Blumeria graminis

Fungal pathogen of grasses

Blumeria graminis (commonly called barley powdery mildew or corn mildew) is a fungus that causes powdery mildew on grasses, including cereals. It is the only species in the genus Blumeria. It has also been called Erysiphe graminis and (by its anamorph) Oidium monilioides or Oidium tritici.

Previously B. graminis was included within the genus Erysiphe, but molecular studies have placed it into a clade of its own. In 1975, it was moved to the new monospecific genus Blumeria. Blumeria differs from Erysiphe in its digitate haustoria and in details of the conidial wall. Blumeria is also considered to be phylogenetically distinct from Erisiphe as it solely infects the true grasses of Poaceae. Eight special forms or formae speciales (ff. spp.) of B. graminis have been distinguished, each of which is parasitic on a particular genus or pareticular genera of grasses. Those that infect crop plants are B. g. f. sp. , which causes powdery mildew of wheat and infects other grasses in the genera Triticum and Aegilops, f. sp. on barley, f. sp. on oats and f. sp. on rye. Other formae speciales are pathogenic on wild grasses, including on grasses in the genera Agropyron and Elymus, on Bromus spp., on Poa spp. and on Lolium spp. (ryegrass).

The mycelium can cover the plant surface almost completely, especially the upper sides of leaves. Ascocarp is dark brown, globose with filamentous appendages, asci oblong. Ascospores hyaline, ellipsoid, 20–30 x 10–13 μm in size. Anamorph produces on hyaline conidiophores catenate conidia of oblong to cylindrical shape, not including fibrosin bodies, 32–44 x 12–15 μm in size. Haustoria are palmate. B. graminis is unique among the Erysiphales by having conidia with a primary germ tube and finger shaped ("digitate") appressoria.

The genus name of Blumeria is in honour of Samuel Blumer (b. 1895), a Swiss botanist (Mycology), Phytopathology, from the University of Bern (Universität Bern). The genus was circumscribed by Golovin ex Speer in Sydowia Vol.27 on page 2 in 1975.

B. graminis asexually produces conidia and sexually forms ascospores. Conidia are mainly distributed by wind, pests, or human activities. The water initiating ascospores are hypothesized to be dispersed not only by wind but also by splashing water droplets. It is biotrophic, and does not grow on synthetic media. Relatively cool and humid conditions are favourable for its growth. Its relatively great genetic variability enables it often to infect previously resistant plant varieties.

The genomes of B. g. f. sp. hordei and B. g. f. sp. tritici have recently been sequenced. Sequencing of the genome of the wheat powdery mildew B. g. f. sp. tritici, has allowed inference of important aspects of its evolution. It has been seen that it is the most repetitive fungal genome sequenced with 90% transposable elements. Additionally, 6540 genes were annotated, from which 437 encoded candidate secretor proteins and 165 for non secreted candidate secretor proteins. These were shown to be subject to positive selection, due to their implication in the gene for gene relationship to defeat plant disease resistance. The ability to infect tetraploid as well as domesticated hexaploid wheat, was seen to be the result of mildew genomes being mosaics of ancient haplogroups that existed before wheat domestication. This has allowed wheat powdery mildew to maintain genetic flexibility, variability and thus a great potential for pathogen variation. It is hypothesized that this mosacisism can be maintained through clonal reproduction in populations with a small effective size or quasi clonal reproduction in populations with large effective size.

Wheat powdery mildew is an obligate biotroph with a poorly understood evolutionary history. Sequencing its genome in 2013, many aspects of the evolution of its parasitism were unveiled. Obligate biotrophy has appeared multiple times in evolution in both ascomycetes like B. graminis and basidiomycetes, thus different selective pressure must have acted in the different organisms through time. It has been seen that B. g. f. sp. triticis genome is a mosaic of haplogroups with different divergence times, which explains its unique pathogen adaptability. Haplogroup Hold (diverged 40 80 mya) allows for the infection of wild tetraploid wheat and Hyoung (diverged 2 10 mya) allows for the infection of both domesticated hexaploid wheat species. Additionally, it has been seen that there is a positive selective pressure acting on genes that code for candidate secretor proteins and non secreted candidate secretor proteins, indicating that these might participate in the gene for gene relationship of plant disease resistance.

Powdery mildew of wheat is relatively easy to diagnose B. g. f. sp. tritici is an obligate parasite which means it only grows on living tissue. Though present throughout wheat growing regions, it especially favors the eastern seaboard of the United States as well as coastal regions of the United Kingdom.

Triticum spp. (wheat) is the only host of B. g. f. sp. tritici. The lower leaf surface corresponding to the mycelial mat will also show chlorosis.

B. g. f. sp. tritici has a polycyclic life cycle typical of its phylum, Ascomycota. Powdery mildew of wheat overwinters as cleistothecia dormant in plant debris. Under warmer conditions, however, the fungus can overwinter as asexual conidia or mycelium on living host plants. It can persist between seasons most likely as ascospores in wheat debris left in the field. Ascospores are sexual spores produced from the cleistothecia. These spores, as well as conidia, serve as the primary inoculum and are dispersed by wind. Neither spore requires free water to germinate, only high relative humidity. Wheat powdery mildew thrives in cool humid conditions and cloudy weather increases chances of disease. When conidia land on a wheat leaf's hydrophobic surface cuticle, they release proteins which facilitate active transport of lightweight anions between leaf and fungus even before germination. This process helps Blumeria recognize that it is on the correct host and directs growth of the germ tube. Both ascospores and conidia germinate directly with a germ tube. Conidia can recognize the host plant and within one minute of initial contact, the direction of germ tube growth is determined. The development of appressoria then begins infection following the growth of a germ tube. After initial infection, the fungus produces haustoria inside of the wheat cells and mycelium grows on the plant's outer surface. These conidia function as secondary inoculum as growth and reproduction repeat throughout the growing season.

Powdery mildew of wheat thrives in cool, humid climates and proliferates in cloudy weather conditions. The pathogen can also be an issue in drier climates if wheat fields are irrigated. Ideal temperatures for growth and reproduction of the pathogen are between and with growth ceasing above Dense, genetically similar plantings provide opportune conditions for growth of powdery mildew. Milk has long been popular with home gardeners and small scale organic growers as a treatment for powdery mildew. Milk is diluted with water (typically 1:10) and sprayed on susceptible plants at the first sign of infection, or as a preventative measure, with repeated weekly application often controlling or eliminating the disease. Studies have shown milk's effectiveness as comparable to some conventional fungicides, and better than benomyl and fenarimol at higher concentrations. Milk has proven effective in treating powdery mildew of summer squash, pumpkins, grapes, and roses. The exact mechanism of action is unknown, but one known effect is that ferroglobulin, a protein in whey, produces oxygen radicals when exposed to sunlight, and contact with these radicals is damaging to the fungus. Another way to control wheat powdery mildew is breeding in genetic resistance, using "R genes" (resistance genes) to prevent infection. There are at least 25 loci on the wheat genome that encode resistance to powdery mildew. If the particular variety of wheat has only one loci for resistance, the pathogen may be controlled only for a couple years. If, however, the variety of wheat has multiple loci for resistance, the crop may be protected for around 15 years. Because finding these loci can be difficult and time consuming, molecular markers are used to facilitate combining resistant genomes. is an HSP70 a family of heat shock proteins in Arabidopsis.

Powdery mildew can be found in all wheat growing areas of the United States but usually will be most severe in the east and southeast. It is more common in areas with a humid or semi arid environment where wheat is grown. Powdery mildew has become a more important disease in some areas because of increased application of nitrogen fertilizer, which favors the development of the fungus. Severe symptoms of powdery mildew can cause stunting of wheat. If unmanaged, this disease can reduce yields significantly by reducing photosynthetic areas and causes non seed producing tillers. Powdery mildew causes reduced kernel size and lower yields. The sooner powdery mildew begins to develop and how high on the plant it develops by flowering the larger the yield loss. Yield Losses up to 45 percent have been shown in Ohio on susceptible varieties when plants are infected early and weather favors disease.