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Table 1 Overview of characterized LPMOs

From: Current understanding of substrate specificity and regioselectivity of LPMOs

FamilyOrganismProtein namesAssociated CBMsPDB codeSubstratesRegioselectivitiesReferences
AA9Aspergillus nidulansAN1602CBM1PASC
Cellohexaose
C4(Jagadeeswaran et al. 2018; Jagadeeswaran et al. 2016)
 Aspergillus nidulansAN3046PASC
Xyloglucan
C1(Jagadeeswaran et al. 2016)
 Gloeophyllum trabeum NBRC 6430LPMO9A-2
GtLPMO9A-2
C-terminal domain with unknown functionPASC
CMC
Xyloglucan
Glucomannan
C1/C4(Kojima et al. 2016)
 Gloeophyllum trabeum KUC 8013Cel61G
GtGH61
GtLPMO9B LPMO9B
PASC
Xyloglucan
C1/C4(Hegnar et al. 2019)
 Heterobasidion irregulare TC 32-1HiLPMO9B5NNSPASCC1(Liu et al. 2018)
  HiLPMO9HCBM1PASCC1(Liu et al. 2017)
  HiLPMO9ICBM1PASC
Glucomannan
C4(Liu et al. 2017)
 Lentinus similisLsAA9A5ACF
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 OR74APMO-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 OR74APMO-03328
LPMO-03328 NcLPMO9F
GH61-6
NCU03328
LPMO9F
4QI8Microcrystalline celluloseC1(Kittl et al. 2012)
(Tan et al. 2015)
 Neurospora crassa OR74APMO-01867
LPMO-01867 NcLPMO9J
GH61-10
NCU01867
B13N4.070
LPMO9J
CBM1Microcrystalline celluloseC1(Kittl et al. 2012)
 Neurospora crassa OR74APMO-3
NcLPMO9M
GH61-13
NcPMO-3
NCU07898
LPMO9M
4EISPASCC1/C4(Li et al. 2012; Vu et al. 2014a)
 Neurospora crassa OR74APMO-02916
LPMO-02916 NcLPMO9C
GH61-3
NCU02916
LPMO9C
CBM14D7U
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 OR74AGH61-2
NCU07760
CBM1PASCC1/C4(Vu et al. 2014a)
 Neurospora crassa OR74AGH61-1
NcLPMO9A NCU02240
CBM15FOHPASCC4(Petrovic et al. 2019; Vu et al. 2014a)
 Neurospora crassa OR74ANCU00836CBM1PASCC1(Vu et al. 2014a)
 Neurospora crassa OR74APMO-08760
LPMO-08760 NcLPMO9E
GH61-5
NCU08760
LPMO9E
CBM1PASCC1(Kracher et al. 2016; Vu et al. 2014a)
 Pestalotiopsis sp. NCi6PsLPMOBPASCC4(Patel et al. 2016)
 Pestalotiopsis sp. NCi6PsLPMOAPASCC1/C4(Patel et al. 2016)
 Phanerochaete chrysosporium K-3GH61D
PcGH61D
PcLPMO9D
4B5QPASC
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 PaLPMO9HCBM1PASC
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 PaLPMO9DPASCC1(Bennati-Granier et al. 2015)
 Podospora anserina S mat+Pa_1_16300 PaLPMO9ECBM1PASCC1(Bennati-Granier et al. 2015; Chabbert et al. 2017; Garajova et al. 2016)
 Podospora anserina S mat+Gh61B
Pa_7_3160 PaLPMO9B
Gh61B
CBM1PASCC1/C4(Bey et al. 2013)
 Podospora anserina S mat+Pa_6_7780
PaLPMO9F
PASCC1(Bennati-Granier et al. 2015)
 Podospora anserina S mat+Gh61A
Pa_2_6530 PaLPMO9A
GH61A
CBM1PASCC1/C4(Bennati-Granier et al. 2015; Bey et al. 2013)
 Thermoascus aurantiacusTaAA9
TaAA9A
TaGH61
TaGH61A
TaLPMO9A
2YET
3ZUD
PASCC1/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 thermophilusMtLPMO9J MYCTH_79765PASC
Cello-oligosaccharides
Xyloglucan
C4(Kadowaki et al. 2018)
 Myceliophthora thermophila/Thermothelomyces thermophilusMtPMO3 MYCTH_92668
MtLPMO9D
5UFVRAC
PASC
C1(Frommhagen et al. 2018; Span et al. 2017; Vu et al. 2014a)
 Myceliophthora thermophila/Thermothelomyces thermophilusMYCTH_112089 MYCTH112089PASCC1(Vu et al. 2014a)
 Myceliophthora thermophila/Thermothelomyces thermophilusMtLPMO9A
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 thermophilusMYCTH_103537
MtLPMO9L
PASC
Avicel
C1(Zhou et al. 2019a)
 Myceliophthora thermophila/Thermothelomyces thermophilusMtLPMO9BRACC1(Frommhagen et al. 2016; Frommhagen et al. 2017a; Frommhagen et al. 2018)
 Myceliophthora thermophila/Thermothelomyces thermophilusMtLPMO9CRAC
β-(1→3,1→4)-d-Glucan
Xyloglucan
C4(Frommhagen et al. 2016; Frommhagen et al. 2017b)
 Myceliophthora thermophila/Thermothelomyces thermophilusMtLPMO9CBM1PASCC1/C4(Karnaouri et al. 2017)
 Trichoderma reeseiLPMO9A
HjLPMO9A
TrCel61A
CBM15O2W
5O2X
PASCC1/C4(Hansson et al. 2017; Tanghe et al. 2015)
 Trichoderma reeseiHjLPMO9B
HjGH61B
Cel61B
2VTC  (Karkehabadi et al. 2008)
 Collariella virescensCvAA9A5NLTPASC
Gluco-oligosaccharides
Xyloglucan
β-(1→3,1→4)-D-Glucan
Glucomannan
Mannohexaose
Xylohexaose
C1/C4(Simmons et al. 2017)
 Aspergillus fumigatusAfAA9B5X6A
6H1Z
6HA5
6HAQ
(Lo Leggio et al. 2018)
 Fusarium graminearumFgLPMO9APASC
Xyloglucan
C1/C4(Nekiunaite et al. 2016b)
 Geotrichum candidumGcLPMO9APASC
Xyloglucan
C1/C4(Ladeveze et al. 2017)
 Geotrichum candidumGcLPMO9BPASC
Xyloglucan
C1/C4(Ladeveze et al. 2017)
 Malbranchea cinnamomeaMcAA9APASC
Xyloglucan
Glucomannan
Cellohexaose
C1/C4
C4 (Xyloglucan)
(Huttner et al. 2019)
 Malbranchea cinnamomeaMcAA9BPASC
Xyloglucan
C1/C4
C4 (Xyloglucan)
(Huttner et al. 2019)
 Malbranchea cinnamomeaMcAA9FPASC
Xyloglucan
Cellohexaose
C1/C4
C4 (Xyloglucan)
(Huttner et al. 2019)A
 Malbranchea cinnamomeaMcAA9HPASC
Xylan
C1 (PASC)
C1/C4 (Xylan)
(Huttner et al. 2019)
 Thielavia terrestrisTtLPMO9E 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 thermophilumCtPMO1PASC
Celloheptaose
C1/C4(Chen et al. 2018)
AA10Bacillus amyloliquefaciensChbB
BaAA10A
BaCBM33
Rbam17540
BAMF_1859
2YOW
2YOX
2YOY
5IJU
α and β chitinC1(Gregory et al. 2016; Hemsworth et al. 2013b)
 Bacillus licheniformisChbB
Cbp
BliCBP
BlAA10A
BLi00521
BL00145
BlLPMO10A
5LW4α and β chitinC1(Courtade et al. 2015; Forsberg et al. 2014b)
 Bacillus thuringiensis ACCC10066Lpmo10A
BtLPMO10A
5WSZα and β chitinC1(Zhang et al. 2015)
 Bacillus thuringiensis ATCC33679Cbp
BtCBP
BtLPMO10A-FL
CBM5
Fibronectin-type III-like domains
β ChitinC1(Manjeet et al. 2019; Manjeet et al. 2013)
 Cellvibrio japonicus Ueda107CjLPMO10A CJA_2191CBM5
CBM73
5FJQα and β chitinC1(Forsberg et al. 2016)
 Cellvibrio japonicus Ueda107CjLPMO10B
CJA_3139
CBM10PASC
Avicel
BMCC
Filter paper
C1(Gardner et al. 2014)
 Enterococcus faecalis V583EfAA10A
EF0362
EfCBM33A
EfaCBM33
4A02
4ALC
4ALE
4ALQ
4ALR
4ALS
4ALT
α and β chitinC1(Gudmundsson et al. 2014; Vaaje-Kolstad et al. 2012)
 Hahella chejuensis KCTC 2396HcAA10-2
HCH_00807
CBM2ΑvicelC1(Ghatge et al. 2015)
 Jonesia denitrificans DSM 20603Jden_1381
JdLPMO10A LPMO10A
CBM5
GH18
5AA7
5VG0
5VG1
α and β chitinC1(Bacik et al. 2017; Mekasha et al. 2016)
 Listeria monocytogenesGbpA
LmLPMO10
Lmo2467
CBM5
Fibronectin-type III-like domain
5L2Vα and β chitinC1(Paspaliari et al. 2015)
 Serratia marcescens BJL200Cbp21
CBP21
Cbp
SmAA10A
SmLPMO10A
2BEM
2BEN
2LHS
α and β chitinC1(Aachmann et al. 2012; Vaaje-Kolstad et al. 2005a; Vaaje-Kolstad et al. 2005b; Vaaje-Kolstad2010)
 Streptomyces ambofaciens ATCC 23877SAML0570
SamLPMO10B
β ChitinC1(Valenzuela et al. 2017)
 Streptomyces ambofaciens ATCC 23877SAML1174
SamLPMO10C
CBM2PASC
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
CBM24OY7PASC
Avicel
C1(Courtade et al. 2018; Forsberg et al. 2014a; Forsberg et al. 2011)
 Streptomyces griseus subsp. griseus NBRC 13350SGR_6855
SgLPMO10F
α and β chitinC1(Nakagawa et al. 2015)
 Streptomyces lividans 1326SliLPMO10E SLI_31825FTZβ ChitinC1(Chaplin et al. 2016)
 Teredinibacter turnerae T7901TtAA10A TERTU_0046CBM106RW7AvicelC1/C4(Fowler et al. 2019)
 Thermobifida fusca YXTfu_1268
E7
TfLPMO10A
4GBOβ Chitin
PASC
Avicel
C1/C4(Forsberg et al. 2014a; Kruer-Zerhusen et al. 2017; Russo et al. 2019)
 Thermobifida fusca YXE8
TfAA10B
Tfu_1665
CBM2
FN3
PASC
Avicel
C1(Forsberg et al. 2014a; Kruer-Zerhusen et al. 2017)
 Vibrio cholerae O1 biovar El Tor str. N16961GbpA
VcGbpA
VcAA10B
VCA0811
GbpAD2
GbpAD3
GbpAD4
2XWXβ ChitinC1(Loose et al. 2014; Wong et al. 2012)
 Tectaria macrodontaTma126IF7Colloidal crab chitin(Shukla et al. 2016; Yadav et al. 2019)
 Anomala cuprea entomopoxvirus CV6MFusolin
ACV034
4YN1
4YN2
4X29
4X27
4OW5
(Chiu et al. 2015)
 Micromonospora aurantiacaMaLPMO10BCBM25OPFPASC
β Chitin
C1/C4
C1 (chitin)
(Forsberg et al. 2018)
 Micromonospora aurantiacaMaLPMO10DCBM2PASC
β Chitin
C1/C4
C1 (chitin)
(Forsberg et al. 2018)
 Bacillus cereusBcLPMO10ACBM5
Fibronectin-type III-like domain
α and β chitinC1(Mutahir et al. 2018)
AA11Aspergillus oryzae RIB40AoLpmo11X2784MAH
4MAI
β ChitinC1(Hemsworth et al. 2014)
 Fusarium fujikuroiFfAA11X278α and β chitinC1(Wang et al. 2018)
AA13Aspergillus nidulans FGSC A4AnAA13
AN5463.2
CBM20Retrograded starchC1(Lo Leggio et al. 2015)
 Aspergillus oryzae RIB40AoAA134OPB
5LSV
5T7J
5T7N
(Frandsen et al. 2017; Lo Leggio et al. 2015)
 Aspergillus terreus NIH2624AtLPMO13A
ATEG_07286
CBM20Wheat starch(Nekiunaite et al. 2016a)
 Magnaporthe oryzaeMoLPMO13ACBM20Binding to wheat starch(Nekiunaite et al. 2016a)
 Neurospora crassa OR74ANcAA13
NCU08746
CBM20Amylose
Amylopectin
Cornstarch
C1(Vu et al. 2014b; Vu et al. 2019)
 Myceliophthora thermophilaMtAA13Amylose
Amylopectin
Cornstarch
 (Vu et al. 2019)
AA14Trametes coccinea CIRM-BRFM 310PcAA14AXylanC1(Couturier et al. 2018)
 Trametes coccinea CIRM-BRFM 310PcAA14B5NO7XylanC1(Couturier et al. 2018)
AA15Thermobia domesticaTdAA15A5MSZβ Chitin
PASC
C1(Sabbadin et al. 2018)
 Thermobia domesticaTdAA15Bα and β chitinC1(Sabbadin et al. 2018)
AA16Aspergillus aculeatus ATCC 16872ASPACDRAFT_74022
AaAA16
PASCC1(Filiatrault-Chastel et al. 2019)