. 2019 Jun 26;24(13):2362. doi: 10.3390/molecules24132362

Table 1.

Enzymes detoxifying mycotoxins and their properties. Molar weight (MW) of enzymes corresponds to the value issued in the article or presented in the UniProt database. Optimal or used conditions for determination of enzyme activity are listed. Catalytic characteristics of enzymes are shown towards specified mycotoxin, if not stated otherwise. Tolerable daily intake (TDI)* of mycotoxins is listed for referential purpose and could vary widely depending on the local legislations.

Enzyme (MW)	Source, Ref.	Conditions	Catalytic Characteristics	Comments
Aflatoxins (aflatoxin B₁, TDI = 0.11–0.19 ng/d/kg)
AKR7A1 with N-His₆ (37 kDa)	Rattusnorvegicus [19]	pH 6.6, 25 °C, 0.2 mM NADPH	K_m = 21 μM, V_max = 0.34 μM/min/mg, 6.4 mg/h/mg	–
AKR7A3 with N-His₆ (37 kDa)	Homosapiens [19]	pH 7.4, 25 °C, 0.2 mM NADPH	K_m = 9 μM, V_max = 0.6 μM/min/mg, 11.2 mg/h/mg	–
AKR7A5 (38 kDa)	Musmusculus [20,21]	pH 6.6, 25 °C, 0.2 mM NADPH	K_m = 90 μM, k_cat = 34 min⁻¹, 17 mg/h/mg	K_m = 1.6 μM (NADPH) Inhibition: valproic acid, ethacrynic acid, quercitin, indomethacin
Laccase (56 kDa)	Trametesversicolor [22]	pH 4.5, 35 °C	K_m = 0.28 mM, 0.37 μg/h/mg	–
BADE (22 kDa)	Bacillusshackletonii [23]	pH 7–8, 37 °C	0.05 μg/h/mg	Activation: Cu²⁺ Inhibition: Mn²⁺ < Li⁺ ≈ Zn²⁺ < Mg²⁺
MnP (18–42 kDa)	Pleurotusostreatus [24]	pH 4–5, 25 °C, 50 μM H₂O₂, 50 μM Mn²⁺	up to 2 mg/h/mg	Inhibition: Cd²⁺, Hg²⁺; ≥1 mM H₂O₂, Cu²⁺
Peroxidase (32–39 kDa)	Armoraciarusticana [25]	pH 7.5, 30 °C, 2.4 mM H₂O₂	K_m = 16 nM, V_max = 6.4 μM/min, 3–15 μg/h/mg **	–
AFO (77 kDa)	Armillariella tabescens [26,27]	pH 5.8–6.0, 30 °C, Cu²⁺	K_m = 0.334 μM, k_cat = 2.7 min⁻¹, 0.66 mg/h/mg	–
ADTZ (52 kDa)	Armillariella tabescens [28,29]	pH 5–8, 35 °C	K_m~ 3 μM **, 0.13 mg/h/mg	Activation: Ba²⁺ Inhibition: Ni²⁺ < Fe^2+/3+ ≈ Cu²⁺ < Mn²⁺ < Cr³⁺ < Co²⁺ < Zn²⁺ ≈ EDTA
MADE (32 kDa)	Myxococcusfulvus [30]	pH 6.0, 35 °C, Mg²⁺	12 μg/h/mg	Activation: Mg²⁺ Inhibition: Li⁺ < Cu²⁺ < Zn²⁺
F₄₂₀H₂-dependent reductases with N-His₆ (14–21 kDa)	Actinomycetales [31]	pH 7.5, 22 °C, 10 μM F₄₂₀	up to K_m = 47 μM, k_cat = 63 min⁻¹, 11 mg/h/mg	K_m = 0.1–0.3 μM (F₄₂₀H₂)
CYP1A2 with N-His₆ (59 kDa)	Homosapiens [32]	pH 7.4, 37 °C, 1 mM NADPH	K_m = 30 μM, k_cat = 0.24 min⁻¹, 0.08 mg/h/mg	–
Sterigmatocystin (TDI = 16 ng/d/kg)
AFO (77 kDa)	Armillariella tabescens [27]	pH 5.8–6.0, 30 °C, Cu²⁺	K_m = 0.106 μM, k_cat = 1.7 min⁻¹, 0.44 mg/h/mg	–
Zearalenone (TDI = 0.5 μg/d/kg)
ZHD (29 kDa)	Clonostachysrosea [33]	pH 10.5, 30 °C	K_m = 34 μM, k_cat = 0.51 s⁻¹, 20 mg/h/mg	Inhibition: PMSF, AEBSF
ZHD with N-eGFP (56 kDa)	Clonostachysrosea [33]	pH 9.5, 30 °C	K_m = 10 μM, k_cat = 0.53 s⁻¹, 11 mg/h/mg	–
CbZHD with N-His₆ (30 kDa)	Cladophialophora bantiana [34]	pH 8.0, 35 °C	13 mg/h/mg	–
Zhd518 with N-His₆ (29 kDa)	Rhinocladiella mackenziei [35]	pH 8.0, 40 °C	12.4 mg/h/mg	Activation: Ca²⁺ Inhibition: Li⁺ < Mn²⁺ = Ni²⁺ < Co²⁺ < Cu²⁺ < Hg²⁺
ZENC (30 kDa)	Neurosporacrassa [36]	pH 8.0, 45 °C; Na⁺, Ca²⁺, or Mg²⁺	K_m = 39 μM, 31.8 mg/h/mg	Inhibition: Zn²⁺ < Mn²⁺ < EDTA < SDS ≈ Cu²⁺
Peroxiredoxin (21 kDa)	Acinetobacter sp., Saccharomyces cerevisiae [37,38]	pH 9.0, 70 °C, 20 mM H₂O₂	K_m = 7.55 mM, 0.14 mg/h/mg	–
Peroxidase (32–36 kDa)	Armoraciarusticana, Oryza sativa [39]	pH 5–6, 25 °C, 2.4 mM H₂O₂	K_m = 9–40 μM, V_max = 11–170 nM/min, up to 1.5 mg/h/mg	–
HvUGT14077 (97 kDa)	Hordeumvulgare [40]	pH 7.5, 37 °C, 10 mM UDP-G	K_m = 3 μM, k_cat = 0.54 s⁻¹, 6.4 mg/h/mg	K_m = 78 μM (UDP-G) Inhibition: UDP
Ochratoxins (ochratoxin A, TDI = 16 ng/d/kg)
OTase (47–51 kDa)	Aspergillusniger [41,42]	pH 7.0, 40 °C, Zn²⁺	K_m = 13 μM, V_max = 0.29 μM/min, 21.8 mg/h/mg	Inhibition: 1,10- phenanthroline
OTA hydrolase (MW is unknown)	Aspergillusniger [43]	pH 7.5, 37 °C	K_m = 0.5 mM, V_max = 0.44 μM/min, 2.16 mg/h/mg	Inhibition: EDTA
Patulin (TDI = 0.4 μg/d/kg)
PGUG (27 kDa)	Meyerozyma guilliermondii [44]	pH 5.0, 28 °C	0.3 μg/h/mg	–
Lipase (25–64 kDa)	porcine pancreas [45,46]	pH 6.0, 40 °C	0.3 μg/h/mg	–
Putative orotate phosphoribosyl- transferase (25 kDa)	Rhodotorula mucilaginosa [47]	pH 4.0, 25 °C	K_m = 16 μM, k_cat = 3.4 × 10⁻⁵ s⁻¹, 0.76 mg/h/mg	–
Fumonisins (fumonisin B₁, TDI = 2 μg/d/kg)
FumD (57 kDa)	Sphingopyxis sp. [48]	pH 8.0, 30 °C	3 μg/h/mg	–
Ergot alkaloids (ergotamine, TDI = 0.6 μg/d/kg)
ErgA (35 kDa)	Rhodococcus erythropolis [49]	pH 8–9, 35 °C	13 mg/h/mg	–
Trichothecenes (deoxynivalenol, TDI = 1 μg/d/kg)
OsUGT79 (51 kDa)	Oryzasativa [50]	pH 8.0, 23 °C, 0.5 mM UDP-G	K_m = 61 μM, k_cat = 1.07 s⁻¹, 22.4 mg/h/mg	–
OsUGT79 with N-His₆-MBP (95 kDa)	Oryzasativa [51]	pH 7.0, 37 °C, 10 mM UDP-G	K_m = 0.23 mM, k_cat = 0.57 s⁻¹, 6.4 mg/h/mg	K_m = 2.2 mM (UDP-G) Activation: Ca²⁺ < Fe²⁺ ≈ Mg²⁺ < Mn²⁺ Inhibition: UDP, DON, Cu²⁺, Zn²⁺
HvUGT13248 with C-His₆ (53 kDa)	Hordeumvulgare [52]	pH 7.0, 25 °C, 1 mM UDP-G	K_m = 1.5 mM, V_max = 0.22 μmol/min/mg, 3.91 mg/h/mg	–
Bradi5g03300 with N-MBP and C-His₆ (96 kDa)	Brachypodium distachyon [52]	pH 7.0, 25 °C, 1 mM UDP-G	K_m = 0.37 mM, V_max = 19 nmol/min/mg, 0.34 mg/h/mg	–
TRI101 (50 kDa)	Fusarium sporotrichioides, Fusarium graminearum [53]	pH 8.0, 25 °C, 1.5 mM acetyl-CoA	K_m = 11.7 μM, k_cat = 13.5 s⁻¹, 288 mg/h/mg	–
Lipase (41 kDa)	Aspergillusniger [54]	pH 8.5, 40 °C	4.3 μg/h/mg	Activation: Ca²⁺ < Fe²⁺ ≈ Mg²⁺ Inhibition: EDTA << Zn²⁺ ≈ Cu²⁺
AKR18A1 with His₆ (37 kDa)	Sphingomonas sp. [55]	pH 9.5, 55 °C, 2 mM NADP⁺	K_m = 1.2 mM, V_max = 26 nmol/min/mg, 0.46 mg/h/mg	K_m = 0.48 mM (NADP⁺)
DepA (62 kDa)	Devosiamutans [56,57]	pH 7.5, Ca²⁺, 0.1 mM PQQ	K_m = 32 μM, k_cat = 4.2 s⁻¹, 72 mg/h/mg	Inhibition: Co²⁺ < Cu²⁺ < Fe²⁺
BdCXE29 (38 kDa)	Brachypodium distachyon [58]	pH 7.5, 25 °C	K_m = 0.42 mM (15-ADON), V_max = 3.4 μmol/min/mg, 69 mg/h/mg	Inhibition: 3-ADON

* Approved TDI or provisional maximal TDI are listed according to Joint FAO/WHO Expert Committee on Food Additives (http://apps.who.int/food-additives-contaminants-jecfa-database/), if not stated otherwise. As being a genotoxicant and carcinogen, aflatoxin B₁ cannot have an established TDI, so the value is an estimated TDI at a cancer risk level of 10⁻⁵ (10 per million) in countries where peoples may in addition be exposed to Hepatitis B infection [14]. Sterigmatocystin being carcinogenic too has only provisional dose of low health concern [17] that was specified in the Table. The limit of ergotamine was proposed by European Food Safety Authority [18] and has not been approved yet (as to 2019). Aflatoxin B₁—(3S,7R)-11-methoxy-6,8,19-trioxapentacyclo [10.7.0.02,9.03,7.013,17]nonadeca-1,4,9,11,13(17)-pentaene-16,18-dione; sterigmatocystin—(3aR,12cS)- 8-hydroxy-6-methoxy-3a,12c-dihydro-7H-furo[3′,2′:4,5]furo[2,3-c]xanthen-7-one; zearalenone—(4S,12E)-16,18-dihydroxy-4-methyl-3-oxabicyclo[12.4.0]octadeca-1(18),12,14,16-tetraene-2,8-dione; ochratoxin A—(2S)-2-[[(3R)-5-chloro-8-hydroxy-3-methyl-1-oxo-3,4-dihydroisochromene- 7-carbonyl]amino]-3-phenylpropanoic acid; patulin—4-hydroxy-4,6-dihydrofuro[3,2-c]pyran-2-one; fumonisin B₁—(2R)-2-[2-[(5R,6R,7S,9S,11R,16R,18S,19S)-19-amino-6-[(3R)-3,4-dicarboxybutanoyl] oxy-11,16,18-trihydroxy-5,9-dimethylicosan-7-yl]oxy-2-oxoethyl]butanedioic acid; ergotamine—(6aR,9R)-N-[(1S,2S,4R,7S)-7-benzyl-2-hydroxy-4-methyl-5,8-dioxo-3-oxa-6,9-diazatricyclo[7.3.0.02,6]dodecan-4-yl]-7-methyl-6,6a,8,9-tetrahydro-4H-indolo[4,3-fg]quinoline-9-carboxamide; deoxynivalenol—(1R,2R,3S,7R,9R,10R,12S)-3,10-dihydroxy-2-(hydroxymethyl)-1,5-dimethylspiro [8-oxatricyclo[7.2.1.02,7]dodec-5-ene-12,2′-oxirane]-4-one. ** The value was calculated using data of referenced authors. Abbreviations: PMSF—phenylmethylsulfonyl fluoride; AEBSF—4-(2-aminoethyl)-benzenesulfonyl fluoride; UDP-G—uridine diphosphate glucose; N-His₆-MBP—hexahistidine tag and maltose-binding protein on the N-terminus of enzyme molecule; C-His₆—hexahistidine tag on the C-terminus of enzyme molecule; N-MBP—maltose-binding protein on the N-terminus of enzyme molecule; PQQ—pyrroloquinoline quinone; 3-ADON—3-acetyl deoxynivalenol, 15-ADON—15-acetyl deoxynivalenol.