Current understanding of substrate specificity and regioselectivity of LPMOs

Zhou, Xiaoli; Zhu, Honghui

doi:10.1186/s40643-020-0300-6

Bioresources and Bioprocessing

Table 1 Overview of characterized LPMOs

From: Current understanding of substrate specificity and regioselectivity of LPMOs

Family	Organism	Protein names	Associated CBMs	PDB code	Substrates	Regioselectivities	References
AA9	Aspergillus nidulans	AN1602	CBM1	–	PASC Cellohexaose	C4	(Jagadeeswaran et al. 2018; Jagadeeswaran et al. 2016)
	Aspergillus nidulans	AN3046	–	–	PASC Xyloglucan	C1	(Jagadeeswaran et al. 2016)
	Gloeophyllum trabeum NBRC 6430	LPMO9A-2 GtLPMO9A-2	C-terminal domain with unknown function	–	PASC CMC Xyloglucan Glucomannan	C1/C4	(Kojima et al. 2016)
	Gloeophyllum trabeum KUC 8013	Cel61G GtGH61 GtLPMO9B LPMO9B	–	–	PASC Xyloglucan	C1/C4	(Hegnar et al. 2019)
	Heterobasidion irregulare TC 32-1	HiLPMO9B	–	5NNS	PASC	C1	(Liu et al. 2018)
		HiLPMO9H	CBM1	–	PASC	C1	(Liu et al. 2017)
		HiLPMO9I	CBM1	–	PASC Glucomannan	C4	(Liu et al. 2017)
	Lentinus similis	LsAA9A	–	5ACF 5ACG 5ACH 5ACI 5ACJ 5N04 5N05 5NKW 5NLN 5NLO 5NLP 5NLQ 5NLR 5NLS	PASC Cello-oligosaccharides	C1/C4(PASC) C4 (Cello-oligosaccharides)	(Frandsen et al. 2016)
	Neurospora crassa OR74A	PMO-2 NcPMO-2 NcLPMO9D GH61-4 NCU01050 LPMO9D	–	4EIR 5TKF 5TKG 5TKH 5TKI	PASC Xyloglucan Glucomannan	C4	(Li et al. 2012; Petrovic et al. 2019)
	Neurospora crassa OR74A	PMO-03328 LPMO-03328 NcLPMO9F GH61-6 NCU03328 LPMO9F	–	4QI8	Microcrystalline cellulose	C1	(Kittl et al. 2012) (Tan et al. 2015)
	Neurospora crassa OR74A	PMO-01867 LPMO-01867 NcLPMO9J GH61-10 NCU01867 B13N4.070 LPMO9J	CBM1	–	Microcrystalline cellulose	C1	(Kittl et al. 2012)
	Neurospora crassa OR74A	PMO-3 NcLPMO9M GH61-13 NcPMO-3 NCU07898 LPMO9M	–	4EIS	PASC	C1/C4	(Li et al. 2012; Vu et al. 2014a)
	Neurospora crassa OR74A	PMO-02916 LPMO-02916 NcLPMO9C GH61-3 NCU02916 LPMO9C	CBM1	4D7U 4D7V	PASC Avicel Steam-exploded spruce Cello-oligosaccharides Xyloglucan β-(1→3,1→4)-D-Glucan Glucomannan	C4	(Agger et al. 2014; Borisova et al. 2015; Courtade et al. 2016; Isaksen et al. 2014; Karnaouri et al. 2017; Kittl et al. 2012; Kojima et al. 2016; Kracher et al. 2018; Nekiunaite et al. 2016b; Varnai et al. 2018; Westereng et al. 2016)
	Neurospora crassa OR74A	GH61-2 NCU07760	CBM1	–	PASC	C1/C4	(Vu et al. 2014a)
	Neurospora crassa OR74A	GH61-1 NcLPMO9A NCU02240	CBM1	5FOH	PASC	C4	(Petrovic et al. 2019; Vu et al. 2014a)
	Neurospora crassa OR74A	NCU00836	CBM1	–	PASC	C1	(Vu et al. 2014a)
	Neurospora crassa OR74A	PMO-08760 LPMO-08760 NcLPMO9E GH61-5 NCU08760 LPMO9E	CBM1	–	PASC	C1	(Kracher et al. 2016; Vu et al. 2014a)
	Pestalotiopsis sp. NCi6	PsLPMOB	–	–	PASC	C4	(Patel et al. 2016)
	Pestalotiopsis sp. NCi6	PsLPMOA	–	–	PASC	C1/C4	(Patel et al. 2016)
	Phanerochaete chrysosporium K-3	GH61D PcGH61D PcLPMO9D	–	4B5Q	PASC Avicel	C1	(Danneels et al. 2019; Danneels et al. 2017; Vuong et al. 2017; Westereng et al. 2013; Westereng et al. 2011; Wu et al. 2013)
	Podospora anserina S mat+	Pa_4_1020 PaLPMO9H	CBM1	–	PASC Gluco-oligosaccharides Xyloglucan Glucomannan Lichenan β-(1→3,1→4)-d-Glucan CMC	C1/C4	(Bennati-Granier et al. 2015; Chalak et al. 2019; Fanuel et al. 2017; Garajova et al. 2016; Villares et al. 2017)
	Podospora anserina S mat+	Pa_4_7570 PaLPMO9D	–	–	PASC	C1	(Bennati-Granier et al. 2015)
	Podospora anserina S mat+	Pa_1_16300 PaLPMO9E	CBM1	–	PASC	C1	(Bennati-Granier et al. 2015; Chabbert et al. 2017; Garajova et al. 2016)
	Podospora anserina S mat+	Gh61B Pa_7_3160 PaLPMO9B Gh61B	CBM1	–	PASC	C1/C4	(Bey et al. 2013)
	Podospora anserina S mat+	Pa_6_7780 PaLPMO9F	–	–	PASC	C1	(Bennati-Granier et al. 2015)
	Podospora anserina S mat+	Gh61A Pa_2_6530 PaLPMO9A GH61A	CBM1	–	PASC	C1/C4	(Bennati-Granier et al. 2015; Bey et al. 2013)
	Thermoascus aurantiacus	TaAA9 TaAA9A TaGH61 TaGH61A TaLPMO9A	–	2YET 3ZUD	PASC	C1/C4	(Cannella et al. 2016; Harris et al. 2010; Kitaoku et al. 2018; Muller et al. 2015; Petrovic et al. 2018; Quinlan et al. 2011; Singh et al. 2019)
	Myceliophthora thermophila/Thermothelomyces thermophilus	MtLPMO9J MYCTH_79765	–	–	PASC Cello-oligosaccharides Xyloglucan	C4	(Kadowaki et al. 2018)
	Myceliophthora thermophila/Thermothelomyces thermophilus	MtPMO3 MYCTH_92668 MtLPMO9D	–	5UFV	RAC PASC	C1	(Frommhagen et al. 2018; Span et al. 2017; Vu et al. 2014a)
	Myceliophthora thermophila/Thermothelomyces thermophilus	MYCTH_112089 MYCTH112089	–	–	PASC	C1	(Vu et al. 2014a)
	Myceliophthora thermophila/Thermothelomyces thermophilus	MtLPMO9A LPMO9A	–	–	RAC PASC Xyloglucan Xylan β-(1→3,1→4)-d-Glucan	C1/C4	(Frommhagen et al. 2016; Frommhagen et al. 2015; Gusakov et al. 2017)
	Myceliophthora thermophila/Thermothelomyces thermophilus	MYCTH_103537 MtLPMO9L	–	–	PASC Avicel	C1	(Zhou et al. 2019a)
	Myceliophthora thermophila/Thermothelomyces thermophilus	MtLPMO9B	–	–	RAC	C1	(Frommhagen et al. 2016; Frommhagen et al. 2017a; Frommhagen et al. 2018)
	Myceliophthora thermophila/Thermothelomyces thermophilus	MtLPMO9C	–	–	RAC β-(1→3,1→4)-d-Glucan Xyloglucan	C4	(Frommhagen et al. 2016; Frommhagen et al. 2017b)
	Myceliophthora thermophila/Thermothelomyces thermophilus	MtLPMO9	CBM1	–	PASC	C1/C4	(Karnaouri et al. 2017)
	Trichoderma reesei	LPMO9A HjLPMO9A TrCel61A	CBM1	5O2W 5O2X	PASC	C1/C4	(Hansson et al. 2017; Tanghe et al. 2015)
	Trichoderma reesei	HjLPMO9B HjGH61B Cel61B	–	2VTC			(Karkehabadi et al. 2008)
	Collariella virescens	CvAA9A	–	5NLT	PASC Gluco-oligosaccharides Xyloglucan β-(1→3,1→4)-D-Glucan Glucomannan Mannohexaose Xylohexaose	C1/C4	(Simmons et al. 2017)
	Aspergillus fumigatus	AfAA9B	–	5X6A 6H1Z 6HA5 6HAQ	–	–	(Lo Leggio et al. 2018)
	Fusarium graminearum	FgLPMO9A	–	–	PASC Xyloglucan	C1/C4	(Nekiunaite et al. 2016b)
	Geotrichum candidum	GcLPMO9A	–	–	PASC Xyloglucan	C1/C4	(Ladeveze et al. 2017)
	Geotrichum candidum	GcLPMO9B	–	–	PASC Xyloglucan	C1/C4	(Ladeveze et al. 2017)
	Malbranchea cinnamomea	McAA9A	–	–	PASC Xyloglucan Glucomannan Cellohexaose	C1/C4 C4 (Xyloglucan)	(Huttner et al. 2019)
	Malbranchea cinnamomea	McAA9B	–	–	PASC Xyloglucan	C1/C4 C4 (Xyloglucan)	(Huttner et al. 2019)
	Malbranchea cinnamomea	McAA9F	–	–	PASC Xyloglucan Cellohexaose	C1/C4 C4 (Xyloglucan)	(Huttner et al. 2019)A
	Malbranchea cinnamomea	McAA9H	–	–	PASC Xylan	C1 (PASC) C1/C4 (Xylan)	(Huttner et al. 2019)
	Thielavia terrestris	TtLPMO9E		–	PASC Avicel	C1 C1/C4(Cannella et al. 2016; Mollers et al. 2017)	(Cannella et al. 2016; Gusakov et al. 2017; Kim et al. 2017; Mollers et al. 2017; Westereng et al. 2015)
	Chaetomium thermophilum	CtPMO1	–	–	PASC Celloheptaose	C1/C4	(Chen et al. 2018)
AA10	Bacillus amyloliquefaciens	ChbB BaAA10A BaCBM33 Rbam17540 BAMF_1859	–	2YOW 2YOX 2YOY 5IJU	α and β chitin	C1	(Gregory et al. 2016; Hemsworth et al. 2013b)
	Bacillus licheniformis	ChbB Cbp BliCBP BlAA10A BLi00521 BL00145 BlLPMO10A	–	5LW4	α and β chitin	C1	(Courtade et al. 2015; Forsberg et al. 2014b)
	Bacillus thuringiensis ACCC10066	Lpmo10A BtLPMO10A	–	5WSZ	α and β chitin	C1	(Zhang et al. 2015)
	Bacillus thuringiensis ATCC33679	Cbp BtCBP BtLPMO10A-FL	CBM5 Fibronectin-type III-like domains	–	β Chitin	C1	(Manjeet et al. 2019; Manjeet et al. 2013)
	Cellvibrio japonicus Ueda107	CjLPMO10A CJA_2191	CBM5 CBM73	5FJQ	α and β chitin	C1	(Forsberg et al. 2016)
	Cellvibrio japonicus Ueda107	CjLPMO10B CJA_3139	CBM10	–	PASC Avicel BMCC Filter paper	C1	(Gardner et al. 2014)
	Enterococcus faecalis V583	EfAA10A EF0362 EfCBM33A EfaCBM33	–	4A02 4ALC 4ALE 4ALQ 4ALR 4ALS 4ALT	α and β chitin	C1	(Gudmundsson et al. 2014; Vaaje-Kolstad et al. 2012)
	Hahella chejuensis KCTC 2396	HcAA10-2 HCH_00807	CBM2	–	Αvicel	C1	(Ghatge et al. 2015)
	Jonesia denitrificans DSM 20603	Jden_1381 JdLPMO10A LPMO10A	CBM5 GH18	5AA7 5VG0 5VG1	α and β chitin	C1	(Bacik et al. 2017; Mekasha et al. 2016)
	Listeria monocytogenes	GbpA LmLPMO10 Lmo2467	CBM5 Fibronectin-type III-like domain	5L2V	α and β chitin	C1	(Paspaliari et al. 2015)
	Serratia marcescens BJL200	Cbp21 CBP21 Cbp SmAA10A SmLPMO10A	–	2BEM 2BEN 2LHS	α and β chitin	C1	(Aachmann et al. 2012; Vaaje-Kolstad et al. 2005a; Vaaje-Kolstad et al. 2005b; Vaaje-Kolstad2010)
	Streptomyces ambofaciens ATCC 23877	SAML0570 SamLPMO10B	–	–	β Chitin	C1	(Valenzuela et al. 2017)
	Streptomyces ambofaciens ATCC 23877	SAML1174 SamLPMO10C	CBM2	–	PASC Flax pulp fibers	C1	(Valenzuela et al. 2017)
	Streptomyces coelicolor A3(2)	ScLPMO10B SCO0643 SCF91.03c	–	4OY6 4OY8	PASC Avicel β Chitin	C1/C4	(Forsberg et al. 2014a)
	Streptomyces coelicolor A3(2)	ScLPMO10C CelS2 ScAA10C SCO1188 SCG11A.19	CBM2	4OY7	PASC Avicel	C1	(Courtade et al. 2018; Forsberg et al. 2014a; Forsberg et al. 2011)
	Streptomyces griseus subsp. griseus NBRC 13350	SGR_6855 SgLPMO10F	–	–	α and β chitin	C1	(Nakagawa et al. 2015)
	Streptomyces lividans 1326	SliLPMO10E SLI_3182	–	5FTZ	β Chitin	C1	(Chaplin et al. 2016)
	Teredinibacter turnerae T7901	TtAA10A TERTU_0046	CBM10	6RW7	Avicel	C1/C4	(Fowler et al. 2019)
	Thermobifida fusca YX	Tfu_1268 E7 TfLPMO10A	–	4GBO	β Chitin PASC Avicel	C1/C4	(Forsberg et al. 2014a; Kruer-Zerhusen et al. 2017; Russo et al. 2019)
	Thermobifida fusca YX	E8 TfAA10B Tfu_1665	CBM2 FN3	–	PASC Avicel	C1	(Forsberg et al. 2014a; Kruer-Zerhusen et al. 2017)
	Vibrio cholerae O1 biovar El Tor str. N16961	GbpA VcGbpA VcAA10B VCA0811	GbpAD2 GbpAD3 GbpAD4	2XWX	β Chitin	C1	(Loose et al. 2014; Wong et al. 2012)
	Tectaria macrodonta	Tma12	–	6IF7	Colloidal crab chitin	–	(Shukla et al. 2016; Yadav et al. 2019)
	Anomala cuprea entomopoxvirus CV6M	Fusolin ACV034	–	4YN1 4YN2 4X29 4X27 4OW5	–	–	(Chiu et al. 2015)
	Micromonospora aurantiaca	MaLPMO10B	CBM2	5OPF	PASC β Chitin	C1/C4 C1 (chitin)	(Forsberg et al. 2018)
	Micromonospora aurantiaca	MaLPMO10D	CBM2	–	PASC β Chitin	C1/C4 C1 (chitin)	(Forsberg et al. 2018)
	Bacillus cereus	BcLPMO10A	CBM5 Fibronectin-type III-like domain	–	α and β chitin	C1	(Mutahir et al. 2018)
AA11	Aspergillus oryzae RIB40	AoLpmo11	X278	4MAH 4MAI	β Chitin	C1	(Hemsworth et al. 2014)
	Fusarium fujikuroi	FfAA11	X278	–	α and β chitin	C1	(Wang et al. 2018)
AA13	Aspergillus nidulans FGSC A4	AnAA13 AN5463.2	CBM20	–	Retrograded starch	C1	(Lo Leggio et al. 2015)
	Aspergillus oryzae RIB40	AoAA13	–	4OPB 5LSV 5T7J 5T7N	–	–	(Frandsen et al. 2017; Lo Leggio et al. 2015)
	Aspergillus terreus NIH2624	AtLPMO13A ATEG_07286	CBM20	–	Wheat starch	–	(Nekiunaite et al. 2016a)
	Magnaporthe oryzae	MoLPMO13A	CBM20	–	Binding to wheat starch	–	(Nekiunaite et al. 2016a)
	Neurospora crassa OR74A	NcAA13 NCU08746	CBM20	–	Amylose Amylopectin Cornstarch	C1	(Vu et al. 2014b; Vu et al. 2019)
	Myceliophthora thermophila	MtAA13	–	–	Amylose Amylopectin Cornstarch		(Vu et al. 2019)
AA14	Trametes coccinea CIRM-BRFM 310	PcAA14A	–	–	Xylan	C1	(Couturier et al. 2018)
	Trametes coccinea CIRM-BRFM 310	PcAA14B	–	5NO7	Xylan	C1	(Couturier et al. 2018)
AA15	Thermobia domestica	TdAA15A	–	5MSZ	β Chitin PASC	C1	(Sabbadin et al. 2018)
	Thermobia domestica	TdAA15B	–	–	α and β chitin	C1	(Sabbadin et al. 2018)
AA16	Aspergillus aculeatus ATCC 16872	ASPACDRAFT_74022 AaAA16	–	–	PASC	C1	(Filiatrault-Chastel et al. 2019)

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