Pseudoperonospora cubensis
Encyclopedia
Pseudoperonospora cubensis is a species of water mould
Water mould
Oömycota or oömycetes form a distinct phylogenetic lineage of fungus-like eukaryotic microorganisms . They are filamentous, microscopic, absorptive organisms that reproduce both sexually and asexually...

 known for causing downy mildew
Downy mildew
Downy mildew refers to any of several types of oomycete microbes that are obligate parasites of plants. Downy mildews exclusively belong to Peronosporaceae. In commercial agriculture, they are a particular problem for growers of crucifers, grapes and vegetables that grow on vines...

 on cucurbits
Cucurbitaceae
The plant family Cucurbitaceae consists of various squashes, melons, and gourds, including crops such as cucumber, pumpkins, luffas, and watermelons...

 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
Leaf
A leaf is an organ of a vascular plant, as defined in botanical terms, and in particular in plant morphology. Foliage is a mass noun that refers to leaves as a feature of plants....

 disease of cucurbits, successful breeding in the mid-twentieth century provided adequate control of downy mildew in cucumber without the use of fungicide
Fungicide
Fungicides are chemical compounds or biological organisms used to kill or inhibit fungi or fungal spores. Fungi can cause serious damage in agriculture, resulting in critical losses of yield, quality and profit. Fungicides are used both in agriculture and to fight fungal infections in animals...

s. 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 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
Spore
In biology, a spore is a reproductive structure that is adapted for dispersal and surviving for extended periods of time in unfavorable conditions. Spores form part of the life cycles of many bacteria, plants, algae, fungi and some protozoa. According to scientist Dr...

 of the pathogen. Magnification of the sporulation reveals the acutely and dichotomously branched sporangiophores bearing lemon-shaped sporangia
Sporangium
A sporangium is an enclosure in which spores are formed. It can be composed of a single cell or can be multicellular. All plants, fungi, and many other lineages form sporangia at some point in their life cycle...

. Eventually, leaves will turn necrotic
Necrosis
Necrosis is the premature death of cells in living tissue. Necrosis is caused by factors external to the cell or tissue, such as infection, toxins, or trauma. This is in contrast to apoptosis, which is a naturally occurring cause of cellular death...

 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 than 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.

Pathogen biology

P. cubensis is an obligate parasite
Obligate parasite
An obligate parasite is a parasitic organism that cannot complete its life cycle without dependence on its host.-See also:*Obligate intracellular parasite*Parasitism*Parasitic plant*Facultative parasite...

 or biotroph
Parasitism
Parasitism is a type of symbiotic relationship between organisms of different species where one organism, the parasite, benefits at the expense of the other, the host. Traditionally parasite referred to organisms with lifestages that needed more than one host . These are now called macroparasites...

, meaning that it requires live host tissue in order to survive and reproduce. Because of this characteristic, the pathogen must overwinter
Overwinter
To overwinter is to pass through or wait out the winter season, or to pass through that period of the year when “winter” conditions make normal activity or even survival difficult or near impossible...

 in an area that does not experience a hard frost, such as southern Florida
Florida
Florida is a state in the southeastern United States, located on the nation's Atlantic and Gulf coasts. It is bordered to the west by the Gulf of Mexico, to the north by Alabama and Georgia and to the east by the Atlantic Ocean. With a population of 18,801,310 as measured by the 2010 census, it...

, 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
Oospore
An oospore is a thick-walled sexual spore that develops from a fertilized oosphere in some algae and fungi. Also the result of plasmogamy/karyogamy in oomycetes, which in turn leads to the development of hyphae, then mycelium....

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. At least five pathotypes of P. cubensis have been described in the U.S. More recent studies in the Czech Republic
Czech Republic
The Czech Republic is a landlocked country in Central Europe. The country is bordered by Poland to the northeast, Slovakia to the east, Austria to the south, and Germany to the west and northwest....

 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
    Cultivar
    A cultivar'Cultivar has two meanings as explained under Formal definition. When used in reference to a taxon, the word does not apply to an individual plant but to all those plants sharing the unique characteristics that define the cultivar. is a plant or group of plants selected for desirable...

    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
    QoI
    Qo inhibitors , or Quinone outside inhibitors are a group of fungicides used in agriculture. They represent the most important development made in fungicides by the chemicals industry...

     and cymoxanil, DuPont),and Gavel (zoxamide and mancozeb, 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
    Chlorothalonil is a polychlorinated aromatic mainly used as a broad spectrum, non-systemic fungicide, with other uses as a wood protectant, pesticide, acaricide, and to control mold, mildew, bacteria, algae. Chlorothalonil-containing products are sold under the names Bravo, Echo, and Daconil. It...

     and mancozeb should be used as mixing partners.

  • Biological Control - Although many biological control
    Biological pest control
    Biological control of pests in agriculture is a method of controlling pests that relies on predation, parasitism, herbivory, or other natural mechanisms...

    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. A 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.

Additional Resources

For more information about downy mildew lifestyle, control, genetics, and genomics:

Downy-Mildew Cucumber Interaction
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