HOST PLANT RESISTANCE: CONCEPT AND SIGNIFICANCE Dr. Muhammad Salim M.Sc. (Hons.)PPR(UAE); M.Sc. ENT (UPLB/IRRI, Philippines & Ph.D. ENT (UPLB/IRRI, Philippines Ex-CSO/ Sr. Director, Plant Sciences Division (PARC) Islamabad HIGHER EDUCATION COMMISSION ISLAMABAD 1 CONTENTS ABOUT THE AUTHOR……………………………………………………………...…..i FOREWORD…………………………………………………………………..……...…iii PREFACE………………………………………………………….............................vii CHAPTER-1 INTRODUCTION.……….………………………………………...…….1 1.1. PLANT –INSECT INTERACTIONS CONCEPT…… ……………………...1 1.1.1. Defence Strategies of Plants………………………………………..1 1.1.2. Insect Strategies to Counter Plant Defence……………………….2 1.2. HISTORY OF HOST PLANT RESISTANCE DEVELOPMENT…………..4 1.3. DEFINITIONS…………………………………………………………….……7 1.4. FUNCTIONAL CATEGORIES………………………………………………..8 1.5. ADVANTAGES…………………………………………………………………9 1.6. LIMITATIONS…………………………………………………………………10 1.7. HOST PLANT RESISTANCE PROGRAMME IMPLEMENTATION…....12 1.7.1. Scientific Manpower………………………………………………..12 1.7.2. Scientific Knowledge……………………………………………….12 1.7.3. Availability of Insect Culture……………………………………….12 1.7.4. Genetic Resources………………………………………………....13 1.7.5. Genetic Aspects of Resistance…………………………………...13 1.7.6. Selection of Breeding Methods……………………………………13 1.8. REFERENCES……………………………………………………………….14 CHAPTER-2 MECHANISM OF HOST PLANT RESISTANCE ………………....25 2.1. ECOLOGICAL RESISTANCE………………………………………………25 2.1.1. Host Evasion/Escape/ Phenological Asynchrony……………….25 2.1.2. Induced Resistance………………………………………….……..27 2 2.2. EVOLUTIONARY RESISTANCE…………………………………………..28 2.2.1. Sympatric Resistance………………………………………….…..28 2.2.2. Allopatric Resistance……………………………………………....28 2.3. TROPHIC LEVEL RESISTANCE…………………………………………..29 2.3.1. Intrinsic Resistance………………………………………………...29 2.3.2. Extrinsic Resistance………………………………………………..29 2.4. MECHANISM BASED ON INHERITANCE OF RESISTANCE………….30 2.5. MECHANISM BASED ON CROP STAGE………………………..……….30 2.6. MECHANISM BASED ON SCREENING CONDITIONS………………...30 2.7. MECHANISM BASED ON BIOTYPE REACTION………………………..31 2.8. MECHANISM BASED ON POPULATION………………………..……….33 2.9. MECHANISM BASED ON NATURE OF GENES………………….……..33 2.10. MECHANISM BASED ON FUNCTIONAL CATEGORIES……….…..….33 2.10.1. Non-preference/Antixenosis……………………………………....33 2.10.2. Antibiosis……………………………………………………….…....36 2.10.3. Tolerance……………………………………………………….…...40 2.11. INTENSITY OF RESISTANCE…………………………………..………....41 2.12. REFERENCES…………………………………………………………........45 CHAPTER-3 BEHAVIOURAL AND PHYSIOLOGICAL COMPONENTS OF HOST PLANT SELECTION………….………55 3.1. HOST HABITAT FINDING…………………………………………………..56 3.2. HOST FINDING……………………………………………………………....56 3.3. HOST RECOGNITION……………………………………………………....57 3.4. HOST ACCEPTANCE…………………………………………………….…58 3.5. HOST SUITABILITY…………………………………………………………59 3.6. REFERENCES…………………………………………………………….…61 CHAPTER-4 HOST-INSECT INTERACTIONS THEORIES……………………...67 4.1. THEORIES OF HOST-PLANT SELECTION……………………………...68 4.1.1. Botanical Instinct Theory…………………………………………..68 4.1.2. Token Stimuli Theory………………………………………….…...69 3 4.1.3. Dual Discrimination Theory………………………………………..70 4.1.4. Non-preference and Antibiosis Theory…………………………...71 4.1.5. Chemosensory Theory……………………………………………..71 4.1.6 Inhibitory Chemical Theory or Negative Stimuli Theory………..73 4.1.7. Thorsteinson’s Theory……………………………………………..74 4.1.8. Nutrients Imbalance Theory……………………………………….75 4.2. THEORIES OF HOST PLANT DEFENCE………………………………...76 4.3. THEORIES OF CO-EVOLUTION………………………………................77 4.3.1. Ehrlich and Raven (1964) Theory……………………………...…77 4.3.2. Sequential Co-evolution…………………………………………...80 4.3.3. Difuse Co-evolution………………………………………………...80 4.3.4. Geographic Mosaic Theory………………………………………..81 4.4. REFERENCES……………………………………………………………….82 CHAPTER-5 MORPHOLOGICAL BASIS OF RESISTANCE IN PLANTS TO INSECTS…………………………………..………..87 5.1 COLOUR AND SHAPE……………………………………………………...87 5.2. THICKENING OF CELL WALLS…………………………………………...88 5.3. STRUCTURE AND TEXTURE OF TISSUES……………………………..89 5.4. INCRUSTATION OF MINERALS IN CUTICLE…………………………...90 5.5 SURFACE WAXES…………………………………………………………..91 5.6. ANATOMICAL ADOPTION OF ORGANS…………………………………92 5.7. TRICHOMES………………………………………………………………....94 5.7.1. Locomotion, Attachment and Related Behaviour……………….94 5.7.2. Insect Feeding and Digestion……………………………………..95 5.7.3. Insect Development and Population……………………………...96 5.7.4. Insect Mortality………………………………………………….…..96 5.7.5. Oviposition…………………………………………………………..96 5.7.6. Insect Infestation or Plant Damage……………………………....97 5.7.7. Pubescence Associated with Allelochemical Factors…………..97 5.8. TRICHOMES AND RESISTANCE………………………………………....98 5.9. REFERENCES……………………………………………………………….99 4 CHAPTER-6 BIOCHEMICAL BASIS OF RESISTANCE IN PLANTS.TO INSECTS………………………………………......115 6.1. NON-PROTEIN AMINO ACIDS…………………………………………...117 6.2. LECTINS…………………………………………………………………….118 6.3. CYANOGENIC GLYCOSIDES (CGs) …………………………………...119 6.4. GLUCOSINULATES……………………………………………………..…119 6.5. ALKALOIDS…………………………………………………………….…...120 6.6. PHENOLICS……………………………………………………..………….122 6.7. TERPENOIDS/ISOPRENOIDS……………………………………………123 6.8. EPICUTICULAR LIPIDS……………………………………………...……125 6.9. TANNINS……………………………………………...…………………….126 6.10. PROLINE……………………………………………...…………………….126 6.11. SUGARS……………………………………………...………………….….126 6.12. BIOCHEMICAL BASIS OF RESISTANCE OF MAJOR CROPS………126 6.12.1. Cotton……………………………………………...……………….126 6.12.2. Sugarcane……………………………………………...………….127 6.12.3. Maize……………………………………………...………………..127 6.12.4. Oilseed Crops……………………………………………...……...128 6.12.5. Rice……………………………………………...………………….128 6.13. REFERENCES……………………………………………...……………...139 CHAPTER-7 FACTORS AFFECTING HOST PLANT RESISTANCE…………153 7.1. TEMPERATURE STRESS……………………………………………...…154 7.2. SALINITY STRESS……………………………………………..............…163 7.3. ALUMINUM STRESS……………………………………………...........…176 7.4. NUTRIENTS…………………………………………….........................…181 7.4.1. Nitrogen (N).……………………………………………...………..182 7.4.2. Phosphorus (P) ……………………………………………...……190 7.4.3. Potassium (K) …………………………………………………..…193 7.4.4. Iron (Fe) ………………………………………………..……….…199 7.4.5. Silica (Si) ……………………………………………................…205 5 7.4. 6. Boron (B).……………………………………………...…………..209 7.4.7. Metal Chelates………………………………………………….…209 7.5. GROWTH REGULATORS……………………………………………...…210 7.6. HERBICIDES……………………………………………...………………..210 7.7. PLANT SPACING………………………………………………………..…210 7.8. ATTACK OF INSECTS AND DISEASES………………………………...211 7.9. LIGHT……………………………………………......................................212 7.10. pH……………………………………………........................................….212 7.11. WATER……………………………………………………………….…...…213 7.12. RELATIVE HUMIDITY (RH) ……………………………………………....213 7.13. INSECTICIDES…………………………………………….............………213 7.14. MISCELLANEOUS……………………………………………...............…218 7.15. REFERENCES…………………………………………………..……….…221 CHAPTER-8 DEVELOPMENT OF INSECT RESISTANT VARIETIES………..235 8.1. NEED FOR RESISTANT VARIETIES……………………………...…….235 8.2. SOURCES OF RESISTANCE……………………………...……………..236 8.3. MULTIDISCIPLINARY TEAM APPROACH……………………………...237 8.4. SETTING PRIORITIES FOR THE DEVELOPMENT OF INSECT RESISTANT VARIETIES…………………………………………………..241 8.5. SCREENING AND EVALUATION OF GERMPLASM FOR INSECT RESISTANCE………………………………………………241 8.5.1. Whitebacked Planthopper, Sogatella furcifera……………..….244 8.5.2. Yellow stem borer, Scirpophaga incertulas………………….....245 and white stem borer,Scirpophaga innotata. 8.5.3. Leaffolder, Cnaphalocrocis medinalis…………………………..247 8.6. MAINTENANCE OF INSECT POPULATION……………….…………...249 8.6.1. Increasing Infestation of Sorghum Shoot Fly..………………....250 Atherigona soccata 8.6.2. Increasing Field Infestation of Rice Gall Midge………….........250 Pachydiplosis oryzae 8.6.3. Increasing Population of Rice Brown Planthopper..………..…250 Nilaparvata lugens 6 8.7. BREEDING METHODS FOR HOST PLANT RESISTANCE…………..251 8.7.1. Introduction of Varieties .…………………………………………252 8.7.2. Selection.………………………………...……………..………….252 8.7.3. Hybridization.………………………………………..…………….252 8.7.3.1. Pedigree selection.………………………………………253 8.7.3.2. Bulk selection.………………………………………..….253 8.7.3.3. Backcross method.……………………………………...254 8.7.4. Mutation Breeding.…………………………………………….….254 8.7.5. Methods of Producing Transgenic Plants.……………………..255 8.7.5.1. Agrobacterium mediated gene transfer.………………255 8.7.5.2. Direct gene transfer.…………………………………….256 8.8. LEVEL OF RESISTANCE: EXAMPLES FRO PAKISTAN……………..257 8.8.1. Cotton.………………………………………..…………...……….257 8.8.2. Sugarcane.………………………………………..……………….260 8.8.3. Pulses.………………………………………..…………...……….261 8.8.4. Rice.………………………………………..…………...………….263 8.8.5. Wheat.………………………………………..…………...……….274 8.8.6. Sunflower.………………………………………..………….....….274 8.8.7. Maize.………………………………………..…………...………..275 8.8.8. Soybean.………………………………………..…………...…….278 8.8.9. Brassica.………………………………………..…………...…….279 8.8.10. Mulberry.………………………………………..…………...…….282 8.8.11. Barley.………………………………………..…………...……….282 8.8.12. Cucumber.………………………………………..…………....….282 8.8.13. Okra.………………………………………..…………...……..….282 8.9. REFERENCES.………………………………………..………………....282 CHAPTER-9 GENETICS OF INSECT RESISTANCE IN CROP PLANTS…………………………………………………...290 9.1. INSECT RESISTANCE IN MAJOR CROPS…………………………….290 9.1.1. Rice………………………………………………………………...290 9.1.1.1 Brown plant hopper (BPH), Nilaparvata lugens……...290 7 9.1.1.2. Whitebacked planthopper, Sogatella furcifera……….291 9.1.2. Wheat……………………………………………………………....292 9.1.3. Maize……………………………………………………………....294 9.1.3.1 European corn borer, Ostrinia nubilalis……………....294 9.1.3.2 Corn earworms, Helicoverpa zea and H. armigera….295 9.1.3.3. Stem borer, Chilo partellus……………………………..295 9.1.3.4 Shoot fly,Atherigona