title: The sunflower downy mildew pathogen Plasmopara halstedii.
abstract: Downy mildew of sunflower is caused by Plasmopara halstedii (Farlow) Berlese & de Toni. Plasmopara halstedii is an obligate biotrophic oomycete pathogen that attacks annual Helianthus species and cultivated sunflower, Helianthus annuus. Depending on the sunflower developmental stage at which infection occurs, the characteristic symptoms range from young seedling death, plant dwarfing, leaf bleaching and sporulation to the production of infertile flowers. Downy mildew attacks can have a great economic impact on sunflower crops, and several Pl resistance genes are present in cultivars to protect them against the disease. Nevertheless, some of these resistances have been overcome by the occurrence of novel isolates of the pathogen showing increased virulence. A better characterization of P. halstedii infection and dissemination mechanisms, and the identification of the molecular basis of the interaction with sunflower, is a prerequisite to efficiently fight this pathogen. This review summarizes what is currently known about P. halstedii, provides new insights into its infection cycle on resistant and susceptible sunflower lines using scanning electron and light microscopy imaging, and sheds light on the pathogenicity factors of P. halstedii obtained from recent molecular data.
  
  
  METHODS
  Kingdom Stramenopila; Phylum Oomycota; Class Oomycetes; Order Peronosporales; Family Peronosporaceae; Genus Plasmopara; Species Plasmopara halstedii.
  
  
  METHODS
  Sunflower seedling damping off, dwarfing of the plant, bleaching of leaves, starting from veins, and visible white sporulation, initially on the lower side of cotyledons and leaves. Plasmopara halstedii infection may severely impact sunflower seed yield.
  In spring, germination of overwintered sexual oospores leads to sunflower root infection. Intercellular hyphae are responsible for systemic plant colonization and the induction of disease symptoms. Under humid and fresh conditions, dissemination structures are produced by the pathogen on all plant organs to release asexual zoosporangia. These zoosporangia play an important role in pathogen dissemination, as they release motile zoospores that are responsible for leaf infections on neighbouring plants.
  
  
  RESULTS
  Disease control is obtained by both chemical seed treatment (mefenoxam) and the deployment of dominant major resistance genes, denoted Pl. However, the pathogen has developed fungicide resistance and has overcome some plant resistance genes. Research for more sustainable strategies based on the identification of the molecular basis of the interaction are in progress.
  
  
  BACKGROUND
  http://www.heliagene.org/HP, http://lipm-helianthus.toulouse.inra.fr/dokuwiki/doku.php?id=start, https://www.heliagene.org/PlasmoparaSpecies (soon available).
authors.affiliation: INRA, Laboratoire des Interactions Plantes-Microorganismes (LIPM), UMR441, F-31326, Castanet-Tolosan, France; CNRS, Laboratoire des Interactions Plantes-Microorganismes (LIPM), UMR2594, F-31326, Castanet-Tolosan, France.
authors.forename: Quentin
authors.initials: Q
authors.lastname: Gascuel
authors.forename: Yves
authors.initials: Y
authors.lastname: Martinez
authors.forename: Marie-Claude
authors.initials: MC
authors.lastname: Boniface
authors.forename: Felicity
authors.initials: F
authors.lastname: Vear
authors.forename: Magalie
authors.initials: M
authors.lastname: Pichon
authors.forename: Laurence
authors.initials: L
authors.lastname: Godiard
mesh_terms: D005659:Fungicides, Industrial / Q000494:pharmacology; D006368:Helianthus / Q000382:microbiology*; D009868:Oomycetes / Q000472:pathogenicity*; D010935:Plant Diseases / Q000382:microbiology / Q000517:prevention & control; D014774:Virulence
chemical_list: D005659:Fungicides, Industrial
keywords: Helianthus annuus; Plasmopara halstedii; effectors; infection modes; life cycle; obligate biotroph oomycete; pathogen virulence