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''Fusarium circinatum'' 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 that causes the serious disease pitch canker on
pine A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanical Garden accep ...
trees and Douglas firs (''
Pseudotsuga menziesii The Douglas fir (''Pseudotsuga menziesii'') is an evergreen conifer species in the pine family, Pinaceae. It is native to western North America and is also known as Douglas-fir, Douglas spruce, Oregon pine, and Columbian pine. There are three va ...
''). The most common hosts of the pathogen include slash pine ('' Pinus elliottii''), loblolly pine (''
Pinus taeda ''Pinus taeda'', commonly known as loblolly pine, is one of several pines native to the Southeastern United States, from East Texas to Florida, and north to southern New Jersey. The wood industry classifies the species as a southern yellow pine. ...
''), Monterey pine (''
Pinus radiata ''Pinus radiata'' ( syn. ''Pinus insignis''), the Monterey pine, insignis pine or radiata pine, is a species of pine native to the Central Coast of California and Mexico (Guadalupe Island and Cedros island). It is an evergreen conifer in the fa ...
''), Mexican weeping pine (''
Pinus patula ''Pinus patula'', commonly known as patula pine, spreading-leaved pine, or Mexican weeping pine, and in Spanish as ''pino patula'' or ''pino llorón'', (''patula'' Latin = “spreading”) is a tree native to the highlands of Mexico. It grows f ...
''), and Douglas fir. Like other ''
Fusarium ''Fusarium'' is a large genus of filamentous fungi, part of a group often referred to as hyphomycetes, widely distributed in soil and associated with plants. Most species are harmless saprobes, and are relatively abundant members of the soil mi ...
'' species in the phylum
Ascomycota Ascomycota is a phylum of the kingdom Fungi that, together with the Basidiomycota, forms the subkingdom Dikarya. Its members are commonly known as the sac fungi or ascomycetes. It is the largest phylum of Fungi, with over 64,000 species. The def ...
, it is the asexual reproductive state of the fungus and has a
teleomorph In mycology, the terms teleomorph, anamorph, and holomorph apply to portions of the life cycles of fungi in the phyla Ascomycota and Basidiomycota: *Teleomorph: the sexual reproductive stage (morph), typically a fruiting body. *Anamorph: an ase ...
, ''Gibberella circinata''.


Distribution

This fungus is believed to have originated in Mexico. It spread to the eastern United States in 1946 and by 1986 had reached the western United States. It was first recorded in Japan in the 1980s, in South Africa in 1990, in Chile First report of the pitch canker fungus, Fusarium circinatum, on pines in Chile
/ref> and Spain in the mid 1990s and in Italy in 2007.Biosecurity New Zealand
/ref>


Host species

In California this canker has been recorded on nine different species of pine (''
Pinus A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanical Garden accep ...
'') and on Douglas fir (''
Pseudotsuga menziesii The Douglas fir (''Pseudotsuga menziesii'') is an evergreen conifer species in the pine family, Pinaceae. It is native to western North America and is also known as Douglas-fir, Douglas spruce, Oregon pine, and Columbian pine. There are three va ...
''). In Europe and Asia it has been recorded on over 30 other ''Pinus'' species. Monterey pine (''
Pinus radiata ''Pinus radiata'' ( syn. ''Pinus insignis''), the Monterey pine, insignis pine or radiata pine, is a species of pine native to the Central Coast of California and Mexico (Guadalupe Island and Cedros island). It is an evergreen conifer in the fa ...
'') seems to be the most susceptible species, and in California 85% of the native Monterey pine forests were initially thought to be threatened by the disease. Because of the activation of
systemic acquired resistance Systemic acquired resistance (SAR) is a "whole-plant" resistance response that occurs following an earlier localized exposure to a pathogen. SAR is analogous to the innate immune system found in animals, and although there are many shared aspects b ...
in native Monterey pine trees, however, the impacts of the disease in California have been mitigated.


Biology

''Fusarium circinatum'' infects the twigs and branches of pine trees, causing a bark canker. Most infection is by
macroconidia A conidium ( ; ), sometimes termed an asexual chlamydospore or chlamydoconidium (), is an Asexual reproduction, asexual, non-motility, motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word for dust, ('). They are al ...
or
microconidia A conidium ( ; ), sometimes termed an asexual chlamydospore or chlamydoconidium (), is an asexual, non-motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word for dust, ('). They are also called mitospores due to th ...
. The macroconidia are 3-septate, with slightly curved walls and the microconidia are single-celled, ovoid, and borne in false heads on aerial polyphialides. The aerial
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 ...
is white or pale violet colour and slightly twisted below the proliferation of microconidiophores. In culture,
perithecia An ascocarp, or ascoma (), is the fruiting body ( sporocarp) of an ascomycete phylum fungus. It consists of very tightly interwoven hyphae and millions of embedded asci, each of which typically contains four to eight ascospores. Ascocarps are mo ...
are readily produced. They are dark purple or black and ovoid. Cylindrical asci are released by oozing. There are eight ascospores which are 1-septate and ellipsoidal to fusiform. Because peritheca have not been observed in the field, it is not thought that ascospores are an important route for infection. The infection is usually carried from tree to tree by the rain, the wind or by bark-feeding insects. These including
weevil Weevils are beetles belonging to the Taxonomic rank, superfamily Curculionoidea, known for their elongated snouts. They are usually small, less than in length, and Herbivore, herbivorous. Approximately 97,000 species of weevils are known. They b ...
s in the genus ''Pityophthorus'' and
bark beetle A bark beetle is the common name for the subfamily of beetles Scolytinae. Previously, this was considered a distinct family (Scolytidae), but is now understood to be a specialized clade of the "true weevil" family (Curculionidae). Although the ...
s in the genera ''Ips'' and ''Conophthorus''. These insects commonly infect pine trees and the adults may disperse the pathogen. Additionally, these insects cause a wound when feeding and this may facilitate entry of the infection. Warmth and moisture encourage the development of the disease whereas cooler drier conditions restrict it. In California it is more severe in coastal areas. Research was undertaken to see whether spores from the teleomorph, ''Gibberella circinata'', might be responsible for spread of the fungus. It was found that very few vegetative compatibility groups existed among the California strains of the pathogen. This implied that
asexual reproduction Asexual reproduction is a type of reproduction that does not involve the fusion of gametes or change in the number of chromosomes. The offspring that arise by asexual reproduction from either unicellular or multicellular organisms inherit the fu ...
predominated and laboratory tests confirmed this.


Hyperparasites

Unusually for
mycovirus Mycoviruses (Ancient Greek: μύκης ' ("fungus") + Latin '), also known as mycophages, are viruses that infect fungi. The majority of mycoviruses have double-stranded RNA (dsRNA) genomes and isometric particles, but approximately 30% have po ...
es, Muñoz-Adalia ''et al.'', 2016 finds that both ''Fusarium circinatum'' mycovirus 1 ( FcMV1) and ''Fusarium circinatum'' mycovirus 2 ( FcMV2) ''increase'' the virulence of ''F. circinatum''. This has been observed to produce increased disease severity on ''P. radiata''.


Symptoms

The various symptoms of ''F. circinatum'' can help identify and distinguish it from other pathogens or common ''Pinus'' diseases. The symptoms are similar to other damping off diseases with seedlings wilting and dying and exuding
resin In polymer chemistry and materials science, resin is a solid or highly viscous substance of plant or synthetic origin that is typically convertible into polymers. Resins are usually mixtures of organic compounds. This article focuses on natu ...
from the root collar areas. Drooping from the resin production and the plant’s resistance mechanism can be observed along with die-back near the apical meristem. A discoloration of the stem and needles is usually present, with plants exhibiting purple or blue shades. Other symptoms include
chlorosis In botany, chlorosis is a condition in which leaves produce insufficient chlorophyll. As chlorophyll is responsible for the green color of leaves, chlorotic leaves are pale, yellow, or yellow-white. The affected plant has little or no ability to ...
of the needles turning a reddish brown color and lesions on the stems, root collars, and tap roots. Host factors that can trigger infection include plant stress with excessive nitrogen in the soil, unbalanced watering cycles, warmer temperatures, and wounds from pruning or insect damage.


Spread

Numerous plant pathologists have noted ''F. circinatum'' as a serious threat to the pine tree species. Due to the high tree mortality rate, reduced growth, and degradation of wood quality, the economic and ecological importance are greatly affected by this disease. Not only can the spread of infection go from branch to branch, but also infect pine seeds, leading to damping-off of younger seedlings and resulting in death from the fungal infection. The environmental interactions that take place to favor the spread and development of this disease play a large role in transmission. Factors such as soil nutrient ratio,
abiotic In biology and ecology, abiotic components or abiotic factors are non-living chemical and physical parts of the environment that affect living organisms and the functioning of ecosystems. Abiotic factors and the phenomena associated with them under ...
stressors, air pollution, temperature, and humidity can all contribute to the spread of this disease. In Chile, the infection was first reported on ''Pinus radiata'' in nurseries and was thought to be due to the import of contaminated seed. Seedlings could also be infected by soil-borne contamination. A few years later, the disease had not spread to mature stands of trees. The same is true in South Africa, where it was reported to infect nursery stock but not forest trees. ''Fusarium circinatum'' is spread locally by wind and insects, but it is slow to spread into new areas. Over large distances it can be transported in contaminated pine seeds or by young plants. Although it could be carried as infected timber, this is considered unlikely especially if the bark has been removed. If timber had been a significant means of infection, the fungus would have spread more rapidly to other parts of the world as there is a considerable trade in pine. The vectors for this disease, such as insects, rely on whether or not the species is indigenous to where the pine is located. There are also different ranges of susceptibility that can interact within the environment. Bishop pine ''(
Pinus muricata ''Pinus muricata'', the bishop pine, is a pine with a very restricted range: mostly in California, including several offshore Channel Islands, and a few locations in Baja California, Mexico. It is always on or near the coast. In San Luis Obispo ...
'') has a more extensive range of susceptibility compared to Monterey pine, which serves as a “host bridge” to more northern locations for susceptible ''Pinus'' species.


Management

Several strategies are currently being used to help decrease the spread of ''F. circinatum''. Irrigation water can be chemically treated with a
chlorination Chlorination may refer to: * Chlorination reaction In chemistry, halogenation is a chemical reaction that entails the introduction of one or more halogens into a compound. Halide-containing compounds are pervasive, making this type of transform ...
system or ozone treatment. Following corrected pH levels in water, it is recommended that 2-3 ppm of chlorine be administered. Proper hygiene practices and surface sterilization are to be maintained throughout management, especially by preventing the accumulation of water nearby to deter increases in insect populations. The use of
insecticide Insecticides are substances used to kill insects. They include ovicides and larvicides used against insect eggs and larvae, respectively. Insecticides are used in agriculture, medicine, industry and by consumers. Insecticides are claimed to b ...
s can reduce the number of vectors present that typically allow the infection to occur. Within pine nurseries, it is common for the dying seedlings to be removed and saturated with a
fungicide Fungicides are biocidal chemical compounds or biological organisms used to kill parasitic fungi or their spores. A fungistatic inhibits their growth. Fungi can cause serious damage in agriculture, resulting in critical losses of yield, quality, ...
. A single, systematic fungicide should not be used continuously as resistance can develop. While pitch canker can cause damage to the pine industry, some cases of infection recover. Susceptible Monterey pine trees that were repeatedly exposed to the pathogen gained resistance over time under controlled conditions, especially where the disease has been present for over 10 years. Removing diseased trees from areas as a conservative approach allows for the possibility of recovery.


Testing

A test has been developed for detecting contamination of seeds of ''Pinus'' species by ''F. circinatum''. It is based on the biological enrichment of a sample followed by a real-time polymerase chain reaction assay. Many countries are imposing quarantine controls on movement of propagating materials and this test can help prevent the introduction of the pathogen through contaminated seed.


See also

*
Forest pathology Forest pathology is the research of both Biotic stress, biotic and Abiotic stress, abiotic maladies affecting the health of a forest ecosystem, primarily Fungal infection in plants, fungal pathogens and their insect vectors. It is a subfield of ...


Notes

The epithet refers to the diagnostic coiled hyphae that are produced by this species.Mycologia 90(1–6): 442 (1998)
/ref>


References

{{Taxonbar, from=Q5509888 circinatum Fungal conifer pathogens and diseases