Armillaria root rot is a fungal
root rot
Root rot is a condition in which anoxic conditions in the soil or potting media around the roots of a plant cause them to rot. This occurs due to excessive standing water around the roots. It is found in both indoor and outdoor plants, althoug ...
caused by several different members of the genus
''Armillaria''. The symptoms are variable depending on the host infected, ranging from stunted leaves to
chlorotic needles and
dieback of twigs and branches. However, all infected hosts display symptoms characteristic of being infected by a white rotting
fungus
A fungus (plural, : fungi or funguses) is any member of the group of Eukaryote, eukaryotic organisms that includes microorganisms such as yeasts and Mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified ...
. The most effective ways of management focus on limiting the spread of the fungus, planting resistant species, and removing infected material. This disease poses a threat to the
lumber industry as well as affecting recreational areas.
Hosts and symptoms
Because this disease is caused by multiple species within the genus ''Armillaria'', it has an extremely broad host range. Hundreds of
tree
In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondary growth, plants that are ...
s and
shrubs are susceptible to root rot to varying degrees. In fact, the only two genera of tree known to be resistant to Armillaria root rot are
larch
Larches are deciduous conifers in the genus ''Larix'', of the family Pinaceae (subfamily Laricoideae). Growing from tall, they are native to much of the cooler temperate northern hemisphere, on lowlands in the north and high on mountains furt ...
and
birch
A birch is a thin-leaved deciduous hardwood tree of the genus ''Betula'' (), in the family Betulaceae, which also includes alders, hazels, and hornbeams. It is closely related to the beech- oak family Fagaceae. The genus ''Betula'' contains 3 ...
. Further investigation is being conducted for additional species, but at this time there is no further evidence leading to the belief that more exist.
While ''Armillaria'' is a significant and damaging pathogen of tree hosts, it also has many agronomic hosts such as grapevines, berries, roses, stone fruits, rhododendron, and rosaceous plants, although the fungus is primarily native to areas where it can use forest trees as a host. On hosts such as these, infection causes death of the cambium and further decay of the xylem. Vines or stems from these plants, once infected, may remain a potential source for inoculum for up to ten years and can infect neighboring plants, often via rhizomorphs.
As a result of the multitude of possible hosts, symptoms also range a great deal from one infection to another. All infected hosts do however display symptoms common to
white rotting fungi. These include light or bleached wood as a result of the degradation of essential cell wall compounds such as
lignin
Lignin is a class of complex organic polymers that form key structural materials in the support tissues of most plants. Lignins are particularly important in the formation of cell walls, especially in wood and bark, because they lend rigidity ...
and
hemicellulose
A hemicellulose (also known as polyose) is one of a number of heteropolymers (matrix polysaccharides), such as arabinoxylans, present along with cellulose in almost all terrestrial plant cell walls.Scheller HV, Ulvskov Hemicelluloses.// Annu Rev ...
. In addition to these, the host will show above-ground symptoms due to fungal infection of the vasculature. These include chlorosis in the needles as well as dieback of twigs and branches. The extent of these symptoms varies with the degree of infection as well as the susceptibility of the host. In addition to these symptoms, the trunks of
conifer
Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All ex ...
s will also exude excess resin in a process known as
resinopsis which results in a layer of resin, debris, and fungal tissue forming around infected roots.
Deciduous trees occasionally develop sunken cankers but most often fail to exhibit these symptoms on the trunk, and will instead simply display the other symptoms.
In addition to these symptoms, signs of the infectious organism are very evident in the host. Upon removal of the bark, white
mycelial mats are visible along with
rhizomorphs, a distinctive
reproductive structure
The reproductive system of an organism, also known as the genital system, is the biological system made up of all the anatomical organs involved in sexual reproduction. Many non-living substances such as fluids, hormones, and pheromones are ...
. Rhizomorphs are black, shoe-string like growth structure that can grow out from the host and grow in the soil to infect nearby hosts. Clusters of
mushroom
A mushroom or toadstool is the fleshy, spore-bearing fruiting body of a fungus, typically produced above ground, on soil, or on its food source. ''Toadstool'' generally denotes one poisonous to humans.
The standard for the name "mushroom" is ...
s will also form at the base of the infected tree, indicating an infection. However, fruiting is not consistent year-to-year and the mushrooms frequently resemble other species to the untrained eye. Therefore, the presence of mushrooms should not constitute the only diagnostic indicator when identifying the disease. As the name suggests, the caps of these mushrooms are a honey, or light brown color, and the gills of the mushroom and spore print are white. The stem is typically a mottled white and brown, with a ring on the stem, too. Honey mushrooms are edible, but because of their tendency to look so similar to other species, only the very experienced should collect them. Also, zone lines of
melanized
Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amino ...
fungal cells may be visible within infected wood.
Disease cycle
Diagramatic illustration of white root rot disease cycle.
Black root-like rhizomorphs on an unidentified European ''Armillaria'' species
For the most part, this fungus exhibits a life cycle characteristic of
basidiomycetes. It reproduces sexually with the mating of
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 or ...
and produces a
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 n ...
at the base of the infected host. This basidiocarp produces basidiospores that will further infect new hosts. However, ''Armillaria'' rarely spreads using this method in nature, possibly as a result of the spores being ineffective. In contrast, it will spread either through rhizomorphs or direct mycelial contact.
In the case of mycelial contact, the roots of an infected host grow near enough to a new host that
mycelia
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 in ...
simply grow onto the new host and infect.
Rhizomorph
Mycelial cords are linear aggregations of parallel-oriented hyphae. The mature cords are composed of wide, empty vessel hyphae surrounded by narrower sheathing hyphae. Cords may look similar to plant roots, and also frequently have similar functi ...
s are string-like masses of hyphae utilized if no new hosts are nearby and spread by probing through the soil towards uninfected roots. ''Armillaria'' can last up to 50 years in stumps of dead trees, infecting new hosts that come up nearby in that time period. In areas of infection, called a root-disease center, can develop.
Environment
''Armillaria'' infects trees in temperate and tropical regions. ''Armillaria'' commonly infects stressed trees that have been weakened by
insect
Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs ...
s, other
pathogen
In biology, a pathogen ( el, πάθος, "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 ger ...
s and/or climate stresses. It can also kill healthy trees especially in dry areas, like coniferous forests in the western
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 U.S. state, states, a Washington, D.C., federal district, five ma ...
. This fungus is found worldwide, but prefers cool soils and climates.
Managements
Plant varieties that are resistant to ''Armillaria'' or species are resistant to other environmental or biological
stressor
A stressor is a chemical or biological agent, environmental condition, external stimulus or an event seen as causing stress to an organism. Psychologically speaking, a stressor can be events or environments that individuals might consider dema ...
s. If the infected area has been cleared of trees, plants that are not vulnerable to the disease should be planted for five or so years until ''Armillaria'' is eradicated.
Stump removal is also an effective management tool but can be expensive. Another way to reduce susceptibility is to maintain
plant health by regular fertilization (if needed), watering during droughts, and trying not to create wounds on the plant.
Fumigation
Fumigation is a method of pest control or the removal of harmful micro-organisms by completely filling an area with gaseous pesticides—or fumigants—to suffocate or poison the pests within. It is used to control pests in buildings (s ...
can also be used to reduce the amount of
inoculum
In biology, inoculum refers to the source material used for inoculation. ''Inoculum'' may refer to:
* In medicine, material that is the source of the inoculation in a vaccine
* In microbiology, propagules: cells, tissue, or viruses that are used to ...
.
Cultural practices can also be effective for preventing the spread of ''Armillaria''. High temperatures interfere with the progression of this disease; if soil temperatures reach , then the growth of ''Armillaria'' in the soil will be limited. Using collar excavation to expose primary roots directly below the crown to the continuous cycle of solar heating and air may reduce the colonization from ''Armillaria'' in trees that are already infected.
This is called root collar excavation (RCE), and while it has been applied to citrus orchards and grapevines, this method has high labor costs and involvement due to the difficulty of carefully exposing these primary roots.
''Armillaria'' often occurs when a host is stressed due to other abiotic or biotic factors. It's often more cost effective to treat the stressor that caused the pathogen rather than treating ''Armillaria'' on its own, as these treatments and practices are not practical for everyday use.
Importance
This disease poses a serious threat to the timber industry in that it affects nearly every cultivated species of
hardwood
Hardwood is wood from dicot trees. These are usually found in broad-leaved temperate and tropical forests. In temperate and boreal latitudes they are mostly deciduous, but in tropics and subtropics mostly evergreen. Hardwood (which comes fro ...
and proves very difficult to remove once it has entered an area. The ability to spread using rhizomorphs as well as through mycelial contact allow the fungus to spread over very large areas and between many individuals. In fact, ''Armillaria'' is recognized as the largest living organism due to its clonal method of spreading. Infection is especially strong where environmental conditions are optimal and a large amount of possible hosts exist. This essentially is anywhere hardwoods are found growing. It has been found in every state in the United States of America as well as on several other continents including
Australia,
Europe, and
Asia
Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an ...
.
References
{{DEFAULTSORT:Armillaria Root Rot
Parasitic fungi
Fungal tree pathogens and diseases