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''Botrytis cinerea'' is a necrotrophic fungus that affects many plant species, although its most notable hosts may be wine grapes. In viticulture, it is commonly known as "botrytis bunch rot"; in horticulture, it is usually called "grey mould" or "gray mold". The fungus gives rise to two different kinds of infections on grapes. The first, grey rot, is the result of consistently wet or humid conditions, and typically results in the loss of the affected bunches. The second, noble rot, occurs when drier conditions follow wetter, and can result in distinctive sweet dessert wines, such as Sauternes or the Aszú of Tokaji/ Grasă de Cotnari. The species name ''Botrytis cinerea'' is derived from the Latin for "grapes like ashes"; although poetic, the "grapes" refers to the bunching of the fungal spores on their conidiophores, and "ashes" just refers to the greyish colour of the spores ''en masse''. The fungus is usually referred to by its
anamorph In mycology, the terms teleomorph, anamorph, and holomorph apply to portions of the life cycles of fungi in the phyla Ascomycota and Basidiomycota: *Teleomorph: the sexual reproductive stage (morph), typically a fruiting body. *Anamorph: an ase ...
(asexual form) name, because the sexual phase is rarely observed. The teleomorph (sexual form) is an
ascomycete Ascomycota is a phylum of the kingdom Fungi that, together with the Basidiomycota, forms the subkingdom Dikarya. Its members are commonly known as the sac fungi or ascomycetes. It is the largest phylum of Fungi, with over 64,000 species. The defi ...
, ''Botryotinia fuckeliana'', also known as ''Botryotinia cinerea'' (see taxonomy box).


Etymology

"
Botrytis Botrytis may refer to: * ''Botrytis'' (fungus), the anamorphs of fungi of the genus ''Botryotinia'' **''Botrytis cinerea'', a mold important in wine making *Botrytis, the cauliflower cultivar group of ''Brassica oleracea ''Brassica oleracea'' is ...
" is derived from the Ancient Greek ''botrys ''(βότρυς) meaning "grapes", combined with the New Latin suffix ''-itis'' for disease. ''Botryotinia fuckeliana'' was named by mycologist Heinrich Anton de Bary in honor of another mycologist,
Karl Wilhelm Gottlieb Leopold Fuckel Karl Wilhelm Gottlieb Leopold Fuckel (3 February 1821 – 8 May 1876) was a German botanist who worked largely on fungi. He worked as an apothecary from 1836 to 1852, afterwards deriving income from a vineyard he owned in Oestrich im Rheingau.< ...
. Synonyms for the sexual stage are: * ''Botrytis fuckeliana'' N.F. Buchw., (1949) * ''Botrytis gemella'' (Bonord.) Sacc., (1881) * ''Botrytis grisea'' (Schwein.) Fr., (1832) * ''Botrytis vulgaris'' (Pers.) Fr., (1832) * ''Haplaria grisea'' Link, (1809) * ''fuckeliana'' de Bary * ''Phymatotrichum gemellum'' Bonord., (1851) * ''Polyactis vulgaris'' Pers., (1809) * ''Sclerotinia fuckeliana'' (de Bary) Fuckel, (1870)


Hosts and symptoms


Hosts

The disease, gray mold, affects more than 200
dicotyledon The dicotyledons, also known as dicots (or, more rarely, dicotyls), are one of the two groups into which all the flowering plants (angiosperms) were formerly divided. The name refers to one of the typical characteristics of the group: namely, t ...
ous plant species and a few monocotyledonous plants found in temperate and subtropical regions, and potentially over a thousand species. Serious economic losses can be a result of this disease to both field and greenhouse grown crops. The causal agent, ''Botrytis cinerea'' can infect mature or senescent tissues, plants prior to harvest, or seedlings. There is a wide variety of hosts infected by this pathogen including protein crops, fiber crops, oil crops, and horticultural crops. Horticultural crops include vegetables (examples are chickpeas, lettuce, broccoli, and beans) and small fruit crops (examples are grape, strawberry, raspberry, and blackberry), these are most severely affected and devastated by gray mold. Plant organs affected include fruits, flowers, leaves, storage organs, and shoots.


Symptoms and signs

Symptoms vary across plant organs and tissues. ''B. cinerea'' is a soft rot that will have a collapsed and water soaked appearance on soft fruit and leaves. Brown lesions may develop slowly on undeveloped fruit. Twigs infected with gray mold will die back. Blossoms will cause fruit drop and injury, such as ridging on developing and mature fruit. Symptoms are visible at wound sites where the fungus begins to rot the plant. Gray masses with a velvety appearance are conidia on the plant tissues are a sign of plant pathogen. These conidia are asexual spores that will continue to infect the plant and surrounding hosts throughout the growing season making this a polycyclic disease. Plants can produce localized lesions when a pathogen attacks. An oxidative burst causes hypersensitive cell death called a hypersensitive response (HR). This soft rot can trigger HR to assist in colonization. ''Botrytis cinerea'', as a necrotrophic pathogen, exploits the dead tissue for its pathogenicity or its ability to cause disease. Susceptible plants cannot use the HR to protect against ''B. cinerea''. See: *
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Biology

''Botrytis cinerea'' is characterized by abundant hyaline
conidia A conidium ( ; ), sometimes termed an asexual chlamydospore or chlamydoconidium (), is an asexual, non-motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word for dust, ('). They are also called mitospores due to the ...
(asexual spores) borne on grey, branching tree-like conidiophores. The fungus also produces highly resistant sclerotia as survival structures in older cultures. It overwinters as sclerotia or intact mycelia, both of which germinate in spring to produce conidiophores. The conidia, dispersed by wind and by rain-water, cause new infections. ''B. cinerea'' performs an asexual cycle over the summer season. Different strains show considerable genetic variability. '' Gliocladium roseum'' is a fungal parasite of ''B. cinerea''. The hypothetical protein BcKMO was shown to positively regulate growth and development. It showed a great similarity to the kynurenine 3-monooxygenase encoding gene in eukaryotes. Overexpression of the gene ' produces altered versions of the transcription factor ''
mrr1 Agriculture is a significant sector in Economy of California, California's economy, producing nearly $50 billion in revenue in 2018. There are more than 400 commodity crops grown across California, including a significant portion of all fruits, ...
'', which in turn confer a multiple fungicide resistance phenotype known as . An even higher overexpression yields ''mrr1'' composed partly of , yielding
MDR1h Agriculture is a significant sector in Economy of California, California's economy, producing nearly $50 billion in revenue in 2018. There are more than 400 commodity crops grown across California, including a significant portion of all fruits, ...
phenotypes with even more anilinopyramidine- and phenylpyrrole- resistance.


Environment

Gray mold favors moist, humid, and warm environmental conditions between . Temperature, relative humidity, and wetness duration produce a conducive environment that is favorable for inoculation of mycelium or
conidia A conidium ( ; ), sometimes termed an asexual chlamydospore or chlamydoconidium (), is an asexual, non-motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word for dust, ('). They are also called mitospores due to the ...
. Controlled environments, such as crop production greenhouses, provide the moisture and high temperatures that favor the spreading and development of the pathogen ''B. cinerea.'' Standing water on plant leaf surfaces provides a place for spores to germinate. Humid conditions can result from improper irrigation practice, plants placed too close together, or the structure of the greenhouse not allowing for efficient ventilation and air flow. Ventilation at night significantly reduces the incidence of gray mold. Melanized sclerotium allows ''B. cinerea'' to survive for years in the soil. Sclerotia and the asexual conidia spores contribute to the widespread infection of the pathogen. A low pH is preferred by the gray mold to perform well. ''B. cinerea'' can acidify its environment by secreting
organic acid An organic acid is an organic compound with acidic properties. The most common organic acids are the carboxylic acids, whose acidity is associated with their carboxyl group –COOH. Sulfonic acids, containing the group –SO2OH, are rel ...
s, like oxalic acid. By acidifying its surroundings, cell wall degrading enzymes (CWDEs) are enhanced, plant-protection enzymes are inhibited,
stoma In botany, a stoma (from Greek ''στόμα'', "mouth", plural "stomata"), also called a stomate (plural "stomates"), is a pore found in the epidermis of leaves, stems, and other organs, that controls the rate of gas exchange. The pore is bor ...
tal closure is deregulated, and pH signaling is mediated to facilitate its pathogenesis.


Viticulture

In the ''Botrytis'' infection known as "noble rot" ('' pourriture noble'' in
French French (french: français(e), link=no) may refer to: * Something of, from, or related to France ** French language, which originated in France, and its various dialects and accents ** French people, a nation and ethnic group identified with Franc ...
, or '' Edelfäule'' in German), the fungus removes water from the grapes, leaving behind a higher percent of solids, such as sugars, fruit acids and minerals. This results in a more intense, concentrated final product. The wine is often said to have an aroma of honeysuckle and a bitter finish on the palate. A distinct fermentation process initially caused by nature, the combination of geology, climate and specific weather led to the particular balance of beneficial fungus while leaving enough of the grape intact for harvesting. The Chateau d'Yquem is the only Premier Cru Supérieur, largely due to the vineyard's susceptibility to noble rot. ''Botrytis'' complicates winemaking by making
fermentation Fermentation is a metabolic process that produces chemical changes in organic substrates through the action of enzymes. In biochemistry, it is narrowly defined as the extraction of energy from carbohydrates in the absence of oxygen. In food ...
more complex. ''Botrytis'' produces an anti-fungal compound that kills yeast and often results in the fermentation stopping before the wine has accumulated sufficient levels of alcohol. Botrytis bunch rot is another condition of grapes caused by ''B. cinerea'' that causes great losses for the wine industry. It is always present on the fruitset, however, it requires a wound to start a bunch rot infection. Wounds can come from insects, wind, accidental damage, etc. To control botrytis bunch rot there are a number of
fungicide Fungicides are biocidal chemical compounds or biological organisms used to kill parasitic fungi or their spores. A fungistatic inhibits their growth. Fungi can cause serious damage in agriculture, resulting in critical losses of yield, quality, ...
s available on the market. Generally, these should be applied at bloom, bunch closure and veraison (the most important being the bloom application). Some winemakers are known to use the German method of fermentation and prefer having a 5% bunch rot rate in their grapes and will usually hold the grapes on the vine a week longer than normal.


Horticulture

''Botrytis cinerea'' affects many other plants.


Strawberries

It is economically important on soft fruits such as
strawberries The garden strawberry (or simply strawberry; ''Fragaria × ananassa'') is a widely grown hybrid species of the genus '' Fragaria'', collectively known as the strawberries, which are cultivated worldwide for their fruit. The fruit is widely ap ...
and bulb crops. Unlike wine grapes, the affected strawberries are not edible and are discarded. To minimize infection in strawberry fields, good ventilation around the berries is important to prevent moisture being trapped among leaves and berries. A number of bacteria have been proven to act as natural antagonists to ''B. cinerea'' in controlled studies.Donmez, M. F.; Esitken, A.; Yildiz, H.; Ercisli, S
Biocontrol of ''Botrytis Cinerea'' on Strawberry Fruit by Plant Growth Promoting Bacteria
''The Journal of Animal & Plant Sciences'', 21(4), 2011: pp. 758-763, ISSN 1018-7081.


Other plants

In greenhouse horticulture, ''Botrytis cinerea'' is well known as a cause of considerable damage in tomatoes. The infection also affects
rhubarb Rhubarb is the fleshy, edible stalks ( petioles) of species and hybrids (culinary rhubarb) of ''Rheum'' in the family Polygonaceae, which are cooked and used for food. The whole plant – a herbaceous perennial growing from short, thick rhizo ...
, snowdrops,
white meadowfoam ''Limnanthes alba'' is a species of flowering plant in the meadowfoam family known by the common name white meadowfoam. It is native to California and Oregon, where it grows in wet grassy habitat, such as vernal pools and moist spots in woodland ...
, western hemlock, Douglas-fir cannabis, and '' Lactuca sativa''. UV-C treatment against ''B. cinerea'' was investigated by Vàsquez ''et al.'', 2017. They find it increases phenylalanine ammonia-lyase activity and production of phenolics. This in turn decreases ''L. sativa''s susceptibility. Potassium bicarbonate-based fungicide may be used.


Human disease

''Botrytis cinerea'' mold on grapes may cause "winegrower's lung", a rare form of hypersensitivity pneumonitis (a respiratory allergic reaction in predisposed individuals).


Mycoviruses of ''Botrytis cinerea''

''Botrytis cinerea'' not only infects plants, it also hosts several
mycovirus Mycoviruses (Ancient Greek: μύκης ' ("fungus") + Latin '), also known as mycophages, are viruses that infect fungi. The majority of mycoviruses have double-stranded RNA (dsRNA) genomes and isometric particles, but approximately 30% have po ...
es itself (see the table/image). A range of phenotypic alterations due to the mycoviral infection have been observed from symptomless to mild impact, or more severe phenotypic changes including reduction in pathogenicity, growth/suppression of mycelia, sporulation and sclerotia production, formation of abnormal colony sectors (Wu et al., 2010) and virulence.


Management

''Botrytis cinerea'' can be managed through cultural, chemical, and biological practices. There are no resistant species to the gray mold rot. Gray mold can be culturally controlled by monitoring the amount and timing of fertilizer applications to reduce the amount of fruit rot. Excessive application of nitrogen will increase the incidence of disease while not improving yields. Not planting cultivars that have an upright or dense growth habit can reduce disease as these limit airflow and are favorable for the pathogen. Spacing of plants so they are not touching will increase airflow allowing the area to dry out and reduce the spread of disease. Pruning or purposeful removal of diseased, dead, or overgrown limbs on a regular schedule can also help to improve air movement. Sanitation by removing dead or dying plant tissue in the fall will decrease inoculum levels as there is no debris for the sclerotium or mycelia to overwinter. Removing debris in the spring will remove inoculum from the site. Disposal of berries during harvest that have signs and symptoms of gray mold will reduce inoculum for the following year. Biochar, a form of charcoal, can be applied as a
soil amendment A soil conditioner is a product which is added to soil to improve the soil’s physical qualities, usually its fertility (ability to provide nutrition for plants) and sometimes its mechanics. In general usage, the term "soil conditioner" is often ...
to strawberry plants to reduce the severity of the fungal disease by stimulating defense pathways within the plant. Gray mold can be chemically controlled with well-timed fungicide applications starting during the first bloom. Timing can reduce the chance of resistance and will save on costs. Biological controls or
microbial antagonist In phytopathology, antagonism refers to the action of any organism that suppresses or interferes with the normal growth and activity of a plant pathogen, such as the main parts of bacteria or fungus, fungi. These organisms can be used for pest co ...
s used for disease suppression, have been successfully used in Europe and Brazil in the form of fungi-like ''
Trichoderma harzianum ''Trichoderma harzianum'' is a fungus that is also used as a fungicide. It is used for foliar application, seed treatment and soil treatment for suppression of fungal pathogens causing various fungal plant diseases. Commercial biotechnological p ...
'' Rifai and ''Clonostachys rosea'' f. ''rosea'' Bainier (syn. ''Gliocladium roseum''). ''Trichoderma'' species especially, have been shown to control gray mold. Multiple fungicide resistance is a problem in many production areas.


See also

* Botrydial


References


External links


Genome information for ''Botrytis cinerea''

Genome analysis of ''Botrytis cinerea''
*
TheWineDoctor.com
* * * {{Authority control Sclerotiniaceae Fungal plant pathogens and diseases Small fruit diseases Fungal strawberry diseases Fungal citrus diseases Fungal grape diseases Oenology Fungi described in 1794