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The fungal species, ''Myxotrichum chartarum'' was discovered by [[Gustav Kunze|Kunze]] in 1823.<ref name=Orr1963/>
The fungal species, ''Myxotrichum chartarum'' was discovered by [[Gustav Kunze|Kunze]] in 1823.<ref name=Orr1963/>


Originally, ''M. chartarum'' was placed in the genus, [[Oncidium]] by Nees (1823) which was previously in use for species of [[orchid]]. ''M. chartarum'' was thereafter reclassified by [[Gustav Kunze|Kunze]].<ref name=Orr1963/><ref name="Currah1985"/> In the same year, the genus, ''Myxotrichum'' was established by [[Gustav Kunze|Kunze]] to include two species, ''M. chartarum'' and ''Myxotrichum murorum''.<ref name=Orr1963/> Additional species were added thereafter.<ref name=Orr1963/> There has been much confusion within the ''Myxotrichum'' genus due to numerous revisions in classification over the years and scarcity of isolations.<ref name=Orr1963/><ref name="Currah1985"/> Eventually, the genus, ''Myxotrichum'' became associated with ascomycota fungi which are [[dematiaceous]] [[hyphomycetes]], commonly known as black yeasts or moulds.<ref name="Currah1985"/> At the microscopic level, these fungi have mesh-like peridium with hooked appendages.<ref name="Currah1985"/>
Originally, ''M. chartarum'' was mistaknly placed in the genus [[Oncidium]] by Nees (1823) which was already in use for species of [[orchid]]. ''M. chartarum'' was thereafter reclassified by [[Gustav Kunze|Kunze]].<ref name=Orr1963/><ref name="Currah1985"/> In the same year, the genus ''Myxotrichum'' was established by [[Gustav Kunze|Kunze]] to include two species, ''M. chartarum'' and ''Myxotrichum murorum''.<ref name=Orr1963/> Additional species were added thereafter.<ref name=Orr1963/> There has been much confusion within the ''Myxotrichum'' genus due to numerous revisions in classification over the years and scarcity of isolations.<ref name=Orr1963/><ref name="Currah1985"/> Eventually, the genus, ''Myxotrichum'' became associated with ascomycota fungi which are [[dematiaceous]] [[hyphomycetes]], commonly known as black yeasts or moulds.<ref name="Currah1985"/> At the microscopic level, these fungi have a mesh-like surrounding structures ([[peridium]]) with hooked appendages.<ref name="Currah1985"/>


In 1838, [[August Carl Joseph Corda|Corda]] classified ''M. chartarum'' as a [[Hyphomycete]] of the family Sporotrichacheae, as species belonging to this family had ornate appendages resembling deer antlers.<ref name="See1919"/>
In 1838, [[August Carl Joseph Corda|Corda]] classified ''M. chartarum'' as a [[Hyphomycete]] of the family Sporotrichacheae, as species belonging to this family had ornate appendages resembling deer antlers.<ref name="See1919"/>
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[[File:10244 sc 36d PH arthroconidia.jpg|thumb|right|Intercalary arthroconidia of ''Myxotrichum chartarum'' (UAMH 10244) from colony grown for 36d, photographed in Phase Contrast microscopy.]]
[[File:10244 sc 36d PH arthroconidia.jpg|thumb|right|Intercalary arthroconidia of ''Myxotrichum chartarum'' (UAMH 10244) from colony grown for 36d, photographed in Phase Contrast microscopy.]]
===Growth in its natural environment===
===Growth in its natural environment===
The ascospores can also be described as being yellow to orange in colour with a rounded football shape having longitudinal striations.<ref name=Orr1963/><ref name="Currah1985"/><ref name="Ellis1988" />
The ascospores can also be described as being yellow to orange in colour with a rounded football shape having longitudinal striations and having a diameter between 6-8 μm on the long side and 5-8 µm on the short side.<ref name=Orr1963/><ref name="Currah1985"/><ref name="Ellis1988" /><ref name="Tribe2002" />
[[ascus|Asci]] appear hyaline, globular, and contain the typical quantity of 8 ascospores each.<ref name=Orr1963/><ref name="Currah1985"/>
[[ascus|Asci]] appear hyaline, globular, and contain the typical quantity of 8 ascospores each, the size of which are 3.5-5.0 µm on the long side and 2.0-3.0 µm on the short side.<ref name=Orr1963/><ref name="Currah1985"/><ref name="Tribe2002" /> When the spores are mature, they are released altogether, producing a cloud of brown dust.<ref name="Tribe2002" />
[[Ascocarp]]s appear dark and spherical with short appendages, where the ascocarp can appear green or copper when filled with [[spores]].<ref name=Orr1963/><ref name="Currah1985"/> The spore mass fills the ascocarps between one-third and three-fourths of the total volume at peak maturity.<ref name=Orr1963/><ref name="Currah1985"/> The surrounding [[ascocarp#gymnothecium|Gymnothecium]] has septate appendages that are straight and elongated. Branching points present as [[uncinate]], or curved spines that are wider at the apices. <ref name=Orr1963/><ref name="Currah1985"/>
[[Ascocarp]]s appear dark and spherical with short appendages, where the ascocarp can appear green or copper when filled with [[spores]].<ref name=Orr1963/><ref name="Currah1985"/> The diameter of the ascocarp is larger than its close relative ''M. carminoparum'' at 150-555μm.<ref name=Orr1963/> The spore mass fills the ascocarps between one-third and three-fourths of the total volume at peak maturity.<ref name=Orr1963/><ref name="Currah1985"/> The surrounding [[ascocarp#gymnothecium|Gymnothecium]] has septate appendages that are straight and elongated. Branching points present as [[uncinate]], or curved spines that are wider at the apices. <ref name=Orr1963/><ref name="Currah1985"/>
Commonly found in ''Myxotrichum'' species are the secondary and tertiary branching of [[peridium|peridial]] [[hyphae]]. These branches can be identified by the lighter colouring in comparison to the ascocarp of the fungi, or truncation that results from the fragility of such branching. These truncations were previously thought to have been the release of [[conidiophore]]s, but no evidence was found on initial conidiophore attachment.<ref name=Orr1963/>
Commonly found in ''Myxotrichum'' species are the secondary and tertiary branching of [[peridium|peridial]] [[hyphae]]. These branches can be identified by the lighter colouring in comparison to the ascocarp of the fungi, or truncation that results from the fragility of such branching. These truncations were previously thought to have been the release of [[conidiophore]]s, but no evidence was found on initial conidiophore attachment.<ref name=Orr1963/>


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===Growth in laboratory culture===
===Growth in laboratory culture===
In culture and grown at 25°C, ''M. chartarum'' appears yellow and fluffy.<ref name="Currah1985"/> Some cultures had areas that were black in appearance, which were due to visible mature ascomata.<ref name="Currah1985"/> However, growth was restricted at this temperature, as fungi in the family Myxotrichaceae have a preference for temperatures below 18°C.<ref name="Currah1985"/><ref name="See1919"/> Optimal growing temperatures were described as being between +5 and +7°C.<ref name=Orr1963/><ref name="Currah1985"/>
In culture and grown at 25°C, ''M. chartarum'' appears yellow and fluffy.<ref name="Currah1985"/> Some cultures had areas that were black in appearance, which were due to visible mature ascomata.<ref name="Currah1985"/> However, growth was restricted at this temperature, as fungi in the family Myxotrichaceae have a preference for temperatures below 18°C.<ref name="Currah1985"/><ref name="See1919"/> At temperatures of 18°C, the fungus grew at 2 cm a month.<ref name="Tribe2002" /> Optimal growing temperatures were described as being between +5 and +7°C.<ref name=Orr1963/><ref name="Currah1985"/>
Production of a red-brown pigment when grown on mycelia and on certain culture media, functions to detoxify the surrounding environment.<ref name="Currah1985"/><ref name="Guiraud1995" /><ref name="Sato2014"/> This is due to a reaction produced by the [[laccase]]s secreted in the presence of [[polyphenol]]s.<ref name="Guiraud1995"/> The presence of pigmentation occurs early on for polyphenol detection.<ref name="Guiraud1995"/> However, this effect weakens when the fungus is exposed to higher levels of polyphenols, indicating metabolic function inhibition caused by the presence of these compounds.<ref name="Guiraud1995"/>
Production of a red-brown pigment when grown on mycelia and on certain culture media, functions to detoxify the surrounding environment.<ref name="Currah1985"/><ref name="Guiraud1995" /><ref name="Sato2014"/> This is due to a reaction produced by the [[laccase]]s secreted in the presence of [[polyphenol]]s.<ref name="Guiraud1995"/> The presence of pigmentation occurs early on for polyphenol detection.<ref name="Guiraud1995"/> However, this effect weakens when the fungus is exposed to higher levels of polyphenols, indicating metabolic function inhibition caused by the presence of these compounds.<ref name="Guiraud1995"/>



Revision as of 16:43, 23 November 2018

Myxotrichum chartarum
Scientific classification
Kingdom:
Phylum:
Subphylum:
Class:
Order:
Family:
Genus:
Myxotrichum
Species:
M. chartarum
Binomial name
Myxotrichum chartarum
Kunze (1823)[1]
Synonyms
  • Oncidium chartarum Kunze (1823)
  • Actinospira chartarum Corda (1854)
  • Myxotrichum carminoparum Robak (1932)

Myxotrichum chartarum is a psychrophilic and cellulolytic fungus first discovered in Germany by Kunze in 1823. Its classification has changed many times over its history to better reflect the information available at the time. Currently, M. chartarum is known to be an gymnothecal ascocarp with ornate appendages and releases release asexual ascospores. The presence of cellulolytic processes are common in fungi within the Myxotrichaceae family. M. chartarum is one of many chartarum species known to degrade paper and paper products. Evidence of M. chartarum "red spot" mold formation, especially on old books, can be found globally. As a result, this fungal species and other cellulolytic molds are endangering old works of art and books. Currently, there is no evidence that suggests that species within the family Myxotrichaceae are pathogenic.

History and taxonomy

The fungal species, Myxotrichum chartarum was discovered by Kunze in 1823.[1]

Originally, M. chartarum was mistaknly placed in the genus Oncidium by Nees (1823) which was already in use for species of orchid. M. chartarum was thereafter reclassified by Kunze.[1][2] In the same year, the genus Myxotrichum was established by Kunze to include two species, M. chartarum and Myxotrichum murorum.[1] Additional species were added thereafter.[1] There has been much confusion within the Myxotrichum genus due to numerous revisions in classification over the years and scarcity of isolations.[1][2] Eventually, the genus, Myxotrichum became associated with ascomycota fungi which are dematiaceous hyphomycetes, commonly known as black yeasts or moulds.[2] At the microscopic level, these fungi have a mesh-like surrounding structures (peridium) with hooked appendages.[2]

In 1838, Corda classified M. chartarum as a Hyphomycete of the family Sporotrichacheae, as species belonging to this family had ornate appendages resembling deer antlers.[3] In 1854, Corda separated M. chartarum from Myxotrichum into the newly established genus, Actinospira because he believed it to produce conidia rather than ascospores.[1] In 1959, Kuehn, among other investigators, reviewed the status of the family Gymnoascaceae and placed M. chartarum into ascomycetous genus, Myxotrichum rather than the newly established genus for conidial forms, Myxotrichella.[1] In 1875, Fuckel declared M. chartarum to be the conidial form of Chaetomium kunzeanum.[3] Fries thought M. chartarum was a conidial form of Chaetomium chartarum.[4] His opinion was supported by Boulanger in 1897.[3] Their revelation was founded because of the high resemblance between Chaetomium and Myxotrichum fungal families due to the presence of ornamental hairs.[3][1] In 1889, Richon thought M. chartarum was the conidial form of Cephalotheca sulfurea, disputing the claim of Fuckel.[3] In 1891, Constantin showed that M. chartarum belonged to the newly established family, ascomycete of the family Gymnoascaceae, for ascomycetes that lacked true cleistothecia or perithecia.[1] In 1892, Rabenhorst classified M. chartarum into the order, Gymnoascaceae.[3] This was later supported by Schroter in 1893.[3] Fischer later recognized the existence of ascospores in M. chartarum, but a distinct lack of conidial structures.[3] In 1893, Schroeter reviewed family Gymnoascaceae fungi and placed species with uncinate appendages within the genus, Myxotrichum without regard for other characteristics.[1]

Different forms of M. chartarum were isolated by Robak and Udagawa that resembled a phenotypically similar species.[2] The isolate discovered by Robak (1932) of M. carminoparum resembled M. chartarum[2] in every way except in the characteristically flattened apical area. Another isolate was discovered by Udagawa (1963) to have flattened appendages, but the size of the ascomata was smaller, resembling those from M. carminoparum.[2] Later, the species, M. chartarum and M. carminoparum were merged due to the high resemblance between the two species.

Growth and morphology

Intercalary arthroconidia of Myxotrichum chartarum (UAMH 10244) from colony grown for 36d, photographed in Phase Contrast microscopy.

Growth in its natural environment

The ascospores can also be described as being yellow to orange in colour with a rounded football shape having longitudinal striations and having a diameter between 6-8 μm on the long side and 5-8 µm on the short side.[1][2][5][6] Asci appear hyaline, globular, and contain the typical quantity of 8 ascospores each, the size of which are 3.5-5.0 µm on the long side and 2.0-3.0 µm on the short side.[1][2][6] When the spores are mature, they are released altogether, producing a cloud of brown dust.[6] Ascocarps appear dark and spherical with short appendages, where the ascocarp can appear green or copper when filled with spores.[1][2] The diameter of the ascocarp is larger than its close relative M. carminoparum at 150-555μm.[1] The spore mass fills the ascocarps between one-third and three-fourths of the total volume at peak maturity.[1][2] The surrounding Gymnothecium has septate appendages that are straight and elongated. Branching points present as uncinate, or curved spines that are wider at the apices. [1][2] Commonly found in Myxotrichum species are the secondary and tertiary branching of peridial hyphae. These branches can be identified by the lighter colouring in comparison to the ascocarp of the fungi, or truncation that results from the fragility of such branching. These truncations were previously thought to have been the release of conidiophores, but no evidence was found on initial conidiophore attachment.[1]

Life cycle

There are few records of the asexual or anamorphic stage within the family Gymnascaceae.[1] Descriptions made by Kuehn (1955) and Robak (1932) described oidia and chlamydospores, though rare in occurrence. However, Benjamin (1956) acknowledged that there were indeed artheroaleurispores and aleuriospores present in Gymnascaceae.[1][2] The anamorphs of M. chartarum may belong to the genera, Malbranchea and Oidiodendron.[2]

Growth in laboratory culture

In culture and grown at 25°C, M. chartarum appears yellow and fluffy.[2] Some cultures had areas that were black in appearance, which were due to visible mature ascomata.[2] However, growth was restricted at this temperature, as fungi in the family Myxotrichaceae have a preference for temperatures below 18°C.[2][3] At temperatures of 18°C, the fungus grew at 2 cm a month.[6] Optimal growing temperatures were described as being between +5 and +7°C.[1][2] Production of a red-brown pigment when grown on mycelia and on certain culture media, functions to detoxify the surrounding environment.[2][7][8] This is due to a reaction produced by the laccases secreted in the presence of polyphenols.[7] The presence of pigmentation occurs early on for polyphenol detection.[7] However, this effect weakens when the fungus is exposed to higher levels of polyphenols, indicating metabolic function inhibition caused by the presence of these compounds.[7]

Habitat and ecology

M. chartarum is known for its inhabitation of paper and paper products. The species epithet for this fungus, "chartarum", originated from the Latin word for paper and is in reference to its initial discovery from paper in books,[6] and its ability to decay these materials through the production of cellulose-degrading enzymes (cellulases).[9] It was deemed a “material pathogen”, since it is able to degrade specific materials as a source of nutrition.[9] It has also been recorded to inhabit other materials, such as drywall, straw, decaying leather, cloth, grouse dung, rabbit dung, bat guano, soil, leaves and fruit.[1][9][10][2][11] Reports of M. chartarum have come from around the world. Known distribution areas are as follows: Germany, Russia, Italy, France, Switzerland, England, Japan, Austria-Hungary, Czechoslovakia, Maine, Massachusetts, Ontario, New York.[1][2] The endemic region of this fungus is currently unknown. The curved spines allow the ascocarp to adhere to the fur of animals, allowing the fungus to disperse to other areas.[3]

It has been observed by multiple researchers that M. chartarum exhibited slow growth between temperatures of 5-7°C, so this fungus is classified as a psychrophilic organism.[1][6][2] However, no growth was observed at 37°C and since it is unable to grow at human body temperature, this fungus is not a disease agent or an opportunistic pathogen.[9] As found by Tribe and Weber (2002),[6] optimal growth in culture can be achieved on mineral salt agar with a sheet of Cellophane as the only carbon source.[6] In basements, M. chartarum has a preference for gypsum board ceilings and building paper on concrete surfaces on the cold side of foundation walls.[10] For optimal growth, it requires a relative humidity of greater than 98%.[10] Salinity and pH preferences are unknown, but it is thought to be halo-tolerant.[10]

References

  1. ^ a b c d e f g h i j k l m n o p q r s t u v w x Orr, G F; Kuehn, H H; Plunkett, O A (1963). "THE GENUS MYXOTRICHUM KUNZE". Can J of Botany. 41 (10): 1457-1480. doi:10.1139/b63-127.
  2. ^ a b c d e f g h i j k l m n o p q r s t u v Currah, R S (1985). . Taxonomy of the Onygenales: Arthrodermataceae, Gymnoascaceae, Myxotrichaceae and Onygenaceae (24 ed.). Mycotaxon. p. 1-216.
  3. ^ a b c d e f g h i j See, P (1919). La florule du papier. - Étude systématique et biologique des champignons chromogènes du papier piqué. Vol. 815. Université de Paris.{{cite book}}: CS1 maint: location missing publisher (link)
  4. ^ Fries, E M (1836–1838). Epicrisis systematis mycologici, seu synopsis Hymenomycetum. p. 346-349.{{cite book}}: CS1 maint: date format (link)
  5. ^ Ellis, M B; Ellis, J P (1988). Microfungi on miscellaneous substrates: an identification handbook (I ed.). Netherlands: Springer Netherlands. ISBN 9780855462482.
  6. ^ a b c d e f g h Tribe, H T; Weber, R W S (2002). "A low-temperature fungus from cardboard, Myxotrichum chartarum". Mycologist. 16 (1): 3-5. doi:10.1017/S0269-915X(02)00614-6.
  7. ^ a b c d Guiraud, P; Steiman, R; Seiglemurandi, F; Benoitguyod, JL (1995). "Comparison of the toxicity of various lignin-related phenolic compounds toward selected fungi perfecti and fungi imperfecti" (PDF). Ecotoxicology and Environmental Safety. 32 (1): 29-33.
  8. ^ Sato, Y; Aoki, M; Kigawa, R (2014). "Microbial deterioration of tsunami-affected paper-based objects" (PDF). National Research Institute for Cultural Properties Tokyo. (2012): 51–65.
  9. ^ a b c d Sterflinger, K; Pinzari, F (2012). "The revenge of time: fungal deterioration of cultural heritage with particular reference to books, paper and parchment". Environ Microbiol. 14 (3): 559-66. doi:10.1111/j.1462-2920.2011.02584.x.
  10. ^ a b c d Nunez, M; Hammer, H (2014). "Microbial specialists in below‐grade foundation walls in Scandinavia". Int J Indoor Env and Health. 24 (5): 543–551. doi:10.1111/ina.12095.
  11. ^ Nováková, A (2009). "Microscopic fungi isolated from the Domica Cave system (Slovak Karst National Park, Slovakia). A review". International Journal of Speleology. 38 (1): 8. doi:10.5038/1827-806X.38.1.8.