Progress and obstacles in the production and application of recombinant lignin-degrading peroxidases

Figures & data

Figure 1. Lignin subunits attacked by enzymes and their most common reaction mechanisms. The bulky lignin polymer structure represents part of an organosolv lignin substrate.⁶⁵ Both superfamilies of lignin-degrading enzymes (heme peroxidases and laccases) can oxidize phenolic lignin subunits. In order to oxidize non-phenolic lignin subunits, laccase requires the presence of a mediator. Lignin peroxidase, versatile peroxidase and DyP-type peroxidase do not require a mediator to attack non-phenolic structures. Manganese peroxidase and versatile peroxidase oxidize phenolic lignin subunits via the oxidation of manganese (Mn²⁺ → Mn³⁺). The inset box shows heme peroxidases reducing H₂O₂ to water to catalyze the oxidation reactions, whereas laccases reduce molecular oxygen to water, which is accompanied by the oxidation of the substrates or mediators. The atoms and bonds of the phenolic and non-phenolic lignin subunits are highlighted in bold within the bulky structure. Abbreviations: med = mediator, sub = substrate.

Table 1. Recombinant fungal lignin-degrading peroxidases. Examples of successfully produced fungal lignin-degrading peroxidases and the substrates that have been used. The activity of the enzymes (if stated) is shown in parentheses. ABTS: 2,2-azinobis (3-ethylbenzothiazoline-6-sulfonate), DCP: 2,4-dichlorophenol, DFAD: 4-[(3,5-difluoro-4-hydroxyphenyl)azo]benzenesulfonic acid, DMP: 2,6-dimethoxyphenol, icb: inclusion body; MHG: methoxyhydroquinone, VA: veratryl alcohol.

enzyme	origin	host organism	localization	substrate (activity)	ref
LiP H8	Phanerochaete chrysosporium	baculovirus	extracellular	VA (20 U mg^−1)	^18
LiP H8	P. chrysosporium	P. chrysosporium	extracellular	VA (Km 89.4 µM, kcat 23.2 s^−1)	^21
LiP H8	P. chrysosporium	E. coli	cytoplasm (ib)	VA (29.7 s^−1) ABTS (27.0 s^−1) DFAD (31.1 s^−1)	^24
LiP H2	P. chrysosporium	E. coli	cytoplasm (ib)	Mn^2+ (14 s^−1) VA (39 s^−1)	^25
LiP H8	P. chrysosporium	E. coli	cytoplasm (ib)	β-0-4 tetrameric lignin model compound	^33
LiP	Trametes cervina	E. coli	cytoplasm (ib)	ABTS VA (Km 3240 µM, Kcat 17.7 s^−1)1,4-dimethoxybenzeneferrocytochrome c	^28
LiP H8	P. chrysosporium	Pichia methanolica	extracellular	VA (α-mating factor: 1933 U L^−1) VA (native leader peptide: 932 U L^−1)	^34
LiP H2 variants	P. chrysosporium	Saccharomyces cerevisiae	extracellular	DCP (78.1 – 81.8 U mg^−1) DCP (kcat 3520 – 3960 min^−1, Km 163 – 190 µM)	^37
MnP H4	P. chrysosporium	baculovirus	extracellular	phenol red (83 mU L^−1) vanillylacetone (201 mU L^−1) guaiacol (94 mU L^−1)	^19
MnP isozyme 1 (mnp1)	P. chrysosporium	P. chrysosporium	extracellular	Mn^2+ (341.3 U mg^−1) DMP (158.3 U mg^−1)	^20
MnP H4	P. chrysosporium	E. coli	cytoplasm (ib)	Mn^2+(140 U mg^−1)	^23
MnP H4 S168W	P. chrysosporium	E. coli	cytoplasm (ib)	Mn^2+ (kcat 260 s^−1; Km 0.11 mM) VA (kcat 11 s^−1; Km 0.49 mM)	^26
MnP	P. chrysosporium	Aspergillus oryzae	extracellular	Mn^2+ (0.33 U mL^−1; kcat 132 ± 15 s^−1) DMP	^40
MnP H4 (mnp1)	P. chrysosporium	Aspergillus niger	extracellular	ABTS (∼66.2 ± 12.1 ΔAbs min^−1 mL^−1)	^41
MnP (mnp1)	P. chrysosporium	Pichia pastoris	extracellular	DMP (160 U mg^−1) Kraft lignin (Kappa number: 49.2 to 42.0, Klason lignin: 7.41% to 6.75%)	^39
MnP6 S168W^*	Ceriporiopsis subvermispora	E. coli	cytoplasm (ib)	Mn^2+ (kcat 60.5 ± 5.0 s^−1; Km 8.5 ± 1.2 µM) ABTS (kcat 1.4 ± 0.1 s^−1; Km 60.2 ± 13.4 µM) VA (kcat 0.54 ± 0.04 s^−1; Km 2740 ± 560 µM) Reactive Black 5 (kcat 7.8 ± 1.1 s^−1; Km 812.6 ± 3 µM)DMP 4-O-methylsyringylglycerol-ß-guaiacyl ether (kcat 0.47 ± 0.0 s^−1; Km 1.1 ± 0.2 µM)	^29
MnP1 – MnP6	Pleurotus ostreatus	E. coli	cytoplasm (ib)	Mn^2+^** ABTS ^** VA ^** Reactive Black 5 ^** DMP ^**	^30
VP (MnPL2 -mnpI2)	Pleurotus eryngii	Aspergillus nidulans	extracellular	Mn^2+ (Km 20 µM; kcat 99 s^−1) VA (Km 1780 µM; kcat 12 s^−1) MHQ (Km 23 µM; kcat 10 s^−1) DMP	^42
versatile MnP2 (mnp2)	P. ostreatus	P. ostreatus	extracellular	Mn^2+ (Km 22.3 µM)guaiacol (7300 U L^−1) VA (Km 5080 µM)	^22
VP	Bjerkandera adusta	E. coli	cytoplasm	Mn^2+ (27.5 mU mg^−1) DMP (0.13 mU mg^−1) VA (0.25 mU mg^−1)1-napthol (70 mU mg^−1)	^31
VP (vpl2)	P. eryngii	E. coli	cytoplasm	Mn^2+ (194 ± 3.1 U mg^−1; kcat 138 ± 5.2 s^−1; Km 78 ± 7.9 µM) ABTS (8.8 ± 0.03 U mg^−1; kcat 5.4 ± 0.1 s^−1; Km 0.7 ± 0.04 µM) VA (7.2 ± 0.6 U mg^−1; kcat 6.4 ± 0.3 s^−1; Km 4090 ± 523 µM) Reactive Black 5 (2.2 ± 0.2 U mg^−1; kcat 2.7 ± 0.08 s^−1; Km 4.8 ± 0.3 µM)	^32
VP1VP2VP3	P. ostreatus	E. coli	cytoplasm (ib)	Mn^2+ ^** ABTS ^** VA ^** Reactive Black 5 ^** DMP ^** 4-ethoxy-3-methoxyphenylglycerol-β-guaiacyl ether synthetic lignin	^30
VP (vpl2)variant	P. eryngii	S. cerevisiae	extracellular	ABTS (717 U mg^−1)	^35
VP (vpl2)MnP (mnp1)LiP (lipH8)	P. eryngii P. chrysosporiumP. chrysosporium	P. chrysosporium	extracellular	Mn^2+(9.084 U mL^−1 culture) DMP (20 U mL^−1 culture) VA (∼3.8 U mL^−1)	^43
DyP	Thanatephorus cucumeris(Geotrichum candidum Dec 1)	A. oryzae	extracellular	(relative substrate specificity) Reactive Blue 5 (100 %) Reactive Blue 19 (100 %) Reactive Black 5 (0.9 %) Reactive Red 33 (1.8 %) VA (0 %)guaiacol (0.17 %) DMP (0.54 %)	^46
DyP	T. cucumeris	A. oryzae	extracellular	1-amino-2-sulfonyl-4-aminomethyl-9,10-anthraquinone sodium salt (600 U mg^−1)	^45
DyP	T. cucumeris	E. coli	periplasm	(relative substrate specificity) Reactive Blue 5 (100 %) Reactive Blue 19 (119 %) Reactive Blue 21 (26 %) Reactive Blue 114 (4.4 %) Reactive Black 5 (1.0 %) Reactive Red 33 (2.3 %) Reactive Red 120 (0.96 %) Reactive Orange 13 (0.22 %) VA (0 %) guaiacol (9.2 %) DMP (9.3 %)	^47

* see Ref. ²⁹ for kinetic constants for recombinant peroxidases from C. subvermispora (MnP6, MnP6-S168W-environment variants), P. ostreatus (VP1), and P. chrysosporium (LiP-H8).

** see Ref. ³⁰ for kinetic constants (K_m, k_cat, k_cat/K_m) for the recombinant peroxidases from the P. ostreatus genome.

Table 2. Recombinant bacterial lignin-degrading peroxidases. Examples of successfully produced recombinant bacterial lignin-degrading peroxidases and the substrates that have been used. The activity of the enzymes (if stated) is shown in parentheses. ABTS: 2,2-azinobis (3-ethylbenzothiazoline-6-sulfonate), DCP: 2,4-dichlorophenol, DMP: 2,6-dimethoxyphenol, DOPA: L-3,4-dihydroxyphenylalanine, VA: veratryl alcohol.

enzyme	origin	host organism	localization	substrate (activity)	Ref
extracellular lignin peroxidase	Streptomyces viridosporus	Pichia pastoris	extracellular	DCP (∼1.1 (AU) min^−1) DOPA	^49
DyP-type (Tfu_3078)	Thermobifida fusca	Escherichia coli	periplasm	Reactive Blue 19 (Km 29 µM, kcat 10 s^−1)	^50
TfuDyP	T. fusca	E. coli	intracellular	ABTS (Km 0.86 ± 0.07 mM, kcat 28.1 ± 1 s^−1) DCP (Km 5.51 ± 0.5 mM, kcat 2.86 ± 0.1 s^−1) guaiacolphenol (Km 0.126 ± 0.01 mM, kcat 0.136 ± 0.03 s^−1) pyrogallol (Km 11.29 ± 1 mM, kcat 6.63 ± 0.25 s^−1) Reactive Blue 4 (Km 0.179 ± 0.01 mM, kcat 1.88 ± 0.07 s^−1) guaiacylglycerol-β-guaiacyl etherKraft lignin	^51
DyP (DyP2)	Amycolatopsis sp. 75iv2	E. coli	extracellular	kcat/ Km (M^−1 s^−1)ABTS ((6.6 ± 0.9) × 10^6)Reactive Blue 5 ((7.1 ± 0.9) × 10^5)Reactive Black 5 ((1.6 ± 0.1) × 10^5)Mn^2+ ((1.2 ± 0.2) × 10^5)guaiacylglycerol-β-guaiacol ether,veratrylglycerol-β-guaiacol ether	^52
DyP-type (dyp Pa)	Pseudomonas aeruginosa	E. coli	intracellular	Reactive Blue 5 (4395 µM mg^−1 min^−1) Reactive Blue 19 (107 µM mg^−1 min^−1) Reactive Black 5 (704 µM mg^−1 min^−1) DMP (786 µM mg^−1 min^−1)	^53
DyP	Anabaena sp strain PCC7120	E. coli	intracellular	Reactive Blue 5 (Km 3.6 M, kcat/Km 1.2 × 10^7 M^−1 s^−1)	^54
PpDyPBsDyP	Pseudomonas putidaBacillus subtilis	E. coli	intracellular	(PpDyP // BsDyP) ABTS (40 ± 0.9 U mg^−1 // 15 ± 1 U mg^−1) Reactive Blue 5 (15 ± 0.2 U mg^−1 // 11 ± 0.6 U mg^−1) guaiacol (0.2 ± 0.004 U mg^−1 // 0.2 ± 0.006 U mg^−1) syringaldehyde (0.11 ± 0.005 U mg^−1 // 0.004 ± 0.0002 U mg^−1) acetosyringone (0.16 ± 0.008 U mg^−1 // 0.012 U mg^−1)	^55
BsDyP	B. subtilis	E. coli	intracellular	ABTS (66.80 U mg^−1) VA (0.13 U mg^−1) Reactive Blue 19 (11.55 U mg^−1) Reactive Black 5 (17.65 U mg^−1) veratrylglycerol-β-guaiacolether (0.086 U mg^−1)	^56
SviDyP	Saccharomonospora viridis	E. coli	intracellular	Reactive Blue 19 (1.29 U mg^−1) Reactive Green 19 (1.32 U mg^−1) Reactive Yellow 2 (4.86 U mg^−1) Reactive Black 5 (0.96 U mg^−1) Reactive Red 120 (0.69 U mg^−1) Brilliant Green (12.24 U mg^−1) Malachite Green (8.4 U mg^−1) Crystal Violet (4.11 U mg^−1) Azure B (1.62 U mg^−1) DMP (0.06 U mg^−1) VA (0.03 U mg^−1) eucalyptus kraft pulp (Kappa decrease 21.8 %)	^57
DyP1B	Pseudomonas fluorescens	E. coli	intracellular	ABTS (Km 1.13 ± 0.1 mM, kcat 13.5 ± 0.4 s^−1) DCP (Km 1.25 ± 0.1 mM, kcat 0.66 ± 0.02 s^−1) guaiacol (Km 0.056 ± 0.006 mM, kcat 0.058 ± 0.001 s^−1) pyrogallol (Km 4.0 ± 0.6 mM, kcat 2.5 ± 0.1 s^−1) Reactive Blue 4 (Km 0.12 ± 0.01 mM, kcat 1.04 ± 0.03 s^−1) Kraft lignin (Km 0.006 ± 0.001 mM, kcat 0.9 ± 0.1 s^−1)	^58
DypB	Rhodococcus jostii	E. coli	intracellular	ABTS (0.0549 ΔAbs min^−1) β-aryl ether lignin model Kraft lignin (0.24 ΔAbs 10 min^−1) nitrated lignin (0.0081 ΔAbs 20 min^−1) wheat straw lignocellulose	^59
DyPA	R. jostii	E. coli	intracellular	ABTS (Km 8.2 mM, kcat 16.83 s^−1)	^60
peroxidase	R. jostii	R. jostii	extracellular	wheat straw lignocellulose (96 mg L^−1 vanillin53 mg L^−1 p-hydroxybenzaldehyde 3-120 mg L^−1 vanillic acid 23-86 mg L^−1 ferulic acid)	^61

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