Xu J, Gong ZZ (2003) Intron requirement for AFP gene
expression in Trichoderma viride. Microbiology
149:3093–3097
Xu W, Chooi Y-H, Choi JW, Li S, Vederas JC, Da Silva
NA, Tang Y (2013) LovG: the thioesterase required
for dihydromonacolin L release and lovastatin
nonaketide synthase turnover in lovastatin biosynthesis. Angew Chem Int Ed 52:6472–6475
Yang S, Kuang Y, Li H, Liu Y, Hui X, Li P, Jiang Z, Zhou
Y, Wang Y, Xu A, Li S, Liu P, Wu D (2013)
Enhanced production of recombinant secretory
proteins in Pichia pastoris by optimizing Kex2
P1’ site. PLoS One 8:1–11
Yoon J, Aishan T, Maruyama JI, Kitamoto K (2010)
Enhanced production and secretion of heterologous proteins by the filamentous fungus Aspergillus oryzae via disruption of vacuolar protein
sorting receptor gene AoVps10. Appl Environ
Microbiol 76:5718–5727
Yoshino-Yasuda S, Karita S, Kato M, Kitamoto N (2012)
Sequence analysis and heterologous expression of
rhamnogalacturonan lyase A gene (AsrglA) from
Shoyu koji mold, Aspergillus sojae KBN1340. Food
Sci Technol Res 18:901–909
Yu CH, Dang Y, Zhou Z, Wu C, Zhao F, Sachs MS, Liu Y
(2015) Codon usage influences the local rate of
translation elongation to regulate co-translational
protein folding. Mol Cell 59:744–754
Yuill E (1950) The numbers of nuclei in conidia of
Aspergilli. Trans Br Mycol Soc 33:324–331
Zadra I, Abt B, Parson W, Haas H (2000) xylP
promoter-based expression system and its use for
antisense downregulation of the Penicillium chrysogenum nitrogen regulator NRE. Appl Environ
Microbiol 66:4810–4816
Zeilinger S, Mach RL, Schindler M, Herzog P, Kubicek
CP (1996) Different inducibility of expression of
the two xylanase genes xyn1 and xyn2 in Trichoderma reesei. J Biol Chem 271:25624–25629
Zeilinger S, Mach R, Kubicek C (1998) Two adjacent
protein binding motifs in the cbh2 (Cellobiohydrolase II-encoding) promoter of the fungus
Hypocrea jecorina (Trichoderma reesei) cooperate
in the induction by cellulose. J Biol Chem
273:34463–34471
Zhang F, Keasling J (2011) Biosensors and their applications in microbial metabolic engineering.
Trends Microbiol 19:323–329
Zhang DX, Lu HL, Liao X, St Leger RJ, Nuss DL (2013)
Simple and efficient recycling of fungal selectable
marker genes with the Cre-loxP recombination
system via anastomosis. Fungal Genet Biol 61:1–9
Zhang XH, Tee LY, Wang XG, Huang QS, Yang SH
(2015) Off-target effects in CRISPR/Cas9-mediated
genome engineering. Mol Ther Nucleic Acids
4:1–8
Zhang C, Meng X, Wei X, Lu L (2016a) Highly efficient
CRISPR mutagenesis by microhomologymediated end joining in Aspergillus fumigatus.
Fungal Genet Biol 86:47–57
Zhang H, Wang S, Zhang XX, Ji W, Song F, Zhao Y, Li J
(2016b) The amyR-deletion strain of Aspergillus
niger CICC2462 is a suitable host strain to express
secreted protein with a low background. Microb
Cell Factories 15:1–11
Zhang J, Zhang G, Wang W, Wei D (2018) Enhanced
cellulase production in Trichoderma reesei RUT
C30 via constitution of minimal transcriptional
activators. Microb Cell Factories 17:1–14
Zhao C, Waalwijk C, de Wit PJGM, Tang D, van der Lee
T (2013) RNA-Seq analysis reveals new gene models and alternative splicing in the fungal pathogen
Fusarium graminearum. BMC Genomics 14:1–16
Zheng YM, Lin FL, Gao H, Zou G, Zhang JW, Wang GQ,
Chen GD, Zhou ZH, Yao XS, Hu D (2017) Development of a versatile and conventional technique
for gene disruption in filamentous fungi based on
CRISPR-Cas9 technology. Sci Rep 7:1–10
Zhou M, Wang T, Fu J, Xiao G, Liu Y (2015) Nonoptimal codon usage influences protein structure in
intrinsically disordered regions. Mol Microbiol
97:974–987
Zhou Z, Dang Y, Zhou M, Li L, Yu C, Fu J, Chen S, Liu Y
(2016) Codon usage is an important determinant
of gene expression levels largely through its effects
on transcription. Proc Natl Acad Sci USA 113:
E6117–E6125
Zhou Z, Dang Y, Zhou M, Yuan H, Liu Y (2018) Codon
usage biases co-evolve with transcription termination machinery to suppress premature cleavage
and polyadenylation. Elife 7:1–29
Zhu H, Choi S, Johnston AK, Wing RA, Dean RA (1997)
A large-insert (130 kbp) bacterial artificial chromosome library of the rice blast fungus Magnaporthe grisea: genome analysis, contig assembly,
and gene cloning. Fungal Genet Biol 21:337–347
Zhu B, Cai G, Hall EO, Freeman GJ (2007) In-Fusion
TM
assembly: seamless engineering of multidomain
fusion proteins, modular vectors, and mutations.
BioTechniques 43:354–359
Zubieta MP, Contesini FJ, Rubio MV, Gonc ¸alves
AEDSS, Gerhardt JA, Prade RA, Damasio ARDL
(2018) Protein profile in Aspergillus nidulans
recombinant strains overproducing heterologous
enzymes. Microb Biotechnol 11:346–358
10 Filamentous Fungi as Hosts for Heterologous Production of Proteins and Secondary. . .
265
expression in Trichoderma viride. Microbiology
149:3093–3097
Xu W, Chooi Y-H, Choi JW, Li S, Vederas JC, Da Silva
NA, Tang Y (2013) LovG: the thioesterase required
for dihydromonacolin L release and lovastatin
nonaketide synthase turnover in lovastatin biosynthesis. Angew Chem Int Ed 52:6472–6475
Yang S, Kuang Y, Li H, Liu Y, Hui X, Li P, Jiang Z, Zhou
Y, Wang Y, Xu A, Li S, Liu P, Wu D (2013)
Enhanced production of recombinant secretory
proteins in Pichia pastoris by optimizing Kex2
P1’ site. PLoS One 8:1–11
Yoon J, Aishan T, Maruyama JI, Kitamoto K (2010)
Enhanced production and secretion of heterologous proteins by the filamentous fungus Aspergillus oryzae via disruption of vacuolar protein
sorting receptor gene AoVps10. Appl Environ
Microbiol 76:5718–5727
Yoshino-Yasuda S, Karita S, Kato M, Kitamoto N (2012)
Sequence analysis and heterologous expression of
rhamnogalacturonan lyase A gene (AsrglA) from
Shoyu koji mold, Aspergillus sojae KBN1340. Food
Sci Technol Res 18:901–909
Yu CH, Dang Y, Zhou Z, Wu C, Zhao F, Sachs MS, Liu Y
(2015) Codon usage influences the local rate of
translation elongation to regulate co-translational
protein folding. Mol Cell 59:744–754
Yuill E (1950) The numbers of nuclei in conidia of
Aspergilli. Trans Br Mycol Soc 33:324–331
Zadra I, Abt B, Parson W, Haas H (2000) xylP
promoter-based expression system and its use for
antisense downregulation of the Penicillium chrysogenum nitrogen regulator NRE. Appl Environ
Microbiol 66:4810–4816
Zeilinger S, Mach RL, Schindler M, Herzog P, Kubicek
CP (1996) Different inducibility of expression of
the two xylanase genes xyn1 and xyn2 in Trichoderma reesei. J Biol Chem 271:25624–25629
Zeilinger S, Mach R, Kubicek C (1998) Two adjacent
protein binding motifs in the cbh2 (Cellobiohydrolase II-encoding) promoter of the fungus
Hypocrea jecorina (Trichoderma reesei) cooperate
in the induction by cellulose. J Biol Chem
273:34463–34471
Zhang F, Keasling J (2011) Biosensors and their applications in microbial metabolic engineering.
Trends Microbiol 19:323–329
Zhang DX, Lu HL, Liao X, St Leger RJ, Nuss DL (2013)
Simple and efficient recycling of fungal selectable
marker genes with the Cre-loxP recombination
system via anastomosis. Fungal Genet Biol 61:1–9
Zhang XH, Tee LY, Wang XG, Huang QS, Yang SH
(2015) Off-target effects in CRISPR/Cas9-mediated
genome engineering. Mol Ther Nucleic Acids
4:1–8
Zhang C, Meng X, Wei X, Lu L (2016a) Highly efficient
CRISPR mutagenesis by microhomologymediated end joining in Aspergillus fumigatus.
Fungal Genet Biol 86:47–57
Zhang H, Wang S, Zhang XX, Ji W, Song F, Zhao Y, Li J
(2016b) The amyR-deletion strain of Aspergillus
niger CICC2462 is a suitable host strain to express
secreted protein with a low background. Microb
Cell Factories 15:1–11
Zhang J, Zhang G, Wang W, Wei D (2018) Enhanced
cellulase production in Trichoderma reesei RUT
C30 via constitution of minimal transcriptional
activators. Microb Cell Factories 17:1–14
Zhao C, Waalwijk C, de Wit PJGM, Tang D, van der Lee
T (2013) RNA-Seq analysis reveals new gene models and alternative splicing in the fungal pathogen
Fusarium graminearum. BMC Genomics 14:1–16
Zheng YM, Lin FL, Gao H, Zou G, Zhang JW, Wang GQ,
Chen GD, Zhou ZH, Yao XS, Hu D (2017) Development of a versatile and conventional technique
for gene disruption in filamentous fungi based on
CRISPR-Cas9 technology. Sci Rep 7:1–10
Zhou M, Wang T, Fu J, Xiao G, Liu Y (2015) Nonoptimal codon usage influences protein structure in
intrinsically disordered regions. Mol Microbiol
97:974–987
Zhou Z, Dang Y, Zhou M, Li L, Yu C, Fu J, Chen S, Liu Y
(2016) Codon usage is an important determinant
of gene expression levels largely through its effects
on transcription. Proc Natl Acad Sci USA 113:
E6117–E6125
Zhou Z, Dang Y, Zhou M, Yuan H, Liu Y (2018) Codon
usage biases co-evolve with transcription termination machinery to suppress premature cleavage
and polyadenylation. Elife 7:1–29
Zhu H, Choi S, Johnston AK, Wing RA, Dean RA (1997)
A large-insert (130 kbp) bacterial artificial chromosome library of the rice blast fungus Magnaporthe grisea: genome analysis, contig assembly,
and gene cloning. Fungal Genet Biol 21:337–347
Zhu B, Cai G, Hall EO, Freeman GJ (2007) In-Fusion
TM
assembly: seamless engineering of multidomain
fusion proteins, modular vectors, and mutations.
BioTechniques 43:354–359
Zubieta MP, Contesini FJ, Rubio MV, Gonc ¸alves
AEDSS, Gerhardt JA, Prade RA, Damasio ARDL
(2018) Protein profile in Aspergillus nidulans
recombinant strains overproducing heterologous
enzymes. Microb Biotechnol 11:346–358
10 Filamentous Fungi as Hosts for Heterologous Production of Proteins and Secondary. . .
265
