Peronosclerospora philippinensis (Philippine downy mildew of maize)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Hosts/Species Affected
- Host Plants and Other Plants Affected
- Growth Stages
- List of Symptoms/Signs
- Biology and Ecology
- Means of Movement and Dispersal
- Seedborne Aspects
- Pathway Causes
- Pathway Vectors
- Plant Trade
- Impact Summary
- Economic Impact
- Environmental Impact
- Social Impact
- Risk and Impact Factors
- Uses List
- Detection and Inspection
- Similarities to Other Species/Conditions
- Prevention and Control
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Peronosclerospora philippinensis (W. Weston) C.G. Shaw, 1978
Preferred Common Name
- Philippine downy mildew of maize
Other Scientific Names
- Sclerospora maydis Reinking
- Sclerospora philippinensis W. Weston
International Common Names
- English: Philippine maize downy mildew
Summary of InvasivenessTop of page
Peronosclerospora philippinensis, a causal pathogen of maize downy mildews, is one of the major maize diseases reported in some maize-growing countries, especially in the Philippines. High disease incidence has been reported in many parts in the country specifically in northern Luzon and in many parts of Mindanao despite breakthroughs in controlling or mitigating the disease using cultural and chemical control (Pascual et al., 2005). P. philippinensis is considered the most virulent of the downy mildew pathogens affecting maize, causing substantial losses to crop production (Murray, 2009). Under normal conditions, a 40-60% yield reduction is observed; however, favourable conditions for disease development can amplify these losses to 80-100% (Exconde and Raymundo, 1974). As the pathogen is able to survive in seeds, is able to spread rapidly and occasionally forms resting spores that can survive for more than 1 year, the pathogen has the potential to become a threat to local maize production in warm temperate and tropical areas.
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Chromista
- Phylum: Oomycota
- Class: Oomycetes
- Order: Peronosporales
- Family: Peronosporaceae
- Genus: Peronosclerospora
- Species: Peronosclerospora philippinensis
Notes on Taxonomy and NomenclatureTop of page
Philippine maize downy mildew, caused by Peronosclerospora philippinensis, is regularly confused with various other downy mildews affecting maize, e.g., P. spontanea, P. australiensis and P. maydis.
The pathogen has distinct morphological characteristics, allowing it to be differentiated from other Peronosclerospora spp. However, P. philippinensis bears great resemblance to P. sacchari. Although isoenzyme comparisons have been used to identify species of the genus Peronosclerospora, there have been difficulties to distinguish between P. philippinensis and isolates identified as P. sacchari (Bonde et al., 1984; Micales et al., 1988). These two species have been suggested to be conspecific (Yao et al., 1991) but a proof of this assumption is so far lacking.
DescriptionTop of page
P. philippinensis is an obligate pathogen which requires a living host for it to grow and proliferate. Infection occurs when airborne conidia from an infected crop attaches to a susceptible crop host, but seedborne infections are probably in important means of spread of the pathogen. A higher rate of infection occurs at temperatures above 16°C (Jepson, 2008). Inside the host, the pathogen produces mycelium which gives rise to conidiophores bearing conidia. Germinating conidia produce germ tubes and penetrate the merismatic tissues or stomata forming haustoria. These haustoria extend throughout the tissue forming mycelium, and the infection continues.
The mycelia are branched, slender (8 µm in diameter) and irregularly constricted. The conidiophores are dichotomously branched, measuring 15-26 x 150-400 µm. The conidia are hyaline, ovoid to round cylindrical, slightly rounded at the apex, 17-21 x 27-39 µm. The haustoria are simple, hyaline and vesiculiform to subdigitate (Purdue University, undated; Weston, 1920; Smith and Renfro, 1999). Oospores, which are rarely produced, are spherical, smooth-walled and approximately 22 µm in diameter.
DistributionTop of page
P. philippinensis is listed as present in Thailand (Murray, 2009). However, a recent study of the morphological characteristics of conidia and conidiophores, as well as molecular identification through ITS1, suggests that P. philippinensis should be removed from the list of maize downy mildew pathogens in Thailand (Janruang and Unartngam (2018). As there are several Peronosclerospora species known to infect maize, the distribution of P. philippinensis remains mostly uncertain, unless sequencing of suitable barcoding loci is done (preferably cox2; see Telle et al., 2011; Choi et al., 2015).
Distribution TableTop of page
The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.Last updated: 30 Jun 2021
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|South Africa||Absent, Unconfirmed presence record(s)|
|China||Present||Present based on regional distribution.|
|India||Present||Present based on regional distribution.|
|Indonesia||Present||Present based on regional distribution.|
|Japan||Absent, Invalid presence record(s)|
History of Introduction and SpreadTop of page
Philippine downy mildew, caused by P. philippinensis, was described from the Philippines in 1918.
Risk of IntroductionTop of page
Philippine downy mildew, caused by P. philippinensis, is one of the most destructive diseases of maize throughout the world. The pathogen can infect cultivated sugarcane, sorghum and other weed species. The primary source of inoculum is infected hosts such as maize, sugarcane and some susceptible grass species. The introduction risk of this pathogen is intermediate because the conidia are short-lived outside of their host and seeds from affected areas are rarely exported. Dissemination of the pathogen is through water and wind. It can also be spread through seeds, when the moisture content of the seed is more than 30% (recommended MC:14%), or through infected vegetative material (Murray, 2009; USDA, 2013).
HabitatTop of page
No variations were observed among the countries where Philippine maize downy mildew has been reported. Optimum conditions for sporulation of P. philippinensis are 18-23°C, and for germination, 10-35°C (Purdue University, undated). According to a study conducted by Dalmacio and Raymundo (1972), the conditions required for the production, germination and infection of conidia in P. philippinensis are between 21 and 26°C.
Hosts/Species AffectedTop of page
P. philippinensis is restricted to host species within the grass tribes Andropogoneae and Maydeae (Bonde and Peterson, 1983). Some of these hosts may serve as inoculum reservoirs.
Secondary hosts include the hybrid, maize x Zea mexicana.
For further information on natural hosts, see Bonde and Peterson (1983) and Pupipat et al. (1975).
Hosts by artificial inoculation include Andropogon gerardii, Bothriochloa ambigua, B. barbinodis, B. decipiens, B. edwardsiana, B. ischaemum var. ischaemum, B. laguroides, B. perforata, B. springfieldii, B. woodrowii, Eulalia fulva, Schizachyrium hirtiflorum, S. microstachyum, S. scoparium, Sorghum plumosum, Tripsacum dactyloides, Zea diploperennis and Zea perennis (Bonde and Peterson, 1983).
Host Plants and Other Plants AffectedTop of page
|Avena sativa (oats)||Poaceae||Other|
|Saccharum officinarum (sugarcane)||Poaceae||Other|
|Saccharum spontaneum (wild sugarcane)||Poaceae||Wild host|
|Sorghum bicolor (sorghum)||Poaceae||Other|
|Sorghum halepense (Johnson grass)||Poaceae||Wild host|
|Zea mays (maize)||Poaceae||Main|
|Zea mays subsp. mexicana (teosinte)||Poaceae||Other|
Growth StagesTop of page
SymptomsTop of page
P. philippinensis causes a systemic infection wherein intense green and yellow stripes are observed along the entire leaf. Thick, white, woolly growth of conidia and conidiophores can be observed underneath these areas (Gupta and Paul, 2002; Magill et.al., 2006). These symptoms may appear as early as 3 days after infection. As the disease progresses, leaves become narrow and abnormally erect, in some cases, the affected leaves can appear somewhat dried out. Tassels may also exhibit malformation hence, interrupting ear formation and sterility of seed (Magill et al., 2006). Diseased stems do not show external symptoms but are usually stunted.
Moderate infection allows the plant to reach maturity but causes small, deformed ears (Weston, 1920; Dalmacio and Raymundo, 1972). In severe cases, infected plants become stunted and weakened, eventually leading to plant death within a month (Murray, 2009).
List of Symptoms/SignsTop of page
|Inflorescence / abnormal leaves (phyllody)|
|Inflorescence / twisting and distortion|
|Leaves / abnormal colours|
|Leaves / abnormal forms|
|Leaves / fungal growth|
|Leaves / necrotic areas|
|Whole plant / discoloration|
|Whole plant / distortion; rosetting|
|Whole plant / dwarfing|
|Whole plant / plant dead; dieback|
Biology and EcologyTop of page
Infection starts when airborne conidia attach to a susceptible crop host. The germinating conidia produce germ tubes that penetrate the stomata thus forming a haustoria. In case of compatible interaction, a biotrophic interface is established nutrients are taken up and the cycle continues once enough nutrients have been taken up to support sporulation (Jepson, 2008). Sporulation can continue for more than 2 months subject to favourable conditions. However, conidiophores and sporangia are only short-lived and mostly vanish throughout the day in sunny conditions. Viable conidia start to germinate in less than 1 hour, thus penetration into the host takes 2 hours (Weston, 1920; Dalmacio and Exconde, 1969; Bonde, 1982). Oospores are also produced, but only rarely in maize, serving as the survival structures of P. philippinensis, and primary infection source from the soil.
ClimateTop of page
|Af - Tropical rainforest climate||> 60mm precipitation per month|
Means of Movement and DispersalTop of page
Philippine maize downy mildew is commonly disseminated by air currents and rain splash. This results in localised spread among neighbouring hosts of the pathogen. The disease could also be spread through seeds, especially if they that have not been properly dried and have a moisture content of more than 30% (Murray, 2009; USDA, 2013; Purdue University, undated).
The movement of infected plant tissue could introduce the disease to a new location (USDA, 2013).
Seedborne AspectsTop of page
A study conducted by Advincula and Exconde (1975) showed that P. philippinensis is seed transmissible. Infection occurs when the seeds are sown immediately after harvest (moisture content of more than 30-43%) (Murray, 2009). The pathogen becomes established in the seed as hyphae and mycelium in the pericarp but not within the endosperm (Murray, 2009). However, Bains and Jhooty (1982) suggested its presence in the embryo of infected seeds.
Effect on Seed Quality
P. philippinensis infects the pericarp, embryo and the endosperm of susceptible seeds (Bains and Jhooty 1982; Murray, 2009). Infected seeds are hard to determine because they do not exhibit external symptoms and the seed quality is not affected (Jepson, 2008).
The conidia of P. philippinensis are short-lived, thus the introduction or transmission of the pathogen is probably achieved through infective vegetative material and seeds. Seeds produced on systemically infected crops mostly cause infection when sown as fresh seeds shortly after harvest. Infected plants exhibit intense green and yellow stripes along the entire leaf area, thereby causing stunting (Jepson, 2008; Magill et al., 20016). Moreover, infected plants mature slower, thus the formation and development of ears are suppressed (Purdue University, undated). A moisture content of 30-43% in seeds is necessary for seed transmission.
Several methods are used to avoid or reduce the transmission of P. philippinensis. Proper drying of seeds to a moisture content of less than 14% is necessary (USDA, 2013). The application of systemic and protectant fungicides to the seeds such as fentin hydroxide, maneb and metalaxyl is also used (Exconde, 1982; Dalmacio et.al., 1987; Murray, 2009).
Pathway CausesTop of page
Pathway VectorsTop of page
|Plants or parts of plants||Introduction of the disease is through the entry of infected vegetative material or through infected seeds||Yes||Murray (2009)|
|Soil, sand and gravel||Oospores which are rarely produced serve as survival structures of the pathogen in the soil.||Yes||USDA (2013)|
|Water||One of the common agents responsible for the spread of P. philippinensis is rain||Yes||Yes||USDA (2013)|
|Wind||Wind dispersal of P. philippinensis to adjacent fields is mostly localized||Yes||USDA (2006)|
Plant TradeTop of page
|Plant parts liable to carry the pest in trade/transport||Pest stages||Borne internally||Borne externally||Visibility of pest or symptoms|
|Leaves||hyphae; spores||Yes||Pest or symptoms not visible to the naked eye but usually visible under light microscope|
|True seeds (inc. grain)||spores||Yes||Pest or symptoms usually invisible|
Impact SummaryTop of page
Economic ImpactTop of page
P. philippinensis is considered the most virulent of the maize downy mildew pathogens. According to Murray (2009), the pathogen may be able to stop maize production in Asia. It causes substantial losses in maize production. Under normal conditions, yield losses incurred are 40-60%, however, favourable conditions for the pathogen may result to losses ranging from 80 to 100% (Exconde and Raymundo, 1974). During the years 1974-1975, national yield reduction in maize in the Philippines was estimated at 205,470 metric tons with a value of Php 178,759,000 (Exconde, 1982). In sugarcane, losses incurred from the disease (tons of cane per hectare) can reach from 9 to 38%. Losses in picul of sugar per ton may range from 2 to 35% while losses in picul of sugar per hectare may be between 10 and 58% (Husmillo, 1982).
Environmental ImpactTop of page
The impact of the disease on the environment is almost negligible. The disease is not likely to affect species composition while it might be able to reduce longevity or competitiveness of wild hosts.
Social ImpactTop of page
Philippine downy mildew causes significant yield losses on economically important crops including maize and sugarcane. This affects the income of both farm owners and growers. The disease is especially detrimental for traditional farmers that collect their own seeds for sowing in the next season, as disease might accumulate by this practice, and subsistence farmers cannot afford commercial seeds. Aside from affecting income and livelihood, the presence of this disease encourages the use of chemicals for control.
Risk and Impact FactorsTop of page
- Invasive in its native range
- Proved invasive outside its native range
- Has a broad native range
- Abundant in its native range
- Highly mobile locally
- Fast growing
- Has high reproductive potential
- Has propagules that can remain viable for more than one year
- Reproduces asexually
- Has high genetic variability
- Host damage
- Negatively impacts agriculture
- Negatively impacts cultural/traditional practices
- Negatively impacts livelihoods
- Damages animal/plant products
- Negatively impacts trade/international relations
- Antagonistic (micro-organisms)
- Pest and disease transmission
- Parasitism (incl. parasitoid)
- Highly likely to be transported internationally accidentally
- Highly likely to be transported internationally deliberately
- Difficult to identify/detect as a commodity contaminant
- Difficult/costly to control
Uses ListTop of page
- Laboratory use
- Research model
- Test organisms (for pests and diseases)
DiagnosisTop of page
P. philippinensis is usually identified and distinguished from other downy mildew pathogens through conidial morphology (except for P. sacchari due to their high resemblance). Symptomatology among downy mildew pathogens is also different: P. philippinensis causes chlorotic streaks whereas P. sacchari induces discrete, chlorotic blotches (Murray, 2009). However, as conidial morphology may be variable and there are several other downy mildew species causing similar symptoms on maize, molecular barcoding using cox2 sequencing (Choi et al., 2015) is strongly recommended to confirm the identity of the pathogen.
The advent of molecular biology has led to the development of techniques that can differentiate between species of Peronosclerospora (Micales et al., 1988). In Australia, the use of DNA sequence information has been instrumental in the identification of several downy mildew pathogens including P. philippinensis (Telle et al., 2011; Shivas et al., 2012; USDA, 2013). In addition to this, isoenzyme comparisons can also be used to identify P. philippinensis (Bonde et al., 1984; Micales et al., 1988).
Detection and InspectionTop of page
Systemic symptoms may be observed in the first true leaf stage as chlorotic stripes or a pale-yellow coloration throughout the entire leaf. Local symptoms (long, chlorotic streaks with a downy growth of conidia and conidiophores) may be present from the two-leaf stage until the appearance of tassels and silks. Tassels may be malformed, producing less pollen, and ears may be aborted. Early-infected plants become stunted and many die. Weston reported the collapse of badly infected cells and the deterioration of chloroplasts, resulting in the characteristic yellow colour of diseased leaves (Weston, 1920).
Similarities to Other Species/ConditionsTop of page
Downy mildew can be confused with other diseases. Symptoms induced by P. philippinensis in maize are similar to various other downy mildews on maize and maize stripe. To differentiate between the two diseases, the underside of the leaves should be inspected for the presence of thick, white, woolly fungal growth early in the morning (Gupta and Paul, 2002; Magill et al, 2006).
P. philippinensis is closely related to P. sacchari on the basis of morphology, symptoms, host range and isoenzyme comparisons. Variations in these factors have been reported by a number of authors and the differences were mostly attributed to host genotype and the existing environmental conditions. In P. sacchari, the length of the conidia shows a positive correlation with temperature, whereas the width is unaffected. The two pathogens differ in the symptoms produced on maize, P. philippinensis produces chlorotic streaks, whereas P. sacchari induces discrete, chlorotic blotches (Murray, 2009). However, there are other downy mildews on maize that are also causing similar symptoms, including P. maydis, P. australiensis and P. spontanea.
Prevention and ControlTop of page
Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.
Movement of infected planting material is the primary reason for the introduction of a disease into a disease-free environment. Quarantine measures should be strictly enforced to prevent the entry of a disease in an area (Sugar Research Australia, undated). Especially a drying of the seeds to water contents of less than 15% seems to be advisable. However, as a precaution, seeds from areas reporting maize downy mildew should not be imported
All infected plants should be collected and destroyed by incineration. Monitoring of infected hosts or alternate hosts to ensure that the pathogen would not be able to re-establish is also a must (Murray, 2009). Disposable equipment, infected planting materials and soil should eradicated through autoclaving, high temperature incineration, or deep burial. Double bagging of equipment used in the infested site should also be done (Murray, 2009).
All infected plants should be eradicated. Any equipment/vehicles used in the infested site should be subjected to movement restrictions. Harvesting of infected crops should not push-through because the dust during harvesting can serve as dispersal agent to neighbouring areas. Clothing and footwear used in the diseased area should be thoroughly disinfected or double bagged preceding disposal. Seeds from the infested site should not be allowed to be used as planting materials, feed for animals or food for consumption (Murray, 2009).
Cultural control and sanitary measures
Several cultural methods are used to control downy mildew. One is crop rotation wherein there should be a 15-17 day interval to reduce soil inoculum potential. Another is the removal of any symptomatic plant parts to decrease the inoculum potential of infected plants. Drying of seeds down to its recommended moisture content (14%) also reduces the likelihood of transmission of the pathogen (Murray, 2009).
Reduction of the seed moisture content to less than 14% may lower the risk of seed transmission. Crop rotation between sorghum or maize for more than 3 years is also recommended to reduce soil-borne infection (Murray, 2009).
Several fungicides are used for the control and eradication of P. philippinensis. Foliar sprays, soil treatments and seed treatments using protectants or eradicants have been studied. Weston (1923) soaked seeds in 70% alcohol for 30 to 60 seconds and then washed with running water for 1 hour. Exconde (1982) reported metalaxyl to be effective as a seed dressing. Other fungicides such as fentin hydroxide and maneb have also proved to be effective in controlling the disease (Murray, 2009).
Host resistance (incl. vaccination)
Philippine downy mildew resistance is determined through multiple gene inheritance and is dictated by additive gene effects (Leon et al., 1993, cited in Murray, 2009). A number of resistant maize varieties have been developed. The majority of the developed lines exhibit slowed systemic infection rates, thereby containing infection to local areas for longer periods of time and lowering overall destructive effects of the disease (Murray, 2009).
A study conducted by Pascual et al. (2005) showed that two inbred lines exhibited a high degree of resistance in two separate experimental areas.
ReferencesTop of page
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ContributorsTop of page
31/03/20 Reviewed by:
Fe M. Dela Cueva, Plant Pathology Laboratory, Institute of Plant Breeding, University of the Philippines Los Baños 4031, Philippines
Alyssa M. de Castro, Plant Pathology Laboratory, Institute of Plant Breeding, University of the Philippines Los Baños 4031, Philippines
Rachele L. De Torres, Plant Pathology Laboratory, Institute of Plant Breeding, University of the Philippines Los Baños 4031, Philippines
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