Urocystis Agropyri
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''Urocystis agropyri'' is a
fungal A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from th ...
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, oomyc ...
that causes flag smut on wheat.


Biology

The flag smut fungus, ''U. agropyri'', is a
basidiomycete Basidiomycota () is one of two large divisions that, together with the Ascomycota, constitute the subkingdom Dikarya (often referred to as the "higher fungi") within the kingdom Fungi. Members are known as basidiomycetes. More specifically, Basi ...
. It produces
basidiospores A basidiospore is a reproductive spore produced by Basidiomycete fungi, a grouping that includes mushrooms, shelf fungi, rusts, and smuts. Basidiospores typically each contain one haploid nucleus that is the product of meiosis, and they are pro ...
and
teliospores Teliospore (sometimes called teleutospore) is the thick-walled resting spore of some fungi ( rusts and smuts), from which the basidium arises. Development They develop in '' telia'' (sing. ''telium'' or ''teliosorus''). The telial host is the p ...
. This pathogen is found globally, but is most problematic in Australia and India.


Hosts

''Urocystis agropyri'' infects wheat (''
Triticum aestivum Common wheat (''Triticum aestivum''), also known as bread wheat, is a cultivated wheat species. About 95% of wheat produced worldwide is common wheat; it is the most widely grown of all crops and the cereal with the highest monetary yield. Ta ...
'' L.), among other grass species. Relatives of ''U. agropyri'' infect other grasses and are frequently identified as the causal organism of flag smut on wheat, but there is debate still as to whether they are the same organism or different strains.


Symptoms

Flag smut is a systemic disease that starts in young tissues. Early symptoms include "leprous" spots and bending or twisting of
coleoptiles Coleoptile is the pointed protective sheath covering the emerging shoot in monocotyledons such as grasses in which few leaf primordia and shoot apex of monocot embryo remain enclosed. The coleoptile protects the first leaf as well as the growing ...
. Older plant leaves have white striations that eventually turn silvery gray, which is evidence of the pathogen's impending sporulation. Additionally, infected plants may have stunted growth, increased leaf production, sterile seeds, and failure to produce heads or have successful leaf expansion.


Diagnosis

Leaves will appear streaked with stripes, which are white, gray or black. The leaf discoloration is due to fungal structures called sori, which are clusters of spore-bearing structures. During sporulation, sori burst through leaves releasing teliospores and cause leaves to appear tattered. These spores are reddish brown, smoothly rounded, and they tend to be in clumps of 5-6 with sterile cells around them. The clumped spores are often referred to as "spore balls" and measure about 20-50 microns. Large quantities of ''U. agropyri'' spores look like brown or black dust.


Disease cycle

''U. agropyri'' produces teliospores, which may be wind dispersed or distributed through soils via machinery or animals. In soil, a
dikaryotic The dikaryon is a cell nucleus, nuclear feature which is unique to certain fungi. (The green alga ''Derbesia'' had been long considered an exception, until the heterokaryotic hypothesis was challenged by later studies.) Compatible cell-types can fus ...
teliospore germinates, meiosis occurs, then mitosis, and this gives rise to up to four basidiospores, each containing a single nucleus. Basidiospores germinate on seedlings, and each hypha undergoes plasmogamy with a compatible hypha. In this, one nucleus transfers to the other hypha, reestablishing the dikaryotic state of the fungus. The hyphae form
appressoria An appressorium is a specialized cell typical of many fungal plant pathogens that is used to infect host plants. It is a flattened, hyphal "pressing" organ, from which a minute infection peg grows and enters the host, using turgor pressure capable ...
which penetrate the coleoptile of an emerging seeds' shoot through the epidermal tissue, then hyphae grow between vascular bundles of the leaves. Some hyphal cells give rise to smut sori, bearing teliospores, which emerge through the leaf tissue for wind dispersal. Teliospores come to rest in soils, and when conditions are right, they give rise to more basidiospores, further spreading the infection. Alternatively, teliospores can form in seeds when the mycelia grows throughout the plant, in which case they germinate within the seed to give rise to new infection, again via basidiospore production. Teliospores overwinter in the soil, senescent plant tissues, and in seeds. These spores maintain germination viability for 3–7 years.


Environmental conditions

This pathogen prefers arid summers, moderate temperatures, and mild winters. Flag smut fungi germinate in dry soils when the temperature ranges from 40-80 °F. Flag smut has been reported in Australia, the United States, Canada, South Africa, China, Japan, India, Egypt, and Pakistan. Cultivation practices that leave plant debris on soil surfaces enhances ''U. agropyri's'' success, as does sowing wheat in winter rather than spring. Mild winters improves the pathogen's ability to establish infections for seeds sown in autumn or winter; spring plantings give the fungus less opportunity to establish.


Disease management

Generally, strategies to prevent flag smut include use of disease resistant cultivars, chemical seed treatments, and crop rotation to reduce amount of inocula present. Since ''U. agropyri'' spores germinate in dry soils, maintaining wetter soils helps to diminish their viability. Carboxin is a commonly used fungicide on seeds, which works well to prevent onset of disease. In addition to seed treatments, application of systemic fungicides early in the growing season and at low doses is effective at controlling the disease. Sowing seeds shallowly in soil also helps to reduce disease occurrence.


Impact


Yield loss

Losses from flag smut are usually considered on a field by field basis. Fields planted with susceptible cultivars that happen to become infected will generally be total losses. Australia is among a handful of places that still suffer losses resulting from flag smut. Average losses in Australia are greater than AUS$50 million. Australia experienced the greatest loss known early in the 20th century due to susceptibility of popular wheat varieties and lack of fungicides at the time. In the 1960s, new varieties were planted, which supported a resurgence of disease. Occasionally, total crop losses still occur, but the typical loss range is 5-20%. Losses may be either direct or indirect, as affected plants may be more susceptible to other biotic or abiotic stresses. India reported losses from flag smut in the 1940s through the 1970s, and this averaged about 15,000 tons each year. India and the United States currently have low incidence of this disease due to deployment of resistant cultivars. The occurrence of the disease in the United States was limited to the Pacific Northwest and was the result of the introduction of susceptible cultivars in the mid-twentieth century.


Historical

''Urocystis agropyri'' is a global problem in wheat-growing regions, especially where environmental conditions suit the fungus. Flag smut was first reported from ''
Agropyron ''Agropyron'' is a genus of Eurasian plants in the grass family), native to Europe and Asia but widely naturalized in North America. Species in the genus are commonly referred to as wheatgrass. ; Species * ''Agropyron badamense'' - Tajikistan, ...
'' spp. in
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
in 1848, misattributed to '' Uredo agropyri''. It was probably present in
South Australia South Australia (commonly abbreviated as SA) is a state in the southern central part of Australia. It covers some of the most arid parts of the country. With a total land area of , it is the fourth-largest of Australia's states and territories ...
before 1868, known as "black rust", but is only definitely identified starting from that year. It was at first identified by Wolff 1873 as '' U. occulta'', but
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, f ...
morphology differentiated it in the opinion of Körnicke 1877. Later, other countries identified it as the pathogen that was introduced to world crops through trading of seeds and infected cultivars. The development of flag smut in Australia in the nineteenth century, followed by the United States in the twentieth century, is likely due to the spread and exchange of contaminated seeds. Quarantine regulations restricted the movement of infected seed, chaff, and farm machinery from endemic areas. They were first enacted in the USA in the twentieth century to effectively inhibit the spread of disease. Similar regulations were later adopted in Belgium, Germany, the Netherlands, the United Kingdom and many other countries. Flag smut is widespread across Australia and was particularly problematic in the 1920s until resistant cultivars were discovered and put into use. Between 1955 and 1971, flag smut damage and distribution increased in the Pacific Northwest of the United States when the use of several susceptible wheat cultivars was coupled with deep seeding in early autumn planting. Through use of resistant cultivars, the Pacific Northwest's flag smut issue no longer poses a significant threat to yields.


References


External links


Index Fungorum

USDA ARS Fungal Database
{{Taxonbar, from=Q7900814 Fungal plant pathogens and diseases Wheat diseases Ustilaginomycotina Fungi described in 1848