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Interactions Between Trichoderma Spp. and Soil-Borne Plant Pathogens; Implications for Biological Control

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Interactions Between Trichoderma Spp. and Soil-Borne Plant Pathogens; Implications for Biological Control INTERACTIONS BETWEEN TRICHODERMA SPP. AND SOIL-BORNE PLANT PATHOGENS; IMPLICATIONS FOR BIOLOGICAL CONTROL by Anya M. B. Alles BSc (Hons) Durham; MSc Reading A thesis submitted for the degree of Doctor of Philosophy of the University of London and for the Diploma of Imperial College Department of Pure and Applied Biology Imperial College of Science and Technology London, S.W.7 August 1984 ABSTRACT The research was designed to evaluate the commercially marketed biological control agent Binab T.f which contains Trichoderma spp., as an alternative to fungicides for the control of soil borne pathogens under greenhouse conditions. A dual inoculation plate method was used to screen a range of fungi for their susceptibility to antagonism in vitro on a nutrient-poor medium (soil extract agar) and nutrient-rich medium (malt extract agar). Various interactions were observed of which the killing of the Oomycete damping-off fungi Pythium ultimum and Pythium debaryanum was the most striking. On soil extract agar the interaction was characterized by progressive cytoplasmic vacuolation, lysis and a zone of clearing which developed after hyphal contact between the two organisms had been established. Similarly on nutrient-rich media Trichoderma acted as an aggressive antagonist competing with and overgrowing a range of fungi; Pythium multisporum, Pythium undulatum, Phytophthora citricolaf Phytophthora cactorum, Rhizoctonia solani, Corticium praticola and the wilt pathogens Verticillium albo-atrum and Fusarium oxysporum f.sp. lycopersici. Subsequent greenhouse trials were employed to assess the extent of antagonism against P.ultimum and P . debaryanum in soil and the potential of the commercial formulation for control of the damping- off disease. Vermiculite soaked with vegetable juice was the most suitable growth medium and it also provided a convenient and efficient means of delivery. In this way Trichoderma inoculum was employed both as a prophylactic, pre-pathogen inoculation treatment and as a simultaneous-inoculation treatment against the pathogens, with varying degrees of success; but in no instance was a commercially acceptable level -3- of disease control achieved. Use of this Trichoderma carrier-growth system was also ineffective against F .oxysporum f.sp. lycopersici in sterile soil. Crude filtrates from Trichoderma cultures grown in soil extract liquid medium and from cultures grown on heat-killed Pythium mycelium or from dual cultures with Pythium, failed to reproduce the distinctive interaction with Pythium observed in paired culture tests. Agar plate tests using cellophane and dialysis membranes demonstrated that the Trichoderma produced an unidentified antibiotic. Trichoderma also produced a cellulase system which degraded both Whatmans CM11 cellulose and" Avicel, but the hyphal lysis of Oomycetes on low-nutrient status agar could not be attributed to either of these metabolites nor to a physical inter­ action. Whatever the underlying mechanism the extent of hyphal lysis is clearly insufficient to cause appreciable suppression of Pythium inoculum in soil. ACKNOWLEDGEMENTS I would like to thank my supervisor Dr. S.A. Archer for his advice and guidance especially during the prepa­ ration of this manuscript. My thanks also to my coUeagues in Plant Pathology for creating such a friendly and enjoyable environment. I wish to express my appreciation and thanks to my parents and family for their unfailing encouragement and practical help and to Kamala for her excellent typing under duress. I am especially grateful to my long-suffering husband Ranjan, for his remarkable patience and support and in particular for generating enthusiasm and optimism when they were most needed. The research was financed by Stokes Bomford Chemicals Limited. -5- CONTENTS Page No. TITLE 1 ABSTRACT 2 ACKNOWLEDGEMENTS 4 CONTENTS 5 LIST OF TABLES 11 LIST OF FIGURES 14 LIST OF PLATES 15 LIST OF ABBREVIATIONS 16 CHAPTER 1 LITERATURE REVIEW 18 1.1 Introduction 18 1.2 Mechanisms of antagonism between 20 microorganisms 1.3 Strategies for disease control 22 1.4 Methods of introducing and 27 enhancing the activity of • antagonists in soil 1.5 Integration with chemical and 31 cultural control methods 1.6 Induction of new biotypes of 33 Trichoderma 1.7 Suppressive soils 35 CHAPTER 2 INTERACTIONS BETWEEN TRC AND SOIL 42 FUNGI ON AGAR MEDIA 2.1 Introduction 42 2.2 Materials and Methods 42 2 .2.1 Fungi 42 43 2 .2.2 Nutrient media 2.2.3 Dual inoculation plate tests 44 2.2.4 Transects 45 RESULTS 45 -6- Page No. 2.3 Dual Inoculation plate tests 45 using Pythium ultimum and Pythium debaryanum as the test organisms 2.3.1 Malt extract agar (MEA) 45 (2% w/v) 2.3.2 Vg- agar 46 2.3.3 Soil extract agar (SEA) 55 2.3.4 Summary 62 2.4 Interactions between TRC and 65 other Oomycetes 2.5 Interactions between TRC and 69 Corticum praticola and Rhizoctonia solani 2.6 Interactions between TRC and 72 Verticillium albo-atrum and Fusarium oxysporum f.sp. lycopersici 2.7 Paired culture tests using staggered 77 inoculation on 2% (w/v) MEA using TRC, T.harzianum and T.polysporum against F.oxysporum f.sp. lycopers ici and V.albo-atrum 2.8 Tests using other Trichoderma 82 cultures against Pythium ultimum on SEA 2.9 Discussion 84 CHAPTER 3 TESTS TO INVESTIGATE THE MECHANISM 92 OF CLEAR ZONE PRODUCTION 3.1 Introduction 92 3.2 Materials and Methods 93 3.2.1 Preparation of cellophane and 93 dialysis membrane 3.2.2 Tests to investigate the effects 93 of TRC metabolites 3.2.3 Dual culture tests using cellophane 94 and dialysis membranes -7- Page No. 3.2.4.1 Culture filtrates 96 3.2.4.2 Freeze drying of culture 96 f iltrate 3.2.5 Assay methods 97 3.2.6 Modified cellulose medium 98 RESULTS 99 3.3 Further studies on the Inter- 99 action occurring between TRC and Pythium on SEA 3.3.1 The effect of soil extract 99 concentration 3.3.2 The effect of glucose 106 concentration 3.4 Experiments to investigate the 113 effects of prior TRC growth on cellophane and dialysis membranes 3.4.1 MEA (2% (w/v)) and SEA 113 3.4.2 MEA (5% (w/v)) MEA 113 3.4.3 Range of activity 116 3.4.4 Tests using six Trichoderma 119 isolates against P.ultimum 3.5 Dual inoculation plate tests 121 using cellophane and dialysis membrane 3.6 Culture Filtrate Experiments 126 3.6.1 Tests using non-lyophilized un- 126 concentrated TRC culture filtrates 3.6.2 Effect of lyophilized and non- 127 lyophilized culture filtrates from mixed growth of TRC and P .ultimum and pure cultures of TRC and P .ultimum 3.6.3 Use of polycarbonate filters for 128 harvesting culture filtrates 3.6.4 Culture filtrates of T.polysporum 131 3.7 Celluloytic Activity 132 3.8 Discussion 134 - 8 - Page No. CHAPTER 4 EVALUATION OF TRICHODERMA AS 142 A BIOCONTROL AGENT UNDER GREENHOUSE CONDITIONS 4.1 Introduction 142 4.2 Materials and Methods 142 A. The damping-off pathogens 142 P.ultimum and P.debaryanum 4.2.1 Preparation of microbial inoculum 143 4.2.2 Incorporation of the fungal 143 cultures into the growth medium 4.2.3 Seed sowing and seedling emergence 144 4.2.4 Use of Trichoderma inoculum as a 145 prophylactic treatment 4.2.5 Final fresh weight analysis 145 4.2.6 Statistical treatment of data 145 4.2.7 Trichoderma selective media 146 4.2.7.1 Elad et al. (1981) 146 4.2.7.2 Papavizas (1981) 146 4.2.7.3 TRC selective media 147 4.2.7.4 Peptone-dextrose-rose bengal agar 147 B. The vascular wilt pathogen F.oxy- 147 sporum f.sp. lycopersici 4.2.8 Growth of tomato plants 147 4.2.9 Preparation of spore suspensions 148 4.2.9.1 F.oxysporum f.sp. lycopersici 148 4.2.9.2 TRC 149 4.2.9.3 T.harzianum and T.polysporum 149 4.2.10 Pathogen inoculation of plants 149 4.2.11 Evaluation of disease symptoms 149 4.2.12 Preparation of vermiculite media 150 and soil inoculation 4.2.13 Serial dilution of macerated stem 150 segments -9- Page No. RESULTS 151 A. The damping-off pathogens 151 P .ultimum and P ♦debaryanum 4.3.1 Selection of host-pathogen 151 combinations in the presence and absence of TRC 4.3.2 Use of TRC as a seed treatment 155 against P .ultimum and P.debaryanum 4.3.3 The effect of ,TRC 165 on disease incidence in non- sterile compost 4.3.4 Effect of TRC on disease inci- 165 dence in a soil-peat growth medium 4.3.5 Use of TRC as a pre-inoculation, 171 prophylactic treatment against the damping-off pathogens 4.3.6 The interaction between TRC and 176 P.ultimum and P .debaryanum in autoclaved compost 4.3.7 The influence of increased volumes 180 of TRC inoculum on disease incidence 4.3.8 Replant experiment to investigate 187 the longevity of P.ultimum and P.debaryanum in the presence of TRC 4.3.9 Use of a low nutrient growth medium 194 for TRC, P .ultimum and P .debaryanum 4.3.10 The interaction between TRC and the 198 damping-off fungi in the presence of a straw amendment 4.4 Quantitative studies of Trichoderma 205 populations 4.5 Discussion 209 B. The vascular wilt pathogen F,oxy- 217 sporum f.sp. lycopersici 4.6.1 Application of Trichoderma to .sail as 217 a spore suspension 4.6.2 Application of TRC by root dipping 219 4.6.3 Application of TRC by injection 220 4.6.4 Soil inoculations using TRC 220 vermiculite ~ VQ inoculum o 4.7 Discussion 223 CHAPTER 5 GENERAL DISCUSSION 229 REFERENCES 236 -11- LIST OF TABLES Page No.
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