the anaerobic fungus Orpinomyces joyonii: characterization of the gene and its product. Can J Microbiol 43(5):477–485. https://www.ncbi.nlm.nih.gov/
pubmed/9165703
Ljungdahl LG (2008) The cellulase/hemicellulase system of the anaerobic fungus Orpinomyces PC2 and aspects of its applied use. Ann N Y Acad
Sci 1125(1):308–321. https://doi.org/10.1196/
annals.1419.030
Lombard V, Ramulu HG, Drula E, Coutinho PM, Henrissat B (2014) The carbohydrate-active enzymes
database (CAZy) in 2013. Nucleic Acids Res 42
(Database issue):D490–D495. https://doi.org/
10.1093/nar/gkt1178
Lo ´pez-Contreras AM, Smidt H, van der Oost J, Claassen
PAM, Mooibroek H, de Vos WM (2001) Clostridium beijerinckii cells expressing Neocallimastix
patriciarum glycoside hydrolases show enhanced
lichenan utilization and solvent production. Appl
Environ Microbiol 67(11):5127–5133. https://doi.
org/10.1128/aem.67.11.5127-5133.2001
Lowe SE, Theodorou MK, Trinci AP, Hespell RB (1985)
Growth of anaerobic rumen fungi on defined and
semi-defined media lacking rumen fluid. J Gen
Microbiol 131(9):2225–2229. http://mic.sgmjournals.org/content/131/9/2225.full.pdf
Marvin-Sikkema FD, Pedro Gomes TM, Gottschal JC,
Prins RA, Marvin-Sikkema FD (1993) Characterization of hydrogenosomes and their role in glucose
metabolism of Neocallimastix sp. L2. Arch Microbiol 160(5). https://doi.org/10.1007/bf00252226
Milne A, Theodorou MK, Jordan MGC, King-Spooner
C, Trinci APJ (1989) Survival of anaerobic fungi in
feces, in saliva, and in pure culture. Exp Mycol 13
(1):27–37. https://doi.org/10.1016/0147-5975(89)
90005-4
Mondo SJ, Dannebaum RO, Kuo RC, Louie KB, Bewick
AJ, LaButti K, Haridas S, Kuo A, Salamov A,
Ahrendt SR, Lau R, Bowen BP, Lipzen A, Sullivan
W, Andreopoulos BB, Clum A, Lindquist E, Daum
C, Northen TR, Kunde-Ramamoorthy G, Schmitz
RJ, Gryganskyi A, Culley D, Magnuson J, James
TY, O’Malley MA, Stajich JE, Spatafora JW, Visel
A, Grigoriev IV (2017) Widespread adenine N6methylation of active genes in fungi. Nat Genet 49
(6):964–968.
https://doi.org/10.1038/ng.3859.
http://www.nature.com/ng/journal/v49/n6/abs/
ng.3859.html#supplementary-information
Montenecourt BS, Eveleigh DE (1977) Preparation of
mutants of Trichoderma reesei with enhanced cellulase production. Appl Environ Microbiol 34
(6):777–782. http://www.ncbi.nlm.nih.gov/pmc/
articles/PMC242747/
Mood SH, Golfeshan AH, Tabatabaei M, Jouzani GS,
Najafi GH, Gholami M, Ardjmand M (2013) Lignocellulosic biomass to bioethanol, a comprehensive review with a focus on pretreatment. Renew
Sustain Energy Rev 27:77–93. https://doi.org/
10.1016/j.rser.2013.06.033
Morrison JM, Elshahed MS, Youssef N (2016a) A multifunctional GH39 glycoside hydrolase from the
anaerobic gut fungus Orpinomyces sp. strain
C1A. PeerJ 4:e2289. https://doi.org/10.7717/
peerj.2289
Morrison JM, Elshahed MS, Youssef NH (2016b)
Defined enzyme cocktail from the anaerobic fungus Orpinomyces sp. strain C1A effectively releases
sugars from pretreated corn stover and switchgrass. Sci Rep 6:29217. https://doi.org/10.1038/
srep29217
Mountfort D, Orpin C (1994) Anaerobic fungi: biology,
ecology, and function. CRC, Boca Raton, FL
Mu ¨ller M, Mentel M, van Hellemond JJ, Henze K,
Woehle C, Gould SB, Yu RY, van der Giezen M,
Tielens AG, Martin WF (2012) Biochemistry and
evolution of anaerobic energy metabolism in
eukaryotes. Microbiol Mol Biol Rev 76(2):444–
495. https://doi.org/10.1128/mmbr.05024-11
Murphy CL, Youssef NH, Hanafy RA, Couger MB, Stajich JE, Wang Y, Baker K, Dagar SS, Griffith GW,
Farag IF, Callaghan TM, Elshahed MS (2019) Horizontal gene transfer as an indispensable driver for
Neocallimastigomycota evolution into a distinct
gut-dwelling fungal lineage. Appl Environ Microbiol. https://doi.org/10.1128/aem.00988-19
Nagler M, Kozjek K, Etemadi M, Insam H, Podmirseg
SM (2019) Simple yet effective: microbial and biotechnological benefits of rumen liquid addition to
lignocellulose-degrading biogas plants. J Biotechnol
300:1–10.
https://doi.org/10.1016/j.jbiotec.2019.05.004
Nielsen BB, Zhu W-Y, Trinci APJ, Theodorou MK
(1995) Demonstration of zoosporangia of anaerobic fungi on plant residues recovered from faeces
of cattle. Mycol Res 99(4):471–474. https://doi.org/
10.1016/S0953-7562(09)80648-5
Nkemka VN, Gilroyed B, Yanke J, Gruninger R, Vedres
D, McAllister T, Hao X (2015) Bioaugmentation
with an anaerobic fungus in a two-stage process
for biohydrogen and biogas production using corn
silage and cattail. Bioresour Technol 185:79–88.
https://doi.org/10.1016/j.biortech.2015.02.100
Obembe OO, Jacobsen E, Timmers J, Gilbert H, Blake
AW, Knox JP, Visser RGF, Vincken J-P (2007)
Promiscuous,
non-catalytic,
tandem
carbohydrate-binding modules modulate the cellwall structure and development of transgenic
tobacco (Nicotiana tabacum) plants. J Plant Res
120(5):605–617. https://doi.org/10.1007/s10265007-0099-7
O’Malley MA, Theodorou MK, Kaiser CA (2012) Evaluating expression and catalytic activity of anaerobic fungal fibrolytic enzymes native topiromyces
sp E2 in Saccharomyces cerevisiae. Environ Progr
Sustain Energy 31(1):37–46. http://onlinelibrary.
wiley.com/doi/10.1002/ep.10614/abstract
434
V. Flad et al.
pubmed/9165703
Ljungdahl LG (2008) The cellulase/hemicellulase system of the anaerobic fungus Orpinomyces PC2 and aspects of its applied use. Ann N Y Acad
Sci 1125(1):308–321. https://doi.org/10.1196/
annals.1419.030
Lombard V, Ramulu HG, Drula E, Coutinho PM, Henrissat B (2014) The carbohydrate-active enzymes
database (CAZy) in 2013. Nucleic Acids Res 42
(Database issue):D490–D495. https://doi.org/
10.1093/nar/gkt1178
Lo ´pez-Contreras AM, Smidt H, van der Oost J, Claassen
PAM, Mooibroek H, de Vos WM (2001) Clostridium beijerinckii cells expressing Neocallimastix
patriciarum glycoside hydrolases show enhanced
lichenan utilization and solvent production. Appl
Environ Microbiol 67(11):5127–5133. https://doi.
org/10.1128/aem.67.11.5127-5133.2001
Lowe SE, Theodorou MK, Trinci AP, Hespell RB (1985)
Growth of anaerobic rumen fungi on defined and
semi-defined media lacking rumen fluid. J Gen
Microbiol 131(9):2225–2229. http://mic.sgmjournals.org/content/131/9/2225.full.pdf
Marvin-Sikkema FD, Pedro Gomes TM, Gottschal JC,
Prins RA, Marvin-Sikkema FD (1993) Characterization of hydrogenosomes and their role in glucose
metabolism of Neocallimastix sp. L2. Arch Microbiol 160(5). https://doi.org/10.1007/bf00252226
Milne A, Theodorou MK, Jordan MGC, King-Spooner
C, Trinci APJ (1989) Survival of anaerobic fungi in
feces, in saliva, and in pure culture. Exp Mycol 13
(1):27–37. https://doi.org/10.1016/0147-5975(89)
90005-4
Mondo SJ, Dannebaum RO, Kuo RC, Louie KB, Bewick
AJ, LaButti K, Haridas S, Kuo A, Salamov A,
Ahrendt SR, Lau R, Bowen BP, Lipzen A, Sullivan
W, Andreopoulos BB, Clum A, Lindquist E, Daum
C, Northen TR, Kunde-Ramamoorthy G, Schmitz
RJ, Gryganskyi A, Culley D, Magnuson J, James
TY, O’Malley MA, Stajich JE, Spatafora JW, Visel
A, Grigoriev IV (2017) Widespread adenine N6methylation of active genes in fungi. Nat Genet 49
(6):964–968.
https://doi.org/10.1038/ng.3859.
http://www.nature.com/ng/journal/v49/n6/abs/
ng.3859.html#supplementary-information
Montenecourt BS, Eveleigh DE (1977) Preparation of
mutants of Trichoderma reesei with enhanced cellulase production. Appl Environ Microbiol 34
(6):777–782. http://www.ncbi.nlm.nih.gov/pmc/
articles/PMC242747/
Mood SH, Golfeshan AH, Tabatabaei M, Jouzani GS,
Najafi GH, Gholami M, Ardjmand M (2013) Lignocellulosic biomass to bioethanol, a comprehensive review with a focus on pretreatment. Renew
Sustain Energy Rev 27:77–93. https://doi.org/
10.1016/j.rser.2013.06.033
Morrison JM, Elshahed MS, Youssef N (2016a) A multifunctional GH39 glycoside hydrolase from the
anaerobic gut fungus Orpinomyces sp. strain
C1A. PeerJ 4:e2289. https://doi.org/10.7717/
peerj.2289
Morrison JM, Elshahed MS, Youssef NH (2016b)
Defined enzyme cocktail from the anaerobic fungus Orpinomyces sp. strain C1A effectively releases
sugars from pretreated corn stover and switchgrass. Sci Rep 6:29217. https://doi.org/10.1038/
srep29217
Mountfort D, Orpin C (1994) Anaerobic fungi: biology,
ecology, and function. CRC, Boca Raton, FL
Mu ¨ller M, Mentel M, van Hellemond JJ, Henze K,
Woehle C, Gould SB, Yu RY, van der Giezen M,
Tielens AG, Martin WF (2012) Biochemistry and
evolution of anaerobic energy metabolism in
eukaryotes. Microbiol Mol Biol Rev 76(2):444–
495. https://doi.org/10.1128/mmbr.05024-11
Murphy CL, Youssef NH, Hanafy RA, Couger MB, Stajich JE, Wang Y, Baker K, Dagar SS, Griffith GW,
Farag IF, Callaghan TM, Elshahed MS (2019) Horizontal gene transfer as an indispensable driver for
Neocallimastigomycota evolution into a distinct
gut-dwelling fungal lineage. Appl Environ Microbiol. https://doi.org/10.1128/aem.00988-19
Nagler M, Kozjek K, Etemadi M, Insam H, Podmirseg
SM (2019) Simple yet effective: microbial and biotechnological benefits of rumen liquid addition to
lignocellulose-degrading biogas plants. J Biotechnol
300:1–10.
https://doi.org/10.1016/j.jbiotec.2019.05.004
Nielsen BB, Zhu W-Y, Trinci APJ, Theodorou MK
(1995) Demonstration of zoosporangia of anaerobic fungi on plant residues recovered from faeces
of cattle. Mycol Res 99(4):471–474. https://doi.org/
10.1016/S0953-7562(09)80648-5
Nkemka VN, Gilroyed B, Yanke J, Gruninger R, Vedres
D, McAllister T, Hao X (2015) Bioaugmentation
with an anaerobic fungus in a two-stage process
for biohydrogen and biogas production using corn
silage and cattail. Bioresour Technol 185:79–88.
https://doi.org/10.1016/j.biortech.2015.02.100
Obembe OO, Jacobsen E, Timmers J, Gilbert H, Blake
AW, Knox JP, Visser RGF, Vincken J-P (2007)
Promiscuous,
non-catalytic,
tandem
carbohydrate-binding modules modulate the cellwall structure and development of transgenic
tobacco (Nicotiana tabacum) plants. J Plant Res
120(5):605–617. https://doi.org/10.1007/s10265007-0099-7
O’Malley MA, Theodorou MK, Kaiser CA (2012) Evaluating expression and catalytic activity of anaerobic fungal fibrolytic enzymes native topiromyces
sp E2 in Saccharomyces cerevisiae. Environ Progr
Sustain Energy 31(1):37–46. http://onlinelibrary.
wiley.com/doi/10.1002/ep.10614/abstract
434
V. Flad et al.