Trichoderma and Gliocladium Volume Trichoderma and Gliocladium Volume Enzymes, biological control and commercial applications Edited by GARY E.HARMAN Cornell University, Geneva, NY, USA and CHRISTIAN P.KUBICEK University of Technology, Vienna, Austria with the capable editorial assistance of Kristen L.Ondik This edition published in the Taylor & Francis e-Library, 2005 “To purchase your own copy of this or any of Taylor & Francis or Routledge’s collection of thousands of eBooks please go to www.eBookstore.tandf.co.uk.” UK Taylor & Francis Ltd, Gunpowder Square, London, EC4A 3DE USA Taylor & Francis Inc., 1900 Frost Road, Suite 101, Bristol, PA 19007 Copyright © Taylor & Francis Ltd 1998 All rights reserved No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, electrostatic, magnetic tape, mechanical, photocopying, recording or otherwise, without the prior permission of the copyright owner British Library Cataloguing in Publication Data A catalogue record for this book is available from the British Library ISBN 0-203-48445-2 Master e-book ISBN ISBN 0-203-79269-6 (Adobe eReader Format) ISBN 0-7484-0572-0 Vol ISBN 0-7484-0805-3 Vol ISBN 0-7484-0806-1 Two volume set Library of Congress Cataloging in Publication Data are available Cover design by Youngs Design in Production Contents Preface Contributors PART ONE Degradation of Polysaccharides and Related Macromolecules 1 Structure-function relationships in Trichoderma cellulolytic enzymes A.Koivula, M.Linder and T.T.Teeri 1.1 Introduction 1.2 Domain structures of cellulolytic enzymes 1.3 Catalytic reaction mechanisms of T reesei cellobiohydrolases 1.4 Role and function of the cellulose-binding domains 12 1.5 Aspects of crystalline cellulose degradation 14 References 16 Enzymology of hemicellulose degradation P.Biely and M.Tenkanen 21 2.1 Introduction 21 2.2 Xylan structure and enzymes required for its hydrolysis 22 2.3 β-Mannan structures and enzymes required for their hydrolysis 32 2.4 Conclusions 36 References 37 Regulation of production of plant polysaccharide degrading enzymes by Trichoderma C.P.Kubicek and M.E.Penttilä 42 3.1 Introduction 42 3.2 Regulation of cellulase expression 43 3.3 Regulation of hemicellulase expression 54 3.4 Conclusions 57 v References 58 Chitinolytic enzymes and their genes M.Lorito 64 4.1 Introduction 64 4.2 Chitinolytic enzymes from Trichoderma and Gliocladium 65 4.3 Genes encoding Chitinolytic enzymes 74 4.4 Roles of Chitinolytic enzymes and their genes 78 4.5 Potential applications and commercial usefulness of Chitinolytic enzymes and their genes 79 Acknowledgments 81 References 81 Glucanolytic and other enzymes and their genes T.Benitez, C.Limón, J.Delgado-Jarana and M.Rey 89 5.1 Introduction 89 5.2 Glucanolytic enzymes and their genes 92 5.3 Other hydrolases and their genes 101 5.4 Biotechnological applications of hydrolytic enzymes 105 Acknowledgments 107 References 107 PART TWO Application of Trichoderma and Gliocladium in Agriculture 114 Trichoderma and Gliocladium in biological control: an overview L.Hjeljord and A.Tronsmo 115 6.1 The need for biologically based fungicides 115 6.2 Suitability of Trichoderma and Gliocladium spp as biological control agents 116 6.3 Antagonistic mechanisms 122 6.4 Integrated control 125 6.5 Formulation and delivery 126 6.6 Conclusions 126 References 127 Mycoparasitism and lytic enzymes I.Chet, N.Benhamou and S.Haran 134 Sequential events involved in mycoparasitism 134 7.1 vi 7.2 Heterologous lytic enzymes as a tool for enhancing biocontrol activity 141 7.3 Ultrastructural changes and cellular mechanisms during the mycoparasitic process 143 7.4 Concluding remarks 148 References 149 The role of antibiosis in biocontrol C.R.Howell 152 8.1 Introduction 152 8.2 Antibiotics associated with disease control by Trichoderma and Gliocladium species 152 8.3 Environmental parameters affecting antibiotic production and activity 155 8.4 Antibiotics as mechanisms in disease biocontrol, and their modes of action 156 8.5 Genetic manipulation of antibiotics and disease control 157 8.6 Concluding remarks 158 References 158 Direct effects of Trichoderma and Gliocladium on plant growth and resistance to pathogens B.A.Bailey and R.D.Lumsden 163 9.1 Introduction 163 9.2 Plant growth inhibition 163 9.3 Plant growth promotion 169 9.4 Unraveling host plant interactions 176 References 177 Industrial production of active propagules of Trichoderma for agricultural uses E.Agosin and J.M.Aguilera 182 10.1 Introduction 182 10.2 Trichoderma propagules for agricultural purposes 182 10.3 Producing high-quality conidial biomass of Trichoderma 183 10.4 Preserving the quality of a dry commercial biopesticide 192 Acknowledgments 197 References 198 Potential and existing uses of Trichoderma and Gliocladium for plant disease control and plant growth enhancement 203 10 11 vii G.E.Harman and T.Björkman 11.1 Historical perspective 203 11.2 Requirements for development of successful biocontrol systems and products 204 11.3 Product examples and concepts of use 206 11.4 Summary 230 References 231 Trichoderma as a weed mould or pathogen in mushroom cultivation D.Seaby 235 12.1 Introduction 235 12.2 Green mould epidemic in Ireland 237 12.3 A Trichoderma harzianum group green mould epidemic in North America 246 12.4 Green mould in Australia 247 12.5 The Trichoderma spp found on mushroom casing 247 12.6 Summary and discussion 248 12.7 Appendix 250 References 251 12 PART THREE Protein Production and Application of Trichoderma Enzymes 13 254 Industrial mutants and recombinant strains of Trichoderma reesei A.Mäntylä, M.Paloheimo and P.Suominen 255 13.1 Introduction 255 13.2 Industrial mutants of Trichoderma reesei 256 13.3 Industrial recombinant strains of Trichoderma reesei 258 13.4 Concluding remarks 267 References 268 Application of Trichoderma enzymes in the textile industry Y.M.Galante, A.De Conti and R.Monteverdi 272 14.1 Introduction and background: why the textile industry? 272 14.2 Enzymatic stonewashing of denim garments with fungal cellulases 273 14.3 Biofinishing and enzymatic defibrillation of cellulosic fibers 277 14.4 Current perspectives and future developments for cellulase applications in textiles 283 14 viii Acknowledgments 284 References 284 Application of Trichoderma enzymes in the food and feed industries Y.M.Galante, A.De Conti and R.Monteverdi 286 15.1 Brewing 286 15.2 Wine making and fruit juices 287 15.3 Olive oil production 291 15.4 Animal feed 296 References 299 Applications of Trichoderma reesei enzymes in the pulp and paper industry J.Buchert, T.Oksanen, J.Pere, M.Siika-aho, A.Suurnäkki and L.Viikari 302 16.1 Introduction 302 16.2 Reasons for modifying the fibre substrates 303 16.3 Potential enzymes for commercial applications 305 16.4 Applications of T reesei enzymes in the pulp and paper industry 308 16.5 Conclusions 314 References 315 Heterologous protein production in Trichoderma M.Penttilä 322 17.1 General features of Trichoderma in protein production 322 17.2 Tools and strategies for heterologous protein production 323 17.3 Examples of heterologous protein production in T reesei 324 17.4 Conclusions and future aspects 332 References 334 15 16 17 Preface In the course of an investigation of damping off of Citrus seedlings, a strain of Trichoderma lignorum has been found to parasitize a number of pathogenic soil fungi in culture, i.e., Rhizoctonia solani, Phytophthora parasitica, Pythium spp., Rhizopus spp., and Sclerotium rolfsii The inhibition and death of the host hyphae were brought about in two ways : (a) in aerial hyphae by close contact or by coiling of Trichoderma around them; (b) in submerged hyphae by action at a little distance more frequently than by contact Weindling, R 1932 Trichoderma lignorum as a parasite of other soil fungi Phytopathology 22:837–845 The above quotation is from the summary of one of Richard Weindling’s landmark papers on the potential of Trichoderma spp to control plant pathogenic fungi This series of papers was among the first indications that these fungi have a wide-ranging commercial potential that is only now being realized These fungi are among the most widely distributed and common fungi in nature They exist in climates ranging from the tundra to the tropics and colonize a wide range of habitats They very commonly occur in soil and frequently are among the most prevalent fungi there, often occurring at levels of 100 to 10000 propagules per gram of soil They also exist in other environments and are frequently found in decaying wood or other cellulosic substrates such as decaying cotton fabrics in the tropics Further, they may be found on aerial, as well as subterranean, plant parts They have a number of mechanisms for survival and proliferation, including physical attack of other fungi and degradation and utilization of complex carbohydrates For the most part they are beneficial to man’s economic interests, especially since they no doubt reduce plant diseases in nearly all agricultural environments However, there are exceptions: for example, as noted in the Preface to Volume 1, some strains degrade cotton fabrics while other specific strains are pathogenic on commercial mushrooms The mechanisms which these remarkable fungi have evolved can be exploited for commercial applications, and many of these are documented in this volume First and foremost, they are extremely efficient producers of a number of enzymes Such enzymes include cellulases and hemicellulases that can be used for many purposes, including fabric treatments, enhancing nutritional composition of animal feeds, and improving properties and enhancing yields of plant oils and fruit juices They also produce quantities of other enzymes, especially chitinases and glucanases, that are strongly fungitoxic There is substantial potential for use of these enzymes as antifungal theraputants in numerous applications Not only are these useful enzymes produced in large quantities, but also there are multiple forms of each class of enzyme HETEROLOGOUS PROTEIN PRODUCTION IN TRICHODERMA 337 SUOMINEN, P.L., MÄNTYLÄ, A.L., KARHUNEN, T., HAKOLA, S., and NEVALAINEN, H 1993 High-frequency one-step gene replacement in Trichoderma reesei II: Effects of deletions of individual cellulase genes Mol Gen Genet 241:523– 530 UUSITALO, J.M., NEVALAINEN, K.M H., HARKKI, A.M., KNOWLES, J.K.C., and PENTTILÄ, M.E 1991 Enzyme production with recombinant Trichoderma reesei strains J Biotechnol 17:35–50 VANHANEN, S 1991 Isolation and characterization of genes involved in basic metabolism of the filamentous fungus Trichoderma reesei Ph D.Thesis, VTT Publications 75, Espoo, Finland, 119 pp VANHANEN, S., PENTTILÄ, M., LEHTOVAARA, P., and KNOWLES, J 1989 Isolation and characterization of the 3-phosphoglycerate kinase gene (pgk) from the filamentous fungus Trichoderma reesei Curr Genet 15:181–186 VANHANEN, S., SALOHEIMO, A., ILMÉN, M., KNOWLES, J.K.C., and PENTTILA, M 1991 Promoter structure and expression of the 3-phosphoglycerate kinase-encoding gene (pgk1) of Trichoderma reesei Gene 106:129–133 VELDHUISEN, G., SALOHEIMO, M., FIERS, M.A., PUNT, P.J., CONTRERAS, R., PENTTILA, M., and VAN DEN HONDEL, C.A.M.J.J 1997 Isolation and analysis of functional homologues of the secretion related SAR1 gene of S cerevisiae from Aspergillus niger and Trichoderma reesei Mol Gen Genet 256:446–455 WARD, M., WILSON, L.J., KODAMA, K.H., REY, M.W., and BERKA, R.M 1990 Improved production of chymosin in Aspergillus by expression as a glucoamylase-chymosin fusion Bio/Technology 8:435–440 Index acetamidases 295, 366 acetamide 295 acetolactate synthases (ALS) 186 acetyl esterases (AE) 27, 35–6, 37, 41 acetylgalactoglucomannan esterases 37 acetylgalactoside 75 acetylglucomannan esterases 37, 41 acetylglucosaminidases 75, 157–9, 169 acetylglucuronoxylan 36 acetylhexosaminidases 74–8, 81, 85–6 acetylxylan esterases (AXE) 26, 27, 35–6, 41, 65, 348 Achbya 103 acid phosphatases 369 actinomycetes 107 Agaricus 267, 271, 277, 279–80 Agaricus bisporus 102, 103, 106, 267–8, 280, 282 Agaricus bitorquis 267 agriculture 118–19, 121, 131–45, 231–61, 205–23 see also mushrooms alkyl pyrone 174 allergenicity 138 Alternaria spp 82 Alternaria alternata 91 Amillaria luteobubalina 142 amino acids 153, 186, 296, 347, 369 antibiosis 175, 178 cellulolytic enzymes 9, 14, 15 chitinolytic enzymes 75, 81, 85, 86, 87 glucanolytic enzymes 107, 111, 114 hemicellulose degradation 27, 28, 33–5, 38, 41 hydrolases 114, 115, 116, 117 polysaccharide degrading enzymes 58–9 amino peptidases 114, 115 amylases 76, 113–14, 120, 312, 327 cellulolytic enzymes 7, feed industry 338, 340 amyloglucosidases 106 antagonism and antagonists 89–90, 139–42 antibiosis 175, 177, 178, 180 biocontrol 131–5, 137–44 chitinolytic enzymes 74, 76, 79–80, 84, 88–92 glucanolytic enzymes 109, 110, 113 hydrolases 116, 119 mushrooms 267 mycoparasitism 153, 155–7, 159–61, 163–9 anthracnose 197, 254 antibiosis and antibiotics 139, 209 biocontrol 135, 136, 139, 140, 173–80 chitinolytic enzymes 76, 80, 82, 83, 84, 89 hydrolases 116 mushrooms 273 mycoparasitism 153, 168 plant growth and disease 186, 189, 235, 247–9, 252, 254, 260 recombinants 295 antibodies 56, 117, 138, 304 heterologous protein production 368–9, 370–4, 376–7 mycoparasitism 155, 159 antifungal properties 116, 119 antibiosis 173, 176, 177, 178–9 biocontrol 139 chitinolytic enzymes 81–4, 90–1 glucanolytic enzymes 102–3, 110–11 mycoparasitism 157, 161, 169 plant growth 186, 260 see also fungicides antigens 370–3 antitumour properties 91, 116 Aphanomyces entiches 239 Arabidopsis thaliana 112 arabinases 66, 336 arabinofuranose 33 arabinofuranosidases 34–5, 50, 65 hemicellulose degradation 26, 27, 33, 34–5, 40 338 INDEX arabinofuranoside 32, 35 arabinofuranosyl 35 arabinogalactan 25, 35 arabinoglucuronoxylan 25, 26, 35 arabinose 49, 52, 65, 66, 207 hemicellulose degradation 25, 31, 33, 35 arabinosidases 348, 352 arabinoxylans 25, 30–1, 33, 35–6, 327, 338 arabitol 52, 56, 66 arbuscular mycorrhizal (AM) fungi 136–7 ascomycetes 49, 102, 103, 111 asialomucin 155 aspartate proteases 114 Aspergillus 120 food industry 329, 330, 335 glucanolytic enzymes 103, 114 heterologous protein production 365, 367, 368, 374 polysaccharide degrading enzymes 58–9, 61, 66 recombinants 301, 304 Aspergillus aculeatus 38, 112, 334 Asperguillus nidulans 58, 62, 87, 102, 295, 370 Aspergillus niger 34, 113, 114, 295, 302, 328 heterologous protein production 366, 367, 369, 375, 376 Aspergillus oryzae 41, 62, 116 Aspergillus saitoi 116 Australian green mould 280–1, 285 autolysis 73, 88, 111 auxotrophy 193, 295 Bacillus 112, 245 Bacillus lautus 112 Bacillus polymyxa 112 Bacillus subtilis 328 backstaining 314–15, 316 basidiomycetes 79, 102, 103, 106, 111 Beauveria bassiana 214 beer and brewing 120, 311, 327–9 benomyl 193, 282, 366 benzimidazole 143, 268, 277, 282 biocontrol 89–90, 119, 131–45, 173–80, 230–60 chitinolytic enzymes 73–4, 76, 83, 84, 89–90, 91 glucanolytic enzymes 101 hydrolase 116, 117 lytic enzymes 161–3 mushrooms 282 mycoparasitism 140, 153, 156–7, 159–63, 165, 168–9 plant growth and disease 186, 188–9, 198, 229–61 339 production of Trichoderma 205–6, 209–12, 212, 216– 23 biofinishing 311, 317–23, 324 biopesticides 205, 216–23 biopolishing 311, 317, 318, 321, 324 biostoning 311, 312–17, 321, 322, 324 bleaching of pulp 345, 350–2, 353, 354–7 Botrytis 132, 139, 141, 142 Botrytis allii 174 Botrytis cinerea 160, 177–8, 255–60, 330 biocontrol 134, 135, 139 chitinolytic enzymes 79, 82, 83 glucanolytic enzymes 106, 108, 109, 111 hydrolases 118, 119 plant growth and disease 197, 251, 255–60 bread 120 Burkholderia solanacearum 112 Butyrivibrio fibrisolvens 112 Caldocellulosiruptor saccharolyticus 38 callose 101 Candida 102 Candida albicans 107, 112 captan 239, 240, 251, 256 carboxylates 9, 11, 14 carboxylic proteases 114 carboxymethylcellulose (CMC) 4, 259, 292 carboxypeptidases 114 carbon catabolite repression 79, 88, 212, 292, 293 glucanolytic enzymes 108, 114 hydrolases 114,115 polysaccharide degrading enzymes 50, 58–66 carbon sources antibiosis 176–7 biocontrol 134, 143 chitinolytic enzymes 78, 80 glucanolytic enzymes 108–9 heterologous protein production 366, 377 mannan 37 mutants and recombinants 292, 299 mycoparasitism 157, 161, 162 plant growth and diseases 193, 197 polysaccharide degrading enzymes 50–3, 55–8, 64–5 production of Trichoderma 206–7 proteases 114 catalytic domain 4–8, 38, 107, 314 cellulolytic enzymes 4–8, 12, 14, 16–18 cDNA 110, 111, 115, 162, 296–7, 299 cell wall degrading enzymes (CWDE) 340 INDEX antibiosis 179, 180 biocontrol 135, 136, 139 chitinolytic enzymes 73, 76, 80, 83, 84 food industry 328, 329 glucanolytic enzymes 101 mycoparasitism 156–7, 161, 165–6, 167, 168 cellobiohydrolases (CBHI, CHBII and CBHIII) 8–14, 60– cellulolytic enzymes 3–19 food industry 328 heterologous protein production 366–78 mutants and recombinants 291, 294, 296–301, 303 polysaccharide degrading enzymes 50, 55, 57–8, 60– 2, 64 proteases 115 pulp and paper industry 346–50, 354 textile industry 314, 316, 317, 321 cellobiose 299, 366 cellulolytic enzymes 4, 9, 17, 18 polysaccharide degrading enzymes 51, 53–6, 59, 61, 63 cellohexaose 14, 17 cellotetraose 13–14, 17 cellulases 16, 51–63, 313–19, 321–4 cellulolytic enzymes 3–5, 9, 14, 16–18 chitinolytic enzymes 76, 80, 91 feed industry 339 food industry 328–32, 334–7 glucanolytic enzymes 107, 108, 112 hemicellulose degradation 36, 38, 40 heterologous protein production 365–7, 373, 376–7 hydrolases 115, 118 mutants and recombinants 291–4, 297–302, 304 mycoparasitism 162 polysaccharide degrading enzymes 49–50, 61–64, 66– pulp and paper industry 343–4, 347–50, 352, 354–5, 357 textile industry 311–24 cellulolytic enzymes 3–19, 66, 76, 101, 291, 347 mycoparasitism 168 nomenclature 75 plant growth and disease 197–8 Cellulomonas fimi 11, 112 cellulose 3–8, 14–19 antibiosis 177 biocontrol 142 cellulolytic enzymes 3–8, 14–19 chitinolytic enzymes 78 food industry 327 glucanolytic enzymes 101, 103, 111 hemicellulose degradation 25, 28, 35–8 heterologous protein production 366, 370, 377 mutants and recombinants 291, 292, 298–9 mycoparasitism 162, 166 plant growth and disease 193, 245–8 polysaccharide degrading enzymes 49, 51–2, 54–9, 61– 3, 65–6 proteases 114 pulp and paper industry 343–5, 347–50, 354, 356 textile industry 311, 314, 317–23 cellulose binding domain (CBD) 4–5, 14–19, 27 hemicellulose degradation 36, 38, 39 heterologous protein production 367, 372 pulp and paper industry 346–7, 348, 349 textile industry 314, 315, 317 Chaetomium sp 191 chemical pulp 344–6 chemotropic growth 140, 153, 154 chitin 73–92 glucanolytic enzymes 106–11 mycoparasitism 157–9, 161, 162, 164–5, 167 proteases 115 chitinases 75, 82, 83, 86, 88, 101, 198 glucanolytic enzymes 107, 108, 110 hydrolases 118 mycoparasitism 156–7, 158–62, 165, 168–9 chitinolytic enzymes 73–92, 119, 158–9, 161, 162, 267 chitobiases 75 chitobiohydrolases 75 chitobiosidases 74–8, 81, 85, 177 chitodextrin 74 chitooligomers 75–6, 79–81, 90 chitosan 78 chitotriose 156 chlamydospores 82, 175, 205–6, 208, 231, 233 biocontrol 134, 140, 143 chlorine 345, 351, 353 cholesterol 176, 333 Chondrostereum purpureum 142, 160 chromatin 377 chymosin 296, 302, 368–71, 374–6 chymotrypsin 115, 372 cinerean 115 cloned genes 85–7, 316–17, 210 mycoparasitism 161, 162 Clostridium 112 Clostridium acetobutylicum 112 INDEX Clostridium cellulovorans 112 Clostridium thermocellum 112 coating and pelleting 235 Cocholiobolus carbonum 107, 108, 112 codon 295, 296 coiling 140, 155–6, 229 mycoparasitism 154, 160, 165–6, 169 Colletotrichum gloeosporioides 209, 257 Colletotrichum graminicola 254 colonization 119 biocontrol 131, 133–4, 136–8, 140–4 mushrooms 271, 275, 277–81, 283 mycoparasitism 160, 161, 165, 168 plant growth and disease 192–4, 196, 198–9, 233–40, 243–7, 250–3, 256–60 competition 140–2, 180 biocontrol 132, 139, 140–2, 144 mycoparasitism 153, 160, 168 plant growth and disease 248, 252, 260 compost 245–8, 252, 268, 270–1, 273–5, 277–85 contamination routes 282–3 conidia and conidiation 57–8, 63, 110, 279 antibiosis 174, 175, 177 biocontrol 134, 138, 140, 143 chitinolytic enzymes 82, 83 mycoparasitism 154, 155 plant growth and disease 199, 231, 233, 236, 241, 250, 252, 257–8 production of Trichoderma 205–16 conidiogenesis 210 conidiospores 206–7, 211, 214 costs food industry 328 mushrooms 282 plant growth and disease 231–2, 238, 243, 244, 260 production of Trichoderma 205, 207, 208, 212, 214 pulp and paper industry 343, 351, 355 recombinants 298, 304 textile industry 316 Cryphouectria parasitica 62 Cryptococcus albidus 337 cutinases 35 cyclic-AMP 60, 63 cycloheximide 157 defibrillation 312, 317–23, 324 de-inking 345, 355, 356 delivery systems 230, 231, 235, 251, 257–8, 260–1 bees 257–8 341 denim 311, 312–17, 322, 324 dental disease 118 deoxyglucose 292 detergents 90, 311, 312, 313, 315–16, 317 deuteromycetes 102, 103 dextrinases diacetylchitobiose 74, 75 dialysates 174 dicarboximides 143, 251, 259 Dictyostelium discoideum 112 diketopiperazines 175–6 dimethylgliotoxin 248 disaccharides 18, 49, 108, 117, 155, 217 polysaccharide degrading enzymes 54, 55 disease of plants 131–45, 161, 175, 178–80, 229–61 resistance 142–5, 185–200, 230, 248, 252, 256, 260 dissolved and colloidal substances (DCS) 355–6 disulfide bridges 5, 9, 15 DNA chitinolytic enzymes 79, 87 drying 218 mushrooms 268, 269–70, 278, 281, 282, 284 mutants and recombinants 291, 295–7, 301 polysaccharide degrading enzymes 62, 64, 65 endo-arabinases 336 endocellulases 8, 314, 316, 322–4, 336 endochitinases 111, 115, 177, 369, 375 chitinolytic enzymes 74–9, 81–2, 85–8, 90–2 mycoparasitism 157–9, 161, 169 endoglucanases (EG) 104, 107–10, 112–13, 120, 162 cellulolytic enzymes 3–12, 16–18 food and feed industries 328, 334, 340 hemicellulose degradation 28, 37, 39 heterologous protein production 373 mutants and recombinants 292, 294, 298–301 polysaccharide degrading enzymes 50, 55, 57, 63 pulp and paper industry 347, 348–9, 350, 352, 354, 356 textile industry 314, 316–17, 322–3 endoglycosidases 375 endohydrolases 114 endomannanases 336 endopectinases 336 endopeptidases 296, 374, 375 endoplasmic reticulum (ER) 292, 296 heterologous protein production 367, 373–4, 378 endopolygalacturonases 337 endoxylanases 31, 63, 351 342 INDEX Enterobacter sp 119, 245, 247 Enterobacter agglomerans 161 Enterobacter cloacae 84, 91 environmental conditions 304 antibiosis 176–8, 180 biocontrol 133–5, 138, 139, 141, 144 plant growth and disease 193, 198, 199, 259, 260 production of Trichoderma 205, 207, 210, 212, 213 enzymes antibiosis 177–80 biocontrol 135, 136, 139 cellulolytic 3–19 chitinolytic 73–92 food and feed industries 311, 327–40 glucanolytic 101–14 hemicellulose degradation 25–42 heterologous protein production 365, 375, 376 hydrolytic 114–21 lytic 153–69 mutants and recombinants 291–2, 294–300, 303–4 plant growth and disease 186, 197–8, 247, 254, 260 polysaccharide degrading 49–67 pulp and paper industry 343–57 textile industry 311–24 Epicoccum purpurascens 257 epidithiodiketopiperazines 186, 188 Erwinia chrysanthemi 112 erythrocytes 154 Escherichia coli 89, 154, 161, 175, 294 heterologous protein production 365, 370, 376 polysaccharide degrading enzymes 60–1, 62, 64 esterases 26–7, 33, 35–7, 41, 65, 329, 348 ethanol 291, 292 ethylene 32, 197 eukaryotes 41, 59, 86, 365 recombinants 296, 297 exocellulases 314, 323, 324 exochitinases 75 exoglucanases 40, 104, 107, 108, 112–13, 166 cellulolytic enzymes 3–4, 6–7, 12, 18 exponential sporulation stage (ESS) 217 expression vectors 294–7 Fab molecules 368, 370–4, 376 fatty acids 247, 334 feed industry 291, 297, 337–40 additives 120, 121 conversion rates 338–40 fibres properties 353, 354–5 fibrillation pulp and paper 344–5, 350, 352, 354 textile industry 312, 317–23, 324 Fibrobacter succinogenes 112 Flavobacterium 245, 247, 248 Flavobacterium balustinum 247, 252 flies on mushrooms 275, 277, 280, 283 fluorescein diacetate (FDA) 154 Fomes annosus 177 food industry 90, 119–20, 291, 311, 327–41, 365 hydrolases 116, 118, 119–20, 121 shelf life 219, 220 fructose 51, 57, 108 fucose 154 fungicides 119, 169, 177 biocontrol 131–2, 134, 142–3, 144–5 chitinolytic enzymes 83–4, 91, 92 mushrooms 267, 268, 277, 282 plant growth and disease 188, 230–2, 238–40, 243–4, 247, 250–1, 253–60 see also antifungal properties fungistasis 134, 143 Fusarium spp 82, 103, 118 biocontrol 132, 135, 136 plant growth and disease 245, 247–50, 252 Fusarium oxysporum 83, 142, 161, 175, 178, 179 Fusarium roseum 191 fuzz and pill formation 312, 317–23 galactoglucomannan 25, 30, 36–7, 39, 40, 41 galactomannan 37–8, 39, 40 galactose 49, 154, 165, 176 hemicellulose degradation 25, 36, 39, 40, 41 polysaccharide degrading enzymes 52, 56, 57, 65–6 galactosaminidases 76 galactosidases 27, 37, 40–1, 56, 65–6, 348 gene copy number 301–2 genetic engineering (bioengineering) 179, 348, 291–304 biocontrol 161, 162 cellulolytic enzymes 4, 14 heterologous protein production 365–6 plant growth and disease 192, 199 textile industry 311, 316, 321, 323 genomic DNA 296–7 gentiobiose 54 Gigaspora margarita 91 Gliocladium 4, 26, 205 antibiosis 173–4, 176, 177, 178–9 biocontrol 131–45 INDEX chitinolytic enzymes 73–6, 78, 80–92 glucanolytic enzymes 101, 104, 108, 111 hydrolases 114, 117, 118, 119, 120 plant growth and disease 185–200, 229–61 polysaccharide degrading enzymes 49, 50 Gliocladium intraradices 136 Gliocladium roseum 76, 134, 174, 257, 260 Gliocladium virens 136, 168 antibiosis 174,175,176 chitinolytic enzymes 76, 80, 85 plant growth and disease 186–9, 197, 229, 235, 248 gliotoxin 83, 89, 139, 174–8 plant growth and disease 186, 188–9, 229, 248–50, 252 gliovirin 139, 176, 178 plant growth and disease 186, 188, 189 Glomus mosseae 136, 137, 192 glucan 49, 51, 52, 83,179 feed industry 338–40 food industry 327, 328, 329, 330 glucanolytic enzymes 101–4, 106, 108, 110–11, 113, 120 hydrolases 115, 118 mycoparasitism 154–6, 162, 166, 168 glucanases 50, 101, 102–3, 104–14, 155, 375 antibiosis 177, 179 applications 118, 119, 120 chitinolytic enzymes 76, 80, 82, 91, 92 feed industry 338–40 food industry 327–30, 335 genes 107–8 hemicellulose degradation 28, 37 mycoparasitism 156–7, 160, 166, 168 plant growth and disease 198 recombinants 298 glucanolytic enzymes 76, 83, 101–14 applications 118–21 glucoamylases 7, 9, 103, 105, 113–14, 296–7 heterologous protein production 366–7, 369, 374–6 glucomannan 25, 37, 39, 49 pulp and paper industry 351, 356 glucooligomers 49 glucopyranosyl 36, 37, 39, 40, 217 glucosamines 156, 157, 165 chitinolytic enzymes 73, 75, 78, 87, 90 glucosaminidases 75, 76, 78, 118 glucosaminide 75 glucose 57, 58–9, 207, 327 antibiosis 176, 178 343 cellulolytic enzymes 3, 4, 9, 12–13, 17 chitinolytic enzymes 79–80, 87, 88 glucanolytic enzymes 101, 105, 106, 108–10, 113–14 hemicellulose degradation 25, 36, 37, 39 heterologous protein production 366, 367, 377 hydrolases 117 mutants and recombinants 291, 292, 296 mycoparasitism 154, 157, 161, 162 polysaccharide degrading enzymes 49–53, 56–9, 61–6 glucosidases 39–40, 83, 314, 348 cellulolytic enzymes 4, glucanolytic enzymes 104, 107, 111, 113, 114 hemicellulose degradation 27, 33, 37, 39–40 hydrolases 119 mutants and recombinants 292, 293–4, 303 polysaccharide degrading enzymes 50–1, 54–5, 57, 66 trehalases 117 glucosides 292 glucosyl 8, 101 glucuronidases 34, 65, 138, 254, 348 hemicellulose degradation 26, 27, 33, 34, 36 glucuronoxylan 25, 26, 30, 31, 32, 34, 41 glycan 372, 378 glycanases 28, 29, 38 glyceollin 136, 196 glyceraldehyde phosphate dehydrogenase 367 glycerol 108, 176, 207, 208, 292 polysaccharide degrading enzymes 51, 53, 55–6, 58, 61, 63, 66 glycogen 101, 113 glycoproteins 154, 155 glycosidases 9, 117, 329 glycosides 119 glycosyl hydrolases 4, 7, 9, 10 hemicellulose degradation 27, 28, 33, 34, 41 glycosylation 108, 114, 197 chitinolytic enzymes 81, 86 heterologous protein production 365, 367, 374–5, 378 gnotobiotic conditions 191, 195, 229 grapes with B.cinerea 257–9, 260, 330 green mould Australia 280–1 Ireland 270–80 North America 280 Hansenula 376 harzianolide 175 harzianopyridone 188 hemicellulases 27, 63–6, 352 344 INDEX feed industry 340 food industry 329–31, 334, 335, 337 mutants and recombinants 291, 294, 304 polysaccharide degrading enzymes 49–50, 63–7 pulp and paper industry 344, 347–8, 350–2, 354–5, 357 hemicellulose 25–42, 101, 329 polysaccharide degrading enzymes 49, 63, 65, 66 pulp and paper industry 343–8, 350–1, 355, 356 heterologous proteins 50, 365–78 recombinants 297, 299, 302–4 heterooligosaccharides 30, 66 heteropolysaccharides 25, 49 hexosaminidases 85, 88 homologous proteins 368, 376, 377 recombinants 297, 299, 301, 304 homopolymers 49, 101 Hordeum vulgaris 112 Hormoconis resinae 114, 296, 369, 374, 375 hormones 241 Humicola insolens 315, 354 humidity see moisture and humidity hydrolases 2, 114–18, 366 applications 118–21 chitinolytic enzymes 81 feed industry 338 glucanolytic enzymes 104, 108, 110–12 hemicellulose degradation 25, 32, 33, 41 polysaccharide degrading enzymes 52–3 pulp and paper industry 343, 346, 354, 356 hydrolytic enzymes 114–21, 139 antibiosis 177, 179 feed industry 338, 340 food industry 327, 328, 334 mycoparasitism 165, 168 pulp and paper industry 346, 356 hydrophobicity 210, 212, 218 hydrophobin 209, 210, 218, 367 hydroxyethylcellulose (HEC) 4, 298, 301 hygromycin 254, 295, 366 hypercellulolytic mutants 292 hyphae biocontrol 134, 136, 140, 143 chitinolytic enzymes 82, 83–4, 88 glucanolytic enzymes 111, 112 heterologous protein production 371, 375, 377–8 hydrolases 118, 121 mushrooms 278 mutants and recombinants 294, 304 mycoparasitism 154–8, 161–2, 164–9 plant growth and disease 193, 229, 231, 233–4, 236, 239, 254, 256 production of Trichoderma 205, 207, 208, 213 hypochlorite 196, 316 Hypocrea 267 indigo 312, 313, 314–17, 318, 324 Indigofera tinctoria 312 inducer paradigms 51, 54 inert support culture (OSC) 212 inocula 160, 213, 304 biocontrol 131, 132, 137, 140, 141 mushrooms 277, 280, 283 plant growth and disease 186, 188–90, 193, 197 insecticides 143, 214 integrated control 142–3 integration site 301–2 interleukin 369, 374 invertases 366 iprodione 251, 254–6, 259 Ireland 270–80 Isatis tinctoria 312 isonitriles 174 isozymes 81, 106, 109, 110, 113 karyotypes 155 Koninginin 188 kraft pulping 344–6, 350–4, 356 laccase 76, 297, 369, 375 lactonases 118 lactose 52, 55–7, 64, 65, 298–300, 302 laminaribiose 54 laminarin 78, 106, 108–10, 115 laundries and laundering 311–14, 316, 317, 320, 324 lectin 79, 80, 140 mycoparasitism 154–8, 165, 168–9 legal considerations 232 Lentinus edodes 267 ligands 8, 11–13, 14 lignicolous ascomycetes 49 lignin 343–5, 349–50, 351, 356, 375 lignin peroxidases 369, 375 linkers 4–5, 16, 38, 367–70, 372, 374 lipases 76, 101, 108, 118, 198, 355 lipids 178 liposomes 178 long-term root protection 232–3, 237–43 INDEX lyases76, 118, 119, 329, 336 Ly coper sicon esculentum 112, 241 lyocell 311, 317, 320–3 lysis 102, 103, 110,111,119 lysozyme 8, 9, 376 lytic enzymes 81–4, 101, 118, 120, 140, 156–63 mycoparasitism 153–69 macerating enzymes 330–2, 334–7 maltooligosaccharides 114 mannan 36–41, 49 hemicellulose degradation 25, 37, 38–41 mannanases 4, 37–9, 40, 118, 299–301, 365 chitinolytic enzymes 76 food industry 336 hemicellulose degradation 27, 30 pulp and paper industry 346, 348, 350–2, 354–7 mannobiose 38, 39 mannooligosaccharides 39, 41 mannopyranosyl 36–7, 39 mannose 25, 36–40, 154, 365 polysaccharide degrading enzymes 49, 52, 53, 57 mannosidases 37, 39–40 mannotriose 38, 39 markers 138, 371, 377 recombinants 295–6, 297–8, 300, 303 mechanical pulping 344–5, 349–50, 355, 356 medicine 90–1, 118, 121, 304 see also pharmaceuticals membrane affecting compounds (MAC) 73, 83, 84 metalloproteases 114 mice 277, 278, 283 Microbispora bispora 112 microcycle conidiation 207, 209 moisture and humidity 133–4, 143–4, 218–19, 330 mushrooms 268, 271, 275 plant growth and disease 235–7, 239–40, 245–6, 259 production of Trichoderma 205, 207–8, 210–11, 213– 21, 223 monosaccharides 7, 25, 49, 57, 155 morphogenesis 73, 88–9, 115 glucanolytic enzymes 102–3, 104, 106, 110–11 mRNA79, 87, 110, 157, 162 heterologous protein production 370, 373, 374, 375 polysaccharide degrading enzymes 50, 55–7, 61, 65 recombinants 295, 296, 297, 299 mucin 155 Mucor 103 murine antibodies 368, 372 345 mushrooms 135–6, 189, 267–85 chitinolytic enzymes 73, 89 mutagenesis 9, 17, 115, 178–9, 366, 376 industrial T reesei 291, 292, 294, 299, 302–4 mutanases 76, 118 mutants and mutation 25, 161, 292–4, 316 antibiosis 178–9, 180 cellulolytic enzymes 11–14, 16, 17 chitinolytic enzymes 90, 91 heterologous protein production 366, 374 hydrolases 115, 119 industrial T reesei 291–304 plant growth and disease 193, 199, 230 polysaccharide degrading enzymes 50, 54–5, 57, 59, 61, 64–6 mycelia antibiosis 175, 177 biocontrol 134, 136, 137, 138, 140 chitinolytic enzymes 78, 80, 87, 89 glucanolytic enzymes 104, 108, 111 heterologous protein production 375, 378 mushrooms 271, 274, 275, 278, 281 mycoparasitism 154, 155, 157, 158, 168 plant growth and disease 197, 198 polysaccharide degrading enzymes 51, 54, 57, 62–5 production of Trichoderma 210, 213 mycoherbicides 199 mycoparasitism 140 antibiosis 173, 180 biocontrol 131–2, 136, 139–40 chitinolytic enzymes 73–4, 78–80, 82–3, 87–8 glucanolytic enzymes 102–4, 106, 109, 111 hydrolases 115, 117 lytic enzymes 153–69 plant growth and disease 185, 247–8, 252, 254, 259– 60 mycorrhizae 135, 136–7, 185, 192, 195 Myxococcus xanthus 107 necrotic tissue 141, 160 nematodes 163, 275 Neolentinus lepideus 177 Neurospora crassa 102, 193, 295 Nicotiana plumbaginifolia 112 Nicotiana tobacum 112 nigeran 108, 109 nitrogen sources 80, 114, 207–8 antibiosis 176–7 biocontrol 134, 143 346 INDEX mushrooms 270, 275 mutants and recombinants 292, 295 plant growth and disease 241, 242–3, 246 non-starch polysaccharides (NSP) 327, 338–40 North America 280 nucleases 116–17 nucleic acid hydrolases 116–17 nucleophiles 9–10, 14, 29 nucleotides 59, 64, 73, 80, 86, 116 nutrient status 133, 134–5 odour 139, 174, 229, 329–30, 334 mushrooms 269, 275, 278 Oerskovia xanthineolytica 112 Olea europea 333 oligogalacturonides 197 oligoglucosides 114 oligomers 79, 91, 105, 156, 165, 347 oligonucleotides 62, 64, 107, 116 oligosaccharides 51, 56, 65, 104, 299, 356, 366 cellulolytic enzymes 4, 8, 11, 13–14, 17 hemicellulose degradation 26–7, 31–5, 37, 39–40 oligotrophs 246 olive oil 311, 329, 332–7 omycetes 101–3, 162 packed-bed bioreactors 214 patents 138, 206, 232, 330 textile industry 311, 313, 316, 317 pathogenesis-related (pr) protein 73, 91, 197 pathogens 135 antibiosis 173–4, 176–9 biocontrol 131–3, 135–44 chitinolytic enzymes 73–4, 82, 84, 90–2 glucanolytic enzymes 101, 113 hydrolases 116, 118, 119 mushrooms 267–85 mycoparasitism 153–5, 157–63, 165–6, 168–9 plant growth and disease 229, 231–2, 236–7, 245–61 resistance 185–200 xylanases 32 pectin 49, 117–18, 139, 260 food industry 329, 334 pectinases 76, 117–18, 119, 355 food industry 329–32, 334–7 pectin esterase 329 pectin lyases 76, 117, 329, 336 Penicillium spp 192, 257 glucanolytic enzymes 102, 109, 113 Penicillium emersonii 328 Penicillium expansum 189 Penicillium italicum 110 pentosan 327, 338 pepstatin 114–15, 302–3 peptaibols 83, 89, 175, 178, 179 peptidases 107, 327 peptides4–5, 15, 38, 85, 87, 175 glucanolytic enzymes 108, 114 proteases 114–15 recombinants 295, 303 peptones 81, 370 permeases 80 Per sea americana 112 pesticides 131, 143 plant growth and disease 244, 251, 256, 258, 260 pH 81, 112–13, 117, 137, 177, 304 food and feed industries 334, 340 heterologous protein production 368, 369–70 mushrooms 271, 275, 281, 284 plant growth and disease 186, 188, 193, 198, 229, 231, 246 production of Trichoderma 206–7, 213, 215–16, 218 pulp and paper industry 349, 352, 355 textile industry 314–15, 316, 317, 319, 320 pharmaceuticals 219, 220, 378 see also medicine Phaseolus vulgaris 112 phenol 248, 280, 281, 282 phenylalanine 15, 176 phialides 269, 275, 278 phialoconidia 212 phialoconidiogenesis 207 phialophores 278 phialospores 269 Phlebia radiata 297, 369 phleomycin 295, 366, 367, 371, 376 Pholiota nameko 267 phorids 283 phosphatases 101, 118 phosphoglycerate kinase 367 phosphorylation 86 phylogenetic tree 112 phytase 338, 369, 375 phytoalexin 136, 196 phytopathogens 42, 91, 104, 109, 113, 206 antibiosis 177 biocontrol 138, 144 mycoparasitism 156, 161, 162 INDEX Phytophthora 132, 175 plant growth and disease 245–6, 247, 249, 250, 252 Phytophthora cinnamomi 117 Phytophthora citrophthora 108, 109, 113, 117 phytosterol 197 phytotoxocity 173, 179, 185–8 antibiosis 174, 176, 179 plant growth and disease 185–8, 189, 190, 199 Pichia 376 plant exudates 141–2 plant growth and disease 185–200, 229–61 biocontrol 132, 136, 137 chitinolytic enzymes 90–1, 92 planter box treatments 235, 237–40, 249 Plebia radiata 375 Pleurotus 267 Pleurotus ostreatus 271 polyethylene glycol (PEG) 207, 208 polygalacturonases 117, 197, 329 polyketides 175 polymerases 65 polymers cellulolytic enzymes 3, 7, 11, 17 chitinolytic enzymes 73 food industry 328 glucanolytic enzymes 101, 110, 113 hemicellulose degradation 32, 34–6, 37, 39–40 hydrolases 115, 118, 119 mycoparasitism 155, 157 polysaccharide degrading enzymes 49, 51, 64, 66 pulp and paper industry 347 textile industry 323 poly phenol 334, 337 polysaccharides 49–67 cellulolytic enzymes 3, 8, 12, 13, 18 feed industry 341 food industry 327, 329, 330 glucanolytic enzymes 101, 113, 120 hemicellulose degradation 25–7, 30–1, 33, 35, 41 mycoparasitism 154, 165 pulp and paper industry 356 polyvinyl alcohols 244 post-fermentation processing 216 preprochymosin 369 prochymosin 369 production methodology 205–23, 230, 231, 251, 260 programmable logic controller 215 prokaryotes 86, 365 promoters 296, 299–301 347 heterologous protein production 366–9, 371, 375–7 recombinants 291, 294–301, 304 propeptides 368 propiconazole 253 proprietary information 232 proteases 15, 101, 114–16, 118, 302–3, 304, 316 chitinolytic enzymes 76 feed industry 338, 340, 341 heterologous protein production 366–7, 369–70, 372, 374, 376–7 mycoparasitism 155, 156, 162 plant growth and disease 198 protein disulfide isomerase (PDI) 374 proteinases 80, 81, 108, 115 mycoparasitism 157, 162, 163 proteins 299–303, 365–78 cellulolytic enzymes 6, 8, 11, 16, 17 chitinolytic enzymes 73, 78, 80, 83, 85–8, 91 food industry 327 glucanolytic enzymes 105, 107–9, 111, 114, 120 hemicellulose degradation 34 hydrolases 114, 117, 118 mutants and recombinants 291–2, 294–304 mycoparasitism 154–5, 159–60, 162–3 plant growth and disease 197–8 polysaccharide degrading enzymes 49–50, 55, 58–65 production of Trichoderma 210, 217, 218 pulp and paper industry 343, 346–7, 349 proteolysis 14, 50, 109, 115, 302–3, 316 proteose 81 proton donors 9–11, 14, 29 protoplasts 90, 106, 112 biocontrol 135, 142 plant growth and disease 193, 199 prototrophy 295 pseudomonads 246 Pseudomonas 245, 247 Pseudomonas fluor escens 112 pulp carbohydrates 347–9, 356 pulp and paper industry 291, 304, 343–57 pustulan 108, 109, 110, 115 Pygmephorus mesembrinae 271, 273 Pyrenochaeta terrestris 190 pyrones 188 pyruvate kinase 367 Pythium 119, 132, 142 mycoparasitism 162, 165–6 plant growth and disease 195, 235–6, 243, 245–7, 249– 50, 252–3 348 INDEX Pythium graminicola 253, 254 Pythium ultimum 135, 141, 162, 165, 166 antibiosis 175, 176, 178, 179 plant growth and disease 191–2, 193, 236–7, 248 rDNA 284, 294, 311 recognition 140, 153–5, 158 recombinants 50, 54, 291–304, 329, 375–6 textile industry 311–12, 316, 323–4 recycling pulp and paper 345, 355, 356 red pepper mites 268, 271, 273–5, 277–81, 283 residual moisture content (RMC) 218–19 rhizobia 185 Rhizobium 219 Rhizobium meliloti 161 Rhizoctonia 132, 154, 245, 250, 252 Rhizoctonia solani 91–2, 135, 138 antibiosis 175, 176, 177, 178–9 glucanolytic enzymes 108, 110, 113 hydrolases 115–19 mycoparasitism 154–9, 161–2, 164–5, 169 plant growth and disease 190, 197, 246–9, 253–4, 259 Rhizopus 103 Rhizopus nigricans 191 Rhizopus niveus 116 rhizosphere competence 192–4, 195, 197, 199, 249–50 biocontrol 144 plant growth and disease 230, 232–3, 237–9, 241–3, 249–50, 253, 260 Ricinus communis 165 ring distortion 12 RNA 53, 56, 65, 109, 116–17, 162, 218 RNases76, 116 Robillarda sp 112 rRNA 117 Ruminococcus albus 112 Saccharomicopsys fibulligera 103 Saccharomyces 376 Saccharomyces cerevisiae 38, 40, 59, 62 glucanolytic enzymes 102–3, 106–7, 109–10, 112, 120 heterologous protein production 365, 367, 370, 374 Salmonella 339 saprophytes 49, 89–90, 104, 110, 119, 190–1 biocontrol 131, 135, 136, 138, 140–1 chitinolytic enzymes 79, 84, 88, 89–90 scaling-down 209 scaling-up 208–9, 212–13, 217 Schizophyllum commune 41, 102–3, 106, 112 sciarids 283 sclerotia 82, 132, 139, 246 mycoparasitism 157, 167, 168 Sclerotinia 132, 141, 249, 252 Sclerotinia homoeocarpa 253, 254, 255 Sclerotium 252 Sclerotium cepivorum 179 Sclerotium rolfsii 135, 249 antibiosis 174, 175, 176 mycoparasitism 154–5, 157–61, 167–9 Sclerotium sclerotiorum 134 secondary metabolites 139, 186, 188, 189 antibiosis 173, 176, 178, 179 secretion signal 296 seed treatment 119, 137, 141, 232–44 serine esterases 35 serine proteases 114, 115, 116 Serratia marcescens 161 sesquiterpenes 176 shelf life 217–33, 331, 337 disease control 231, 244, 260 production of Trichoderma 205, 207, 210, 216, 217– 23 sialidases single-chain antibodies (SCA) 373 slurry and film coating 235–9, 244 Solanum lycopersicoides 241 solid-state fermentation (SSF) 210–16 sophorose 28, 299, 366 polysaccharide degrading enzymes 51, 53–8, 61–4, 66 sorbitol 51, 52–3, 55–6, 58, 61, 66 sorption isotherm 221 specific energy consumption 350 sporangia 141, 175 sporulation 57, 80, 87, 217 biocontrol 132, 141 glucanolytic enzymes 102, 103, 107 mushrooms 267, 275–8, 281, 283 production of Trichoderma 206, 207, 210, 213 Stachybotrys 102 Staphylococcus aureus 175 starvation 101, 206 chitinolytic enzymes 79, 80, 87, 89 stationary sporulation stage (SSS) 217 steroids 176 sterol 73, 179, 186, 188, 189 stirred bioreactors 214 strawberries wth B.cinerea 256–7, 258, 259, 260 Streptomyces 112, 245 INDEX Streptomyces halstedii 112 Streptomyces lividans 36, 38, 112 stress protectants 217–18 submerged fermentation 207–9, 210, 211–13 substrates 192, 344–6 antibiosis 176, 177 biocontrol 132, 140, 142 cellulolytic enzymes 3, 4, 8–9, 11–12, 14, 16–17 chitinolytic enzymes 76, 77, 80–1, 85, 90 feed industry 340 glucanolytic enzymes 105, 106, 107 hemicellulose degradation 29, 31, 32, 34, 40 hydrolases 117 mushrooms 268 mutants and recombinants 292, 294, 296, 300 mycoparasitism 154 polysaccharide degrading enzymes 50, 51, 57, 66 production of Trichoderma 206, 210–16 pulp and paper industry 343, 344–6, 347, 349, 352 textile industry 319 sucrose 79, 292 sulphite pulping 344–6 sugars 64, 120, 153, 298, 316, 340, 347 cellulolytic enzymes 8, 12, 13–14 mushrooms 267 plant growth and disease 198, 245, 247, 248 production of Trichoderma 207, 217, 218 sulfur sources 114 superior biocontrol strains 230–1, 251 targeted integration 297 taxa 278–9 mushrooms 268–9, 271–3, 275–85 teleomorphs 49 temperature 81, 133, 177, 273, 304 biocontrol 133, 134, 144 feed industry 339 food industry 330, 334–5 mushrooms 268, 270–1, 273, 275, 277–9, 282–5 plant growth and disease 193, 198, 235, 238, 245–6, 257, 259 production of Trichoderma 205, 207, 211, 213–16, 218–23 pulp and paper industry 349 textile industry 314, 315, 319, 320 terminators 295, 296 terpenes 119, 329 textile industry 90, 291, 311–24 Thelaviopsis 249, 252 349 Thelaviopsis basicola 249 thermomechanical pulping 349–50, 356 Thermonospora fusca 112 thigmotropism 153 thiol 114 tochopherol 334, 337 toxicity 138, 365 plant growth and disease 229, 232, 239–40, 253, 256 transglycosylation 51, 54, 57 tray bioreactors 213–14 trehalases 117 trehalose 117, 217 triadimefon 254, 255 Trichoderma antibiosis 173–9 biocontrol 131–45 cellulolytic enzymes 3–19 chitinolytic enzymes 73–92 food and feed industries 327–40 glucanolytic enzymes 101–2, 104–14 hemicellulose degradation 25–42 heterologous protein production 365–78 hydrolases 114–20 identification 268–70, 275, 283, 284 industrial production 205–23 mushrooms 267–85 mutants and recombinants 291–304 mycoparasitism 153–68 plant growth and disease 185–200, 229–61 polysaccharide degrading enzymes 59–67 pulp and paper industry 343–4, 347, 350–1, 354–7 textile industry 311–24 Trichoderma atroviride 76, 86 mushrooms 268, 269, 276, 278, 281, 283 Trichoderma aureoviride 76, 110, 137, 192 Trichoderma citrinoviride 269 Trichoderma hamatum 119, 138, 154, 155, 174 chitinolytic enzymes 76, 85, 86, 87 mushrooms 267, 281 plant growth and disease 190, 197, 246–8, 252 Trichoderma harzianum 77, 82, 249–50, 273–81 antibiosis 174, 175, 177, 179 biocontrol 134–9, 142 chitinolytic enzymes 76, 77, 78–9, 80, 82–9, 91–2 food industry 330 glucanolytic enzymes 104–12, 113, 118–20 hemicellulose degradation 28, 29, 37 heterologous protein production 369, 375 hydrolases 114–17 350 INDEX industrial production 206–7, 210–12, 217–18, 221–2 mushrooms 267–9, 271–85 mycoparasitism 154–69 plant growth and disease 188–9, 191–9, 230, 233–9, 241–3, 245–6, 248–61 polysaccharide degrading enzymes 58, 59 pulp and paper industry 352, 353 Trichoderma inhamatum 281, 284 Trichoderma koningii antibiosis 174, 175, 176, 177 biocontrol 136, 137, 138 chitinolytic enzymes 76, 86 glucanolytic enzymes 106, 109, 113 hemicellulose degradation 28, 34 mushrooms 267, 269, 281, 284 plant growth and disease 188, 190–1, 193, 195, 197 polysaccharide degrading enzymes 50, 60 Trichoderma lignorum 28, 118, 174, 189, 229 Trichoderma longibrachiatum 28, 119, 162, 284, 353 chitinolytic enzymes 76,86 glucanolytic enzymes 105, 106, 108, 109, 113 polysaccharide degrading enzymes 50, 60 Trichoderma longipilis 76 Trichoderma minutisporum 76 Trichoderma parceramosum 281 Trichoderma polysporum 174, 175, 193, 281 Trichoderma pseudokoningii 28, 76, 110, 284 Trichoderma reesei 56–7, 367–76 cellulolytic enzymes 3–19 chitinolytic enzymes 76, 81, 86, 91 food and feed industries 328, 339, 340 glucanolytic enzymes 103, 105–6, 109, 111–14, 120 hemicellulose degradation 25–41 heterologous protein production 365–78 hydrolases 114–15, 117, 118 mutants and recombinants 291–304 plant growth and disease 193 polysaccharide degrading enzymes 50–66 pulp and paper industry 343–57 textile industry 315–17, 319, 321–2 Trichoderma virens antibiosis 174–9 chitinolytic enzymes 76, 80, 82, 83, 85, 86 glucanolytic enzymes 106, 108, 109, 111, 113 plant growth and disease 186, 197, 229, 245, 248–9, 252–3 Trichoderma viride 190–1 antibiosis 174, 176 biocontrol 134, 137, 142 chitinolytic enzymes 76, 86 glucanolytic enzymes 105–6, 109–10, 112–14, 120 hemicellulose degradation 28, 29, 32, 34, 35 hydrolases 115–17 mushrooms 267–9, 277, 278, 281, 284 plant growth and disease 188–91, 193, 197, 229, 257 polysaccharide degrading enzymes 50 trichodermapepsin 302, 303 trichodermin 174 trichorzianines 83, 175, 177–9, 210 Trichothecium roseum 257 tryptophan 8–9, 12–14, 15, 17 turf diseases 251–5 tyrosines 12, 15 Uncinula necator 82 Ustilago avenae 82 Venturia inequalis 91 Verticillium chlamydosporium 137 Vinca 242–3 viridin 139, 174, 176, 186, 188, 189 viridiol 186, 188, 189, 248 antibiosis 174, 177, 179 wine 119, 311, 327, 329–32, 334 wounds 131, 142, 143, 160 Xantomonas campestris 112 xylan 25, 26–36, 37, 300 plant growth and disease 193, 197–8 polysaccharide degrading enzymes 49, 51–3, 56, 63–6 pulp and paper industry 347, 351–2, 354, 356 xylanases 4, 8, 26, 28–32, 197–8, 374 feed industry 338–40 hemicellulose degradation 26–7, 28–32, 33–7 polysaccharide degrading enzymes 50, 63, 64–6 promoters 64–5 pulp and paper industry 346, 348–57 recombinants 299–300, 304 xylitol 52 xylobiose 28, 31, 35, polysaccharide degrading enzymes 51, 53, 56, 63, 65– xylooligosaccharides 31–5, 41, 65 xylopyranosides 31, 32, 35, 36 xylopyranosyl 31, 32–6 xylose 12, 25, 31–3, 35, 177, 197 polysaccharide degrading enzymes 49, 52, 63–4, 66 INDEX xylosidases 32–3, 40, 50, 65–6 hemicellulose degradation 26–7, 32–3, 34–6 pulp and paper industry 348, 352 zoospores 175, 239 zygomycetes 103 351 ... D.Seaby 23 5 12. 1 Introduction 23 5 12. 2 Green mould epidemic in Ireland 23 7 12. 3 A Trichoderma harzianum group green mould epidemic in North America 24 6 12. 4 Green mould in Australia 24 7 12. 5 The Trichoderma. .. mushroom casing 24 7 12. 6 Summary and discussion 24 8 12. 7 Appendix 25 0 References 25 1 12 PART THREE Protein Production and Application of Trichoderma Enzymes 13 25 4 Industrial mutants and recombinant.. .Trichoderma and Gliocladium Volume Trichoderma and Gliocladium Volume Enzymes, biological control and commercial applications Edited by GARY E.HARMAN Cornell University, Geneva, NY, USA and