Heterobasidion Annosum
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''Heterobasidion annosum'' 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 ...
fungus in the family
Bondarzewiaceae The Bondarzewiaceae are a family of fungi in the order Russulales. The type species for both its genus and the family as a whole, ''Bondarzewia montana'', closely resembles members of Polyporales (and was formerly placed there), but has ornament ...
. It is considered to be the most economically important forest pathogen in the Northern Hemisphere. ''Heterobasidion annosum'' is widespread in forests in the United States and is responsible for the loss of one billion U.S. dollars annually. This fungus has been known by many different names. First described by
Fries French fries (North American English), chips (British English), finger chips (Indian English), french-fried potatoes, or simply fries, are '' batonnet'' or ''allumette''-cut deep-fried potatoes of disputed origin from Belgium and France. The ...
in 1821, it was known by the name ''Polyporus annosum''. Later, it was found to be linked to conifer disease by
Robert Hartig Robert Hartig (born: Heinrich Julius Adolph Robert Hartig, 30 May 1839, in Braunschweig – died 9 October 1901, in Munich) was a German forestry scientist and mycologist. Biography He was educated at the Collegium Carolinum of Braunschweig, ...
in 1874, and was renamed ''Fomes annosus'' by H. Karsten. Its current name of ''Heterobasidion annosum'' was given by Brefeld in 1888. ''Heterobasidion annosum'' causes one of the most destructive diseases of conifers.Adomas A, Asiebu FO, Stenlid J. (2005). Conifer root and butt rot caused by ''Heterobasidion annosum'' (Fr.) Bref. s.l. ''Molecular Plant Pathology'' 6(4): 395–409 The disease caused by the fungus is named annosus root rot.


Description

The fruiting bodies of the fungus, which are also known as
basidiocarp In fungi, a basidiocarp, basidiome, or basidioma () is the sporocarp of a basidiomycete, the multicellular structure on which the spore-producing hymenium is borne. Basidiocarps are characteristic of the hymenomycetes; rusts and smuts do not ...
s, are normally
brackets A bracket is either of two tall fore- or back-facing punctuation marks commonly used to isolate a segment of text or data from its surroundings. Typically deployed in symmetric pairs, an individual bracket may be identified as a 'left' or 'r ...
which are whitish around the margins and dark brown on the uneven, knobbly upper surface. They can blacken in age. They can also take a resupinate form, consisting only of a white crust which corresponds to the underside of the bracket. Basidiocarps are up to about 40 cm in diameter and 3.5 cm thick. The fertile surface of the fruiting body is white, easily bruising brown, and has barely visible pores, with 3-4 per mm. The flesh, which has a strong fungus smell, is elastic when young but becomes woody when older. Sexual spores called basidiospores are created in the fertile layer on the lower surface of the basidiocarps, whilst conidiospores occur in the asexual stage and are produced on microscopic "conidiophores" which erupt through the surface of the host tree. Conidiospores and basidiospores are both produced by this fungus, the latter being more important for infecting the conifers. The species is inedible.


Ecology and life cycle

In the summertime,
basidiospore 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 ...
s, the primary infective propagules, are released. These basidiospores are carried long distances by wind currents. They infect trees (usually conifers) through damage such as freshly cut stumps. Once on the stump the fungus colonizes and moves into the root via
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 substrate (biology), substrates. A typical single ...
. ''Heterobasidion annosum'' moves short distances from the roots of an infected stump through root grafts with other trees. It can also spread through insects that feed on roots. Since this fungus can not move very far through soil, it relies on tree roots to help it infect neighbouring trees. In these roots, it can grow 0.1–2.0 m per year. This results in a spread of the fungus and disease gaps in the forest. These disease gaps are produced when the trees dies and falls, creating gaps in the forest canopy. These gaps affect the moisture and sunlight available, altering the habitats for plants and animals on the forest floor. ''Spiniger meineckellus'', the name for the asexual stage of this fungus, is produced on stumps when the conditions are moist, and the conidiospores that are produced will be able to live in the soil for up to ten months. The role of conidiospores is unknown in the infection process and is not thought to be important.


Symptoms and signs of disease

Symptoms and signs of fungus disease are often found underground. The ''H. annosum'' infections cause an abnormal change in structure in the roots that climbs up to the butt of the tree. More than half the tree may be killed before any symptoms appear to the human eye. Basidiocarps can take up to one and a half or even three years to be visible. This infections causes the trees to have abnormal needle growth, pale yellow barks, and the trees to wither and die. This root disease typically causes the tree to have a thin crown from bottom up and inside out. Trees will eventually die. A landscape scale symptom is the rings of dead trees in various stages of decay and death, with the oldest at the center and progressively younger moving outward. The
white rot A wood-decay or xylophagous fungus is any species of fungus that digests moist wood, causing it to rot. Some species of wood-decay fungi attack dead wood, such as brown rot, and some, such as ''Armillaria'' (honey fungus), are parasitic and col ...
fungus found in the roots is the sign of telling whether the tree has been affected by ''H. annosum''. The bark changes colors as the stages progress, they go from pale yellow, to a crusty light brown, and finally in its advanced stage it turns white with the signature of ''Fomes annosus''―a sprinkled streak of black spots. Another sign is the leaking part of the root that causes a compact mass to form between it and the sand.


Isolation

There are several ways to isolate ''H. annosum''. Water agar could be used with infected host tissue to produce conidiophores which a simple or branched part hypha of a fungus to eliminate ''H. annosum''. Another way of isolating ''H. annosum'' is by using the thin disks of living sapwood from ''
Picea abies ''Picea abies'', the Norway spruce or European spruce, is a species of spruce native to Northern Europe, Northern, Central Europe, Central and Eastern Europe. It has branchlets that typically hang downwards, and the largest cones of any spruce, ...
''. By cutting the thin disks into petri dishes which is used to culture bacteria and placing them on moist filter paper, this technique allows spores to be captured from the air, and result in the asexual stage of the fungus forming on the disks.


Range

The species is
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...
to the
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...
. It was introduced into
Italy Italy ( it, Italia ), officially the Italian Republic, ) or the Republic of Italy, is a country in Southern Europe. It is located in the middle of the Mediterranean Sea, and its territory largely coincides with the homonymous geographical re ...
during the
Second World War World War II or the Second World War, often abbreviated as WWII or WW2, was a world war that lasted from 1939 to 1945. It involved the vast majority of the world's countries—including all of the great powers—forming two opposin ...
, vectored in wooden
crate A crate is a large shipping container, often made of wood, typically used to transport or store large, heavy items. Steel and aluminium crates are also used. Specialized crates were designed for specific products, and were often made to be reusa ...
material.


Management

Theoretically, a root can be suppressed during all stages of its life cycle. There are three ways of managing ''Heterobasidion annosum'': silvicultural control measures, chemical methods and biological control. Silvicultural control involves planting species with low susceptibility. This could lower the root rot problem and free an infected site from inoculum. A more extreme measure is to remove the inoculum from the infected site. Proper planning and mixture schemes produce a better yield than a pure plantation (i.e. containing only one species). An alternative species could always be used for protection against ''H. annosum''. Chemical methods include prophylactic stump treatment with a solution of urea immediately after the infection. This protects the stump by hydrolysis of the compound by the enzyme
urease Ureases (), functionally, belong to the superfamily of amidohydrolases and phosphotriesterases. Ureases are found in numerous bacteria, fungi, algae, plants, and some invertebrates, as well as in soils, as a soil enzyme. They are nickel-containin ...
in the living wood tissue, which results in formation of
ammonia Ammonia is an inorganic compound of nitrogen and hydrogen with the formula . A stable binary hydride, and the simplest pnictogen hydride, ammonia is a colourless gas with a distinct pungent smell. Biologically, it is a common nitrogenous was ...
and a rise in pH to a level that ''H. annosum'' at which mycelia are unable to survive. Biological control is another alternative. Currently, a number of fungal species such as '' Phlebiopsis gigantea'', ''
Bjerkandera adusta ''Bjerkandera adusta'', commonly known as the smoky polypore or smoky bracket, is a species of fungus in the family Meruliaceae. It is a plant pathogen that causes white rot in live trees, but most commonly appears on dead wood. It was first d ...
'' and ''
Fomitopsis pinicola ''Fomitopsis pinicola'', is a stem decay fungus common on softwood and hardwood trees. Its conk (fruit body) is known as the red-belted conk. The species is common throughout temperate Europe and Asia. It is a decay fungus that serves as a smal ...
'' have been tested on stumps as competitors and antagonists against ''H. annosum''. However, among these, only ''Phlebiopsis gigantea'' shows good results of eliminating ''H. annosum''.


References


External links


A plant pathogen profile
{{DEFAULTSORT:Heterobasidion Annosum Fungi described in 1841 Fungal tree pathogens and diseases Russulales Fungi of Europe Fungi of North America Inedible fungi Taxa named by Elias Magnus Fries