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''Erysiphe alphitoides'' is a species of
fungus A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
which causes
powdery mildew Powdery mildew is a fungus, fungal disease that affects a wide range of plants. Powdery mildew diseases are caused by many different species of Ascomycota, ascomycete fungi in the order Erysiphales. Powdery mildew is one of the easier plant disea ...
on
oak An oak is a hardwood tree or shrub in the genus ''Quercus'' of the beech family. They have spirally arranged leaves, often with lobed edges, and a nut called an acorn, borne within a cup. The genus is widely distributed in the Northern Hemisp ...
trees.


Discovery and taxonomy

Today oak powdery mildew is one of the most common diseases in European forests but it is thought to not always have been present in these forests. The first reports of the disease were made in Paris and other regions of France, Spain, Luxembourg and the Netherlands in 1907. By 1908 the disease had spread into many other European countries, becoming an epidemic. By 1909 it had reached Russia and Turkey, followed by Brazil in 1912 and soon after was distributed around the world. The sudden outbreak of the disease puzzled
mycologist Mycology is the branch of biology concerned with the study of fungi, including their taxonomy, genetics, biochemical properties, and use by humans. Fungi can be a source of tinder, food, traditional medicine, as well as entheogens, poison, and ...
s as they were unsure about the origin of the pathogen. Its anamorph was easily distinguished from '' Phyllactinia guttata'' which had previously been reported to cause powdery mildew on oaks in Europe at low intensity. It did however share morphological similarities with '' Oïdium quercinum'', '' Calocladia penicillata'' and '' Microsphaera penicillata'' which had previously been reported to cause powdery mildew on oaks in Europe. The sudden appearance and high incidence of the disease made it unlikely that it was caused by any of these species however and instead some authors hypothesised that it was caused by the introduction of a new species from outside Europe. Powdery mildew had already been reported from North America so it was thought that the new disease could have been imported on American oaks into Europe. The absence of the disease on American red oaks growing in France however, made this unlikely. Four years after the initial outbreak, a teleomorph was found in south-east France which Arnaud and Fox identified as an American species, ''Microsphaera quercina'' (now included in '' Microsphaera alni''). A detailed morphological study by Griffon and Maublanc in 1912 suggested that the species was different from all previously described species and proposed the name of ''Microsphaera alphitoides'', referring to the
flour Flour is a powder made by Mill (grinding), grinding raw grains, List of root vegetables, roots, beans, Nut (fruit), nuts, or seeds. Flours are used to make many different foods. Cereal flour, particularly wheat flour, is the main ingredie ...
y appearance of the abundant white sporulation. This identification remained controversial until the 1940s. The origin of the disease was still uncertain however, in 1927, Raymond suggested that the disease may have been caused by a fungus identified in 1877 in Portugal that had been imported from Portuguese colonies. In 1980, Boesewinkel demonstrated that the same species was responsible for powdery mildew on ''
Quercus robur ''Quercus robur'', the pedunculate oak, is a species of flowering plant in the beech and oak family, Fagaceae. It is a large tree, native plant, native to most of Europe and western Asia, and is widely cultivated in other temperate regions. It ...
'' and on a
mango A mango is an edible stone fruit produced by the tropical tree '' Mangifera indica''. It originated from the region between northwestern Myanmar, Bangladesh, and northeastern India. ''M. indica'' has been cultivated in South and Southeast As ...
species from
New Zealand New Zealand () is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and List of islands of New Zealand, over 600 smaller islands. It is the List of isla ...
, supporting the hypothesis that the fungus had shifted its host. Recently,
molecular biology Molecular biology is a branch of biology that seeks to understand the molecule, molecular basis of biological activity in and between Cell (biology), cells, including biomolecule, biomolecular synthesis, modification, mechanisms, and interactio ...
has provided new tools for identifying species and elucidating their relatedness. A study of
ribosomal DNA The ribosomal DNA (rDNA) consists of a group of ribosomal RNA encoding genes and related regulatory elements, and is widespread in similar configuration in all domains of life. The ribosomal DNA encodes the non-coding ribosomal RNA, integral struc ...
from 33 samples of oak powdery mildew from Europe showed that ''E. alphitoides'' contains identical sequences to '' Oïdium mangiferae'', a major disease of mango in several tropical regions and '' Oïdium heveae'', an economically important pathogen of the
para rubber tree ''Hevea brasiliensis'', the Pará rubber tree, ''sharinga'' tree, seringueira, or most commonly, rubber tree or rubber plant, is a flowering plant belonging to the spurge family, Euphorbiaceae, originally native to the Amazon basin, but is now p ...
. These results support the findings of Boesewinkel and suggest that ''E. alphitoides'' and ''O. mangiferae'' could be
conspecific Biological specificity is the tendency of a characteristic such as a behavior or a biochemical variation to occur in a particular species. Biochemist Linus Pauling stated that "Biological specificity is the set of characteristics of living organism ...
. The origin of the species is still uncertain but Mougou ''et al.'' (2008) reported that ''E. alphitoides'' likely originated from the tropics and expanded its range to oak after being introduced to Europe.


Pathology

Only young developing leaves are susceptible to colonisation by ''E. alphitoides''. It only induces
necrosis Necrosis () is a form of cell injury which results in the premature death of cells in living tissue by autolysis. The term "necrosis" came about in the mid-19th century and is commonly attributed to German pathologist Rudolf Virchow, who i ...
when infection occurs very early during leaf development. Because the pathogen develops late in spring, after the first leaves of oak seedlings have developed, it is more prevalent on the second and third flushes of leaves that develop in July and August. This habit also reduces the severity of the disease on mature trees.


Effects on host

The disease can be very severe on ''
Quercus robur ''Quercus robur'', the pedunculate oak, is a species of flowering plant in the beech and oak family, Fagaceae. It is a large tree, native plant, native to most of Europe and western Asia, and is widely cultivated in other temperate regions. It ...
'' and ''
Quercus petraea ''Quercus petraea'', commonly known as the sessile oak, Welsh oak, Cornish oak, Irish oak or durmast oak, is a species of oak tree native to most of Europe and into Anatolia and Iran. The sessile oak is the national tree of Ireland, and an un ...
'' in Europe, particularly on young trees. In mature trees the disease is generally less damaging, but in combination with other factors such as defoliation by insects can contribute to tree decline. A study of the effects of ''E. alphitoides'' on ''Quercus robur'' found it decreased
stomatal conductance Stomatal conductance, usually measured in mmol m−2 s−1 by a porometer, estimates the rate of gas exchange (i.e., carbon dioxide uptake) and transpiration (i.e., water loss as water vapor) through the leaf stomata as determined by the degree of ...
by 15–30%, did not affect the leaf mass to area ratio, decreased the nitrogen content of the leaf and increased dark respiration.
Carbon fixation Biological carbon fixation, or сarbon assimilation, is the Biological process, process by which living organisms convert Total inorganic carbon, inorganic carbon (particularly carbon dioxide, ) to Organic compound, organic compounds. These o ...
was also reduced in infected leaves by about 40–50% in fully infected leaves and those that were heavily infected were shed earlier than those not infected. Despite it affecting the ability of leaves to
photosynthesis Photosynthesis ( ) is a system of biological processes by which photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical energy necessary to fuel their metabo ...
e, the effect is low and this is thought to explain why the disease has only moderate consequences for tree health despite heavy infections. Because the infection makes the plants less
shade tolerant In ecology, shade tolerance is a plant's ability to tolerate low light levels. The term is also used in horticulture and landscaping, although in this context its use is sometimes imprecise, especially in labeling of plants for sale in nursery (ho ...
, it may lead to the death of seedlings growing in shade.


Susceptible species

*''
Aesculus hippocastanum ''Aesculus hippocastanum'', the horse chestnut, is a species of flowering plant in the maple, Sapindus, soapberry and lychee family Sapindaceae. It is a large, deciduous, synoecious (hermaphroditic-flowered) tree. It is also called horse-chestnu ...
'' (horse chestnut) *''
Quercus An oak is a hardwood tree or shrub in the genus ''Quercus'' of the beech family. They have spirally arranged leaves, often with lobed edges, and a nut called an acorn, borne within a cup. The genus is widely distributed in the Northern Hemisp ...
'' species *''
Wisteria ''Wisteria'' is a genus of flowering plants in the legume family, Fabaceae (Leguminosae). The genus includes four species of woody twining vines that are native to China, Japan, Korea, Vietnam, southern Canada, the Eastern United States, and nor ...
''


References

{{Taxonbar, from=Q136225 alphitoides Fungal tree pathogens and diseases Fungi described in 1912 Fungus species