''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, Basid ...
fungus in the family
Bondarzewiaceae. It is considered to be the most economically important forest
pathogen
In biology, a pathogen (, "suffering", "passion" and , "producer of"), in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a Germ theory of d ...
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, or simply fries, also known as chips, and finger chips (Indian English), are ''List of culinary knife cuts#Batonnet, batonnet'' or ''Julienning, julienne''-cut deep frying, deep-fried potatoes of disputed origin. They are prepa ...
in 1821, it was known by the name ''Polyporus annosum''. Later, it was found to be linked to conifer disease by
Robert Hartig 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
The sporocarp (also known as fruiting body, fruit body or fruitbody) of fungi is a multicellular structure on which spore-producing structures, such as basidia or asci, are borne. The fruitbody is part of the sexual phase of a fungal life cyc ...
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 no ...
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. They come in four main pairs of shapes, as given in the box to the right, which also gives their n ...
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
Ploidy () is the number of complete sets of chromos ...
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 (: 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 ...
. ''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 die and fall, 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 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
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" and " tengusa". As found in nature, agar is a mixture of two components, t ...
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 only 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 foun ...
to the
United States
The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
. It was introduced into
Italy
Italy, officially the Italian Republic, is a country in Southern Europe, Southern and Western Europe, Western Europe. It consists of Italian Peninsula, a peninsula that extends into the Mediterranean Sea, with the Alps on its northern land b ...
during the
Second World War
World War II or the Second World War (1 September 1939 – 2 September 1945) was a World war, global conflict between two coalitions: the Allies of World War II, Allies and the Axis powers. World War II by country, Nearly all of the wo ...
, 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 reus ...
material.
Management
Theoretically, a root can be suppressed during all stages of its life cycle. There are three ways of managing ''Heterobasidion annosum'':
silvicultural
Silviculture is the practice of controlling the growth, composition/structure, as well as quality of forests to meet values and needs, specifically timber production.
The name comes from the Latin ('forest') and ('growing'). The study of forests ...
control measures, chemical methods and
biological control
Biological control or biocontrol is a method of controlling pests, whether pest animals such as insects and mites, weeds, or pathogens affecting animals or plants by using other organisms. It relies on predation, parasitism, herbivory, or o ...
. 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, Archaea, fungi, algae, plants, and some invertebrates. Ureases are nickel-containing metalloenzymes of high ...
in the living wood tissue, which results in formation of
ammonia
Ammonia is an inorganic chemical compound of nitrogen and hydrogen with the chemical formula, formula . A Binary compounds of hydrogen, stable binary hydride and the simplest pnictogen hydride, ammonia is a colourless gas with a distinctive pu ...
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'' and ''
Fomitopsis pinicola'' 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
Fungus species