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{{italic title}}
{{ course assignment | course = Education Program:University of Toronto/HMB436H - Human and Veterinary Mycology (2013 Q3) | term = Fall 2013 }}
{{italic title}}{{Taxobox
{{Taxobox
| name = ''Myceliophthora thermophila''
| name = ''Myceliophthora thermophila''
| regnum = [[Fungi]]
| regnum = [[Fungi]]
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*''Thielavia heterothallica'' <small>A. von Klopotek, 1976</small>
*''Thielavia heterothallica'' <small>A. von Klopotek, 1976</small>
*''Corynascus heterothallicus'' <small>(A. von Klopotek) J.A. von Arx ''et al.'', 1983</small>
*''Corynascus heterothallicus'' <small>(A. von Klopotek) J.A. von Arx ''et al.'', 1983</small>
| synonyms_ref = &nbsp;<ref name=fungal_genome_project/>
| synonyms_ref = &nbsp;<ref>{{cite web |url=http://fungalgenomics.concordia.ca/fungi/Mthe.php |title=''Myceliophthora thermophila'' |work=Fungal Genome Project |publisher=[[Concordia University]] |date=April 5, 2005 |accessdate=April 24, 2013}}</ref>
}}
}}
'''''Myceliophthora thermophila''''' is an [[ascomycete]] fungus which grows optimally at {{convert|45|-|50|C|F}}. It efficiently degrades [[cellulose]] and is of interested in the production of [[biofuel]]s. The genome has recently been sequenced,<ref>{{Cite journal | doi = 10.1038/nbt.1976 | title = Comparative genomic analysis of the thermophilic biomass-degrading fungi ''Myceliophthora thermophila'' and ''Thielavia terrestris'' | year = 2011 | last1 = Berka | first1 = Randy M | last2 = Grigoriev | first2 = Igor V | last3 = Otillar | first3 = Robert | last4 = Salamov | first4 = Asaf | last5 = Grimwood | first5 = Jane | last6 = Reid | first6 = Ian | last7 = Ishmael | first7 = Nadeeza | last8 = John | first8 = Tricia | last9 = Darmond | first9 = Corinne | journal = [[Nature Biotechnology]] | volume = 29 | issue = 10 | pages = 922–927 | pmid = 21964414}}</ref> revealing the full range of enzymes this organism uses for the degradation of [[plant cell wall]] material.
'''''Myceliophthora thermophila''''' is an [[ascomycete]] fungus that grows optimally at {{convert|45|-|50|C|F}}. It efficiently degrades [[cellulose]] and is of interest in the production of [[biofuel]]s. The genome has recently been sequenced,<ref name=berka2011/> revealing the full range of enzymes this organism uses for the degradation of [[plant cell wall]] material.

==Taxonomy==
''Myceliophthora thermophila'' has a wide range of synonyms over the history of its classification and distinction of sexual states. ''Myceliophthora thermophila'' was originally described as ''Sporotrichum thermophilum'' in 1963<ref name=apnis1963/>, but it was later found that the species lacked clamp connections characteristic of the basidiomycetous genus, ''Sporotrichum''. It was reclassified to the ascomyceteous genus, ''Chrysosporium'', and became known as ''C. thermophilum''. The genus ''Myceliophthora'' was not used to describe this species until 1977, since the genus ''Chrysosporium'' formerly encompassed the genus ''Myceliophthora'',<ref name=vanoorschot1980/>

The [[teleomorph]] to ''M. thermophila'' first described as ''Thielavia heterothallica'' before the genus ''Corynascus'' was introduced by von Arx in 1983. It has since been known as ''Corynascus heterothallicus'', which has been observed through phylogenetic analysis to bear very strong DNA sequence homology to ''M. thermophila''<ref name=vandenbrink2011/><ref name=mycobank/>.

==Ecology==
''Myceliophthora thermophila'' is a [[thermophilic]] fungus, growing optimally at 38-45°C but not above 60°C.<ref name=maheshwari2000/>. ''Myceliophthora thermophila'' colonies have been commonly isolated from composts, where they generate high temperatures from cellular activities. Moist, sun-heated soils and hay provide ideal places for ''M. thermophila'' growth because they do not easily dissipate heat and help insulate the colony<ref name=jennings1999/>. Due to the scarcity of soluble carbon sources at high temperatures, this species is well adapted to utilizing insoluble carbon sources for energy, such as cellulose and hemicellulose<ref name=maheshwari2011/>.

==Morphology==
Colonies of ''M. thermophila'' initially appear cottony-pink, but rapidly turn cinnamon-brown and granular in texture. It can be distinguished from the closely-related ''Myceliophthora lutea'' by the thermophilic character of the former, and its more darkly-pigmented, markedly obovate conidia<ref name=kane1997/>. Microscopic examination reveals septate hyphae with several obovoidal to pyriform conidia arising singly or in small groups from conidiogenous cells. Conidia are typically 3.0-4.5μm x 4.5-11.0μm in size, hyaline, smooth, and thick-walled. Occasionally a secondary conidium can form at the distal tip of primary conidium.<ref name=bourbeau1992/><ref name=farina1998/>

==Human disease==
''Myceliopthora thermophila'' is rarely implicated in human disease; however, there have been several reported cases of ''M. thermophila'' causing disseminated infections in people with pre-existing immunodeficiency such as [[myeloblastic leukemia]].<ref name=bourbeau1992/><ref name=morio2011/> Infections can occur by direct inoculation into the body by contaminated surgical or garden tools, and tend to manifest themselves in cardiovascular and respiratory systems.<ref name=farina1998/><ref name=destino2006/> Voriconazole is an effective treatment for the infection, however, misdiagnoses for ''M. thermophila'' are possible due to its tendency to test positive on invasive [[aspergillosis]] screens.<ref name=morio2011/><ref name=destino2006/>

==Industrial uses==
The genome of ''M. thermophila'' encodes a number of [[thermostable]] enzymes with important industrial applications. Because of its ability to grow at high temperature, its enzyme yield is greater with fewer contaminants than many mesophilic fungi.<ref name=coutts1976/>

[[Cellulase]]s are rapidly synthesized by ''M. thermophila'' and can be used to degrade cellulose into simple carbohydrates as a food source for livestock.<ref name=coutts1976/> Also expressed by this species are broad-specificity [[phytase]]s that are efficient in breaking down phytic acid to be used for supplementing livestock feed with phosphorus.<ref name=maheshwari2000/><ref name=wyss1999/>

''Myceliophthora thermophila'' expresses [[laccase]]s that can act as clean substitutes for harmful chemical reagents used in the paper and pulp industry and textile dyes<ref name=berka1997/>. They are also useful in ecological restoration through soil bioremediation and ability to degrade rubber.<ref name=couto2006/><ref name=ismail2013/> Furthermore, laccases have shown to have the ability to polymerize lignin from waste material from the [[kraft process]]. The homogeneous lignin polymer may be used as raw materials for other products.<ref name=gouveia2013/>


==References==
==References==
{{Reflist}}
{{Reflist|30em|refs=

<ref name=apnis1963>{{cite journal|last=Apnis|first=A.E.|title=Occurrence of thermophilic microfungi in certain alluvid soils near Nottingham|journal=Nova. Hed.|year=1963|volume=5|pages=57-78}}</ref>

<ref name=berka1997>{{cite journal|last=Berka|first=R.M.|coauthors=Schneider, P.; Golighty, E.J.; Brown, S.H.; Madden, M.; Brown, K.M.; Halkier, T.; Mondorf, K.; Xu, F.;|title=Characterization of the gene encoding extracellular laccase of Myceliophthora thermophila and analysis of the recombinant enzyme expressed in Aspergillus oryzae|journal=Appl. Environ. Microbiol.|year=1997|volume=63|issue=8|pages=3151-3157}}</ref>

<ref name=berka2011>{{Cite journal | doi = 10.1038/nbt.1976 | title = Comparative genomic analysis of the thermophilic biomass-degrading fungi ''Myceliophthora thermophila'' and ''Thielavia terrestris'' | year = 2011 | last1 = Berka | first1 = Randy M | last2 = Grigoriev | first2 = Igor V | last3 = Otillar | first3 = Robert | last4 = Salamov | first4 = Asaf | last5 = Grimwood | first5 = Jane | last6 = Reid | first6 = Ian | last7 = Ishmael | first7 = Nadeeza | last8 = John | first8 = Tricia | last9 = Darmond | first9 = Corinne | journal = [[Nature Biotechnology]] | volume = 29 | issue = 10 | pages = 922–927 | pmid = 21964414}}</ref>

<ref name=bourbeau1992>{{cite journal|last=Bourbeau|first=P.|coauthors=McGough, D.A.; Fraser, H.; Shah, N.; Rinaldi, M.G.|title=Fatal disseminated infection caused by Myceliophthora thermophila, a new agent of mycosis: case history and laboratory characteristics|journal=J. Clin. Microbiol.|year=1992|volume=30|pages=3019-3023}}</ref>

<ref name=couto2006>{{cite journal|last=Rodríguez Couto|first=Susana|coauthors=Toca Herrera, José Luis|title=Industrial and biotechnological applications of laccases: A review|journal=Biotechnology Advances|date=1 September 2006|volume=24|issue=5|pages=500–513|doi=10.1016/j.biotechadv.2006.04.003}}</ref>

<ref name=coutts1976>{{cite journal|last=Coutts|first=A.D.|coauthors=Smith, R.E.|title=Factors influencing the Production of Cellulases by Sporotrichum thermophile|journal=Appl. Environ. Microbiol.|year=1976|volume=31|issue=6|pages=819-826}}</ref>

<ref name=destino2006>{{cite journal|last=Destino|first=Lauren|coauthors=Sutton, Deanna A; Helon, Anna L; Havens, Peter L; Thometz, John G; Willoughby, Rodney E; Chusid, Michael J|journal=Annals of Clinical Microbiology and Antimicrobials|date=1 January 2006|volume=5|issue=1|pages=21|doi=10.1186/1476-0711-5-21}}</ref>

<ref name=farina1998>{{cite journal|last=FARINA|first=C.|coauthors=GAMBA, A.; TAMBINI, R.; BEGUIN, H.; TROUILLET, J. L.|title=Fatal aortic Myceliophthorathermophila infection in a patient affected by cystic medial necrosis|journal=Medical Mycology|date=1 April 1998|volume=36|issue=2|pages=113–118|doi=10.1046/j.1365-280X.1998.00135.x}}</ref>

<ref name=fungal_genome_project>{{cite web |url=http://fungalgenomics.concordia.ca/fungi/Mthe.php |title=''Myceliophthora thermophila'' |work=Fungal Genome Project |publisher=[[Concordia University]] |date=April 5, 2005 |accessdate=April 24, 2013}}</ref>

<ref name=gouveia2013>{{cite journal|last=Gouveia|first=S.|coauthors=Fernández-Costas, C.; Sanromán, M.A.; Moldes, D.|title=Polymerisation of Kraft lignin from black liquors by laccase from Myceliophthora thermophila: Effect of operational conditions and black liquor origin|journal=Bioresource Technology|date=1 March 2013|volume=131|pages=288–294|doi=10.1016/j.biortech.2012.12.155}}</ref>

<ref name=ismail2013>{{cite journal|last=Ismail|first=Mady A.|coauthors=Mohamed, Nadia H.; Shoreit, Ahmed A.M.|title=Degradation of Ficus elastica rubber latex by Aspergillus terreus, Aspergillus flavus and Myceliophthora thermophila|journal=International Biodeterioration & Biodegradation|date=1 March 2013|volume=78|pages=82–88|doi=10.1016/j.ibiod.2012.12.009}}</ref>

<ref name=jennings1999>{{cite book|last=Lysek|first=D.H. Jennings, G.|title=Fungal biology : undertanding the fungal lifestyle.|year=1999|publisher=Springer|location=Nueva York|isbn=0387915931|edition=2nd ed.}}</ref>

<ref name=kane1997>{{cite book|last=Kane|first=Julius; Summerbell, Richard; Sigler, Lynne; Krajden, Sigmund.; Land, Geoffrey|title=Laboratory handbook of dermatophytes : a clinical guide and laboratory handbook of dermatophytes and other filamentous fungi from skin, hair, and nails|year=1997|publisher=Star Pub.|location=Belmont, CA|isbn=978-0898631579}}</ref>

<ref name=maheshwari2000>{{cite journal|last=Maheshwari|first=R.|coauthors=Bharadwaj, G.; Bhat, M. K.|title=Thermophilic Fungi: Their Physiology and Enzymes|journal=Microbiology and Molecular Biology Reviews|date=1 September 2000|volume=64|issue=3|pages=461–488|doi=10.1128/MMBR.64.3.461-488.2000}}</ref>

<ref name=maheshwari2011>{{cite book|last=Maheshwari|first=Ramesh|title=Fungi : experimental methods in biology|year=2011|publisher=CRC Press|location=Boca Raton|isbn=978-1439839034|edition=2nd ed.}}</ref>

<ref name=morio2011>{{cite journal|last=Morio|first=F|coauthors=Fraissinet, F; Gastinne, T; Le Pape, P; Delaunay, J; Sigler, L; Gibas, CF; Miegeville, M|title=Invasive Myceliophthora thermophila infection mimicking invasive aspergillosis in a neutropenic patient: a new cause of cross-reactivity with the Aspergillus galactomannan serum antigen assay.|journal=Medical mycology : official publication of the International Society for Human and Animal Mycology|date=2011 Nov|volume=49|issue=8|pages=883-6|pmid=21619496}}</ref>

<ref name=mycobank>{{cite web|title=Myceliophthora thermophila|url=http://www.mycobank.org/BioloMICS.aspx?Link=T&TableKey=14682616000000063&Rec=11793&Fields=All|work=MycoBank|accessdate=17 October 2013}}</ref>

<ref name=vandenbrink2011>{{cite journal|last=van den Brink|first=Joost|coauthors=Samson, Robert A.; Hagen, Ferry; Boekhout, Teun; Vries, Ronald P.|title=Phylogeny of the industrial relevant, thermophilic genera Myceliophthora and Corynascus|journal=Fungal Diversity|date=28 May 2011|volume=52|issue=1|pages=197–207|doi=10.1007/s13225-011-0107-z}}</ref>

<ref name=vanoorschot1980>{{cite journal|last=van Oorschot|first=C.A.N.|title=A revision of Chrysosporium and allied genera|journal=Studies in Mycology|year=1980|volume=20|url=http://www.cbs.knaw.nl/publications/Sim20/full%20text.htm}}</ref>

<ref name=wyss1999>{{cite journal|last=Wyss|first=M.|coauthors=Brugger, R.; Kronenberger, A.; Remy, R.; Fimbel, R.; Oesterhelt, G.; Lehmann, M.; van Loon, A.|title=Biochemical Characterization of Fungal Phytases (myo-Inositol Hexakisphosphate Phosphohydrolases) Catalytic Properties|journal=Appl. Environ. Microbiol.|year=1999|volume=65|issue=2|pages=367-373}}</ref>
}}


[[Category:Sordariales]]
[[Category:Sordariales]]
[[Category:Fungi described in 1977]]
[[Category:Fungi described in 1977]]



{{Sordariomycetes-stub}}
{{Sordariomycetes-stub}}

Revision as of 05:35, 21 November 2013

Myceliophthora thermophila
Scientific classification
Kingdom:
Phylum:
Class:
Order:
Family:
Genus:
Species:
M. thermophila
Binomial name
Myceliophthora thermophila
(A.E. Apinis) C.A. van Oorschot, 1977
Synonyms [1]
  • Chrysosporium thermophilum (A.E. Apinis) A. von Klopotek, 1974
  • Sporotrichum thermophile A.E. Apinis, 1963
  • Thielavia heterothallica A. von Klopotek, 1976
  • Corynascus heterothallicus (A. von Klopotek) J.A. von Arx et al., 1983

Myceliophthora thermophila is an ascomycete fungus that grows optimally at 45–50 °C (113–122 °F). It efficiently degrades cellulose and is of interest in the production of biofuels. The genome has recently been sequenced,[2] revealing the full range of enzymes this organism uses for the degradation of plant cell wall material.

Taxonomy

Myceliophthora thermophila has a wide range of synonyms over the history of its classification and distinction of sexual states. Myceliophthora thermophila was originally described as Sporotrichum thermophilum in 1963[3], but it was later found that the species lacked clamp connections characteristic of the basidiomycetous genus, Sporotrichum. It was reclassified to the ascomyceteous genus, Chrysosporium, and became known as C. thermophilum. The genus Myceliophthora was not used to describe this species until 1977, since the genus Chrysosporium formerly encompassed the genus Myceliophthora,[4]

The teleomorph to M. thermophila first described as Thielavia heterothallica before the genus Corynascus was introduced by von Arx in 1983. It has since been known as Corynascus heterothallicus, which has been observed through phylogenetic analysis to bear very strong DNA sequence homology to M. thermophila[5][6].

Ecology

Myceliophthora thermophila is a thermophilic fungus, growing optimally at 38-45°C but not above 60°C.[7]. Myceliophthora thermophila colonies have been commonly isolated from composts, where they generate high temperatures from cellular activities. Moist, sun-heated soils and hay provide ideal places for M. thermophila growth because they do not easily dissipate heat and help insulate the colony[8]. Due to the scarcity of soluble carbon sources at high temperatures, this species is well adapted to utilizing insoluble carbon sources for energy, such as cellulose and hemicellulose[9].

Morphology

Colonies of M. thermophila initially appear cottony-pink, but rapidly turn cinnamon-brown and granular in texture. It can be distinguished from the closely-related Myceliophthora lutea by the thermophilic character of the former, and its more darkly-pigmented, markedly obovate conidia[10]. Microscopic examination reveals septate hyphae with several obovoidal to pyriform conidia arising singly or in small groups from conidiogenous cells. Conidia are typically 3.0-4.5μm x 4.5-11.0μm in size, hyaline, smooth, and thick-walled. Occasionally a secondary conidium can form at the distal tip of primary conidium.[11][12]

Human disease

Myceliopthora thermophila is rarely implicated in human disease; however, there have been several reported cases of M. thermophila causing disseminated infections in people with pre-existing immunodeficiency such as myeloblastic leukemia.[11][13] Infections can occur by direct inoculation into the body by contaminated surgical or garden tools, and tend to manifest themselves in cardiovascular and respiratory systems.[12][14] Voriconazole is an effective treatment for the infection, however, misdiagnoses for M. thermophila are possible due to its tendency to test positive on invasive aspergillosis screens.[13][14]

Industrial uses

The genome of M. thermophila encodes a number of thermostable enzymes with important industrial applications. Because of its ability to grow at high temperature, its enzyme yield is greater with fewer contaminants than many mesophilic fungi.[15]

Cellulases are rapidly synthesized by M. thermophila and can be used to degrade cellulose into simple carbohydrates as a food source for livestock.[15] Also expressed by this species are broad-specificity phytases that are efficient in breaking down phytic acid to be used for supplementing livestock feed with phosphorus.[7][16]

Myceliophthora thermophila expresses laccases that can act as clean substitutes for harmful chemical reagents used in the paper and pulp industry and textile dyes[17]. They are also useful in ecological restoration through soil bioremediation and ability to degrade rubber.[18][19] Furthermore, laccases have shown to have the ability to polymerize lignin from waste material from the kraft process. The homogeneous lignin polymer may be used as raw materials for other products.[20]

References

  1. ^ "Myceliophthora thermophila". Fungal Genome Project. Concordia University. April 5, 2005. Retrieved April 24, 2013.
  2. ^ Berka, Randy M; Grigoriev, Igor V; Otillar, Robert; Salamov, Asaf; Grimwood, Jane; Reid, Ian; Ishmael, Nadeeza; John, Tricia; Darmond, Corinne (2011). "Comparative genomic analysis of the thermophilic biomass-degrading fungi Myceliophthora thermophila and Thielavia terrestris". Nature Biotechnology. 29 (10): 922–927. doi:10.1038/nbt.1976. PMID 21964414.
  3. ^ Apnis, A.E. (1963). "Occurrence of thermophilic microfungi in certain alluvid soils near Nottingham". Nova. Hed. 5: 57–78.
  4. ^ van Oorschot, C.A.N. (1980). "A revision of Chrysosporium and allied genera". Studies in Mycology. 20.
  5. ^ van den Brink, Joost (28 May 2011). "Phylogeny of the industrial relevant, thermophilic genera Myceliophthora and Corynascus". Fungal Diversity. 52 (1): 197–207. doi:10.1007/s13225-011-0107-z. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  6. ^ "Myceliophthora thermophila". MycoBank. Retrieved 17 October 2013.
  7. ^ a b Maheshwari, R. (1 September 2000). "Thermophilic Fungi: Their Physiology and Enzymes". Microbiology and Molecular Biology Reviews. 64 (3): 461–488. doi:10.1128/MMBR.64.3.461-488.2000. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  8. ^ Lysek, D.H. Jennings, G. (1999). Fungal biology : undertanding the fungal lifestyle (2nd ed. ed.). Nueva York: Springer. ISBN 0387915931. {{cite book}}: |edition= has extra text (help)CS1 maint: multiple names: authors list (link)
  9. ^ Maheshwari, Ramesh (2011). Fungi : experimental methods in biology (2nd ed. ed.). Boca Raton: CRC Press. ISBN 978-1439839034. {{cite book}}: |edition= has extra text (help)
  10. ^ Kane, Julius; Summerbell, Richard; Sigler, Lynne; Krajden, Sigmund.; Land, Geoffrey (1997). Laboratory handbook of dermatophytes : a clinical guide and laboratory handbook of dermatophytes and other filamentous fungi from skin, hair, and nails. Belmont, CA: Star Pub. ISBN 978-0898631579.{{cite book}}: CS1 maint: multiple names: authors list (link)
  11. ^ a b Bourbeau, P. (1992). "Fatal disseminated infection caused by Myceliophthora thermophila, a new agent of mycosis: case history and laboratory characteristics". J. Clin. Microbiol. 30: 3019–3023. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  12. ^ a b FARINA, C. (1 April 1998). "Fatal aortic Myceliophthorathermophila infection in a patient affected by cystic medial necrosis". Medical Mycology. 36 (2): 113–118. doi:10.1046/j.1365-280X.1998.00135.x. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  13. ^ a b Morio, F (2011 Nov). "Invasive Myceliophthora thermophila infection mimicking invasive aspergillosis in a neutropenic patient: a new cause of cross-reactivity with the Aspergillus galactomannan serum antigen assay". Medical mycology : official publication of the International Society for Human and Animal Mycology. 49 (8): 883–6. PMID 21619496. {{cite journal}}: Check date values in: |date= (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)
  14. ^ a b Destino, Lauren (1 January 2006). Annals of Clinical Microbiology and Antimicrobials. 5 (1): 21. doi:10.1186/1476-0711-5-21. {{cite journal}}: Missing or empty |title= (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)CS1 maint: unflagged free DOI (link)
  15. ^ a b Coutts, A.D. (1976). "Factors influencing the Production of Cellulases by Sporotrichum thermophile". Appl. Environ. Microbiol. 31 (6): 819–826. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  16. ^ Wyss, M. (1999). "Biochemical Characterization of Fungal Phytases (myo-Inositol Hexakisphosphate Phosphohydrolases) Catalytic Properties". Appl. Environ. Microbiol. 65 (2): 367–373. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  17. ^ Berka, R.M. (1997). "Characterization of the gene encoding extracellular laccase of Myceliophthora thermophila and analysis of the recombinant enzyme expressed in Aspergillus oryzae". Appl. Environ. Microbiol. 63 (8): 3151–3157. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)CS1 maint: extra punctuation (link)
  18. ^ Rodríguez Couto, Susana (1 September 2006). "Industrial and biotechnological applications of laccases: A review". Biotechnology Advances. 24 (5): 500–513. doi:10.1016/j.biotechadv.2006.04.003. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  19. ^ Ismail, Mady A. (1 March 2013). "Degradation of Ficus elastica rubber latex by Aspergillus terreus, Aspergillus flavus and Myceliophthora thermophila". International Biodeterioration & Biodegradation. 78: 82–88. doi:10.1016/j.ibiod.2012.12.009. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  20. ^ Gouveia, S. (1 March 2013). "Polymerisation of Kraft lignin from black liquors by laccase from Myceliophthora thermophila: Effect of operational conditions and black liquor origin". Bioresource Technology. 131: 288–294. doi:10.1016/j.biortech.2012.12.155. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)