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Development of Myrothecium Verrucaria As a Mycopesticide for the Control of Plant Pathogenic Fungi and Insects

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Development of Myrothecium Verrucaria As a Mycopesticide for the Control of Plant Pathogenic Fungi and Insects Development of Myrothecium verrucaria as a mycopesticide for the control of plant pathogenic fungi and insects A Thesis submitted to the University of Pune For the Degree of Doctor of Philosophy In Microbiology By Medha Prashant Kulkarni Division of Biochemical Sciences National Chemical Laboratory Pune-411 008, India February 2007 1 Table of contents Sr.No. Contents Page No. List of tables 7-8 List of figures 9-10 Acknowledgement 11 Declaration 12 List of abbreviations 13 Abstract 14-23 1. Chapter 1 Introduction 24-72 1.1 Introduction 25 1.1.1 Pests and pathogens in agriculture 25 1.2 Control of fungal pathogens and insect pests 27 1.2.1 Chemical pest control 27 1.2.1.1 Fungicides 27 1.2.1.2 Insecticides 29 1.2.1.3 Disadvantages of insecticides 31 1.2.2 Biological control agents 32 1.2.2.1 Viruses as biological control agents 33 1.2.2.2 Bacteria as biocontrol agents 34 1.2.2.3 Fungi as biocontrol agents 36 1.3 Host-pathogen interaction 40 1.3.1 Structure of fungal cell wall and insect cuticle 40 1.3.2 Fungus-fungus interaction 43 1.3.3 Fungus-insect interaction 44 1.3.4 Killing components 45 1.3.4.1 Mycolytic and cuticle degrading enzyme complexes 45 1.3.4.2 Antibiotics and toxins 47 1.4 Development of mycopesticide 49 1.4.1 Strain 49 1.4.1.1 Isolation 49 1.4.1.2 Habitat association 51 1.4.1.3 Criteria for strain selection 52 2 1.4.1.4 Strain improvement 53 1.4.1.5 Preservation of fungi 54 1.4.2 Production of fungal biocontrol agents 54 1.4.2.1 Living propagules 54 1.4.2.2 Fungal metabolites 57 1.4.3 Formulation 62 1.4.3.1 Types of formulations 62 1.4.3.2 Storage and shelf life 65 1.4.4 Application 65 1.4.5 Risk assessment 67 1.4.6 Registration and commercialization 68 1.5 Present investigations and thesis organization 69 1.5.1 Myrothecium verrucaria 69 1.5.2 Objectives of this study 70 1.5.3 Thesis organization 71 2. Chapter 2 Materials and methods 73-92 2.1 Materials 74 2.2 Organisms 74 2.2.1 Media 75 2.2.2 Maintenance 76 2.2.3 Growth of M. verrucaria on different media 76 2.3 Hypha-yeast transition in M. verrucaria 77 2.3.1 Morphological triggers 77 2.3.2 Spore germination under yeast- and hypha-favouring 77 conditions 2.4 Aerial conidia production using solid- state fermentation 78 2.5 Cuticle degrading/mycolytic enzyme complex of M. 78 verrucaria 2.5.1 Shake flask studies 78 2.5.2 Fermenter studies 78 2.5.3 Concentration of cuticle degrading/mycolytic enzyme 79 complex 2.6 Plate assays to study antagonistic interaction 79 3 2.6.1 Dual culture plate assay 79 2.6.2 Viability of sclerotia 80 2.6.3 Effect of cuticle degrading/mycolytic enzyme preparation 80 on germination of conidia 2.6.4 Peanut germination trial 81 2.7 Sudan black B staining of yeast-like cells 81 2.8 Enzyme assays 81 2.8.1 Chitinase 81 2.8.2 In situ estimation of endochitinase and N- 82 acetylglucosaminidase 2.8.3 Chitosanase assay 83 2.8.4 Protease assay 83 2.8.5 Lipase assay 83 2.8.6 βββ-1,3-Glucanase assay 84 2.8.7 Mannanase assay 84 2.8.8 Amylase assay 84 2.8.9 Cellulase assay 85 2.8.10 NAD(P)-glutamate dehydrogenase assay 85 2.8.11 Nitrate reductase assay 85 2.8.12 Mandelonitrile lyase assay 86 2.9 Estimation of reducing sugars 86 2.10 Protein estimation 87 2.11 Thin layer chromatography 87 2.12 Native polyacrylamide gel electrophoresis and activity 87 staining of NAD- and NADP- glutamate dehydrogenases 2.13 18s rRNA gene sequence analysis 88 2.14 Immobilization of yeast-like cells in alginate beads 88 2.15 Formulation of conidia with different carriers 89 2.16 Biocontrol of fungal pathogens of plants (Pot 89 experiments) 2.17 Insect bioassay 90 2.18 Field trial 92 4 3. Chapter 3 Morphological, cultural, biochemical and 93-119 molecular characterization of M. verrucaria 3.1 Introduction 94 3.2 Results and discussion 95 3.2.1 Morphological and cultural characteristics of M. 95 verrucaria 3.2..2 Morphological transition in M. verrucaria 98 3.2.2.1 Nature of unicellular growth 98 3.2.2.2 H-Y reversible transition 100 3.2.2.3 Morphological triggers 102 3.2.2.4 Y-H transition in M. verrucaria 109 3.2.2.5 Spore germination under H and Y form triggering conditions 111 3.2.3 Biochemical characterization of M. verrucaria 111 3.2.3.1 Endochitinase and N -acetylglucosaminidase 111 3.2.3.2 NAD- and NADP-dependent glutamate dehydrogenases and 114 nitrate reductase 3.2.4 Molecular characterization of M. verrucaria 117 4. Chapter 4 Mycopesticide production 120-147 4A Production of living propagules, conidia and yeast-like 121 cells of M. verrucaria 4A.1 Introduction 121 4A.2 Results and discussion 122 4A.2.1 Production of conidia using solid-state fermentation 122 4A.2.1.1 Selection of the solid substrate 122 4A.2.1.2 Effect of water content 124 4B.2.1.3 Effect of incubation temperature 125 4A.2.1.4 Effect of an inoculum 126 4A.2.2 Production of yeast-like cells of M. verrucaria 128 4A.2.2.1 Effect of nature of inoculum 128 4A.2.2.2 Production of mycolytic enzymes using yeast-like cells 129 4 B Production of cuticle degrading and mycolytic enzyme 132 complex of M. verrucaria 4B.1 Introduction 132 5 4B.2 Results and discussion 133 4B.2.1 Shake flask studies 133 4B.2.1.1 Effect of different chitin sources 133 4B.2.1.2 Effect of corn steep liquor and molasses 133 4B.2.1.3 Effect of controlled pH 134 4B.2.1.4 Effect of an inoculum 135 4B.2.1.5 Effect of agitation 137 4B.2.1.6 Effect of cultivation volume 138 4B.2.1.7 Time course for production of cuticle degrading/mycolytic 139 enzyme complex in shake flask 4B.2.2 Fermenter studies 141 4B.2.2.1 Effect of agitation 141 4B.2.2.2 Effect of aeration 143 4B.2.2.3 Time course for production of cuticle degrading/mycolytic 145 enzyme complex by M. verrucaria 5. Chapter 5 Biocontrol efficacy of living propagules and 148-167 enzymic components of M. verrucaria against fungal and insect pests 5.1 Introduction 149 5.2 Results and discussion 150 5.2.1 Formulation studies 150 5.2.1.1 Conidial formulation 150 5.2.1.2 Yeast-like cells formulation 152 5.2.1.3 Cuticle degrading/mycolytic enzyme preparation 154 5.2.2 Control of Sclerotium rolfsii infection in peanut 155 5.2.2.1 Plate assays 155 5.2.2.2 Pot experiments 157 5.2.3 Control of insect pests 160 5.2.3.1 Insect bioassays 160 5.2.3.2 Field trial 165 5.2.4 Toxicity studies 167 6 A Appendix 1 Development of a new method for estimation 168-175 of chitinase A 1 Introduction 169 A 2 Results and discussion 169 A 2.1 Substrate preparation 169 A 2.2 Effect of substrate concentration 170 A 2.3 Effect of incubation time 171 A 2.4 Effect of enzyme concentration 171 A 2.5 Thin layer chromatography 172 A 2.6 Definition of unit 174 A 2.7 Validation and specificity of the assay 174 6. Chapter 6 Conclusion 176-180 7. Chapter 7 References 181-199 8. List of publications, patents and presentation 200-201 7 List of tables Sr.No. Titles Page No. 1.1 Major fungal pathogens and insect pests of cash crops 26 1.2 Fungi as biocontrol agents against plant pathogenic fungi and 38 insect pests 1.3 Production of fungal chitinases 60 2.1 Sources of chemicals and materials 74 2.2 List of organisms used 75 2.3 Media 75 3.1 Effect of carbon source on H-Y transition in M. verrucaria 102 3.2 Effect of nitrogen source on H-Y transition in M. verrucaria 104 3.3 Effect of initial pH of the medium on morphology of M. 107 verrucaria 3.4 Effect of continuous maintenance of initial pH on H-Y transition 108 in M. verrucaria 3.5 Differential expression of NADP- and NAD-dependent glutamate 115 dehydrogenases and nitrate reductase during H-Y reversible transition in M. verrucaria 4.1 Production of M. verrucaria conidia on various solid substrates 123 4.2 Effect of nature of inoculum on the production of M. verrucaria 129 yeast-like cells 4.3 Production of mycolytic enzymes using different morphologies 130 of M. verrucaria as inoculum 4.4 Effect of addition of corn steep liquor and molasses in medium 134 on chitinase production by M. verrucaria 5.1 Concentration of cuticle degrading/mycolytic enzyme preparation 154 using different methods 5.2 Control of pre-emergence rot in peanut using various 158 formulations containing M. verrucaria conidia or yeast-like cells 5.3 Control of pre-emergence rot in peanut using mycolytic 160 enzyme preparation of M. verrucaria 8 5.4 Effect of M. verrucaria cuticle degrading enzyme preparation on 161 3rd instar larvae of H. armigera, S.
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