pc GH7 cellobiohydrolase Carries a CBM1; Plant cell wall cbh1-4 (6); cel7d (4)

Transcription

1 Supplementary Table 1 Putative carbohydrate- and lignin-active enzymes identified in the P. chrysosporium genome. Model Designation Predicted function Comment (Ref.) Biological target Carbohydrate active enzymes pc CE1 putative acetylxylan esterase Carries a CBM1 Plant cell wall pc CE1 putative acetylxylan esterase Carries a CBM1 Plant cell wall pc CE1 putative acetylxylan esterase Plant cell wall gx CE1 putative esterase Plant cell wall pc CE4 putative chitin deacetylase Fungal cell wall pc CE4 putative chitin deacetylase Fungal cell wall pc CE4 putative chitin deacetylase Fungal cell wall pc CE4 putative esterase Fungal cell wall pc CE4 putative chitin deacetylase Fungal cell wall gx CE8 putative pectin methylesterase Plant cell wall gx CE9 putative GlcNAc-6-P deacetylase gw CE9 putative esterase gx CE9 putative esterase pc CE14 putative esterase gx GH1 putative β-glucosidase gx GH1 putative β-glucosidase gx GH2 putative β-mannosidase gx GH2 putative β-mannosidase gx GH3 β-glucosidase Plant cell wall pc GH3 putative β-xylosidase Plant cell wall pc GH3 putative β-glucosidase Plant cell wall pc GH3 putative β-glucosidase Plant cell wall gx GH3 putative β-glucosidase Plant cell wall pc GH3 putative β-glucosidase Plant cell wall pc GH3 putative β-glucosidase Plant cell wall gx GH3 putative β-glycosidase Plant cell wall 1

2 gx GH3 putative β-glucosidase Carries a CBM1 (1) Plant cell wall pc GH3 putative β-glycosidase Plant cell wall gx GH3 putative β-glycosidase Plant cell wall gx GH3 putative β-glucosidase Plant cell wall pc GH5 putative β-glycosidase Plant or fungal cell wall pc GH5 putative β-1.3-mannanase Plant or fungal cell wall pc GH5 putative β-1,3-glucanase Plant or fungal cell wall pc GH5 putative β-1,3-glucanase Plant or fungal cell wall pc GH5 putative β-1,3-glucanase Plant or fungal cell wall pc GH5 putative β-1,3-glucanase Plant or fungal cell wall pc GH5 putative β-1,3-glucanase Plant or fungal cell wall gx GH5 putative β-1,3-glucanase Plant or fungal cell wall pc GH5 putative β-1,3-glucanase Plant or fungal cell wall pc GH5 putative β-1,3-glucanase Plant or fungal cell wall pc GH5 putative endoglucanase Carries a CBM1 Plant cell wall gw GH5 putative endoglucanase Carries a CBM1 Plant cell wall gx GH5 putative β-mannanase Carries a CBM1 Plant cell wall pc GH5 putative β-mannanase Carries a CBM1 Plant cell wall gw GH5 putative β-glycosidase gx GH5 putative β-glycosidase pc GH5 putative β-glycosidase gx GH5 putative β-mannanase Plant cell wall pc GH6 cellobiohydrolase II Carries a CBM1; Plant cell wall cel6a (2) pc GH7 cellobiohydrolase Carries a CBM1; Plant cell wall cbh1-5 (3); cel7e (4) ug GH7 putative cellobiohydrolase Carries a CBM1; Plant cell wall cbh1-2 (5); cel7b (4) pc GH7 putative cellobiohydrolase cbh1-1 (5); cel7a (4) Plant cell wall pc GH7 cellobiohydrolase Carries a CBM1; cbh1-3 (5); cel7c (4) Plant cell wall 2

3 pc GH7 cellobiohydrolase Carries a CBM1; Plant cell wall cbh1-4 (6); cel7d (4) pc GH7 cellobiohydrolase Carries a CBM1; Plant cell wall cbh1-6 (3); cel7f (4) pc GH9 putative endoglucanase cel9a (7) Plant cell wall pc GH10 putative xylanase Plant cell wall gw GH10 xylanase A Carries CBM1; Plant cell wall (AAG44993) pc GH10 putative xylanase Carries a CBM1 Plant cell wall pc GH10 putative xylanase Carries a CBM1 Plant cell wall gx GH10 putative xylanase Plant cell wall gx GH10 putative xylanase Carries a CBM1 Plant cell wall pc GH11 xylanase B Carries a CBM1; Plant cell wall (AAG44995) pc GH12 putative xyloglucanase/endoglucanase Plant cell wall pc GH12 putative xyloglucanase/endoglucanase Plant cell wall pc GH13 putative α-glucosidase Starch/glycogen pc GH13 putative α-glucosidase Starch/glycogen pc GH13 putative glycogen branching enzyme Starch/glycogen gw GH13 putative glycogen debranching enzyme Starch/glycogen gx GH13 putative α-amylase Starch/glycogen pc GH13 putative α-amylase Carries CBM20 Starch/glycogen pc GH13 putative α-amylase Starch/glycogen pc GH13 putative α-amylase Starch/glycogen pc GH13; GT5 putative α-glucan synthase Modular protein Starch/glycogen pc GH15 putative glucoamylase Carries a CBM20 Starch/glycogen pc GH15 putative α-glycosidase Starch/glycogen pc GH16 putative β-1,3-glucanase Plant or fungal cell wall pc GH16 putative β-1,3-glucanase Plant or fungal cell wall pc GH16 putative β-1,3-glucanase Plant or fungal cell wall pc GH16 putative β-1,3-glucanase Plant or fungal cell wall pc GH16 putative β-1,3-glucanase Plant or fungal cell wall 3

4 pc GH16 putative β-1,3-glucanase Plant or fungal cell wall pc GH16 putative β-1,3-glucanase Plant or fungal cell wall pc GH16 putative β-1,3-glucanase Plant or fungal cell wall pc GH16 putative β-1,3-glucanase Plant or fungal cell wall pc GH16 putative β-1,3-glucanase Plant or fungal cell wall pc GH16 putative β-1,3-glucanase Plant or fungal cell wall pc GH16 putative β-1,3-glucanase Plant or fungal cell wall pc GH16 putative β-1,3-glucanase Plant or fungal cell wall pc GH16 putative β-1,3-glucanase Plant or fungal cell wall pc GH16 putative β-1,3-glucanase Plant or fungal cell wall gw GH16 putative β-1,3-glucanase Plant or fungal cell wall gw GH16 putative β-1,3-glucanase Plant or fungal cell wall gw GH16 putative β-1,3-glucanase Plant or fungal cell wall gx GH16 putative β-1,3-glucanase Plant or fungal cell wall pc GH16 putative β-1,3-glucanase Plant or fungal cell wall pc GH17 putative β-glycosidase pc GH17 putative β-glycosidase pc GH18 putative chitinase Fungal cell wall pc GH18 putative chitinase Fungal cell wall gx GH18 putative chitinase Fungal cell wall pc GH18 putative chitinase Fungal cell wall gx GH18 putative chitinase Fungal cell wall pc GH18 putative chitinase Fungal cell wall pc GH18 putative chitinase Fungal cell wall pc GH18 putative chitinase Fungal cell wall pc GH18 putative chitinase Fungal cell wall pc GH18 putative chitinase Fungal cell wall pc GH18 putative chitinase Fungal cell wall gx GH18 putative chitinase Fungal cell wall gx GH20 putative β-hexosaminidase pc GH20 putative β-hexosaminidase pc GH20 putative β-hexosaminidase pc GH25 putative lysozyme 4

5 pc GH27 α-galactosidase (8) gx GH27 putative α-galactosidase gx GH27 putative α-galactosidase gx GH28 putative polygalacturonase Plant cell wall gw GH28 putative polygalacturonase Plant cell wall gx GH28 putative rhamnogalacturonase Carries a CBM1 Plant cell wall pc GH28 putative polygalacturonase gw GH28 putative polygalacturonase pc GH30 putative β-1,6-glucanase gx GH31 putative α-glucosidase Starch/glycogen pc GH31 putative α-glucosidase Starch/glycogen gw GH31 putative α-glucosidase Starch/glycogen pc GH31 putative α-glucosidase Starch/glycogen pc GH31 putative α-glucosidase Starch/glycogen pc GH31 putative α-glucosidase Starch/glycogen pc GH35 putative β-galactosidase gw GH35 putative β-galactosidase gx GH37 putative trehalase Trehalose gx GH37 putative trehalase Trehalose gw GH38 putative α-mannosidase gx GH43 putative β-xylosidase Plant cell wall pc GH43 putative β-glycosidase Plant cell wall pc GH43 putative β-glycosidase Plant cell wall gx GH47 putative α-mannosidase pc GH47 putative α-mannosidase pc GH47 putative α-mannosidase pc GH47 putative α-mannosidase pc GH47 putative α-mannosidase gw GH51 putative α-l-arabinofuranosidase Plant cell wall pc GH51 putative α-l-arabinofuranosidase Plant cell wall pc GH53 putative β-1,4-galactanase Plant cell wall gx GH55 putative exo-1,3-β-glucanase Plant or fungal cell wall pc GH55 putative exo-1,3-β-glucanase Plant or fungal cell wall 5

6 pc GH61 putative endoglucanase Plant cell wall pc GH61 putative endoglucanase Plant cell wall pc GH61 putative endoglucanase Plant cell wall pc GH61 putative endoglucanase Carries a CBM1 Plant cell wall pc GH61 putative endoglucanase Carries a CBM1 (7) Plant cell wall pc GH61 putative endoglucanase Carries a CBM1 Plant cell wall pc GH61 putative endoglucanase Plant cell wall pc GH61 putative endoglucanase Carries a CBM1 Plant cell wall pc GH61 putative endoglucanase Plant cell wall gw GH61 putative endoglucanase Carries a CBM1 Plant cell wall gx GH61 putative endoglucanase Plant cell wall gx GH61 putative endoglucanase Plant cell wall pc GH61 putative endoglucanase Plant cell wall gx GH61 putative endoglucanase Plant cell wall gw GH61 putative endoglucanase Plant cell wall pc GH61 putative endoglucanase Plant cell wall pc GH61 putative endoglucanase Plant cell wall gx GH63 putative processing α-glucosidase gx GH71 putative α-1,3-glucanase Fungal cell wall pc GH71 putative α-1,3-glucanase Fungal cell wall pc GH71 putative α-1,3-glucanase Fungal cell wall pc GH72 putative β-1,3-glucanosyltransferase Fungal cell wall pc GH74 putative xyloglucanase/endoglucanase Plant cell wall pc GH74 putative xyloglucanase/endoglucanase Carries a CBM1 Plant cell wall pc GH85 putative endo-β-n-acetylglucosaminidase gx GH88 putative delta4,5 unsaturated glucuronyl hydrolase pc GH89 putative α-n-acetylglucosaminidase pc GH89 putative α-n-acetylglucosaminidase gx GH92 putative α-1,2-mannosidase pc GH92 putative α-1,2-mannosidase pc GH92 putative α-1,2-mannosidase pc GH92 putative α-1,2-mannosidase pc GH92 putative α-1,2-mannosidase 6

7 pc GH92 putative α-1,2-mannosidase pc GT1 putative β-glycosyltransferase gx GT1 putative UDP-glucose:sterol glucosyltransferase gx GT1 putative UDP-glucose:sterol glucosyltransferase pc GT1 putative β-glycosyltransferase gx GT1 putative β-glycosyltransferase gw GT2 putative chitin synthase Fungal cell wall gw GT2 putative chitin synthase Fungal cell wall gx GT2 putative β-glycosyltransferase gw GT2 putative chitin synthase Fungal cell wall pc GT2 putative chitin synthase Fungal cell wall pc GT2 putative chitin synthase Fungal cell wall gx GT2 putative chitin synthase Fungal cell wall pc GT2 putative chitin synthase Fungal cell wall pc GT2 putative chitin synthase Fungal cell wall pc GT2 putative dolichyl-phosphate β-glucosyltransferase Fungal cell wall pc GT2 putative dolichol-phosphate β-mannosyltransferase Fungal cell wall pc GT3 putative glycogen synthase Glycogen gx GT4 putative trehalose phosphorylase Trehalose pc GT4 putative α-glycosyltransferase pc GT4 putative GPI-anchor biosynthesis protein (PigA) Glycosylphosphatidylinositol pc GT4 putative α-mannosylltransferase pc GT8 putative α-glycosyltransferase pc GT8 putative α-glycosyltransferase pc GT8 putative α-glycosyltransferase gx GT8 putative glycogenin Glycogen pc GT8 putative α-glycosyltransferase pc GT15 putative α-mannosyltransferase gw GT15 putative GDP-mannose α-1,2-mannosyltransferase pc GT17 putative β-n-acetylglucosaminyltransferase pc GT20 putative trehalose-phosphate synthase (Tps1) Trehalose pc GT20 putative trehalose-5-phosphate synthase/phosphatase Trehalose 7

8 gx GT20 putative trehalose-6-phosphate Trehalose synthase/phosphatase pc GT21 putative UDP-glucose ceramide Glycolipids glucosyltransferase pc GT22 putative dolichyl phosphate mannose : α- mannosyltransferase (Smp3) gx GT22 putative dolichyl phosphate mannose : α- mannosyltransferase (Alg9) gx GT22 putative dolichyl phosphate mannose : α- mannosyltransferase (Alg12) gx GT22 putative dolichyl phosphate mannose : α- Glycosylphosphatidylinositol mannosyltransferase (PigB) pc GT24 putative UDP-glucose glycoprotein α- glucosyltransferase pc GT31 putative glycosyltransferase pc GT32 putative α-1,6-mannosyltransferase (Och1) pc GT33 putative β-mannosyltransferase gx GT35 putative glycogen phosphorylase Glycogen pc GT39 putative dolichyl phosphate mannose : α- mannosyltransferase gw GT39 putative dolichyl phosphate mannose : α- mannosyltransferase gw GT39 putative dolichyl phosphate mannose : α- mannosyltransferase pc GT41 putative UDP-GlcNAc : peptide N- acetylglucosaminyltransferase pc GT48 putative β-1,3-glucan synthase gx GT48 putative β-1,3-glucan synthase Fungal cell wall pc GT50 putative dolichyl phosphate mannose : α- Glycosylphosphatidylinositol mannosyltransferase pc GT57 putative dolichyl phosphate glucose : α- glucosyltransferase (Alg8) 8

9 gw GT57 putative dolichyl phosphate glucose : α- glucosyltransferase (Alg6) pc GT58 putative dolichyl phosphate mannose : α- mannosyltransferase (Alg3) gx GT59 putative dolichyl-phosphate-glucose : α- glucosyltransferase (Alg1) pc GT66 putative oligosaccharyl transferase subunit similar to S. cerevisiae STT3 pc GT69 putative GDP-mannose α-mannosyltransferase similar to CAP59 from C. neoformans pc GT69 putative GDP-mannose α-mannosyltransferase Pc GT69 putative GDP-mannose α-mannosyltransferase Extracellular oxidative enzymes Peroxidases* pc lipe Lignin peroxidase (9, 10) Lignin pc lipa Lignin peroxidase Major isozyme(11-13) Lignin pc lipb Lignin peroxidase (14-16) Lignin pc lipc Lignin peroxidase (14, 17) Lignin pc lipj Lignin peroxidase (11, 18) Lignin pc liph Lignin peroxidase (19) Lignin pc lipg Lignin peroxidase (18) Lignin ug lipi Lignin peroxidase Transposon remnant Lignin (11, 20) pc lipd Lignin peroxidase Major isozyme (17) Lignin pc lipf Lignin peroxidase (21) Lignin pc nop1 Hybrid peroxidase Similar to AB Lignin? Gypsy retroelement inserted 100 nt upstream. pc mnp2 Manganese peroxidase Isozyme H4 (22) Lignin pc mnp3 Manganese peroxidase Isozyme H3 (23) Lignin pc mnp1 Manganese peroxidase (24, 25) Lignin 9

10 pc mnp4 Manganese peroxidase Lignin pc mnp5 Manganese peroxidase Pulp enzyme Lignin A61147(26) pc cpo1 chloroperoxidase Similar to CAC03461 Possibly membrane bound.? Copper radical oxidases** pc glx1 Glyoxal oxidase Major isozyme(27,28) Methyl glyoxal & related aldehydes pc cro3 Copper radical oxidase Methyl glyoxal & related aldehydes gw cro4 Copper radical oxidase unknown pc cro5 Copper radical oxidase unknown pc cro2 Copper radical oxidase Similar to M86819 Possible galactose oxidase pc cro1 Copper radical oxidase unknown pc cro6 Copper radical oxidase unknown FAD oxidases pc cdh1 Cellobiose dehydrogenase (29-31) Cellobiose gw Pyranose 2-oxidase Similar to D73369 Aldopyranoses (C2 oxidation) pc gox1 Glucose oxidase Similar to AJ Glucose pc Alcohol oxidase Similar to AF Possible aryl alcohol oxidase pc Alcohol oxidase Similar to AF Possible aryl alcohol oxidase pc Alcohol oxidase Similar to AF Possible aryl alcohol oxidase pc Alcohol oxidase Similar to AF Possible aryl alcohol oxidase Multicopper oxidases pc pc mco1 Multicopper oxidase (32) Possible iron metabolism cdna sequence AY ug mco2 Multicopper oxidase Possible iron metabolism ug mco3 Multicopper oxidase Gypsy element within Possible iron metabolism ug th intron pc mco4 Multicopper oxidase Possible iron metabolism Glycoside hydrolases (GH), carbohydrate esterases (CE) and glycosyltransferases (GT) families according to the CAZy database ( /CAZY). * Lignin peroxidases and manganese peroxidase nomenclature as described by Gaskell et al (33) and Martinez (34), respectively. ** Copper radical oxidases defined by conserved catalytic residues (35). 10

11 References 1. Li, B. & Renganathan, V. Gene cloning and characterization of a novel cellulose-binding β-glucosidase from Phanerochaete chrysosporium. Appl. Environ. Microbiol. 64, (1998). 2. Tempelaars, C., Birch, P., Sims, P. & Broda, P. Isolation, characterization, and analysis of the cbhii gene of Phanerochaete chrysosporium. Appl. Environ. Microbiol. 60, (1994). 3. Covert, S., Bolduc, J. & Cullen, D. Genomic organization of a cellulase gene family in Phanerochaete chrysosporium. Curr. Genet. 22, (1992a). 4. Munoz, I.G. et al. Family 7 cellobiohydrolases from Phanerochaete chrysosporium: crystal structure of the catalytic module of Cel7D (CBH58) at 1.32 A resolution and homology models of the isozymes. J. Mol. Biol. 314, (2001). 5. Covert, S., Vanden Wymelenberg, A. & Cullen, D. Structure, organization and transcription of a cellobiohydrolase gene cluster from Phanerochaete chrysosporium. Appl. Environ. Microbiol. 58, (1992b). 6. Vanden Wymelenberg, A., Covert, S. & Cullen, D. Identification of the gene encoding the major cellobiohydrolase of the white-rot fungus Phanerochaete chrysosporium. Appl. Environ. Microbiol. 59, (1993). 7. Wymelenberg, A.V. et al. Transcript analysis of genes encoding a family 61 endoglucanase and a putative membrane-anchored family 9 glycosyl hydrolase from Phanerochaete chrysosporium. Appl. Environ. Microbiol. 68, (2002). 8. Hart, D.O. et al. Identification of Asp-130 as the catalytic nucleophile in the main α-galactosidase from Phanerochaete chrysosporium, a family 27 glycosyl hydrolase. Biochem. 39, (2000). 9. Reiser, J., Walther, I., Fraefel, C. & Fiechter, A. Methods to investigate the expression of lignin peroxidase genes by the whiterot fungus Phanerochaete chrysosporium. Appl. Environ. Microbiol. 59, (1993). 10. Ritch, T.G. et al. Lignin peroxidase from the basidiomycete Phanerochaete chrysosporium is synthesized as a preproenzyme. Gene 107, (1991). 11. Schalch, H., Gaskell, J., Smith, T.L. & Cullen, D. Molecular cloning and sequences of lignin peroxidase genes of Phanerochaete chrysosporium. Mol. Cell. Biol. 9, (1989). 12. Tien, M. & Tu, C.-P.D. Cloning and sequencing of a cdna for a ligninase from Phanerochaete chrysosporium. Nature 326, (1987). 13. Smith, T.L., Schalch, H., Gaskell, J., Covert, S. & Cullen, D. Nucleotide sequence of a ligninase gene from Phanerochaete chrysosporium. Nucl. Acids Res. 16, 1219 (1988). 14. Gaskell, J., Dieperink, E. & Cullen, D. Genomic organization of lignin peroxidase genes of Phanerochaete chrysosporium. Nucleic Acids Res. 19, (1991). 11

12 15. Huoponen, K. et al. Characterisation of lignin peroxidase-encoding genes from lignin-degrading basidiomycetes. Gene 89, (1990). 16. Broda, P. et al. in Enzyme Systems for Lignocellulose Degradation. (ed. M.P. Couglan) (Elsevier Applied Science, London; 1989). 17. de Boer, H.A., Zhang, Y.Z., Collins, C. & Reddy, C.A. Analysis of nucleotide sequences of two ligninase cdnas from a white-rot filamentous fungus, Phanerochaete chrysosporium. Gene 60, (1987). 18. Stewart, P. & Cullen, D. Organization and differential regulation of a cluster of lignin peroxidase genes of Phanerochaete chrysosporium. J. Bact. 181, (1999). 19. Brown, A., Sims, P.F.G., Raeder, U. & Broda, P. Multiple ligninase-related genes from Phanerochaete chrysosporium. Gene 73, (1988). 20. Gaskell, J., Vanden Wymelenberg, A. & Cullen, D. Structure, inheritance, and transcriptional effects of Pce1, an insertional element within Phanerochaete chrysosporium lignin peroxidase gene lipi. Proc. Natl. Acad. Sci. USA 92, (1995). 21. Padmavathy, S.N., Zhang, Y. & Reddy, C.A. Characterization of a new lignin peroxidase gene (GLG6) from Phanerochaete chrysosporium. Biochem. Biophys. Res. Commun. 173, (1991). 22. Pease, E.A., Andrawis, A. & Tien, M. Manganese-dependent peroxidase from Phanerochaete chrysosporium. Primary structure deduced from complementary DNA sequence. J. Biol. Chem. 264, (1989). 23. Orth, A., Rzhetskaya, M., Cullen, D. & Tien, M. Characterization of a cdna encoding a manganese peroxidase from Phanerochaete chrysosporium: genomic organization of lignin and manganese peroxidase genes. Gene 148, (1994). 24. Godfrey, B.J., Mayfield, M.B., Brown, J.A. & Gold, M. Characterization of a gene encoding a manganese peroxidase from Phanerochaete chrysosporium. Gene 93, (1990). 25. Pribnow, D., Mayfield, M.B., Nipper, V.J., Brown, J.A. & Gold, M.H. Characterization of a cdna encoding a manganese peroxidase, from the lignin-degrading basidiomycete Phanerochaete chrysosporium. J. Biol. Chem. 264, (1989). 26. Datta, A., Bettermann, A. & Kirk, T.K. Identification of a specific manganese peroxidase among ligninolytic enzymes secreted by Phanerochaete chrysosporium during wood decay. Appl. Environ. Microbiol. 57, (1991). 27. Kersten, P. & Cullen, D. Cloning and characterization of a cdna encoding glyoxal oxidase, a peroxide-producing enzyme from the lignin-degrading basidiomycete Phanerochaete chrysosporium. Proc. Nat. Acad. Sci. USA 90, (1993). 28. Kersten, P.J., Witek, C., Vanden Wymelenberg, A. & Cullen, D. Phanerochaete chrysosporium glyoxal oxidase is encoded by two allelic variants: structure, genomic organization and heterologous expression of glx1 and glx2. J. Bact. 177, (1995). 29. Li, B., Nagalla, S.R. & Renganathan, V. Cloning of a cdna encoding cellobiose dehydrogenase, a hemoflavoenzyme from Phanerochaete chrysosporium. Appl. Environ. Microbiol. 62, (1996). 12

13 30. Li, B., Nagalla, S.R. & Renganathan, V. Cellobiose dehydrogenase from Phanerochaete chrysosporium is encoded by two allelic variants. Appl. Environ. Microbiol. 63, (1997). 31. Raices, M. et al. Cloning and characterization of a cdna encoding a cellobiose dehydrogenase from the white rot fungus Phanerochaete chrysosporium. FEBS Lett. 369, (1995). 32. Larrondo, L., Salas, L., Melo, F., Vicuna, R. & Cullen, D. A novel extracellular multicopper oxidase from Phanerochaete chrysosporium with ferroxidase activity. Appl. Environ. Microbiol. 69, (2003). 33. Gaskell, J. et al. Establishment of genetic linkage by allele-specific polymerase chain reaction: application to the lignin peroxidase gene family of Phanerochaete chrysosporium. Bio/Technology 12, (1994). 34. Martinez, A.T. Molecular biology and structure-function of lignin-degrading heme peroxidases. Enzyme Microb. Technol. 30, (2002). 35. Whittaker, M.M., Kersten, P.J., Cullen, D. & Whittaker, J.W. Identification of catalytic residues in glyoxal oxidase by targeted mutagenesis. J. Biol. Chem. 274, (1999). 13