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Characterization of Mechanisms of Resistance to Common Insecticides

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Characterization of Mechanisms of Resistance to Common Insecticides Aus dem Institut für Phytomedizin Universität Hohenheim Fachgebiet: Angewandte Entomologie Prof. Dr. C.P.W. Zebitz Characterization of mechanisms of resistance to common insecticides in noctuid pest species and resistance risk assessment for the new lepidopteran specific compound flubendiamide Dissertation zur Erlangung des Grades eines Doktors der Agrarwissenschaften vorgelegt der Fakultät Agrarwissenschaften von Stefanie Konanz aus Pforzheim 2009 Die vorliegende Arbeit wurde am 3. Juni 2009 von der Fakultät Agrarwissenschaften der Universität Hohenheim als “Dissertation zur Erlangung des Grades eines Doktors der Agrarwissenschaften“ angenommen. Tag der mündlichen Prüfung: 10. Juli 2009 1. Prodekan: Professor Dr. W. Bessei 1. Prüfer, Berichterstatter: Professor Dr. C.P.W. Zebitz 2. Prüfer, Mitberichterstatter: Professor Dr. W. Schwack 3. Prüfer: Professor Dr. R. Blaich TABLE OF CONTENTS 1 INTRODUCTION 1 1.1 BIOLOGY LEPIDOPTERA: NOCTUIDAE 1 1.2 SPODOPTERA EXIGUA , HELICOVERPA ARMIGERA AND HELIOTHIS VIRESCENS 2 1.3 LEPIDOPTERAN SPECIES – IMPORTANT INSECTICIDES 3 1.3.1 History – Development of the lepidopteran insecticide market.......................................................3 1.3.2 General overview of the world market for insecticides ...................................................................4 1.3.3 “Old Chemistry”..............................................................................................................................6 1.3.3.1 Pyrethroids 6 1.3.3.2 Carbamates and organophosphates 7 1.3.3.3 Cyclodienes 8 1.3.3.4 Benzoylphenyl ureas 9 1.3.4 “New Chemistry” .......................................................................................................................... 10 1.3.4.1 Indoxacarb 10 1.3.4.2 Emamectin 10 1.3.4.3 Spinosad 11 1.3.4.4 Chlorfenapyr 12 1.3.4.5 Moulting accelerating compounds (diacylhydrazines) 12 1.3.4.6 Flubendiamide 13 1.3.4.7 Flubendiamide a new tool on the insecticide market 14 1.4 RESISTANCE 15 1.5 OVERVIEW OF RESISTANCE MECHANISMS IN INSECTS 16 1.5.1 Penetration and behavioural resistance.......................................................................................... 17 1.5.2 Metabolic resistance ...................................................................................................................... 17 1.5.2.1 Glutathione S-transferases 17 1.5.2.2 Carboxylesterases 18 1.5.2.3 Cytochrome P 450 -dependent monooxygenases 19 1.5.3 Altered target-site resistance.......................................................................................................... 20 1.5.3.1 GABA receptor: target-site resistance to cyclodienes 20 1.5.3.2 Insensitive AChE: target-site resistance to carbamates and organophosphates 21 1.5.3.3 Nerve insensitivity: target-site resistance to pyrethroids 22 1.6 RESISTANCE MANAGEMENT 24 1.6.1 Cross-resistance............................................................................................................................. 24 1.6.2 Integrated pest and insecticide resistance management................................................................. 25 2 OBJECTIVE 27 3 MATERIAL AND METHODS 29 3.1 CHEMICALS AND INSECTICIDES 29 3.1.1 Chemicals and reagents ................................................................................................................. 29 3.1.2 Insecticides and synergists............................................................................................................. 29 3.2 LEPIDOPTERAN STRAINS 29 3.3 BIOASSAYS 30 3.3.1 Larval spray bioassay .................................................................................................................... 30 3.3.2 Larval artificial diet bioassay......................................................................................................... 31 3.3.3 Worldwide monitoring .................................................................................................................. 31 3.3.4 Application and synergism studies ................................................................................................ 32 3.3.5 Pharmacokinetic studies ................................................................................................................ 33 3.4 BIOCHEMISTRY 35 3.4.1 Protein determination .................................................................................................................... 35 3.4.2 Glutathione S-transferases............................................................................................................. 35 3.4.2.1 Standard photometric assay using 1-chloro-2,4-dinitrobenzene as substrate 35 3.4.2.2 Fluorometric assay using monochlorobimane as substrate 36 3.4.3 Carboxylesterases .......................................................................................................................... 36 3.4.3.1 Photometric assay with 1-naphthyl acetate 36 3.4.3.2 nPAGE and inhibition studies 37 3.4.4 Cytochrome P 450 -dependent monooxygenases .............................................................................. 38 3.4.4.1 Fluorometric assay to measure 7-ethoxycoumarin O-deethylation 38 3.4.5 Acetylcholinesterases .................................................................................................................... 39 3.4.5.1 Photometric determination of acetylcholinesterase activity 39 3.4.5.2 Inhibition studies and determination of kinetic parameters 40 3.4.6 Affinity purification of acetylcholinesterases................................................................................ 40 3.5 MOLECULAR BIOLOGY 41 3.5.1 RNA extraction from Spodoptera exigua for sodium channel amplification ................................ 41 3.5.2 Synthesis of first-strand cDNA of Spodoptera exigua strains ....................................................... 42 3.5.3 Polymerase chain reaction (PCR).................................................................................................. 42 3.5.4 Cloning and sequencing of sodium channel from Spodoptera exigua ........................................... 44 3.5.5 Pyrosequencing.............................................................................................................................. 44 3.5.5.1 The principle of pyrosequencing 44 3.5.5.2 Pyrosequencing 45 3.6 STATISTICS AND CALCULATIONS 47 4 RESULTS 48 4.1 BIOASSAYS 48 4.1.1 Baseline susceptibility of H. armigera and H. virescens to different insecticides......................... 48 4.1.2 Resistance pattern in Spodoptera exigua strain SPOD-EX E-98................................................... 50 4.2 EFFICACY OF FLUBENDIAMIDE 54 4.2.1 Baseline susceptibility and stage-specific activity......................................................................... 54 4.2.2 Worldwide monitoring .................................................................................................................. 55 4.3 APPLICATION AND SYNERGISM STUDIES 58 4.4 BIOCHEMICAL ASSAYS 59 4.4.1 Glutathione S-transferases............................................................................................................. 60 4.4.1.1 Photometric assay in larvae mass homogenates 60 4.4.1.2 Frequency distribution of enzyme activity of single larvae 60 4.4.1.3 Fluorometric assay with monochlorobimane 61 4.4.2 Carboxylesterases .......................................................................................................................... 62 4.4.2.1 Standard assay in larvae mass homogenates 62 4.4.2.2 Frequency distribution of enzyme activity of single larvae 63 4.4.2.3 Carboxylesterase iso-enzyme analysis and inhibition visualised by nPAGE 64 4.4.3 Cytochrome P 450 -dependend monooxygenases ............................................................................. 67 4.4.3.1 Fluorometric assay with 7-ethoxycoumarin in second-instar larvae 67 4.4.3.2 Monooxygenase activity in isolated guts of fifth-instar larvae 68 4.4.4 Acetylcholinesterases .................................................................................................................... 69 4.4.4.1 Standard assay in larvae mass homogenates 69 4.4.4.2 Frequency distribution of enzyme activity of single larvae 70 4.4.4.3 Michaelis-Menten kinetics 71 4.4.4.4 Acetylcholinesterase inhibition studies in vitro 72 4.4.5 Partial purification of acetylcholinesterases .................................................................................. 73 4.4.5.1 Michaelis-Menten kinetics 75 4.4.5.2 Inhibition studies with purified acetylcholinesterases 75 4.4.5.3 SDS-PAGE 76 4.5 MOLECULAR BIOLOGY 77 4.5.1 Cloning and sequencing of sodium channel from Spodoptera exigua ........................................... 77 4.5.2 Pyrosequencing.............................................................................................................................. 80 4.6 PHARMACOKINETIC STUDIES IN SPODOPTERA EXIGUA 82 5 DISCUSSION 86 5.1 BIOASSAYS 86 5.1.1 Endosulfan resistance .................................................................................................................... 86 5.1.2 Indoxacarb resistance .................................................................................................................... 87 5.1.3 Carbamate and organophosphate resistance .................................................................................. 88 5.1.4 Benzoylphenyl urea resistance ...................................................................................................... 88 5.1.5 Pyrethroid resistance.....................................................................................................................
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