Tilletia Tritici
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''Tilletia caries'' (
synonymous A synonym is a word, morpheme, or phrase that means precisely or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are a ...
with ''Tilletia tritici'') 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, Basid ...
that causes
common bunt Common bunt, also known as hill bunt, Indian bunt, European bunt, stinking smut or covered smut, is a disease of both spring and winter wheats. It is caused by two very closely related fungi, '' Tilletia tritici'' (syn. '' Tilletia caries'') and ' ...
of wheat. The common names of this disease are stinking bunt of wheat and stinking smut of wheat. This pathogen infects wheat, rye, and various other grasses. ''T. caries'' is economically and agriculturally important because it reduces both the wheat yield and grain quality.


Life cycle

Infection of the wheat occurs during
germination Germination is the process by which an organism grows from a seed or spore. The term is applied to the sprouting of a seedling from a seed of an angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the spores of fungi, ...
of the plant seed and is favored by cool, wet conditions. Optimum conditions for spore germination are soil temperatures in the range of . Bunt fungi overwinter as dikaryotic
teliospore Teliospore (sometimes called teleutospore) is the thick-walled resting spore of some fungi (Rust (fungus), rusts and Smut (fungus), smuts), from which the basidium arises. Development They develop in ''telium, telia'' (sing. ''telium'' or ''telio ...
s typically on seed and occasionally in soil. The fungus infects the shoots of wheat seedlings before the plants emerge from the soil. After
karyogamy Karyogamy is the final step in the process of fusing together two haploid eukaryotic cells, and refers specifically to the fusion of the two cell nucleus, nuclei. Before karyogamy, each haploid cell has one complete copy of the organism's genome. ...
, the teliospores germinate to form a
basidium A basidium (: basidia) is a microscopic spore-producing structure found on the hymenophore of reproductive bodies of basidiomycete fungi. The presence of basidia is one of the main characteristic features of the group. These bodies are also ...
, on which 8–16 haploid
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 pr ...
(primary
sporidia ''Sporidia'' are result of homokaryotic smut fungi (which are not pathogenic), asexual reproduction through the process of budding. Thus far, this has only been observed ''in vitro ''In vitro'' (meaning ''in glass'', or ''in the glass'') ...
) will develop. There are two mating types of basidiospores (+ and -) and they fuse to form H-shaped structures to establish a
dikaryon The dikaryon (karyogamy) is a cell nucleus feature that 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 c ...
. This dikaryon then will yield infectious
hyphae A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one o ...
which can either produce more hyphae or more secondary sporidia. The pathogen grows within the terminal
meristem In cell biology, the meristem is a structure composed of specialized tissue found in plants, consisting of stem cells, known as meristematic cells, which are undifferentiated cells capable of continuous cellular division. These meristematic c ...
via
mycelium Mycelium (: mycelia) is a root-like structure of a fungus consisting of a mass of branching, thread-like hyphae. Its normal form is that of branched, slender, entangled, anastomosing, hyaline threads. Fungal colonies composed of mycelium are fo ...
and completes its life cycle by transforming the mycelial cells into teliospores. The smutted wheat kernels that are full of teliospores break open and release upon harvest, which allows for the teliospores to overwinter on the seed and are blown away by currents onto the soil, thus completing the life cycle.


Morphology

Teliopsores are thick-walled, globiose, reticulate and 13–23 μm in diameter.


Hosts

''
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 ...
'' (wheatgrass), ''
Bromus ''Bromus'' is a large genus of grasses, classified in its own tribe Bromeae. They are commonly known as bromes, brome grasses, cheat grasses, or chess grasses. Estimates in the scientific literature of the number of species have ranged from 100 ...
'' (bromegrasses), '' Elymus'' (wildrye), ''
Festuca ''Festuca'' (fescue) is a genus of flowering plants belonging to the grass family Poaceae (subfamily Pooideae). They are evergreen or herbaceous perennial tufted grasses with a height range of and a cosmopolitan distribution, occurring on ...
'' (fescues), ''
Hordeum ''Hordeum'' is a genus of annual and perennial plants in the grass family. The species are native throughout the temperate regions of Africa, Eurasia, and the Americas. Taxonomy Species Species include: * '' Hordeum aegiceras'' – Mongo ...
'' (barleys), ''
Lolium ''Lolium'' is a genus of tufted grasses in the bluegrass subfamily (Pooideae). It is often called ryegrass, but this term is sometimes used to refer to grasses in other genera. They are characterized by bunch-like growth habits. ''Lolium'' ...
'' (ryegrasses), ''
Poa ''Poa'' is a genus of about 570 species of Poaceae, grasses, native to the temperate regions of both hemispheres. Common names include meadow-grass (mainly in Europe and Asia), bluegrass (mainly in North America), tussock (some New Zealand spe ...
'' (meadow grass), ''
Secale cereale Rye (''Secale cereale'') is a grass grown extensively as a grain, a cover crop and a forage crop. It is grown principally in an area from Eastern and Northern Europe into Russia. It is much more tolerant of cold weather and poor soil than o ...
'' (rye),
Triticale Triticale (; × ''Triticosecale'') is a hybrid of wheat (''Triticum'') and rye (''Secale'') first bred in laboratories during the late 19th century in Scotland and Germany. Commercially available triticale is almost always a second-generation ...
, ''
Triticum Wheat is a group of wild and domesticated grasses of the genus ''Triticum'' (). They are cultivated for their cereal grains, which are staple foods around the world. Well-known wheat species and hybrids include the most widely grown comm ...
'' spp. (wheats) – including '' T. aestivum'' (common wheat), '' T. dicoccum'' (hulled wheat), '' T. turgidum'' (durum wheat) – and other
Poaceae Poaceae ( ), also called Gramineae ( ), is a large and nearly ubiquitous family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos, the grasses of natural grassland and species cultivate ...
(other grasses).


Signs and symptoms

It is difficult to detect ''Tilletia caries'' early in its infection period; by the time symptoms are able to be detected, the pathogen is systemic throughout the plant. Plants that are infected will be stunted anywhere from a few centimeters/inches below average to half the average height of a healthy plant. Additionally, the heads are slender and remain green longer than healthy heads. A symptom that is indicative of ''T. caries'' is the replacement of yellow heads with grey bunt balls in the head of infected plants. The infected bunt balls are about the same shape and size as normal kernels. When the mature kernels are broken, they are full of a black, powdery mass of the fungal spores. These fungal spores give off a distinctive fishy smell and are oily to the touch.


History and significance

From the late 1800s until the 1930s, stinking smut was a devastating disease of wheat. For example, infection levels over 20% were common in
Washington State Washington, officially the State of Washington, is a state in the Pacific Northwest region of the United States. It is often referred to as Washington State to distinguish it from the national capital, both named after George Washington ...
in the early 1900s. One of the most extreme cases was in Kansas in 1890 where the yield was reduced 20–50% because of ''Tilletia caries''. Only when seed treatments became available after 1930 did losses from smut drop to much lower levels. Today, losses from smut rarely occur unless a grower chooses not to plant treated seed, but, if left untreated, bunt can reduce yield by more than 50%. In modern agriculture, if an infection occurs, losses are 5–10%. If there is a significant ''T. caries'' infection, the dusty and oily spore masses released during harvest can lead to combine explosions.
Static electricity Static electricity is an imbalance of electric charges within or on the surface of a material. The charge remains until it can move away by an electric current or electrical discharge. The word "static" is used to differentiate it from electric ...
that develops around the combine machinery ignites the teliospore dust released from the combine.


Use as a biological weapon

It was used as a
biological weapon Biological agents, also known as biological weapons or bioweapons, are pathogens used as weapons. In addition to these living or replicating pathogens, toxins and Toxin#Biotoxins, biotoxins are also included among the bio-agents. More than 1,2 ...
by
Iraq Iraq, officially the Republic of Iraq, is a country in West Asia. It is bordered by Saudi Arabia to Iraq–Saudi Arabia border, the south, Turkey to Iraq–Turkey border, the north, Iran to Iran–Iraq border, the east, the Persian Gulf and ...
against
Iran Iran, officially the Islamic Republic of Iran (IRI) and also known as Persia, is a country in West Asia. It borders Iraq to the west, Turkey, Azerbaijan, and Armenia to the northwest, the Caspian Sea to the north, Turkmenistan to the nort ...
during the
Iran–Iraq War The Iran–Iraq War, also known as the First Gulf War, was an armed conflict between Iran and Iraq that lasted from September 1980 to August 1988. Active hostilities began with the Iraqi invasion of Iran and lasted for nearly eight years, unti ...
in the 1980s.


Management

The most effective and widely used management strategy for common bunt is to treat seed with
fungicide Fungicides are pesticides used to kill parasitic fungi or their spores. Fungi can cause serious damage in agriculture, resulting in losses of yield and quality. Fungicides are used both in agriculture and to fight fungal infections in animals, ...
before planting. According to the
University of Nebraska-Lincoln A university () is an institution of tertiary education and research which awards academic degrees in several academic disciplines. ''University'' is derived from the Latin phrase , which roughly means "community of teachers and scholars". Uni ...
Institute of Agriculture and Natural Resources, it is recommended to buy certified, fungicide-treated seed or have it cleaned and treated by a commercial seed conditioner. There are ways that farmers can manipulate the severity of the infection to a certain extent. For example, they can plant the seed when the soil temperature is higher than what is ideal for teliospore germination, e.g., above . For winter wheat this means planting in early fall and for spring wheat planting in late spring. This tactic can reduce the amount of smut that occurs, but it typically does not eliminate the disease. Typically, smut poses more of a problem in winter wheat than in spring wheat because in autumn, when winter wheat is planted, there is a longer period of more favorable temperatures for teliospore germination than compared to the planting season for spring wheat. There are no current wheat cultivars on the market with good resistance to common bunt. However, there have been research efforts that utilize DNA markers for resistant cultivars in the attempt to understand the specific genes that code for resistance against common bunt. This may be applied for future breeding of commercially available resistant wheat crop.


See also

*
Common bunt Common bunt, also known as hill bunt, Indian bunt, European bunt, stinking smut or covered smut, is a disease of both spring and winter wheats. It is caused by two very closely related fungi, '' Tilletia tritici'' (syn. '' Tilletia caries'') and ' ...


Notes


References

* * * * * * {{DEFAULTSORT:Tilletia Caries Ustilaginomycotina Fungal plant pathogens and diseases Wheat diseases Fungi described in 1775 Fungus species