originally coined the genus Peronospora. All isolates pathogenic on brassicas were described initially as P. parasitica but Gäumann (1918) classified isolates from different brassicaceous hosts distinctly and thus defined 52 new species based on conidial dimensions and host range. After much debate it was decided to revert to the aggregate species of P. parasitica for all brassica-infecting downy mildews, whilst recognizing that these show some isolate-specific differences (Yerkes and Shaw, 1959). The latest re-examination of P. parasitica by Constantinescu and Fatehi (2002) has placed isolates of P. parasitica and five other downy mildew species in a clear new subgroup on the basis of their hyaline conidiospores, recurved conidiophore branch tips and ITS1, ITS2 and 5.8S rDNA sequence comparisons; meriting the coining of the new genus 'Hyaloperonospora Constant'. The class Oomycetes in the Kingdom Chromista (Straminipila) comprises fungus-like organisms with heterokont zoospores (i.e. possessing two types of flagellae, whiplash and tinsel). The Oomycetes have non-septate hyphae with cellulose-based walls containing very little or no chitin. The latter is regarded as a major distinction separating the Oomycetes from the true fungi, and reports of the presence of chitin had generally been regarded as due to small amounts of contamination (Gams et al., 1998). However, in view of recent studies by Werner et al. (2002) showing a chitin synthase gene in an Oomycete and demonstrating the presence of the polymer itself by an interaction with wheat germ agglutinin (WGA), it is perhaps safe to say that we have not seen the last taxonomic revision which will affect this group! The families within the Oomycetes show a clear evolutionary trend to a lesser absolute dependence on an aqueous environment and some members of the Peronosporales, e.g. H. parasitica, have no zoosporic stage in the life cycle.

HOST RANGE: Isolates infecting Arabidopsis thaliana have so far proven to be non-pathogenic on other crucifers tested but exist in a clear gene-for-gene relationship with different host ecotypes. Disease symptoms: Infections are first apparent to the naked eye as a carpet or 'down' of conidiophores covering the upper and lower surfaces of leaves and petioles. This symptom is characteristic of this group of diseases and lends it its name.

USEFUL WEBSITES: (links to references on Oomycetes), (TAIR, The Arabidopsis Information Resource).