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plant pathology fourth edition
plant pathology fourth edition

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part one GENERAL ASPECTS 3

chapter oneINTRODUCTION 3

Plant Pathology: The Science 3

The Concept of Disease in Plants 4

Types of Plant Diseases 4

History of Plant Pathology 7

The Role o f Fungi in Plant Disease 10

Discovery o f Other Pathogens as Causes of Plant Disease 12

Serious Plant Diseases o f Unknown Etiology 13

Early Development o f Plant Pathology 13

The Search for Control o f Plant Diseases 14

Interest in the Mechanisms by Which PathogensCause Disease 17

The Concept o f Genetic Inheritance o f Resistance and Pathogenicity 18

Epidemiology o f Plant Disease Comes o f Age 19

Molecular Plant Pathology 19

Development o f Plant Pathology Worldwide 20

Trends in Teaching and Training in Plant Pathology 22

Education and Information Technology in Plant Pathology 23

Plant Disease Clinics 23

Plant Pathology: The Practice 23

Certification of Professional Plant Pathologists 25

Significance of Plant Diseases 25

Kinds and Amounts o f Losses 25

Some Historical and Present Examples of Losses Caused by Plant Diseases 28

Plant Diseases and World Crop Production 28

Effects o f Changes in Agricultural Methods and in Human Society on the Development and Spread of Plant Diseases 34

Basic Procedures in the Diagnosis of Plant Diseases 37

Pathogen or Environment? 37

Infectious Diseases 37

Noninfectious Diseases 39

Identification of a Previously Unknown Disease:Koch’s Rules 39

chapter two PARASITISM AND DISEASE DEVELOPMENT 43

Parasitism and Pathogenicity 43

Host Range of Pathogens 44

Development of Disease in Plants 45

Stages in the Development of Disease:The Disease Cycle 45

Inoculation 45

Prepenetration Phenomena 46

Penetration 48

Infection 53

Invasion 54

Growth and reproduction of the pathogen(colonization) 54

Dissemination o f the Pathogen 56

Overwintering and/or Oversummering of Pathogens 59

Relationships between Disease Cycles and Epidemics 61

chapter three HOW PATHOGENS ATTACK PLANTS 63

Mechanical Forces Exerted by Pathogens on Host Tissues 63

Chemical Weapons of Pathogens 65

Enzymes in Plant Disease 65

Microbial Toxins in Plant Disease 73

Growth Regulators in Plant Disease 77

Polysaccharides 81

Suppressors of Plant Defense Responses 81

chapter four EFFECTS OF PATHOGENS ON PLANT PHYSIOLOGICAL FUNCTIONS 83

Effects of Pathogens on Photosynthesis 83

Effect of Pathogens on Translocation of Water and Nutrients in the Host Plant 85

Interference with Upward Translocation o f Water and Inorganic Nutrients 85

Interference with the Translocation o f Organic Nutrients through the Phloem 86

Effect of Pathogens on Host Plant Respiration 87

Respiration o f Diseased Plants 88

Effect of Pathogens on Permeability of Cell Membranes 89

Effects of Pathogens on Transcription and Translation 90

Effect on Transcription 90

Effect on Translation 91

chapter five HOW PLANTS DEFEND THEMSELVES AGAINST PATHOGENS 93

Preexisting Structural and Chemical Defenses 93

Preexisting Defense Structures 93

Preexisting Chemical Defenses 94

Induced Structural and Biochemical Defenses 96

Recognition o f the Pathogen by the Host Plant 96

Transmission o f the Alarm Signal to Host Defense Providers: Signal Transduction 96

Induced Structural Defenses 96

Cytoplasmic defense reaction 97

Cell wall defense structures 97

Histological defense structures 98

Necrotic defense reaction:defense through the hypersensitive response 100

Induced Biochemical Defenses 100

The hypersensitive response 100

Active oxygen radicals,lipoxygenases,and disruption of cell membranes 103

Reinforcement of host cell walls with strengthening molecules 103

Production of antimicrobial substances in attacked host cells 104

Detoxification of pathogen toxins 108

Immunization of plants against pathogens 108

Local and systemic acquired resistance 108

Defense through genetically engineering disease resistant plants 112

chapter six GENETICS OF PLANT DISEASE 115

Introduction 115

Genes and Disease 116

Variability in Organisms 117

Mechanisms of Variability 118

General Mechanisms of Variability 118

Specialized Mechanisms of Variability in Pathogens 119

Stages of Variation in Pathogens 121

Types of Plant Resistance to Pathogens 122

True Resistance 122

Apparent Resistance 124

Genetics of Virulence in Pathogens and of Resistance in Ho st Plants 125

The Gene-for-Gene Concept 126

The Nature o f Resistance to Disease 128

Genetics of Resistance through the Hypersensitive Response 129

Breeding of Resistant Varieties 135

Natural Variability in Plants 135

Effects o f Plant Breeding on Variability in Plants 135

Plant Breeding for Disease Resistance 136

chapter seven ENVIRONMENTAL EFFECTS ON THE DEVELOPMENT OF INFECTIOUS PLANT DISEASE 144

Effect of Temperature 144

Effect of Moisture 146

Effect of Wind 148

Effect of Light 149

Effect of Soil pH and Soil Structure 149

Effect of Host-Plant Nutrition 149

Effect of Herbicides 151

chapter eight PLANT DISEASE EPIDEMIOLOGY 153

The Elements of an Epidemic 153

Host Factors That Affect the Development of Epidemics 155

Pathogen Factors That Affect Development of Epidemics 156

Environmental Factors That Affect Development of Epidemics 158

Effect o f Human Cultural Practices and Control Measures 158

Measurement of Plant Disease and of Yield Loss 159

Patterns of Epidemics 160

Comparison of Epidemics 162

Development of Epidemics 163

Modeling of Plant Disease Epidemics 165

Computer Simulation of Epidemics 166

Forecasting Plant Disease Epidemics 167

Assessment o f Initial Inoculum and of Disease 167

Monitoring Weather Factors That Affect Disease Development 167

Examples o f Plant Disease Forecast Systems 168

Disease-Warning Systems 170

Development and Use o f Expert Systems in Plant Pathology 171

chapter nine CONTROL OF PLANT DISEASES 174

Control Methods That Exclude the Pathogen from the Host 174

Quarantines and Inspections 174

Evasion or Avoidance of Pathogen 175

Use o f Pathogen-Free Propagating Material 175

Control Methods That Eradicate or Reduce Pathogen Inoculum 177

Cultural Methods That Eradicate or Reduce the Inoculum 178

Biological Methods That Eradicate or Reduce the Inoculum 181

Physical Methods That Eradicate or Reduce the Inoculum 188

Chemical Methods That Eradicate or Reduce the Inoculum 190

Disease Control by Immunizing, or Improving the Resistance of, the Host 192

Cross Protection 193

Induced Resistance:Systemic Acquired Resistance 193

Plant Defense Activators 193

Improving the Growing Conditions of Plants 194

Use o f Resistant Varieties 194

Direct Protection of Plants from Pathogens 195

Direct Protection by Biological Controls 195

Fungal antagonists 196;Bacterial antagonists 197

Biological Control of Weeds 199

Direct Protection by Chemical Controls 200

Methods of application of chemicals for plant disease control 203

Types of chemicals used for plant disease control 208

Inorganic chemicals used for plant disease control 208

Organic chemicals:contact protective fungicides 209

Organic compounds: systemic fungicides 210

Miscellaneous organic fungicides 212

Antibiotics 212

Plant oils and petroleum oils 213

Growth regulators 213

Nematicides 213

Mechanisms of action of chemicals used to control plant diseases 214

Resistance of pathogens to chemicals 215

Restrictions on chemical control of plant diseases 216

Integrated Control of Plant Diseases 216

Integrated Control in a Perennial Crop 217

Integrated Control in an Annual Crop 218

part two SPECIFIC PLANT DISEASES 225

chapter ten ENVIRONMENTAL FACTORS THAT CAUSE PLANT DISEASES 225

Introduction 225

General Characteristics 225

Diagnosis 225

Control 226

Temperature Effects 226

High-Temperature Effects 227

Low-Temperature Effects 228

Low-Temperature Effects on Indoor Plants 228

Mechanisms of Low- and High-Temperature Injury to Plants 230

Moisture Effects 230

Low Soil Moisture Effects 230

Low Relative Humidity Effects 231

High Soil Moisture Effects 231

Inadequate Oxygen 232

Light 232

Air Pollution 233

Air Pollutants and Kinds of injury to Plants 233

Main Sources o f Air Pollutants 234

How Air Pollutants Affect Plants 234

Acid Rain 236

Nutritional Deficiencies in Plants 236

Soil Minerals Toxic to Plants 238

Herbicide Injury 240

Other Improper Agricultural Practices 241

The Often Confused Etiology of Stress Diseases 242

chapter eleven PLANT DISEASE CAUSED BY FUNGI 245

Introduction 245

Characteristics of Plant Pathogenic Fungi 245

Morphology 245

Reproduction 246

Ecology 246

Dissemination 248

Classification of Plant Pathogenic Fungi 248

Fungallike Organisms or Pseudofungi 248

The True Fungi 249

Identification 254

Symptoms Caused by Fungi on Plants 254

Isolation o f Fungi (and Bacteria) 255

Preparing for Isolation 255

Isolating the Pathogen 257

Life Cycles of Fungi 258

Control of Fungal Diseases of Plants 259

Diseases Caused by Fungallike Organisms 260

Diseases Caused by Myxomycota (Myxomycetes) 260

Diseases Caused by Plasmodiophoromycetes 263

Clubroot of crucifers 263

Diseases Caused by the Chromistan Fungallike Oomycetes 266

Pythium Seed Rot,Damping-off, Root Rot,and Soft Rot 266

Phytophthora Diseases 270

Phytophthora root and stem rots 270

Late blight of potatoes 274

The Downy Mildews 278

Downy mildew of grape 280

Diseases Caused by True Fungi 283

Diseases Caused by Chytridiomycota(Chytridiomycetes) 283

Diseases Caused by Zygomycetes 283

Rhizopus soft rot of fruits and vegetables 285

Diseases Caused by Ascomycetes and Imperfect (Asexual)Fungi 286

Sooty Molds 289

Leaf Curl Diseases Caused by Tapbrina 293

The Powdery Mildews 295

Powdery mildew of rose 297

Foliar Diseases Caused by Ascomycetes and Imperfect Fungi 298

Alternaria diseases 300

Rice blast disease 303

Black rot of grape 304

Needle casts and blights of conifers 304

Mycospbaerella diseases 307

Banana leaf spot,or Sigatoka disease 307

Cucurbit gummy stem blight 307

Cerospora diseases 309

Septoria diseases 310

Cochliobolus and Pyrenophora diseases of cereals and grasses 310

Stem and Twig Cankers Caused by Ascomycetes and Imperfect Fungi 317

Dothichiza canker of poplar 318

Black knot of plum and cherry 319

Chestnut blight 319

Nectria canker 319

Leucostoma canker 321

Antbracnose Diseases Caused by Ascomycetes and Imperfect Fungi 324

Black spot of rose 324

Glomerella diseases 325

Gnomonia anthracnose and leaf spot diseases328;Colletotrichum (Gloeosporium) diseases 329

Fruit and General Diseases Caused by Ascomycetes and Imperfect Fungi 331

Ergot of cereals and grasses 332

Apple scab 334

Brown rot of stone fruits 336

Botrytis diseases 339

Vascular Wilts Caused by Ascomycetes and Imperfect Fungi 342

Fusarium wilts 343

Fusarium wilt of tomato 343

Verticillium wilts 346

Ophiostoma (Ceratocystis)wilt of elm trees:Dutch elm disease 346

Root and Stem Rots Caused by Ascomycetes and Imperfect Fungi 350

Gibberella diseases 351

Fusarium root and stem rots of nongrain crops 352

Take-all of wheat 354

Sclerotinia diseases 355

Pbymatotrichum root rot 358

Postharvest Diseases o f Plant Products Caused by Ascomycetes and Imperfect Fungi 359

Postharvest decays of fruits and vegetables 361

Alternaria 361

Botrytis 362

Fusarium,Geotrichuni,Penicillium,Sclerotinia 363

Control of postharvest decays of fresh fruits and vegetables 364

Postharvest decays of grain and legume seeds364;Mycotoxins and mycotoxicoses365; Aflatoxins 365

Fusarium toxins 366

Other Aspergillus toxins andPenicillium toxins 366

Diseases Caused by Basidiomycetes 368

The Rusts 368

Cereal rusts 371

Stem rust of wheat and other cereals 371

Cedar-apple rust 375

Coffee rust 375

Rusts of forest trees 377

White pine blister rust 378

Fusiform rust 380

The Smuts 382

Corn smut 382

Kernel smuts of small grains 384

Loose smut of cereals 384

Covered smut,or bunt,of wheat 387

Root and Stem Rots Caused by Basidiomycetes 389

Root and stem rot diseases caused by the “sterile fungi”Rhizoctonia and Sclerotium 390

Rhizoctonia diseases 390

Sclerotium diseases 395

Root rots of trees 397

Armillaria root rot of fruit and forest trees 397

Wood Rots and Decays Caused by Basidiomycetes 399

Mycorrhizae 404

Ectomycorrhizae 404

Endonnycorrhizae 404

chapter twelve PLANT DISEASES CAUSED BY PROKARYOTES:BACTERIA AND MOLLICUTES 407

Introduction 407

Plant Diseases Caused by Bacteria 408

Characteristics of Plant Pathogenic Bacteria 409

Morphology 409

Reproduction 411

Ecology and Spread 412

Identification of Bacteria 412

Symptoms Caused by Bacteria 415

Control o f Bacterial Diseases of Plants 415

Bacterial Spots and Blights 416

Wildfire of tobacco 417

Bacterial blights of bean 418

Angular leaf spot of cucumber 419

Angular leaf spot of cotton 420

Bacterial leaf spots and blights of cereals and grasses 421

Bacterial spot of tomato and pepper 421

Bacterial speck of tomato 422

Bacterial spot of stone fruits 422

Bacterial Vascular Wilts 423

Bacterial wilt of cucurbits 424

Fire blight of pear and apple 426

Ring rot of potato 429

Bacterial canker and wilt of tomato 430

Southern bacterial wilt of solanaceous plants and Moko disease of banana 431

Black rot of crucifers 433

Bacterial Soft Rots 434

Bacterial soft rots of vegetables 435

Bacterial Galls 438

Crown gall 438

Bacterial Cankers 441

Bacterial canker and gummosis of stone fruit trees 442

Citrus canker 445

Bacterial Scabs 449

Common scab of potato 449

Root Nodules of Legumes 451

Plant Diseases Caused by Fastidious Vascular Bacteria 453

Pierce’s disease of grape 455

Citrus variegated chlorosis 456

Ratoon stunting of sugarcane 456

Plant Diseases Caused by Mollicutes:Phytoplasmas and Spiroplasmas 457

Properties of True Mycoplasmas 457

Phytoplasmas 458

Spiroplasmas 460

Other Organisms That Resemble Mollicutes:L-Forms of Bacteria 462

Examples of Plant Diseases Caused by Mollicutes 462

Aster yellows 462

Lethal yellowing of coconut palms 464

Elm yellows(phloem necrosis) 465

Peach X-disease 465

Pear decline 467

Citrus stubborn disease 467

Corn stunt disease 469

chapter thirteen PLANT DISEASES CAUSED BY PARASITIC HIGHER PLANTS 471

Introduction 471

Dodder 471

Witchweed 473

Broomrapes 475

Dwarf Mistletoes of Conifers 475

True or Leafy Mistletoes 477

chapter fourteen PLANT DISEASES CAUSED BY VIRUSES 479

Introduction 479

Characteristics of Plant Viruses 479

Detection 480

Morphology 480

Composition and Structure 480

Satellite Viruses and Satellite RNAs 484

The Biological Function of Viral Components:Coding 485

Virus Infection and Virus Synthesis 485

Translocation and Distribution of Viruses in Plants 488

Symptoms Caused by Plant Viruses 489

Physiology of Virus-Infected Plants 491

Transmission of Plant Viruses 491

Transmission of Viruses by Vegetative Propagation 492

Mechanical Transmission o f Viruses through Sap 492

Seed Transmission 494

Pollen Transmission 494

Insect Transmission 494

Mite Transmission 495

Nematode Transmission 495

Fungus Transmission 496

Dodder Transmission 496

Epidemiology of Plant Viruses and Viroids 496

Purification of Plant Viruses 497

Serology of Plant Viruses 497

Nomenclature and Classification of Plant Viruses 500

Detection and Identification of Plant Viruses 503

Economic Importance of Plant Viruses 505

Control of Plant Viruses 505

Diseases Caused by Rigid Rod-Shaped Viruses 508

Diseases Caused by Tobamoviruses:Tobacco Mosaic Virus 508

Diseases Caused by Tobraviruses 510

Diseases Caused by Furoviruses 510

Diseases Caused by Hordeiviruses 511

Diseases Caused by Filamentous Viruses 512

Diseases Caused by Potexviruses 512

Diseases Caused by Carlaviruses 512

Diseases Caused by Capilloviruses and Trichoviruses 512

Diseases Caused by Potyviridae 513

Diseases Caused by Potyviruses 513

Bean common mosaic and bean yellow mosaic 515

Lettuce mosaic 515

Papaya ringspot 516

Plum pox 517

Potato virus Y 517

sugarcane mosaic 518

Tobacco etch 518

Turnip mosaic 519

Watermelon mosaic 519

Zucchini yellow mosaic 519

Diseases Caused by Rymoviruses and Bymoviruses 519

Diseases Caused by Closteroviruses 519

Beet yellows 519

Citrus tristeza 519

Lettuce infectious yellows 521

Diseases Caused by Isometric Single-Stranded RNA Viruses 523

Diseases Caused by Waikaviruses 523

Rice tungro 523

Maize chlorotic dwarf 525

Diseases Caused by Luteoviruses 525

Beet western yellows 525

Potato leafroll 525

Barley yellow dwarf 525

Diseases Caused by Comoviridae 527

Diseases Caused by Comoviruses 527

Diseases Caused by Nepoviruses 528

Tomato ring spot 528

Cherry leaf roll 528

Grapevine fanleaf 529

Raspberry yellow dwarf,raspberry ring spot,and raspberry leaf curl 529

Diseases Caused by Brommoviridae 530

Diseases Caused by Cucumoviruses 530

Cucumber mosaic 531

Diseases Caused by Ilarviruses 532

Prunus necrotic ring spot 533

Diseases Caused by Isometric Double-Stranded RNA Viruses 534

Diseases Caused by Reoviridae 536

Diseases Caused by Negative RNA[(一)ssRNA]Viruses 538

Plant Diseases Caused by Rhabdoviruses 538

Plant Diseases Caused by Tospoviruses 539

Plant Diseases Caused by Tenuiviruses 541

Diseases Caused by Double-Stranded DNA Viruses 545

Diseases Caused by Caulimoviruses 545

Diseases Caused by Badnaviruses 546

Diseases Caused by Single-Stranded DNA Viruses 547

Plant Diseases Caused by Geminiviruses 547

Maize streak 551

Beet curly top 551

African cassava mosaic 551

Bean golden mosaic 552

Squash leaf curl 553

Tomato mottle 553

Tomato yellow leaf curl 554

Plant Diseases Caused by Isometric Single-Stranded DNA Viruses 554

Banana bunchy top 555

Viroids 556

Plant Diseases Caused by Viroids 556

Potato spindle tuber 560

Citrus exocortis 560

Coconut cadang-cadang 561

chapter fifteen PLANT DISEASES CAUSED BY NEMATODES 565

Introduction 565

Characteristics of Plant Pathogenic Nematodes 565

Morphology 565

Anatomy 565

Life Cycles 566

Ecology and Spread 566

Classification 568

Isolation of Nematodes 569

Isolation of Nematodes from Soil 569

Isolation of Nematodes from Plant Material 570

Symptoms Caused by Nematodes 571

How Nematodes Affect Plants 571

Interrelationships between Nematodes and Other Plant Pathogens 572

Control of Nematodes 573

Root-Knot Nematodes:Meloidogyne 574

Cyst Nematodes:Heterodera and Globodera 577

Soybean cyst nematode:Heterodera glycines 577

Sugar beet nematode:Heterodera schachtii 579

The Citrus Nematode:Tylenchulus semipenetrans 581

Lesion Nematodes:Pratylenchus 581

The Burrowing Nematode:Radopbolus 583

Stem and Bulb Nematode:Ditylenchus 586

Seed-Gall Nematodes: Anguina 588

Foliar Nematodes:Aphelencboides 590

Pine Wilt and Palm Red Ring Diseases:Bursaphelenchus 592

Pine wilt nematode:Bursaphelenchus xylopbilus 592

Red ring nematode:Bursapbelencus cocopbilus 593

Stubbly-Root Nematodes:Paratricbodorus and Tricbodorus 596

chapter sixteen PLANT DISEASES CAUSED BY FLAGELLATE PROTOZOA 599

Introduction 599

Phloem Necrosis of Coffee 601

Hartrot of Coconut Palms 604

Sudden Wilt (Marchitez)of Oil Palm 604

Empty Root of Cassava 605

Glossary 607

Index 619

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