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| species = '''''M. chartarum'''''
| species = '''''M. chartarum'''''
| binomial = ''Myxotrichum chartarum''
| binomial = ''Myxotrichum chartarum''
| binomial_authority = [[Gustav Kunze|Kunze]] (1823)
| binomial_authority = [[Gustav Kunze|Kunze]] (1823)<ref name=Orr1963/>
| synonyms = *''Oncidium chartarum'' <small>[[Gustav Kunze|Kunze]] (1823)</small>
| synonyms = *''Oncidium chartarum'' <small>[[Christian Gottfried Daniel Nees von Esenbeck|Nees]] (1823)</small>
*''Actinospira chartarum'' <small>[[August Carl Joseph Corda|Corda]] (1854)</small>
*''Actinospira chartarum'' <small>[[August Carl Joseph Corda|Corda]] (1854)</small>
*''Myxotrichum carminoparum'' <small>Robak (1932)</small>
*''Myxotrichum carminoparum'' <small>Robak (1932)</small>
}}
}}


'''''Myxotrichum chartarum''''' is a [[psychrophilic]] and [[cellulolyti]]c [[fungus]]<ref name=Nunez2014/> first discovered in [[Germany]] by [[Gustav Kunze|Kunze]] in 1823.<ref name=Orr1963/> ''M. chartarum'' was previously classified as ''Oncidium chartarum'', ''Actinospira chartarum'' and ''Myxotrichum carminoparum''. Its classification has since changed to better reflect the information available at the time. The presence of cellulolytic processes are common in fungi within the Myxotrichaceae family.<ref name="Currah1985"/><ref name=Tribe2002/> ''M. chartarum'' is one of many ''chartarum'' species known to degrade [[paper]] and paper products.<ref name=Tribe2002/> Evidence of ''M. chartarum'' "red spot" mold formation, especially on old books, can be found globally.<ref name=Orr1963/><ref name="Currah1985"/><ref name="Sato2014"/> As a result, this fungal species and other cellulolytic molds are endangering old works of art and books.<ref name="Sterf2012" /> It has not been shown that Myxotrichaceae fungi are a pathogenic family. <ref name="Currah1985"/>
'''''Myxotrichum chartarum''''' is a [[psychrophilic]] and [[cellulolytic]] [[fungus]] described by the [[Germany|German]] mycologist [[Gustav 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 [[ascomycete]] surrounded by a [[ascocarp#gymnothecium|gymnothecium]] composed of ornate spines and releases asexual [[ascospores]]. The production of [[cellulolytic]] enzymes is common in members of the family Myxotrichaceae. ''M. chartarum'' is one of many 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==
==History and taxonomy==
''Myxotrichum chartarum'' was first discovered by [[Christian Gottfried Daniel Nees von Esenbeck|Nees]] in 1823 and assigned to the genus ''[[Oncidium]]''.<ref name=Orr1963/> The species epithet "chartarum" is derived from the Latin word for paper, ''[[wikt:charta|charta]]'', in turn derived from the ancient Greek, [[wikt:χάρτης|χάρτης]] (khártēs, “papyrus, paper”).<ref name=Tribe2002/> That same year, [[Gustav Kunze|Kunze]] created the genus ''[[Myxotrichum]]'' to accommodate it because the name assigned by Nees was already in use for a genus of [[orchid]].<ref name=Orr1963/><ref name="Currah1985"/> [[Gustav Kunze|Kunze]] include two species, ''M. chartarum'' and ''Myxotrichum murorum''.<ref name=Orr1963/> Additional species were added thereafter.<ref name=Orr1963/>
The fungal species, ''Myxotrichum chartarum'' was discovered by [[Gustav Kunze|Kunze]] in 1823.<ref name=Orr1963/>


There has been much confusion on the taxonomic placement of the genus ''Myxotrichum'' resulting in numerous revisions in classification over the years.<ref name=Orr1963/><ref name="Currah1985"/> In 1838, [[August Carl Joseph Corda|Corda]] classified ''M. chartarum'' as in the [[hyphomycete]] family, Sporotrichaceae, characterized by ornate appendages resembling [[deer]] [[antler]]s.<ref name="See1919"/> In 1854, [[August Carl Joseph Corda|Corda]] segregated ''M. chartarum'' into the newly established genus, ''Actinospira'' because he believed it to produce [[conidia]] rather than [[ascospores]].<ref name=Orr1963/> In 1959, Kuehn, among other investigators, reviewed the status of members of the family Gymnoascaceae, and maintained this species in the genus, ''Myxotrichum'' rather than establishing a new genus, ''Myxotrichella'', for conidial forms.<ref name=Orr1963/> In 1875, Fuckel declared ''M. chartarum'' to be the conidial form of ''[[Chaetomium]] kunzeanum''.<ref name="See1919"/> Fries similarly thought ''M. chartarum'' was a conidial form of ''Chaetomium chartarum''.<ref name="Fries1836"/> His opinion was supported by Boulanger in 1897.<ref name="See1919"/> This conclusion is thought to be due to the substantial resemblance between members of the genera ''Chaetomium'' and ''Myxotrichum'' with respect to ornamental hairs.<ref name="See1919"/><ref name=Orr1963/> In 1889, Richon thought ''M. chartarum'' was the conidial form of ''[[Cephalotheca]] sulfurea'', disputing the claim of Fuckel.<ref name="See1919"/> In 1891, Constantin showed that ''M. chartarum'' belonged to the newly established family, [[ascomycota|ascomycete]] of the family [[Gymnoascaceae]], for [[ascomycete]]s that lacked true [[cleistothecia]] or [[perithecia]].<ref name=Orr1963/> In 1892,[[Gottlob Ludwig Rabenhorst|Rabenhorst]] classified ''M. chartarum'' in the [[Gymnoascaceae]].<ref name="See1919"/> This was later supported by Schroter in 1893.<ref name="See1919"/> Fischer later recognized the existence of ascospores in ''M. chartarum'', but a distinct lack of conidial structures.<ref name="See1919"/> In 1893, Schroeter reviewed family Gymnoascaceae and placed species with hooked appendages in the genus, ''Myxotrichum'' without regard for other characteristics.<ref name=Orr1963/>
Originally, ''M. chartarum'' was placed in the genus, [[Oncidium]] 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/> 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 fungi which are dematiaceous hyphomycetes, to reticuloperidial ascomycetes with hooked appendages.<ref name="Currah1985"/>


Different forms of ''M. chartarum'' were isolated by Robak and Udagawa that resembled a phenotypically similar species.<ref name="Currah1985"/> The isolate discovered by Robak (1932) of ''M. carminoparum'' resembled ''M. chartarum''<ref name="Currah1985"/> 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''.<ref name="Currah1985"/> Later, the species, ''M. chartarum'' and ''M. carminoparum'' were merged due to the high resemblance between the two species. 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 1854, [[August Carl Joseph Corda|Corda]] separated ''M. chartarum'' from ''Myxotrichum'' into the newly established genus, ''Actinospira'' because he believed it to produce [[conidia]] rather than [[ascospores]].<ref name=Orr1963/> In 1959, Kuehn, among other investigators, reviewed the status of ''M. chartarum'' and declared the fungus to be an [[ascomycota|ascomycete]] of the [[Gymnoascaceae]] family, for [[ascomycete]]s that lacked true [[cleistothecia]] or [[perithecia]].<ref name=Orr1963/>
In 1875, Fuckel declared ''M. chartarum'' to be the conidial form of ''[[Chaetomium]] kunzeanum''.<ref name="See1919"/>
Fries thought ''M. chartarum'' was a state of ''Chaetomium chartarum''. His opinion was supported by Boulanger in 1897.<ref name="See1919"/> Their revelation was founded because of the high resemblance between ''Chaetomium'' and ''Myxotrichum'' fungal families due to the presence of ornamental hairs.<ref name="See1919"/>
In 1889, Richon thought ''M. chartarum'' was the conidial form of ''[[Cephalotheca]] sulfurea'', disputing the claim of Fuckel.<ref name="See1919"/>
In 1892, Rabenhorst classified ''M. chartarum'' into the order, [[Gymnoascaceae]].<ref name="See1919"/> This was later supported by Schroter in 1893.<ref name="See1919"/> Fischer later recognized the existence of ascospores in ''M. chartarum'', but lack of conidial structures.<ref name="See1919"/>

Different forms of ''M. chartarum'' were isolated by Robak and Udagawa that resembled a phenotypically similar species.<ref name="Currah1985"/> The isolate discovered by Robak (1932) of ''M. carminoparum'' resembled ''M. chartarum''<ref name="Currah1985"/> 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''.<ref name="Currah1985"/> Later, the species, ''M. chartarum'' and ''M. carminoparum'' were merged due to the high resemblance between the two species.


==Growth and morphology==
==Growth and morphology==
[[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===
[[Ascocarp]]s appear dark and spherical with short appendages, that can also 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"/> [[ascocarp#gymnothecium|Gymnothecium]] have appendages that are straight and elongated, with septa. 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 is the secondary and tertiary branching of [[peridium|peridial]] [[hyphae]]. These branches are identified by their lighter colour from the centrum of the fungi, or truncation, due to the fragility of such branching. These truncations were previously thought to have been the release of [[conidiophore]]s, but no evidence was found on conidiophore attachment.<ref name=Orr1963/>
The centrally-located centrum is initially pale-white and turns yellow as it matures. Contained within are [[ascus|asci]], which are hyaline, globular, and contain 8 spores each which is typical of ascospores.<ref name=Orr1963/><ref name="Currah1985"/>
The [[ascospore]]s can also be described as being yellow to orange in colour with a fusoid shape having longitudinal striations.<ref name=Orr1963/><ref name="Currah1985"/><ref name="Ellis1988" />
The curved spines allow the ascocarp to adhere to the fur of animals, allowing the fungus to [[Biological dispersal#animal dispersal|disperse]] to other areas.<ref name="See1919"/>


===Life cycle===
===Natural environment===
The ascospores can also be described as being yellow to orange in colour with a rounded football shape having longitudinal striations and 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" />
In the [[asexual reproduction|asexual]] or [[anamorph]] stage, [[arthroconidium|artheroaleurispores]] and [[aleuriospore]]s are present.<ref name=Orr1963/><ref name="Currah1985"/> These anamorphs may belong to the genii, ''Malbranchea'' and ''Oidiodendron''.<ref name="Currah1985"/>
[[ascus|Asci]] appear hyaline, globular, and contain the typical quantity of 8 ascospores each, the size of which are 3-5 µm on the long side and 2-3 µm on the short side.<ref name=Orr1963/><ref name="Currah1985"/><ref name="Tribe2002" /> When the spores mature, they are released en masse, producing a cloud of brown-coloured dust.<ref name="Tribe2002" /> [[Ascocarp]]s are dark and spherical with short appendages, and when filled with its yellow to orange [[spores]], the ascocarp can appear green or copper.<ref name=Orr1963/><ref name="Currah1985"/> The diameter of the ascocarp is 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 or flattened 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/>


===Growth in laboratory culture===
===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 Myxotrichaceae family have a preference for temperatures below 18°C.<ref name="Currah1985"/><ref name="See1919"/> Optimal growing temperatures were described as being between +5°C and +7°C.<ref name=Orr1963/><ref name="Currah1985"/>
In culture and grown at 25°C, ''M. chartarum'' is yellow and fluffy.<ref name="Currah1985"/> Cultures can appear black due to presence of visibly maturing ascomata.<ref name="Currah1985"/> Growth tends to be restricted at 25 C 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 grows at a rate of 2 cm per month.<ref name="Tribe2002" /> Optimal growing temperatures are between 5-7°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" /> 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"/>
There is little morphological variability in [[asexual reproduction|asexual]] reproduction in the family Gymnascaceae.<ref name=Orr1963/> Descriptions by Kuehn (1955) and Robak (1932) included oidia and chlamydospores, though rare in occurrence. However, Benjamin (1956) acknowledged that there were indeed [[arthroconidium|arthrospores]] and [[aleuriospore]]s present in members of the Gymnascaceae.<ref name=Orr1963/><ref name="Currah1985"/> The anamorphs of ''M. chartarum'' belong to the genera, ''Malbranchea'' and ''Oidiodendron''.<ref name="Currah1985"/>


==Habitat and ecology==
==Habitat and ecology==
''M. chartarum'' has been recorded to inhabit these materials: paper, [[cardboard]], [[drywall]], [[straw]], rotting [[wood]], decaying [[leather]], [[cloth]], [[grouse]] [[dung]], [[rabbit]] dung, bat [[guano]], [[soil]], leaves and fruit.<ref name=Orr1963/><ref name=Sterf2012/><ref name=Nunez2014/><ref name="Currah1985"/><ref name="Novakova2009" />
''M. chartarum'' is known for its inhabitation of paper and its ability to decay these materials through the production of cellulose-degrading enzymes (cellulases).<ref name=Sterf2012/> It was deemed a “material pathogen”, since it is able to degrade specific materials for a source of nutrition.<ref name=Sterf2012/> 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.<ref name=Orr1963/><ref name=Sterf2012/><ref name=Nunez2014/><ref name="Currah1985"/><ref name="Novakova2009" /> 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]].<ref name=Orr1963/><ref name="Currah1985"/> The endemic region of this fungus is currently unknown. The presence of curved spines allow the fungus to adhere to the fur of animals, allowing the fungus to [[Biological dispersal#animal dispersal|disperse]] to other areas.<ref name="See1919"/>

The known distribution areas are as follows: [[Germany]], [[Russia]], [[Italy]], [[France]], [[Switzerland]], [[England]], [[Japan]], [[Austria-Hungary]], [[Czechoslovakia]], [[Maine]], [[Massachusetts]], [[Ontario]], [[New York]].<ref name=Orr1963/><ref name="Currah1985"/>

This fungus was named ''chartarum'' because it was discovered on the paper of old books,<ref name=Tribe2002/> and its ability to decay them through cellulolytic processes.<ref name=Sterf2012/> It was deemed a “material pathogen”, since it is able to degrade specific materials as a source of nutrition.<ref name=Sterf2012/>


It has been observed by multiple researchers that ''M. chartarum'' exhibited slow growth at 5-7°C and no growth at 37°C, so this fungus is classified as a [[psychrophilic]] organism.<ref name=Orr1963/><ref name=Tribe2002/><ref name="Currah1985"/> Since it cannot grow at human body temperature, this fungus is not an [[opportunistic pathogen|opportunistic]] pathogen.<ref name=Sterf2012/>
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.<ref name=Orr1963/><ref name=Tribe2002/><ref name="Currah1985"/> However, no growth was observed at 37°C and since it is unable to grow at the human body temperature, this fungus is not a disease agent or an [[opportunistic pathogen]].<ref name=Sterf2012/>
As found by Tribe and Weber (2002)<ref name="Tribe2002" />, optimal growth can be achieved on mineral salt [[agar]] with a sheet of [[Cellophane]] as the only [[carbon]] source.<ref name=Tribe2002/> In basements, it has a preference for [[drywall|gypsum board]] ceilings and building paper on concrete surfaces on the cold side of foundation walls.<ref name=Nunez2014/> For optimal growth, it requires a relative [[humidity]] of greater than 98%.<ref name=Nunez2014/> [[Salinity]] and [[pH]] preferences are unknown, but it is probably [[halotolerance|halo-tolerant]].<ref name=Nunez2014/>
As found by Tribe and Weber (2002),<ref name="Tribe2002" /> optimal growth in culture can be achieved on mineral salt [[agar]] with a sheet of [[Cellophane]] as the only [[carbon]] source.<ref name=Tribe2002/> In basements or cellars, ''M. chartarum'' has a preference for [[drywall|gypsum board]] ceilings and building paper on concrete surfaces on the cold side of foundation walls.<ref name=Nunez2014/> For optimal growth, it requires a relative [[humidity]] of greater than 98%.<ref name=Nunez2014/> [[Salinity]] and [[pH]] preferences are unknown, but it is thought to be [[halotolerance|halo-tolerant]].<ref name=Nunez2014/>


==References==
==References==

Latest revision as of 20:37, 3 January 2019

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 Nees (1823)
  • Actinospira chartarum Corda (1854)
  • Myxotrichum carminoparum Robak (1932)

Myxotrichum chartarum is a psychrophilic and cellulolytic fungus described by the German mycologist Gustav 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 ascomycete surrounded by a gymnothecium composed of ornate spines and releases asexual ascospores. The production of cellulolytic enzymes is common in members of the family Myxotrichaceae. M. chartarum is one of many 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

[edit]

Myxotrichum chartarum was first discovered by Nees in 1823 and assigned to the genus Oncidium.[1] The species epithet "chartarum" is derived from the Latin word for paper, charta, in turn derived from the ancient Greek, χάρτης (khártēs, “papyrus, paper”).[2] That same year, Kunze created the genus Myxotrichum to accommodate it because the name assigned by Nees was already in use for a genus of orchid.[1][3] Kunze include two species, M. chartarum and Myxotrichum murorum.[1] Additional species were added thereafter.[1]

There has been much confusion on the taxonomic placement of the genus Myxotrichum resulting in numerous revisions in classification over the years.[1][3] In 1838, Corda classified M. chartarum as in the hyphomycete family, Sporotrichaceae, characterized by ornate appendages resembling deer antlers.[4] In 1854, Corda segregated M. chartarum 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 members of the family Gymnoascaceae, and maintained this species in the genus, Myxotrichum rather than establishing a new genus, Myxotrichella, for conidial forms.[1] In 1875, Fuckel declared M. chartarum to be the conidial form of Chaetomium kunzeanum.[4] Fries similarly thought M. chartarum was a conidial form of Chaetomium chartarum.[5] His opinion was supported by Boulanger in 1897.[4] This conclusion is thought to be due to the substantial resemblance between members of the genera Chaetomium and Myxotrichum with respect to ornamental hairs.[4][1] In 1889, Richon thought M. chartarum was the conidial form of Cephalotheca sulfurea, disputing the claim of Fuckel.[4] 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 in the Gymnoascaceae.[4] This was later supported by Schroter in 1893.[4] Fischer later recognized the existence of ascospores in M. chartarum, but a distinct lack of conidial structures.[4] In 1893, Schroeter reviewed family Gymnoascaceae and placed species with hooked appendages in 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.[3] The isolate discovered by Robak (1932) of M. carminoparum resembled M. chartarum[3] 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.[3] Later, the species, M. chartarum and M. carminoparum were merged due to the high resemblance between the two species. At the microscopic level, these fungi have a mesh-like surrounding structures (peridium) with hooked appendages.[3]

Growth and morphology

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

Natural environment

[edit]

The ascospores can also be described as being yellow to orange in colour with a rounded football shape having longitudinal striations and a diameter between 6-8 μm on the long side and 5-8 µm on the short side.[1][3][6][2] Asci appear hyaline, globular, and contain the typical quantity of 8 ascospores each, the size of which are 3-5 µm on the long side and 2-3 µm on the short side.[1][3][2] When the spores mature, they are released en masse, producing a cloud of brown-coloured dust.[2] Ascocarps are dark and spherical with short appendages, and when filled with its yellow to orange spores, the ascocarp can appear green or copper.[1][3] The diameter of the ascocarp is 150-555μm.[1] The spore mass fills the ascocarps between one-third and three-fourths of the total volume at peak maturity.[1][3] The surrounding Gymnothecium has septate appendages that are straight and elongated. Branching points present as uncinate, or curved spines that are wider or flattened at the apices.[1][3] 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]

Laboratory culture

[edit]

In culture and grown at 25°C, M. chartarum is yellow and fluffy.[3] Cultures can appear black due to presence of visibly maturing ascomata.[3] Growth tends to be restricted at 25 C as fungi in the family Myxotrichaceae have a preference for temperatures below 18°C.[3][4] At temperatures of 18°C, the fungus grows at a rate of 2 cm per month.[2] Optimal growing temperatures are between 5-7°C.[1][3] Production of a red-brown pigment when grown on mycelia and on certain culture media, functions to detoxify the surrounding environment.[3][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]

There is little morphological variability in asexual reproduction in the family Gymnascaceae.[1] Descriptions by Kuehn (1955) and Robak (1932) included oidia and chlamydospores, though rare in occurrence. However, Benjamin (1956) acknowledged that there were indeed arthrospores and aleuriospores present in members of the Gymnascaceae.[1][3] The anamorphs of M. chartarum belong to the genera, Malbranchea and Oidiodendron.[3]

Habitat and ecology

[edit]

M. chartarum is known for its inhabitation of paper 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 for 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][3][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][3] The endemic region of this fungus is currently unknown. The presence of curved spines allow the fungus to adhere to the fur of animals, allowing the fungus to disperse to other areas.[4]

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][2][3] However, no growth was observed at 37°C and since it is unable to grow at the human body temperature, this fungus is not a disease agent or an opportunistic pathogen.[9] As found by Tribe and Weber (2002),[2] optimal growth in culture can be achieved on mineral salt agar with a sheet of Cellophane as the only carbon source.[2] In basements or cellars, 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

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  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 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.
  3. ^ a b c d e f g h i j k l m n o p q r s t u Currah, R S (1985). . Taxonomy of the Onygenales: Arthrodermataceae, Gymnoascaceae, Myxotrichaceae and Onygenaceae (24 ed.). Mycotaxon. p. 1-216.
  4. ^ 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)
  5. ^ Fries, E M (1836–1838). Epicrisis systematis mycologici, seu synopsis Hymenomycetum. p. 346-349.{{cite book}}: CS1 maint: date format (link)
  6. ^ Ellis, M B; Ellis, J P (1988). Microfungi on miscellaneous substrates: an identification handbook (I ed.). Netherlands: Springer Netherlands. ISBN 9780855462482.
  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.
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