Peronosclerospora philippinensis

Commonly known as Philippine downy mildew, this disease is caused by the species Peronosclerospora philippinensis of the fungal-like protist class Oomycetes, which also has members such as water molds and Phytophthora infestans, which caused the potato blight that led to the Great Irish famine.

Peronosclerospora philippinensis
Scientific classification
Clade: SAR
Phylum: Oomycota
Order: Peronosporales
Family: Peronosporaceae
Genus: Peronosclerospora
Species:
P. philippinensis
Binomial name
Peronosclerospora philippinensis
(Weston) C. G. Shaw 1978 [1]
Synonyms

Sclerospora philippinensis (Weston 1920)[2]

Its hosts are typically grasses, cultivated and wild sorghum, sugarcane, and maize found in most of South and Southeast Asia, including India, China, and the Philippines. This disease results in reduced yield in crops, a withered appearance of the leaves, abortive reproductive structures, and a grayish down on the surfaces.

Historically, there have been cases of epidemics in which the species infected cultivars of maize and sugarcane and decimated the populations, causing up to $23 million US dollars’ worth of damage. Because of this, the USDA Animal and Plant Health Inspection Service has included Philippine downy mildew on its Plant Pathogen Select Agent List, being a pathogen that poses a great threat for American plants, both wild and cultivated.

History

Philippine downy mildew was first briefly mentioned by Professor Baker of the College of Agriculture of the University of the Philippines in 1916 and then again in 1918 by Dr. Reinking, also of the College of Agriculture. His studies showed that the species had been present in the Philippines for more than 10 years already. It was not until 1920 when William Weston named it as Sclerospora philippinensis in a report, marking its similarities to its close relative Sclerospora sacchari.[2] It continued to be known as Sclerospora philippinensis until 1980, when it was renamed Peronosclerospora philippinensis according to the shifts in the classification.[3]

Yearly, there are 1 to 2 citations in biological journals. Research proves to be continual although scarce. Because of its designation as a Select Agent, American scientists have difficulty in obtaining samples within the United States to study the disease. In terms of gene sequences at GenBank of the National Center for Biotechnology Information, there are five, one complete sequence of an internal transcribed spacer and four partial sequences of internal transcribed spacers.[4]

Pathophysiology

Philippine downy mildew has been most studied on maize for its economic significance, but it shares similar symptoms with other hosts. A characteristic chlorosis, which develops from the loss of chlorophyll, and the downiness that gives the genus its popular name, are the two most common symptoms. The species also cause the growth of new leaves to be narrow and rigid, a stunting of shoots and roots, decreased production of pollen, and sterile and malformed ears. Mycelium extends throughout the entire plant except for the roots.[2] When the disease is transferred to a new host, there is a delay in virulence, known as parasitic fitness.[5]

In maize, the reproductive tassels are malformed. The disease seems to stimulate growth while suppressing it in other areas of development: the anthers are atrophied while the female structure development is blocked. The resulting pollen is hollow and wrinkled.[6]

Hosts

Socioeconomically important hosts are Zea mays and the Saccharum and Sorghum species due to their cultivation for crops and export. Other species of interest are the Graminaea species[2] and many of the Andropogon, Bothriochloa, Schizachyrium, and Zea genera, which amounts to over 30 species. Most of these species are common grasses. Over 60 species, though, of the previously mentioned genera are resistant. The susceptible Andropogon and Bothriochloa species are significant because they are common across the United States, especially areas where corn is grown agriculturally, like Illinois.[7]

Effects on fitness

Infection by Philippine downy mildew results in a lack of viable reproductive structures and often earlier death. In addition, the infected plant is more likely to be infected by a secondary pathogen, such as a species of Pythium, Helminthosporium, or Fusarium.[2]

At the same time, when a section of the infected plant is transplanted to the soil, there is pronounced growth. This action seems to be by influence of the fungus-like protist to produce a large-enough host capable of transmitting spores.[6]

Life cycle

Infection begins when conidia from nearby infected plants or those carried by the wind enter through the stomata by landing and developing a germ tube or hypha. The conidia germinate optimally in the early morning and with a high relative humidity. The haustoria extend from the hyphae of the mycelium and systemically invade the rest of the plant, draining nutrients.

Eventually, from the stomata, hyphae extend out and elongate, branching into sterigmata or conidiophores (spore-bearing structures). These then asexually bud into conidium spores. These structures create the characteristic grayish-white down-like appearance of downy mildew. Unlike other oomycetes, or Peronosclerospora, P. philippinensis is not known to produce oospores.[2] There is no known sexual stage of the life cycle for P. philippinensis.

In dry seasons, there is low infection. Maximum sporulation occurs in the late hours of the night and early morning, typically at 2-3AM with high humidity and moisture or dew. The conidia are then spread via air currents or wind.[8] Infection occurs most in the summer, peaking in July.[9]

Transmission via seeds occurs with infection in the pericarp.[10]

Epidemiology

Philippine downy mildew can be found in China, India, Indonesia, Nepal, Pakistan, Thailand, and in the Philippines. Currently, there are no known cases of the disease occurring within the United States.[11]

In Nepal, there was a large epidemic in 1967, followed by a couple more in 1970 and 1973.[12] A large epidemic in the Philippines from 1974-1975 devastated the national agricultural maize crop, costing the country millions in yield.[10]

There is limited information on diseases of P. philippinensis.

Diagnosis

Diagnosis as downy mildew can be characterized by the appearance of a down-like covering, the conidiophores, on the leaves, indicating a member of Peronosclerospora. To discriminate between species, a closer examination must be followed. Conidiophores of P. philippinensis are longer and thicken much less gradually for this disease than of other species of Peronosclerospora, and the conidia are also more elongated than the elliptical spores of P. sacchari. Spore size and shape are the most accurate method in differentiating between Peronosclerospora species.[10]

Prevention

Drying seeds and limiting the amount of moisture that accumulates on seeds and plants is shown to control the development of the protist from infected seeds and young plants. The preemptive use of systemic protectant fungicides prevents the progression of P. philippinensis. Less sprayings in the dry season has proven to be more effective than more sprayings in the wet season. Also, combining mineral oil with fungicide reduces yield loss while still being cost effective.[10] An alternative method would be to limit secondary hosts, such as grasses and weeds.[9] There have been many attempts to develop identify genes that code for resistance and to breed strains for those genes. Studies show some progress, but the degree of gene dominance of the resistance and the inheritance is still questionable.[3][13]

Treatment

The early application of fungicide to infected plants is the most common form of treatment. Using fungicides that are systemic, like Apron 35, which give protection within the plant, are the most reliable. These can be applied in the soil to the roots or as a spray to be taken up via transpiration.[14] Other fungitoxicants, like Dithane M-45 and Dithane Z-78, are also effective.[15] At high inoculation, the disease may be extended too systemically to be combatted.

Human relevance

Corn is an important staple food for the world, especially for the Philippines, being the primary crop for 20% of the population.[16] In the Philippine epidemic from 1974-1975, Philippine downy mildew cost the nation 8% of the total yield, accounting to 205,470 metric tons, or $22,601,700.[10] Because of its high destruction and high percentages of infection, P. philippinensis threatens agriculture worldwide. Funding has been focused on studies to identifying the minimal effective use of fungicide.[14] Other scientists have looked at inheritable resistance as a preventive effort[3] or its effects on other crop exports, like sugarcane.[17]

As of 2010, this disease is on the USDA APHIS Plant Pathogen Select Agent List. There are provisions for the USDA to treat the introduction of P. philippinensis into the country as an act of bioterrorism because of the possible effect it could have on natural flora and on national agriculture. Because of this status, there is currently no budget for studies of this disease in the United States.[18]

Research

A number of experts on the disease are members of the U.S. Department of Agriculture or are of Texas A&M University. Much research has been put into the phylogenetic relationship between P. sacchari and P. philippinensis. Experiments have shown that of twenty-six species, only three species that can be infected by P. philippinensis show little infection by P. sacchari. This difference is minimalized when comparing host ranges of isolates of the same species. One isolate of P. philippinensis can infect seven more than another.[7] There is no significant difference in the number of nuclei per spore for these two species,[19] and they have almost identical banding patterns when analysed through a Southern blot.[20] When looking at their genomes, using simple sequence repeats as markers, there is shown to be lack of uniqueness between P. philippinensis and P. sacchari.[21] Because of these similarities, many scientists question that initial distinction that Weston placed on them in 1920.

More studies can be expected on a possible sexual stage as well as information on inherited resistance to P. philippinensis in the host plants.

The species can germinate in water and nutrient solutions, including those with 1% agar.[2] In most experiments, isolates are extracted from the disease containment greenhouse or nursery. The conidia are collected by washing the spores from leaves, and then filtered through a screen. This solution is then sprayed onto the test plants which are incubated in dew chambers for 6–7 hours at about 21 °C before they are moved to the laboratory.[22]

See also

References
  1. Shaw, Charles Gardner (1978). "Peronosclerospora species and other downy mildews of the Gramineae". Mycologia. 70 (3): 594–604. doi:10.1080/00275514.1978.12020263.
  2. Weston, William H. Jr (May 1920). "Philippine downy mildew of maize". Journal of Agricultural Research. 19: 97–146.
  3. Kaneko, K; Aday BA (1980). "Inheritance of Resistance to Philippine Downy Mildew of Maize, Peronosclerospora philippinensis". Crop Science. 20 (5): 590–594. doi:10.2135/cropsci1980.0011183x002000050011x.
  4. "NCBI".
  5. Duck, NB; Bonde MR; Peterson GL; Bean GA (1986). "Sporulation of Peronosclerospora sorghi, P. sacchari, and P. philippinensis of Maize". Phytopathology. 77 (3): 438–441. doi:10.1094/phyto-77-438.
  6. Weston, WH Jr (1929). "Observations on tassels of teosinte malformed by Sclerospora". Journal of Agricultural Research. 39: 817–836.
  7. Bonde, MR; Peterson GL (1983). "Comparison of host ranges of Peronosclerospora philippinensis and P. sacchari". Phytopathology. 73 (6): 875–878. doi:10.1094/phyto-73-875.
  8. Bonde, MR (1982). "Epidemiology of downy dildew disease of maize, sorghum and pearl millet". Tropical Pest Management. 28: 49–60. doi:10.1080/09670878209370674.
  9. Bains, SS; Jhooty JS (1982). "Distribution, spread and perpetuation of Peronosclerospora philippinensis in Punjab". India Phytopathology. 35: 566–570.
  10. Exconde, OR (1976). "Philippine corn downy mildew: assessment of present knowledge and future research needs". Kasetsart Journal: Natural Science. 10: 94–99.
  11. Magill, C; et al. (2006). "Recovery Plan for Philippine Downy Mildew and Brown Stripe Downy Mildew of Corn" (PDF): 1–13. Cite journal requires |journal= (help)
  12. Shah, SM (1976). "Downy mildew of maize in Nepal". Kasetsart Journal: Natural Science. 10: 137–142.
  13. Mochizuki, N; Carangal VR; Aday BA (1974). "Diallel analysis of host resistance to Philippine downy mildew of maize caused by Sclerospora philippinensis". JARQ. 8: 185–187.
  14. Molina AB, AB Jr; Exconde OR (1981). "Efficacy of Apron 35 SD (Metalaxyl) against Philippine corn downy mildew. I. Effects of seed-dressing methods and rainfall frequency". The Philippine Agriculturist. 64: 99–105.
  15. Sharma, SC; Khehra AS; Bains SS; Malhi NS (1981). "Efficacy of fungitoxicant sprays and seed treatment against Philippine downy mildew of maize Peronosclerospora philippinensis, Zea mays". India Phytopathology. 34: 498–499.
  16. Suliek, GP; Tangonan NG (2010). "Resistance evaluation of newly-developed USM yellow corn lines against Peronosclerospora philippinensis causing downy mildew disease". USM Research and Development Journal. 18: 143–145.
  17. Husmillo, FR (1982). "Assessment of yield loss due to downy mildew of sugarcane by Peronosclerospora philippinensis (Weston) C.G. Shaw". Sugarcane Pathologists' Newsletter. 28: 17–24.
  18. Fletcher, J; et al. (2010). "Emerging infectious plant diseases". Emerging Infections 9. Emerging Infectious Diseases. pp. 337–366. doi:10.1128/9781555816803.ch18. ISBN 9781555815257.
  19. Pawar, MN (1986). "Pathogenic variability and sexuality in Peronosclerospora sorghi (Weston and Uppal) Shaw, and comparative nuclear cytology of Peronosclerospora species". Texas A & M University: 1–107.
  20. Yao, C (1991). "Classification and detection of Peronosclerospora species on the basis of DNA Southern hybridization and the polymerase chain reaction". Texas A & M University: 1–71.
  21. Perumal, R; Nimmakayala, P.; Erattaimuthu, S. R.; No, E. G.; Reddy, U. K.; Prom, L. K.; et al. (2008). "Simple sequence repeat markers useful for sorghum downy mildew (Peronosclerospora sorghi) and related species". BMC Genetics. 9: 77–91. doi:10.1186/1471-2156-9-77. PMC 2620352. PMID 19040756.
  22. Bonde, MR; Peterson, GL; Kenneth, RG; Vermeulen, HD; Sumartini; Bustaman, M (1992). "Effect of Temperature on Conidial Germination and Systemic Infection of Maize by Peronosclerospora Species". Phytopathology. 82: 104–109. doi:10.1094/phyto-82-104.

Further reading
  • Baker, CF (1916). "Additional notes on Philippine plant disease". Philip. Agric. Forester. 5: 73–78.
  • Reinking, OA (1918). "Philippine economic plant diseases". Philip. J. Sci. 13: 165–174.
  • Weston, WH Jr (1923). "Production and dispersal of conidia in the Philippine Sclerosporas of maize". J. Agric. Res. 23: 239–277.
  • Chona, BL; Suryanarayana (1955). "The occurrence of Sclerospora philippinensis Weston on "Kans grass" (Saccharum spontaneum L.) in India". Indian Phytopath. 8: 209–210.
  • Carangal, V; Claudio, M; Sumayao, M (1970). "Breeding for resistance to maize downy mildew caused by Sclerospora philippinensis in the Philippines". Indian Phytopathol. 23: 285–305.
  • Exconde, OR (1970). "Philippine corn downy mildew". Indian Phytopathology. 23: 275–284.
  • Exconde, OR; Raymundo, AD (1974). "Yield loss caused by Philippine corn downy mildew". The Philippine Agriculturist. 58: 115–120.
  • Fuentes FD. 1976. Dynamics of Philippine corn downy mildew caused by Sclerospora philippinensis Weston. Ph.D. thesis. University of the Philippines at Los Banos. 194.
  • Josue AR. 1976. Virulence pattern and gross morphology of isolates of S. philippinensis from Bukidnon. Plant Pathology Seminar, Univ. of Phil., Los Banos, Philippines.
  • Telle, S; Shivas, RG; Ryley, MJ; Thines, M (2011). "Molecular phylogenetic analysis of Peronosclerospora Oomycetes) reveals cryptic species and genetically distinct species parasitic to maize". European Journal of Plant Pathology. 130 (4): 521–528. doi:10.1007/s10658-011-9772-8. S2CID 27542739.
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