Rantasalo A, Vitikainen M, Paasikallio T, Ja ¨ntti J, Landowski CP, Mojzita D (2019) Novel genetic tools
that enable highly pure protein production in
Trichoderma reesei. Sci Rep 9:1–12
Read G, Vining LC (1968) The biosynthesis of terreic
acid. Chem Commun 16:935–937
Record E, Punt PJ, Chamkha M, Labat M, Van Den
Hondel CAMJJ, Asther M (2002) Expression of
the Pycnoporus cinnabarinus laccase gene in
Aspergillus niger and characterization of the
recombinant enzyme. Eur J Biochem 269:602–609
Rendsvig JKH, Workman CT, Hoof JB (2019) Bidirectional histone-gene promoters in Aspergillus:
characterization and application for multi-gene
expression. Fungal Biol Biotechnol 6:1–14
Riquelme M, Sa ´nchez-Leo ´n E (2014) The Spitzenko ¨rper: a choreographer of fungal growth and morphogenesis. Curr Opin Microbiol 20:27–33
Rogers JK, Taylor ND, Church GM (2016) Biosensorbased engineering of biosynthetic pathways. Curr
Opin Biotechnol 42:84–91
Rokas A, Wisecaver JH, Lind AL (2018) The birth,
evolution and death of metabolic gene clusters in
fungi. Nat Rev Microbiol 16:731–744
Roller S, Covill N (1999) The antifungal properties of
chitosan in laboratory media and apple juice. Int J
Food Microbiol 47:67–77
Romero B, Turner G, Olivas I, Laborda F, Ramo ´n De
Lucas J (2003) The Aspergillus nidulans alcA promoter drives tightly regulated conditional gene
expression in Aspergillus fumigatus permitting
validation of essential genes in this human pathogen. Fungal Genet Biol 40:103–114
Roze LV, Chanda A, Linz JE (2011) Compartmentalization and molecular traffic in secondary metabolism: a new understanding of established cellular
processes. Fungal Genet Biol 48:35–48
Ryan MD, King AMQ, Thomas GP (1991) Cleavage of
FMDV polyprotein is mediated by residues located
within a 19 amino acid sequence. J Gen Virol
72:2727–2732
Sakai K, Kinoshita H, Shimizu T, Nihira T (2008) Construction of a citrinin gene cluster expression system in heterologous Aspergillus oryzae. J Biosci
Bioeng 106:466–472
Sandermann H Jr (1992) Plant metabolism of xenobiotics. Trends Biochem Sci 17:82–84
Schale ´n M, Anyaogu DC, Hoof JB, Workman M (2016)
Effect of secretory pathway gene overexpression
on secretion of a fluorescent reporter protein
in Aspergillus nidulans. Fungal Biol Biotechnol
3:1–14
Scha ¨pe P, Kwon MJ, Baumann B, Gutschmann B, Jung
S, Lenz S, Nitsche B, Paege N, Schu ¨tze T, Cairns
TC, Meyer V (2019) Updating genome annotation
for the microbial cell factory Aspergillus niger
using gene co-expression networks. Nucleic
Acids Res 47:559–569
Schuetze T, Meyer V (2017) Polycistronic gene expression in Aspergillus niger. Microb Cell Factories
16:1–8
Shi TQ, Gao J, Wang WJ, Wang KF, Xu GQ, Huang H, Ji
XJ (2019) CRISPR/Cas9-based genome editing in
the filamentous fungus Fusarium fujikuroi and its
application in strain engineering for gibberellic
acid production. ACS Synth Biol 8:445–454
Shimizu T, Ito T, Kanematsu S (2012) Transient
and multivariate system for transformation of a
fungal plant pathogen, Rosellinia necatrix, using
autonomously replicating vectors. Curr Genet
58:129–138
Shoji J, Maruyama J, Arioka M, Kitamoto K (2005)
Development of Aspergillus oryzae thiA promoter
as a tool for molecular biological studies. FEMS
Microbiol Lett 244:41–46
Sibthorp C, Wu H, Cowley G, Wong PWH, Palaima P,
Morozov IY, Weedall GD, Caddick MX (2013)
Transcriptome analysis of the filamentous fungus
Aspergillus nidulans directed to the global identification of promoters. BMC Genomics 14:1–18
Song Z, Bakeer W, Marshall JW, Yakasai AA, Khalid
RM, Collemare J, Skellam E, Tharreau D, Lebrun
M-H, Lazarus CM, Bailey AM, Simpson TJ, Cox RJ
(2015) Heterologous expression of the avirulence
gene ACE1 from the fungal rice pathogen Magnaporthe oryzae. Chem Sci 6:4837–4845
Stajich JE, Dietrich FS, Roy SW (2007) Comparative
genomic analysis of fungal genomes reveals
intron-rich ancestors. Genome Biol 8:1–13
Steinberg G, Pen ˜alva MA, Riquelme M, Wo ¨sten HA,
Harris SD (2017) Cell biology of hyphal growth.
Microbiol Spectr 5:231–265
Sticha KRK, Sieg CA, Bergstrom CP, Hanna PE, Wagner
CR (1997) Overexpression and large-scale purification of recombinant hamster polymorphic arylamine N-acetyltransferase as a dihydrofolate
reductase fusion protein. Protein Expr Purif
10:141–153
Stone PJ, Makoff AJ, Parish JH, Radford A (1993) Cloning and sequence analysis of the glucoamylase
gene of Neurospora crassa. Curr Genet 24:205–211
Stricker AR, Grosstessner-Hain K, Wu ¨rleitner E, Mach
RL (2006) Xyr1 (xylanase regulator 1) regulates
both the hydrolytic enzyme system and d-xylose
metabolism in Hypocrea jecorina. Eukaryot Cell
5:2128–2137
Sun A, Peterson R, Te’o J, Nevalainen H (2016) Expression of the mammalian peptide hormone obestatin
in Trichoderma reesei. New Biotechnol 33:99–106
Tada S, Gomi K, Kitamoto K, Takahashi K, Tamura G,
Hara S (1991) Construction of a fusion gene comprising the Taka-amylase a promoter and the
Escherichia coli b-glucuronidase gene and analysis
of its expression in Aspergillus oryzae. Mol Gen
Genet 229:301–306
Tanaka M, Tokuoka M, Gomi K (2014) Effects of codon
optimization on the mRNA levels of heterologous
10 Filamentous Fungi as Hosts for Heterologous Production of Proteins and Secondary. . .
263
that enable highly pure protein production in
Trichoderma reesei. Sci Rep 9:1–12
Read G, Vining LC (1968) The biosynthesis of terreic
acid. Chem Commun 16:935–937
Record E, Punt PJ, Chamkha M, Labat M, Van Den
Hondel CAMJJ, Asther M (2002) Expression of
the Pycnoporus cinnabarinus laccase gene in
Aspergillus niger and characterization of the
recombinant enzyme. Eur J Biochem 269:602–609
Rendsvig JKH, Workman CT, Hoof JB (2019) Bidirectional histone-gene promoters in Aspergillus:
characterization and application for multi-gene
expression. Fungal Biol Biotechnol 6:1–14
Riquelme M, Sa ´nchez-Leo ´n E (2014) The Spitzenko ¨rper: a choreographer of fungal growth and morphogenesis. Curr Opin Microbiol 20:27–33
Rogers JK, Taylor ND, Church GM (2016) Biosensorbased engineering of biosynthetic pathways. Curr
Opin Biotechnol 42:84–91
Rokas A, Wisecaver JH, Lind AL (2018) The birth,
evolution and death of metabolic gene clusters in
fungi. Nat Rev Microbiol 16:731–744
Roller S, Covill N (1999) The antifungal properties of
chitosan in laboratory media and apple juice. Int J
Food Microbiol 47:67–77
Romero B, Turner G, Olivas I, Laborda F, Ramo ´n De
Lucas J (2003) The Aspergillus nidulans alcA promoter drives tightly regulated conditional gene
expression in Aspergillus fumigatus permitting
validation of essential genes in this human pathogen. Fungal Genet Biol 40:103–114
Roze LV, Chanda A, Linz JE (2011) Compartmentalization and molecular traffic in secondary metabolism: a new understanding of established cellular
processes. Fungal Genet Biol 48:35–48
Ryan MD, King AMQ, Thomas GP (1991) Cleavage of
FMDV polyprotein is mediated by residues located
within a 19 amino acid sequence. J Gen Virol
72:2727–2732
Sakai K, Kinoshita H, Shimizu T, Nihira T (2008) Construction of a citrinin gene cluster expression system in heterologous Aspergillus oryzae. J Biosci
Bioeng 106:466–472
Sandermann H Jr (1992) Plant metabolism of xenobiotics. Trends Biochem Sci 17:82–84
Schale ´n M, Anyaogu DC, Hoof JB, Workman M (2016)
Effect of secretory pathway gene overexpression
on secretion of a fluorescent reporter protein
in Aspergillus nidulans. Fungal Biol Biotechnol
3:1–14
Scha ¨pe P, Kwon MJ, Baumann B, Gutschmann B, Jung
S, Lenz S, Nitsche B, Paege N, Schu ¨tze T, Cairns
TC, Meyer V (2019) Updating genome annotation
for the microbial cell factory Aspergillus niger
using gene co-expression networks. Nucleic
Acids Res 47:559–569
Schuetze T, Meyer V (2017) Polycistronic gene expression in Aspergillus niger. Microb Cell Factories
16:1–8
Shi TQ, Gao J, Wang WJ, Wang KF, Xu GQ, Huang H, Ji
XJ (2019) CRISPR/Cas9-based genome editing in
the filamentous fungus Fusarium fujikuroi and its
application in strain engineering for gibberellic
acid production. ACS Synth Biol 8:445–454
Shimizu T, Ito T, Kanematsu S (2012) Transient
and multivariate system for transformation of a
fungal plant pathogen, Rosellinia necatrix, using
autonomously replicating vectors. Curr Genet
58:129–138
Shoji J, Maruyama J, Arioka M, Kitamoto K (2005)
Development of Aspergillus oryzae thiA promoter
as a tool for molecular biological studies. FEMS
Microbiol Lett 244:41–46
Sibthorp C, Wu H, Cowley G, Wong PWH, Palaima P,
Morozov IY, Weedall GD, Caddick MX (2013)
Transcriptome analysis of the filamentous fungus
Aspergillus nidulans directed to the global identification of promoters. BMC Genomics 14:1–18
Song Z, Bakeer W, Marshall JW, Yakasai AA, Khalid
RM, Collemare J, Skellam E, Tharreau D, Lebrun
M-H, Lazarus CM, Bailey AM, Simpson TJ, Cox RJ
(2015) Heterologous expression of the avirulence
gene ACE1 from the fungal rice pathogen Magnaporthe oryzae. Chem Sci 6:4837–4845
Stajich JE, Dietrich FS, Roy SW (2007) Comparative
genomic analysis of fungal genomes reveals
intron-rich ancestors. Genome Biol 8:1–13
Steinberg G, Pen ˜alva MA, Riquelme M, Wo ¨sten HA,
Harris SD (2017) Cell biology of hyphal growth.
Microbiol Spectr 5:231–265
Sticha KRK, Sieg CA, Bergstrom CP, Hanna PE, Wagner
CR (1997) Overexpression and large-scale purification of recombinant hamster polymorphic arylamine N-acetyltransferase as a dihydrofolate
reductase fusion protein. Protein Expr Purif
10:141–153
Stone PJ, Makoff AJ, Parish JH, Radford A (1993) Cloning and sequence analysis of the glucoamylase
gene of Neurospora crassa. Curr Genet 24:205–211
Stricker AR, Grosstessner-Hain K, Wu ¨rleitner E, Mach
RL (2006) Xyr1 (xylanase regulator 1) regulates
both the hydrolytic enzyme system and d-xylose
metabolism in Hypocrea jecorina. Eukaryot Cell
5:2128–2137
Sun A, Peterson R, Te’o J, Nevalainen H (2016) Expression of the mammalian peptide hormone obestatin
in Trichoderma reesei. New Biotechnol 33:99–106
Tada S, Gomi K, Kitamoto K, Takahashi K, Tamura G,
Hara S (1991) Construction of a fusion gene comprising the Taka-amylase a promoter and the
Escherichia coli b-glucuronidase gene and analysis
of its expression in Aspergillus oryzae. Mol Gen
Genet 229:301–306
Tanaka M, Tokuoka M, Gomi K (2014) Effects of codon
optimization on the mRNA levels of heterologous
10 Filamentous Fungi as Hosts for Heterologous Production of Proteins and Secondary. . .
263