Botrytis allii
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''Botrytis allii'' is a
plant pathogen Plant pathology (also phytopathology) is the scientific study of diseases in plants caused by pathogens (infectious organisms) and environmental conditions (physiological factors). Organisms that cause infectious disease include fungi, oomy ...
, a fungus that causes neck rot in stored onions ('' Allium cepa'') and related crops. Its teleomorph is unknown, but other species of ''
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 ...
'' are
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 a ...
s of ''
Botryotinia ''Botryotinia'' is a genus of ascomycete fungi causing several plant diseases. The anamorphs of ''Botryotinia'' are mostly included in the " imperfect fungi" genus ''Botrytis''. The genus contains 22 species and one hybrid. Plant diseases caus ...
'' species. The species was first described scientifically by
Mancel Thornton Munn Mancel Thornton Munn (January 31, 1887 in Plainwell, Michigan – November 16, 1956 in Arcadia, California) was a New York State botanist, an Agronomy, agronomist, and an expert in crop seed testing who pioneered some of the early United States o ...
in 1917.


Biology

There are seven different species of ''Botrytis'' associated with onions in storage, but the rot induced by ''B. allii'' and ''B. aclada'' causes the greatest commercial loss. The two can be distinguished microscopically; the
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 t ...
of ''B. allii'' have a maximum length of 15 μm and mean size of 10.2 × 5.7 
μm The micrometre ( international spelling as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer ( American spelling), also commonly known as a micron, is a unit of length in the International System of Uni ...
, while the conidia of ''B. aclada'' have a maximum length of 12 μm and mean size of 8.6 × 4.6 μm. The infection is present in the field but does not manifest itself until after harvest, however there may be a falling off of vigour while the crop is still growing, particularly in cool, moist weather. In the stored crop, the rot typically starts in the neck of the
bulb In botany, a bulb is structurally a short stem with fleshy leaves or leaf basesBell, A.D. 1997. ''Plant form: an illustrated guide to flowering plant morphology''. Oxford University Press, Oxford, U.K. that function as food storage organs dur ...
but can occur in other parts if there is physical injury. The scales inside the bulb become progressively translucent and watery and a
mycelium Mycelium (plural mycelia) is a root-like structure of a fungus consisting of a mass of branching, thread-like hyphae. Fungal colonies composed of mycelium are found in and on soil and many other substrates. A typical single spore germinates ...
develops between them. A mass of grey
conidiophore 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 ...
s and
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 t ...
develop on the mycelium, and blackish sclerotia form at the site of the initial infection. In onion crops grown for the production of seed, ''Botrytis allii'' can cause spotting and
girdling Girdling, also called ring-barking, is the complete removal of the bark (consisting of cork cambium or "phellogen", phloem, cambium and sometimes going into the xylem) from around the entire circumference of either a branch or trunk of a woody ...
of the stipe (stem) and develop on the sheath that protects the
inflorescence An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are forme ...
and on the flowers themselves. Concentric grey rings may form as the fungus sporulates and the crop may lodge (become flattened). It has been shown that a major source of the pathogen is infected seed. In 1973, 71% of commercially available seed was found to be contaminated, and the infection was found to persist for over three years in seeds being stored. In the seedling, infection with ''B. allii'' does not produce any symptoms, but the fungus spreads between plants as the conidiophores release
spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, ...
s into the air. The leaf tips are invaded first, the infection spreading down the leaves and into the neck of the bulb, where its presence only becoming apparent when the foliage dies down at the end of the season.


Hosts

''Botrytis allii'' grows on ''Allium'' species, including onion (''A. cepa''), aggregating onion (''A. cepa'' var. ''aggregatum''),
shallot The shallot is a botanical variety (a cultivar) of the onion. Until 2010, the (French red) shallot was classified as a separate species, ''Allium ascalonicum''. The taxon was synonymized with '' Allium cepa'' (the common onion) in 2010, as the ...
(''A. cepa'' var. ''ascalonicum''),
garlic Garlic (''Allium sativum'') is a species of bulbous flowering plant in the genus '' Allium''. Its close relatives include the onion, shallot, leek, chive, Welsh onion and Chinese onion. It is native to South Asia, Central Asia and northeas ...
(''A. sativum'') and
leek The leek is a vegetable, a cultivar of '' Allium ampeloprasum'', the broadleaf wild leek ( syn. ''Allium porrum''). The edible part of the plant is a bundle of leaf sheaths that is sometimes erroneously called a stem or stalk. The genus '' Al ...
(''A. porrum''). It may also infect wild ''Allium'' species and can grow saprophytically on decaying crop residues such as
cereal A cereal is any grass cultivated for the edible components of its grain (botanically, a type of fruit called a caryopsis), composed of the endosperm, germ, and bran. Cereal grain crops are grown in greater quantities and provide more food ...
,
pea The pea is most commonly the small spherical seed or the seed-pod of the flowering plant species ''Pisum sativum''. Each pod contains several peas, which can be green or yellow. Botanically, pea pods are fruit, since they contain seeds and d ...
and bean straw. It is capable of colonizing and producing spores on sterilized
poppy straw Poppy straw (also known as opium straw, mowed opium straw, crushed poppy capsule, poppy chaff, or poppy husk) is derived from opium poppies (''Papaver somniferum'') that are harvested when fully mature and dried by mechanical means, minus the rip ...
(''
Papaver somniferum ''Papaver somniferum'', commonly known as the opium poppy or breadseed poppy, is a species of flowering plant in the family Papaveraceae. It is the species of plant from which both opium and poppy seeds are derived and is also a valuable orname ...
'').


Commercial importance

Neck rot caused by ''Botrytis allii'' occurs in regions all over the world, but, it is most prevalent in temperate regions due to their conducive climates.Toit, Lindsey Du. "Botrytis Species Associated with Neck Rot, Scape Blight, and Umbel Blight of Onion." ''Detection and Identification of Botrytis Species Associated with Neck Rot, Scape Blight, and Umbel Blight of Onion''.
Plant Health Progress The American Phytopathological Society (APS) is an international scientific organization devoted to the study of plant diseases (phytopathology). APS promotes the advancement of modern concepts in the science of plant pathology and in plant healt ...
, 27 Aug. 2006. Web. 17 Nov. 2016.
''Botrytis allii'' used to cause significant losses in the onion bulb industry in the UK. Fortunately, by utilization of better harvesting and curing methods, the disease prevalence has decreased. It is important that the US utilize these methods as well because the onion is a highly produced agronomic crop. In 2005, the US grew 65,000
hectare The hectare (; SI symbol: ha) is a non-SI metric unit of area equal to a square with 100- metre sides (1 hm2), or 10,000 m2, and is primarily used in the measurement of land. There are 100 hectares in one square kilometre. An acre i ...
s of onions with a
farm gate value The farm gate value of a cultivated product in agriculture and aquaculture is the market value of a product minus the selling costs (transport costs, marketing costs). The market value is not the same as the price farmers get for their product, ...
of $922 million. ''Botrytis allii'' poses a threat to that value because it can potentially cause a 50% reduction in yield if left unchecked. These losses caused by the disease may be even greater than reported. Detection of this disease is difficult due to the morphologically indistinguishable characteristics of ''Botrytis'' species cultured on
agar Agar ( or ), or agar-agar, is a jelly-like substance consisting of polysaccharides obtained from the cell walls of some species of red algae, primarily from ogonori (''Gracilaria'') and "tengusa" (''Gelidiaceae''). As found in nature, agar i ...
. Higher quality detection methods of ''Botrytis allii'' are starting to be implemented to better differentiate it from other closely related ''Botrytis'' species. Within the last 15 years, scientists have been able to get a more accurate picture of ''Botrytis allii'' distribution through the use of PCR- RFLP detection methods.


Management

''Botrytis allii’s'' main agronomic host is the onion (''Allium cepa)''. In order to control this disease, most agriculturalists utilize the
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 ...
benomylMaude, R.B., M.R. Shipway, A.H. Presly, and D. O'Connor. "The Effects of Direct Harvesting and Drying Systems on the Incidence and Control of Neck Rot (Botrytis Allii) in Onions." ''- MAUDE''. N.p., 1984. Web. 27 Oct. 2016.''.'' Benomyl is applied directly to the seed, as a pretreatment, before planting. However, the pretreatment must be used in addition to correct farming practices to further minimize the spread of ''Botrytis allii'' spores. For instance, benomyl works best if you dry the onions in a environment after harvest. Lower drying temperatures, such as , during the post harvest period have been known to increase disease presence. In addition, removal of the onions from the field in the first 48 hours after mechanical removal of the top has also shown to lower post harvest disease. Moreover, it is important to pay attention to your fertilization schedule. High levels of
nitrogen fertilizer A fertilizer (American English) or fertiliser (British English; see spelling differences) is any material of natural or synthetic origin that is applied to soil or to plant tissues to supply plant nutrients. Fertilizers may be distinct from ...
during the growing season may cause delayed maturity. This is an issue because the onions that have the best chance of avoiding disease are the ones that have reached full maturity by the time they are harvested.Mcdonald, Mary Ruth, Maria De Los Angeles Jaime, and Marilyn H.y. Hovius. "Management of Diseases of Onions and Garlic." ''Diseases of Fruits and Vegetables: Volume II'' (2004): 149-200. Web. 15 Nov. 2016.
Crop rotation Crop rotation is the practice of growing a series of different types of crops in the same area across a sequence of growing seasons. It reduces reliance on one set of nutrients, pest and weed pressure, and the probability of developing resistant ...
s can also help control disease outbreak. The rotation should take place over at least two years and crops not related to onions should be planted. Onion fields should be separated from each other because the spores of ''Botrytis allii'' are able to travel far distances.


Pathogenesis

''Botrytis allii'' is able to use
polysaccharide Polysaccharides (), or polycarbohydrates, are the most abundant carbohydrates found in food. They are long chain polymeric carbohydrates composed of monosaccharide units bound together by glycosidic linkages. This carbohydrate can react with w ...
-degrading
enzyme Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products ...
s to enter their onion host.Mankarios, A.T., and J. Friend. "Polysaccharide-degrading Enzymes of Botrytis Allii and Sclerotium Cepivorum. Enzyme Production in Culture and the Effect of the Enzyme on Isolated Onion Cell Walls." ''Cell Wall Plant Polysaccharide-degrading Enzymes in Autolysis of Botrytis Cinerea''.
Science Direct ScienceDirect is a website which provides access to a large bibliographic database of scientific and medical publications of the Dutch publisher Elsevier. It hosts over 18 million pieces of content from more than 4,000 academic journals and 30 ...
, July 1980. Web. 14 Nov. 2016.
These enzymes are able to degrade onion
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering mec ...
components such as sodium polypectate, citrus pectin, lupin galactan, araban,
xylan Xylan (; ) ( CAS number: 9014-63-5) is a type of hemicellulose, a polysaccharide consisting mainly of xylose residues. It is found in plants, in the secondary cell walls of dicots and all cell walls of grasses. Xylan is the third most abundan ...
and
carboxymethyl cellulose Carboxymethyl cellulose (CMC) or cellulose gum is a cellulose derivative with carboxymethyl groups (-CH2-COOH) bound to some of the hydroxyl groups of the glucopyranose monomers that make up the cellulose backbone. It is often used as its sodiu ...
. The most active enzyme that ''Botrytis allii'' uses is
polygalacturonase Endo-polygalacturonase (EC 3.2.1.15, pectin depolymerase, pectolase, pectin hydrolase, and poly-α-1,4-galacturonide glycanohydrolase; systematic name (1→4)-α-D-galacturonan glycanohydrolase (endo-cleaving)) is an enzyme that hydrolyzes the α- ...
, which is used to degrade sodium polypectate. When the fungus attempts to penetrate the epidermal cells of the onion, there is an observed accumulation of granular deposits, called reaction material, that are found between the cell wall and
plasma membrane The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of all cells from the outside environment (t ...
.McLusky, S., M. Bennett, M. Beale, M. Lewis, P. Gaskin, and J. Mansfield. "Cell Wall Alterations and Localized Accumulation of Feruloyl-3′-methoxytyramine in Onion Epidermis at Sites of Attempted Penetration by Botrytis Allii Are Associated with Actin Polarisation, Peroxidase Activity and Suppression of Flavonoid Biosynthesis." ''
Wiley Online Library John Wiley & Sons, Inc., commonly known as Wiley (), is an American multinational publishing company founded in 1807 that focuses on academic publishing and instructional materials. The company produces books, journals, and encyclopedias, in ...
''.
The Plant Journal ''The Plant Journal'' is a peer-reviewed scientific journal of plant science published by Wiley-Blackwell for the Society for Experimental Biology. It was established in 1991 and is currently edited by Lee Sweetlove. The journal is published twi ...
, Mar. 1999. Web. 26 Oct. 2016.
The reaction material correlates with reduced fungal growth between the cell walls where it is found. During infection, the onion will also produce hydroxycinnamic acid
amide In organic chemistry, an amide, also known as an organic amide or a carboxamide, is a compound with the general formula , where R, R', and R″ represent organic groups or hydrogen atoms. The amide group is called a peptide bond when it i ...
, which is a common plant defense response. There may also be some
cytoskeletal The cytoskeleton is a complex, dynamic network of interlinking protein filaments present in the cytoplasm of all cells, including those of bacteria and archaea. In eukaryotes, it extends from the cell nucleus to the cell membrane and is compo ...
rearrangements within the onion so that it can provide a better route to deliver the phenolic products to the fungal penetration site.


References

{{Taxonbar, from=Q10433881 Fungal plant pathogens and diseases Monocot diseases Fungi described in 1850 Sclerotiniaceae