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Peronospora
	P. manshurica on soybean leaf
Scientific classification
Domain:	Eukaryota
Clade:	Diaphoretickes
Clade:	SAR
Clade:	Stramenopiles
Phylum:	Oomycota
Order:	Peronosporales
Family:	Peronosporaceae
Genus:	Peronospora; Corda

Peronospora is a genus of oomycetes that are obligate plant pathogens of many eudicots.^[1] Most species in this group produce a downy mildew disease, which can cause severe damage to many different cultivated crops, as well as wild and ornamental plants.^[2] There are 19 genera that produce downy mildew, and Peronospora has been placed alongside Pseudoperonospora in the group of downy mildews with coloured conidia.^[3] Peronospora has far more species than any other genus of the downy mildews.^[3] However, many species have been moved from this genus to be reclassified to other or new genera.^[4] Among these was the most famous Peronospora species, formerly known as Peronospora parasitica, and now known as Hyaloperonospora parasitica.^[4] Now, the Peronospora species of most importance is likely the Peronospora tabacina.^[5] Peronospora tabacina causes blue mold on tobacco plants and can severely reduce yields of this economically important crop to the point where it has been classified as a bioweapon.^[5]^[3]

History

Peronospora was first described in 1837 by August Carl Joseph Corda, a Czech mycologist and physician, in his first of six volumes of his Icones fungorum hucusque cognitorum.^[6] Since then, many of the species originally placed in Peronospora have been allocated to other genera or given rise to new genera based on new techniques such as molecular genetics.^[4]

There was an epidemic in 1960 of Peronospora tabacina affecting tobacco plants leading to $25 million in losses across eleven countries, which was about 30 percent of the tobacco plants.^[5] Another epidemic that was caused by Peronospora destructor reduced the yield of sweet onions by 25 percent in Georgia, USA in 2012, and led to an estimated $18.2 million in losses.^[7]

Habitat and ecology

Most of the Peronospora species are highly specific to their hosts and can generally be found anywhere the host plant grows, or is being cultivated.^[3] A large portion of their life cycle is spent inside their host plant. Many species of Peronospora are seedborne pathogens, so the worldwide spread of Peronospora crop-plant pathogens is likely to be a result of unknowingly trading infected seeds to new areas.^[3] There are also many Peronospora species that are spread by wind currents, which allows them to disperse over large distances.^[3] Peronospora species prefer humid air and cool temperatures.^[5]

General form and structure

The first stage in the Peronospora life history is the sporangia.^[5] The sporangia are small spore-like structures about 65 um long that germinate a germ-tube when they are near a leaf stoma.^[8]^[5] A germ tube will come from the sporangium and penetrate the leaf cell where it will form a haustorium.^[5] The haustorium absorbs nutrients from the leaf, while hyphae invade the intercellular space, and the leaf will eventually develop a lesion.^[5] These lesions often start out yellow and then turn brown as the leaf starts to undergo necrosis.^[5] From here, Peronospora can undergo either asexual reproduction or sexual reproduction.^[5] Asexual reproduction occurs when the air outside is moist making for favourable conditions.^[5] During asexual reproduction, hyphae on the host plant will form sporangiophores, which will produce conidia.^[5] The conidia will be dispersed by the wind is able to infect other plants.^[5] The asexual cycle only takes five to seven days to complete.^[5] Sexual reproduction occurs when the conditions are unfavourable and it needs to withstand harsh environmental conditions.^[5] During sexual reproduction, the hyphae will undergo meiosis forming antheridia and oogonia, the only haploid structures in the Peronospora life history.^[5] The antheridia will fuse to the oogonia, initiating plasmogamy and then karyogamy, and will result in the production of many oospores.^[5] The oospores can then be dispersed by the wind to infect more plants.^[5]

Both Peronospora and Pseudoperonospora are characterized by their ability to produce melanized sporangia, but Pseudoperonospora produces zoospores whereas Peronospora cannot.^[3]

Practical importance

The model oomycete pathogen, Peronospora parasitica, used to be included in this genus, however it has been reclassified to the genus Hyaloperonospora.^[3]

Some species of Peronospora have been considered for their use as a bioweapon or have been classified as potential bioweapons.^[3] Peronspora somniferi was considered for its ability to devastate fields of the opium poppy, which could have targeted areas that depend on the crop.^[3] The United States has classified Peronospora tabacina as a possible bioweapon, because if it were used to target the US tobacco industry, it would lead to major economic loss.^[3]

Genomics and genetics

Only one species in the genus Peronospora has had its genome sequenced and assembled. In 2015, Derevnina et al. performed a de novo sequence assembly of the genome of two Peronospora tabacina isolates using Illumina sequencing.^[9] They estimated the genome size to be 68 Mb with a mitochondrial genome of 43 kb.^[9] The two assemblies had 61.8x and 128.9x coverage for the nuclear genomes and 6,824x and 43,225x coverage for the mitochondrial genomes.^[9] The mitochondrial genome only differed by seven single nucleotide polymorphisms, three small indels, and one copy number variant.^[9] Using a program to predict gene models, they found 18,000 potential protein coding genes.^[9]