Pseudoperonospora cubensis
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''Pseudoperonospora cubensis'' is a species of
water mould Oomycota forms a distinct phylogenetic lineage of fungus-like eukaryotic microorganisms, called oomycetes (). They are filamentous and heterotrophic, and can reproduce both sexually and asexually. Sexual reproduction of an oospore is the r ...
known for causing
downy mildew Downy mildew refers to any of several types of oomycete microbes that are obligate parasites of plants. Downy mildews exclusively belong to the Peronosporaceae family. In commercial agriculture, they are a particular problem for growers of cruc ...
on
cucurbits The Cucurbitaceae, also called cucurbits or the gourd family, are a plant family consisting of about 965 species in around 95 genera, of which the most important to humans are: *''Cucurbita'' – squash, pumpkin, zucchini, some gourds *''Lagena ...
such as cantaloupe, cucumber, pumpkin, squash and watermelon. This water mould is an important pathogen of all these crops, especially in areas with high humidity and rainfall, such as the eastern United States. In most years the disease is an annual, late-season problem on squash and pumpkin in the eastern and central United States, however, since 2004 it has become one of the most important diseases in cucumber production. Considered a highly destructive
foliar A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, ste ...
disease of cucurbits, successful breeding in the mid-twentieth century provided adequate control of downy mildew in cucumber without the use of fungicides. The resurgence in virulence has caused growers great concern and substantial economic losses, while downy mildew in other cucurbit crops continues to be a yearly hindrance.


Symptoms and signs

The pathogen causes angular
chlorotic In botany, chlorosis is a condition in which leaves produce insufficient chlorophyll. As chlorophyll is responsible for the green color of leaves, chlorotic leaves are pale, yellow, or yellow-white. The affected plant has little or no ability to ...
lesions on the foliage. These lesions appear angular because they are bound by leaf veins. During humid conditions, inspection of the underside of the leaf reveals gray-brown to purplish-black fungal growth. This downy material is the
sporulation In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, ...
of the pathogen. Magnification of the sporulation reveals the acutely and dichotomously branched sporangiophores bearing lemon-shaped
sporangia A sporangium (; from Late Latin, ) is an enclosure in which spores are formed. It can be composed of a single cell or can be multicellular. Virtually all plants, fungi, and many other lineages form sporangia at some point in their life cyc ...
. Eventually, leaves will turn
necrotic Necrosis () is a form of cell injury which results in the premature death of cells in living tissue by autolysis. Necrosis is caused by factors external to the cell or tissue, such as infection, or trauma which result in the unregulated dige ...
and curl upwards. The disease is sometimes called wildfire because of how rapidly it progresses, as if the crop were burned by fire. Symptoms on watermelon and cantaloupe are different from on other cucurbits; leaf spots are typically not angular and turn brown to black in color. Often, an exaggerated upward leaf-curling will occur. Regardless of which cucurbit is involved, only the leaves are infected, not fruit, flowers, stems or roots. Disease of the leaves results in three major effects: 1. reduced yields, 2. a greater proportion of misshapen fruit, especially in cucumber, and 3. sunscalded fruit, due to increased exposure to direct sunlight, especially in watermelon and
winter squash Winter squash is an annual fruit representing several squash species within the genus ''Cucurbita''. Late-growing, less symmetrical, odd-shaped, rough or warty varieties, small to medium in size, but with long-keeping qualities and hard rinds, are ...
.


Pathogen biology

''Pseudoperonospora cubensis'' is an obligate parasite or biotroph, meaning that it requires live host tissue in order to survive and reproduce. Because of this characteristic, the pathogen must
overwinter Overwintering is the process by which some organisms pass through or wait out the winter season, or pass through that period of the year when "winter" conditions (cold or sub-zero temperatures, ice, snow, limited food supplies) make normal acti ...
in an area that does not experience a hard frost, such as southern
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and to ...
, and where wild or cultivated cucurbits are present. The spores are dispersed via wind to neighboring plants and fields and often over long distances. Symptoms appear 4 to 12 days after infection. The pathogen thrives under cool and moist conditions, but can do well under a wide range of conditions. Optimum conditions for sporulation are 59 °F (15 °C) with 6 to 12 hours of moisture present, often in the form of morning dew. Even when high daytime temperatures are not favorable for the pathogen (>95 °F or >35 °C), nighttime temperatures may be very suitable.
Oospore An oospore is a thick-walled sexual spore that develops from a fertilized oosphere in some algae, fungi, and oomycetes. They are believed to have evolved either through the fusion of two species or the chemically-induced stimulation of mycelia ...
s (thick-walled, resting spores) of ''P. cubensis'' are rare and their role in nature is unknown.


Host specificity, pathotypes

Within the cucurbit family, ''P. cubensis'' isolates will exhibit a specific host range, that is, it will infect certain cucurbits and not others. For example, if cucumber and squash are grown side-by-side sometimes only the cucumber is diseased. When a pathogen exhibits this type of host specificity within a plant family, it is referred to as a
pathotype A pathovar is a bacterial strain or set of strains with the same or similar characteristics, that is differentiated at infrasubspecific level from other strains of the same species or subspecies on the basis of distinctive pathogenicity to one o ...
. At least five pathotypes of ''P. cubensis'' have been described in the U.S. More recent studies in the
Czech Republic The Czech Republic, or simply Czechia, is a landlocked country in Central Europe. Historically known as Bohemia, it is bordered by Austria to the south, Germany to the west, Poland to the northeast, and Slovakia to the southeast. The ...
have revealed more pathotypes.


Disease management

Controlling downy mildew requires the use of resistant cultivars, fungicide applications and early detection. * Resistant cultivars - Host resistance is an important tool in disease control and should be used whenever possible.
Cultivar A cultivar is a type of cultivated plant that people have selected for desired traits and when propagated retain those traits. Methods used to propagate cultivars include: division, root and stem cuttings, offsets, grafting, tissue culture ...
s resistant to downy mildew have been developed for cucumber and cantaloupe and to a lesser extent for squash and pumpkin. Although cucumber downy mildew has been severe on resistant cultivars, these cultivars are more effective than susceptible cultivars in delaying infection. * Chemical control - Chemical control is highly recommended because downy mildew is an aggressive and destructive disease and satisfactory control without the use of fungicides is unlikely. Both protectant and systemic products should be applied. Fungicides are most effective when applied prior to infection and reapplied at 5- to 7-day intervals. The following products have proven to be the most effective fungicides in cucumber downy mildew control in North Carolina trials conducted annually from 2004 to 2008: Presidio ( fluopicolide, Valent), Ranman ( cyazofamid, FMC), Previcur Flex ( propamocarb, Bayer), Curzate ( cymoxanil, DuPont), Tanos ( famoxadone and cymoxanil, DuPont), and Gavel ( zoxamide and
mancozeb Mancozeb is a dithiocarbamate non-systemic agricultural fungicide with multi-site, protective action on contact. It is a combination of two other dithiocarbamates: maneb and zineb. The mixture controls many fungal diseases in a wide range of f ...
, Dow AgroSciences). These products should be applied in a program to prevent pathogen resistance, that is, rotated with fungicides of a different mode of action. Protectant fungicides such as
chlorothalonil Chlorothalonil (2,4,5,6-tetrachloroisophthalonitrile) is an organic compound mainly used as a broad spectrum, nonsystemic fungicide, with other uses as a wood protectant, pesticide, acaricide, and to control mold, mildew, bacteria, algae. Ch ...
and mancozeb should be used as mixing partners. * Biological control - Although many
biological control Biological control or biocontrol is a method of controlling pests, such as insects, mites, weeds, and plant diseases, using other organisms. It relies on predation, parasitism, herbivory, or other natural mechanisms, but typically also i ...
products have been evaluated for their ability to control downy mildew, none have proven effective and are not recommended for use. * Early detection & forecasting - Many growers have lost the battle against downy mildew by waiting until they could clearly see the disease before initiating sprays. Early detection of downy mildew and immediate or preventative fungicide application is imperative for the control of this disease.
forecasting system
exists to assist growers in timing their fungicide applications for maximum benefit. The system tracks outbreaks of the disease and provides a forecast or risk assessment for future outbreaks.


References


Further reading

*


External links


Downy-Mildew Cucumber Interaction
{{Taxonbar, from=Q3924916 Water mould plant pathogens and diseases Vegetable diseases Fruit diseases Peronosporales Leaf diseases