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EPIDEMIOLOGICAL ASPECTS of MBC RESISTANCE in Monilinia Fructicola (Wint.) Honey and MECHANISMS of RESISTANCE

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EPIDEMIOLOGICAL ASPECTS of MBC RESISTANCE in Monilinia Fructicola (Wint.) Honey and MECHANISMS of RESISTANCE Apothecia of Monilinia fructicola (Wint. ) Honey under controlled conditions. And so, from hour to hour we ripe and ripe, And then from hour to hour we rot and rot, And thereby hangs a tale. As You Like It: Act II, Scene vii (William Shakespeare) EPIDEMIOLOGICAL ASPECTS OF MBC RESISTANCE IN Monilinia fructicola (Wint.) Honey AND MECHANISMS OF RESISTANCE A thesis submitted in fulfilment of the requirements for the Degree of Doctor of Philosophy in Plant Pathology at Lincoln University New Zealand by Niwat Sanoamuang Lincoln University 1992 DEDICATION This thesis is dedicated to my mother, CHANTR and my father, SAIYOOD SANOAMUANG for their love, wish and encouragement. Lincoln University Te Whare Wanaka O Aoraki P. O. Box 84 Lincoln University Canterbury New Zealand Telephone: Christchurch (03)252 811 Vice Chancellor & Registry (64)(03)252 965 Library & Departments (64)(03)252 944 New telephone no: (03) 325 2811 Fax no: (03) 325 3843 REG: JGL DEPARTMENT OF PLANT SCIENCE 8 July 1992 CERTIFICATE OF SUPERVISION I certify that the work reported in this thesis was planned, executed and described by the candidate under my direct supervision. Reader in Plant Pathology i Abstract of a thesis submitted in fulfilment of the requirements for the degree of Doctor of Philosophy. EPIDEMIOLOGICAL ASPECTS OF MBC RESISTANCE IN Monilinia fructicola (wint.) Honey AND MECHANISMS OF RESISTANCE by N. Sanoamuang Isolates of Monilinia fructicola (Wint.) Honey obtained from stone fruit orchards in Hawkes Bay, North Island and from Californian fruit exported to New Zealand, were tested for resistance to methyl benzimidazole carbamate (MBC). Resistant isolates from the North Island had EC50 values of >30,000, and most isolates from the imported fruit had of values approximately 1.5 mg a.i./l carbendazim. Sensitive isolates failed to grow on 1 mg a.i./l carbendazim. A detached peach shoot system was used in controlled conditions for estimation of values for incubation period, latent period and rate of spore production on flowers (cv Glohaven). The same variables and the rate of colonisation of host tissue were measured on fruit (cv Fantasia) in controlled conditions. An inoculum density of 1x104 spore/flower or fruit greatly increased fitness in vivo compared to an inoculum density of 1x102 spore/flower (fruit). Isolates varied considerably, but there was no consistent relationship between the degrees of resistance and fitness. This was in contrast to earlier studies with dicarboximide resistant strains of M. fructicola. The survival in the field of 10 isolates resistant or sensitive to MBC or dicarboximide fungicides on twig cankers and mummified fruit was compared. The ability to produce conidia on twig cankers inoculated in late spring 1989 was maintained by all sensitive and MBC resistant isolates for at least 1 year. The production of conidia on mummified fruit inoculated in February 1990 decreased after 2-3 months in the field but some conidia were still produced on ii all fruit in the following spring. Dicarboximide resistant isolates produced less conidia than either the MBC resistant and the sensitive isolates. The pathogenicity and fitness of all isolates were similar to the original values after survival for 1 year. A technique was developed to produce apothecia reliably from inoculated peach (cv Black Boy) and nectarine (cv Fantasia) fruit in controlled conditions in the laboratory. The fruit were inoculated with resistant or sensitive isolates, or combinations, and were incubated for 8 weeks at 25 0 C (±1 0 C) with 12 hours photoperiod of fluorescent light (Sylvania 2x65 W, daylight) to produce mummified fruit. The fruit were then buried in moist autoclaved peat moss for 10 weeks at 25 0 C (±1 °C) in the dark to form stromata. These fruit were then 3 hydrated with running tap-water (total hardness (CaCO3 ) = 47 g/m and conductivity at 20 0 C = 12. 7 mS/m) for 72 hours. The hydrated mummified fruit were placed in moist peat moss and were incubated for 13-14 weeks at 8 0 C (±0. 5 °C) in the dark. At the end of this period, stipe initials were visible. Differentiation of stipe initials into mature apothecia occurred within 15-20 days after transfer to 12 0 C (±2 °C) with a 12 hour photoperiod of fluorescent and incandescent light. All isolates produced apothecia when treated in this way. A technique for isolation of ascospore sets in linear arrangement was developed for tetrad analysis of the inheritance of resistance. At least 3 hours of fluorescent and incandescent light at 12 0 C (±2 °C) was essential to allow ascospore ejection from individual asci taken from apothecia previously maintained in a 12 hour photoperiod at 12 0 C (±1 °C). A water film on the surface of water agar was necessary to hold a set of ejected ascospores in linear sequence. Single ascospores were obtained in sequence with the aid of a micromanipulator. Genetic analysis of MBC resistant isolates was carried out on ascospores derived from apothecia produced in the laboratory. Analysis of ascospore sets in linear arrangement and ascospore populations indicated that resistance to iii >30,000 mg a.i./l carbendazim (high-resistant) is governed by a single major gene and is affected by gene conversion mechanisms. Crossing over was frequent, suggesting that recombination of resistance with other characters, such as pathogenicity and fitness, may occur readily. The segregation ratio (1:1) from most resistant isolates revealed that heterokaryons containing both resistant and sensitive alleles were common in resistant populations and that resistance is dominant. Allozyme analysis of ascospore progeny through electrophoresis revealed a narrow genetic base of M. fructicola in New Zealand. The technique for reliable apothecial production in controlled conditions developed in this study provided an important step for the determination of the biology of M. fructicola strains resistant to MBC fungicides, and the complexity of its life cycle. Genetic heterogeneity in field populations can be conserved in one isolate through heterokaryosis, thus providing for adaptability of the pathogen to the changing environmental conditions. Knowledge on genetic variability, overwintering ability, pathogenicity and fitness factors may be useful for future management strategies of stone fruit brown rot. Special emphasis should be made in particular to prevent primary infection on blossoms, which would delay the establishment of recombinant strains of M. fructicola and the onset of brown rot epidemics. KEYWORDS: Methyl benzimidazole carbamate; carbendazim; resistance; nectarine; peach; stone fruit; Monilinia fructicola; brown rot; epidemiology; virulence; fitness; competition; survival; twig cankers; mummified fruit; genetic; apothecia; ascospores; mechanisms of resistance; inheritance. iv CONTENTS ABSTRACT i CONTENTS iv LIST OF TABLES vii LIST OF FIGURES ix LIST OF APPENDK x CHAPTER 1 PAGE INTRODUCTION 1 1. 1 General introduction 1 1. 2 Fungicidal mode of action and resistance 2 1. 3 Epidemiological aspects of fungicide resistance.. 4 1. 4 Genetics of fungicide resistance 10 1. 5 Biology of the fungus 15 1. 6 Objectives 20 CHAPTER 2 PARASITIC FITNESS AND COMPETITIVE ABBLITY OF MBC RESISTANT STRAINS... 23 2. 1 Introduction 23 2. 2 Materials and methods 26 2. 2. 1 Isolation and sensitivity test 26 2. 2. 2 Parasitic fitness investigation 28 2. 2. 3 Competitive ability on host tissues 32 PAGE 2. 3 Results 34 2. 3. 1 The incidence of resistance 34 2. 3. 2 Parasitic fitness in vivo 36 2. 3. 3 Competitive ability in vivo 39 2. 4 Discussion 42 CHAPTER 3 SURVIVAL OF M. fructicola ISOLATES RESISTANT TO MBC AND DICARBOXIMIDE FUNGICIDES 46 3. 1 Introduction 46 3. 2 Materials and methods 48 3. 3 Results 53 3. 4 Discussion 60 CHAPTER 4 APOTHECLAL INDUCTION AND SINGLE ASCOSPORE ISOLATION 63 4. 1 Introduction 63 4. 2 Materials and methods 65 4. 3 Results 68 4. 4 Discussion 74 vi CHAPTER 5 PAGE INHERITANCE OF RESISTANCE AND MECHANISMS OF RESISTANCE TO MBC FUNGICIDE 80 5. 1 Introduction 80 5. 2 Materials and methods 83 5. 3 Results 86 5. 4 Discussion 94 CHAPTER 6 GENERAL DISCUSSION 99 6. 1 Current status of M. fructicola resistance to MBC fungicide in New Zealand 99 6. 2 Techniques for detailed studies of the sexual reproduction and the genetic basis of resistance to fungicide 100 6. 3 Epidemiological aspects of MBC and dicarboximide resistant and sensitive isolates... 102 6. 4 Genetic basis of MBC resistance in M. fructicola 107 6. 5 Management of resistance in M. fructicola 110 6. 6 Future research 113 ACKNOWLEDGEMENTS 116 REFERENCES 118 APPENDIX 135 vii LIST OF TABLES TABLE PAGE 1. 1 Modes of action of selective fungicides in target organisms 3 2. 1 Isolations, and phenotype to fungicides of M. fructicola used for determination of fitness attributed 29 2. 2 Fungicide resistance in isolates of M. fructicola from commercial orchards and imported fruit in a supermarket 35 2. 3 The fitness components, incubation period, latent period and rate of spore production on blossom for M. fructicola field isolates resistant and sensitive to MBC fungicide 37 2. 4 The fitness components, incubation period, latent period, rate of colonization and rate of spore production on fruit of M. fructicola field isolates resistant and sensitive to MBC fungicide 38 2. 5 The competitive ability of four M B C resistant M. fructicola field isolates with sensitive isolates on peach blossom (cv Glohaven) and on fruit (nectarine cv Fantasia) at 1x102 and 1x104 inoculum concentrations 41 3. 1 Experimental designs for survival investigations and subsequent pathogenicity on host tissues 52 3. 2 Mean canker length (mm) on nectarine twigs (cv Fantasia) by resistant and sensitive isolates to MBC and dicarboximide fungicides at different intervals after inoculation 54 viii TABLE PAGE 3. 3 Number of spores produced on twig cankers under conducive conditions in the laboratory of M.
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