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Plant Pathology
Third Edition
 About the Authors

R. S. Mehrotra
A Ph.D. from the University of Saugor, (now Dr. Hari Singh Gour University),
Sagar, Dr. R. S. Mehrotra has a teaching and research experience of more than
35 years. He retired from Kurukshetra University as Professor in 1997. He was
a Postdoctoral Fellow at the University of Western Ontario, London, Ontario,
Canada (1968-1970) and a Fulbright Fellow at the University of California,
Riverside Campus (1979). He is a Fellow of National Academy of Sciences,
Allahabad, and was elected President of the Indian Phytopathological Society
(1990), President of Indian Society of Plant Pathologists (1992), President of
the Mycological Society of India (1996), and President of the Botanical Sec-
tion of Indian Science Congress (1997). He was the recipient of the Birbal Sahni Gold Medal of the
Indian Botanical Society in 1998. Professor Mehrotra delivered the Dastur Memorial Lecture of the
Indian Phytopathological Society in 2010. He was awarded the I.P.S. recognition award by the Indian
Phytopathological Society for the year 2014. He has published around 200 research papers in national
and international journals and has authored 5 books—Fundamentals of Plant Pathology (McGraw Hill
Education, India), An Introduction to Mycology, A Monograph on Phytophthora Diseases in India,
Principles of Microbiology (McGraw Hill Education, India), and Fungal Diversity and Biotechnology.

Ashok Aggarwal
Ashok Aggarwal is currently working as Professor of Botany and teaches
Plant Pathology, Mycology and Microbiology at the Department of Botany,
Kurukshetra University, Kurukshetra, where he has been a faculty member for
the past 3 decades. He is also a member of several scientific organizations and
has published more than 190 research papers and review articles in reputed
journals of high impact factors. He has co-authored two monographs on the
topic entitled, Phytophthora Diseases in India, in 2001, and Endomycorrhizal
Diversity on Ethnomedicinal Plants of Himachal Pradesh, VDM Verlag Pub. Germany, in 2013. In ad-
dition, he has written a textbook entitled Fundamentals of Plant Pathology, published by McGraw Hill
Education (India) Private Ltd., with Professor Mehrotra. He has also edited few other books—Glimpses
of Plant Sciences, in 2002; and Mycorrhizal Fungi, with Dr. Kuldeep Yadav, in 2017. Dr. Aggarwal has
guided 10 M.Phil and 25 Ph.D. students on diverse aspects of mycorrhizal fungi.
 Plant Pathology
Third Edition

R. S. Mehrotra
Retired Professor of Botany
Former Dean, Faculty of Science
Kurukshetra University
Kurukshetra

Ashok Aggarwal
Professor and Former Chairman
Department of Botany
Kurukshetra University
Kurukshetra

McGraw Hill Education (India) Private Limited

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Plant Pathology, 3e
Copyright © 2017 by McGraw Hill Education (India) Private Limited
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 Dedicated to
Professor S. B. Saksena
For his inspiration as a teacher
(On his centenary birth anniversary)
 Preface to the
Third Edition

Overview
Anyone who ever grows a plant or crop not only enjoys the rewards of beautiful flowers, fruits, seeds,
vegetables, etc., but also faces the disappointment when the plants or crops become diseased. Plant
diseases cause great losses annually throughout the world and reduce the yield even with the use
of improved varieties and technologies. The prospects of increasing production of various crops by
increasing land under cultivation are limited. Because of dependence of humans for food, fiber and other
resources, understanding plant diseases and their control is vital for our survival, and there is a need to
increase productivity of various crops to meet the future demand and to achieve the goal of sustainable
agriculture development.
We all know that newer researches and findings in the field of sciences regularly enhance the
knowledge-base and dimensions of the subject. Nature has empowered us with the intellect and curiosity
to think, to question, and to go beyond the limits of general perceptions. Science gives us a direction
to discover, experiment and reveal the hidden truths of the Universe for further studying and exploring
them. A theory or a notion without queries and research becomes out of date very soon. In the field of
Plant Pathology also, knowledge of techniques and newer researches are always necessary to remain up
to date.

Aim of the Book

Keeping in view these facts, this new edition of Plant Pathology is written to provide recent information
and advances in the field, mainly for the students of plant pathology, agriculture and plant sciences. The
present book is a joint effort made after years of teaching and research and months of focused thinking
so that it may help students to develop a deep understanding of the subject.
We have presented the latest information, leavened the lessons learned from our own experiences as
well as colleagues, students and others, on precise and well-defined topics in a concise manner.

Salient Features
● The textbook covers approximately 132 diseases of crop plants, providing details of symptoms,
latest nomenclature of causal organisms and their control measures.
viii Preface to the Third Edition

● It attempts to provide postgraduate and professional readers, already familiar with the subject, with
a means to acquire latest and deep knowledge on various aspects of plant pathology.
● Profusely illustrated with more than 191 figures in the form of diagrams and photographs. Some
diagrams are redrawn so as to improve the quality of the book.
● Most of the chapters have been revised with recent information like Chapter 2 (Food safety, Biological
warfare, Bioterrorism); Chapter 4 (Cercosporin toxin), Chapter 5 (Effect of polysaccharides and
growth regulators on pathogen/ diseases); Chapter 6 (Defence through genes); Chapter 7 (Genetics
of virulence in pathogens and resistance in host plants); Chapter 8 (Effect of silicon on nutrition and
disease development); Chapter 9 (Overwintering of pathogens and their role in epidemics); Chapter
10 (Host physiology/Factors based prediction, role of molecular tools, GIS, GPS and remote sensing
based prediction); Chapter 11 (Control of diseases through use of transgenic plants); Chapter 12
(recent classification of fungi by Kirk et al., 2008); Chapter 13 (Collar rot of apple); Chapter 17
(Fusarium Head Blight of wheat, Bakanae or foot rot disease of rice); Chapter 20 (Post harvest
decay of grains and legumes, different mycotoxins, methods of preservations); Chapter 26 (Recent
classification of nematodes); Chapter 29 (role of innovative tools like monoclonal, recombinant
antibodies, flow cytometry, FISH, PCR, DNA microarray, gene pyramiding in the field of plant
pathology); and Chapter 30 (maintenance and preservation of microbial cultures).

Chapter Organization
The book is unique as it contains information on general concepts as well as information on specific
diseases. From chapters 1 to 10, the authors have discussed the host pathogen interactions in detail,
and in the later chapters, have given a detailed account of diseases caused by fungi, bacteria, viruses,
phytoplasmas and nematodes.
The book adopts multidisciplinary approach in addressing both basic and applied aspects of plant
diseases management. Latest information regarding disease management has been added from journals,
review articles and books to make the subject up to date in this edition.
R. S. MehRotRa
aShok aggaRwal

Publisher’s Note:
McGraw Hill Education (India) invites suggestions and comments, all of which can be sent to
info.india@mheducation.com (kindly mention the title and author name in the subject line).
Piracy-related issues may also be reported.
 Preface to the
First Edition

Plant pathology is intimately related to the health of plant and the production of crop yields. It has
special relevance in agricultural countries like India which are fast developing, and where much still
needs to be done in the modernization of control measures against plant diseases. Rough estimates have
suggested that in India, on an average, there is a loss of about 10 per cent in crop yield due to plant
diseases. This loss would certainly amount to hundreds of crores of rupees annually, taking into account
the output of all the cultivated crops. Hence, a book on plant pathology is always welcome.
During the last quarter of the century, there have been enormous developments in the science of plant
pathology. Many new techniques and chemicals have been evolved to fight plant diseases. Efforts have
been made to understand the nature of diseases at molecular levels and the relationship between the host
and parasite has been put, in many cases, on a sound biochemical footing. It is a stupendous task to pres-
ent this knowledge in a volume of suitable size to a young post graduate or research student.
The book has several important features, one of which is the “all-in-one” approach which will be
useful to postgraduate students. To the best of my knowledge there is no book with both generalized and
specialized topics discussed in parallel which caters to the needs of undergraduate, postgraduate and
research students at the same time. Most of the important aspects of plant pathology, such as physiol-
ogy of diseased plants, root diseases, seed-borne diseases, post harvest diseases of fruits and vegetables,
diseases caused by fungi, bacteria, viruses, mycoplasmas, nematodes and non-parasitic diseases have all
been dealt with suitable detail for the benefit of students. Besides, the book should also serve as a good
reference for research workers.
It is hoped that this work will enrich the background of those who wish to study plant pathology in-
tensively. Suggestions for improving the book will be gratefully received and acknowledged.

R. S. MehRotRa
 Acknowledgements

The authors are thankful to Prof. M. P. Srivastava, Retired Professor of Plant Pathology, CCS HAU,
Hisar, and Late Dr. Y. R. Sarma, Ex-Director, National Centre for Spices, Calicut, for providing useful
information, suggestions and encouragement. Thanks are also due to Prof. T. S. Thind, Punjab Agri-
cultural University, Ludhiana, and Prof. Ram Singh, Rice Research Station, HAU, Kaul, for providing
photographs of different diseased symptoms. The authors thank Dr. B.B. Nagaich, former Director,
Central Potato Research Institute, Shimla and Dr. S.M. Paul Khurana, former Director C.P.R.I., Shimla,
for providing photographs of diseases of potato; Dr. J.C. Majumdar of BASF India Limited, for permis-
sion to reproduce some photographs; Late Dr. M.M. Payak, for providing photographs of diseases of
maize; Late Dr. R.K. Grover, former Professor and Head, Department of Plant Pathology, CCS Haryana
Agriculture University, Hisar, for providing photographs of downy mildew of bajra and angular leaf spot
of cotton; Dr. R.N. Gupta, for providing photographs of the germinating chlamydospore of Protomyces
macrosporous, and Dr. D.K. Chakravarti, Department of Horticulture, N.D. University of Agriculture
and Technology, Faizabad, Uttar Pradesh, for providing the photographs of mango malformation and
providing relevant information on this disease.
Acknowledgements are also due to the following:
● Agrios, G.N. (1969) Plant Pathology, Academic Press, New York, for reproducing Figures 3.1, 4.3,
4.4, 13.1, 13.3, 15.2, 15.6, 15.22, 17.1, 17.5, 25.1, 26.1, 26.5, 26.6, and 26.7.
● Bateman, D.F. and H.G. Basham (1976), Physiological Plant Pathology, edited by R. Heitefuss and
P.H. Williams, Springer Verlag, Berlin, Heidelberg, New York, for giving permission to reproduce
Figure 4.2 from their article.
● Dickinson, C.H. and J.A. Lucas (1977), Plant Pathology and Plant Pathogens, Blackwell Scientific
Publications, Oxford, London, Edinburg, Melbourne, for giving permission to reproduce seven fig-
ures (Figures 5.2, 5.3, 5.4, 5.5, 5.6, 5.8, and 5.10).
● Raychaudhuri, S.P. and T.K. Nariani (1977), Virus and Mycoplasmal Diseases of Plants in India,
Oxford and IBH Publication Co., New Delhi, Bombay, Calcutta, for granting permission to repro-
duce the photographs of papaya leaf curl.
We are also thankful to our students, especially Dr. Kuldeep Yadav and Esha for their help in the
completion of the book.
It must be emphasized that it would not have been possible to complete this book without the will-
ing help rendered to the authors by their colleagues and research associates. Last but not the least, the
authors express their appreciation and thanks to their family members for their unwavering support and
encouragement.
R. S. MehRotRa
aShok aggaRwal
 Contents

Preface to the Third Edition vii

Preface to the First Edition ix
Acknowledgements xi

1. Introduction 1
Nature and Concept of Plant Diseases 1
Cause of Diseases 3
Various Levels of Parasitism 6
Integrated Disease/Pest Management 10
Importance of Plant Diseases 10
Effect of Plant Diseases on Human Affairs 10
Classification of Plant Diseases 16
Trends in Teaching and Research in Plant Pathology Worldwide 17
Selected References 18
Questions 20

2. History of Plant Pathology 21

The Herbalists 22
The Systematists 22
The Beginning of Modern Period 23
Confirmation of Prevost’s Work 24
Doctrine of Spontaneous Generation 24
Forest Pathology 26
Discovery of Bordeaux Mixture 26
Plant Pathology in The Twentieth Century 27
Physiological Plant Pathology 27
Genetics of The Host and The Pathogen 28
Fungicidal Research 28
Environment in Relation to Plant Disease 29
Nature of Disease Resistance 29
Biochemistry and Physiology of The Diseased Host Plant 30
xiv Contents

Molecular Biology of Pathogenesis and Induced Systemic Resistance 30

Tissue Culture in Plant Pathology 30
Biotechnological Approach Towards Disease Control 31
Ecological Studies of Soil-Borne Fungal Plant Pathogens 31
Bacteriology 31
Virology (Viruses, Viroids and Prions) 32
Prions 33
Mollicutes 33
Nematology 34
Protozoans as Plant Pathogens 34
History of The Development of Plant Pathology in India 35
Trends in Teaching and Training in Plant Pathology in India 38
Holistic Approaches for Plant Health Management 39
Plant Disease Clinics 40
Education and Information Technology in Plant Pathology 41
Innovations in Teaching Plant Pathology 41
Some Aspects of Applied Plant Pathology 42
Food Safety 43
Biological Warfare, Bioterrorism 43
Plant Pathology Today and Future Directions: Molecular Approaches 44
Selected References 45
Questions 51

3. Pathogenesis 52
Penetration and Entry by Plant Pathogens 52
Host Penetration and Infection 55
Prepenetration 56
Entry Through Natural Openings 58
Direct Penetration 60
Entry Through Wounds 65
Wounds Caused By Other Fungi 66
Wounds Caused by Nematodes 66
Entry Through Root Hairs 66
Entry Through Buds 67
Development Inside Host Tissue 67
Selected References 69
Questions 71

4. Enzymes and Toxins in Plant Diseases 72

Enzymes in Plant Diseases 72
Composition of Cell Wall Materials and Middle Lamella 73
Enzymes for Waxes and Cutins 78
Pectic Enzymes 80
Macerating Enzymes 81
 Contents xv

Cellulolytic Enzymes 83
Hemicellulases 85
Lignolytic Enzymes 85
Proteolytic Enzymes 86
Lipolytic Enzymes 86
Inactivation of Enzymes 87
Toxins and Plant Diseases 89
Classification of Toxins 90
Selected References 104
Questions 109

5. Alteration in Plant Physiological Functions Due to Plant Pathogen Interaction 110

Permeability Changes in Diseased Plants 110
Effect of Plant Pathogens on Translocation of Water and Nutrients in Host Plant 112
Effect of Pathogens on Translocation of Nutrients and Water 113
Interference with Upward Translocation of Water and Inorganic Nutrients 114
Photosynthesis 114
Photosynthesis as Influenced by Virus Infection 117
Photosynthesis as Influenced by Bacterial Pathogens 118
Photosynthesis as Influenced by Fungi 119
Respiration 122
Mechanism of Respiratory Increase 124
Uncoupling of Oxidative Phosphorylation 125
Augmented Synthetic Processes 126
Changes in Respiratory Pathways 126
Increased Activity of Terminal Oxidases 127
Relationship of Resistance and Susceptibilty 128
Effect on Transpiration 128
Changes in Nitrogen Metabolism of Infected Plants 129
Changes in Protein Metabolism 130
Phenols 131
Phenol Metabolism 134
Importance of Phenol Oxidation in Plant Diseases 135
Growth Regulators in Plant Diseases 137
Transcription and Translation in Diseased Plants 144
Polysaccharides 144
Detoxification of Antimicrobial Molecules 145
Promotion of Bacterial Virulence by Avr Genes 145
Effect of Type III Secretion In Bacterial Pathogenesis 145
Suppressors of Plant Defense Responses 146
Pathogenicity and Virulence Factors in Viruses and Viroids 146
Interference with Translocation of Organic Nutrients Through The Phloem 146
Effect of Pathogens on Plant Growth 147
Selected References 147
Questions 154
xvi Contents

6. How Plants Defend Themselves Against Infection 155

Morphological or Structural Defense Mechanism 155
Biochemical Defense 160
Defense Through Induced Synthesis of Proteins and Enzymes 172
Defense Through Formation of Substrates Resisting the Enzymes of Pathogens 175
Defense Through Detoxification of Pathogen Toxin 176
Defense Through Altered Respiration 176
Defense Through Hypersensitive Reaction 176
The Concept of Phytoncides 177
Defense Through Genes 177
Lack of Recognition Between Host and Pathogen 178
Defense Through Lack of Host Receptors and Sensitive Sites for Toxins 179
Defense Through Recognition of Pathogen by the Host Plant 179
Defense Through Transmission of the Alarm Signal to Host Defense System 179
Induced Biochemical Defense in Hypersensitive Response (Race-Specific, Monogenic,
R-Gene Or Vertical Resistance) 180
Defense Involving Bacterial Type III-Effector Proteins 182
Reinforcement of Host Cell Walls with Strengthening Molecules 184
Production of Antimicrobial Substances (Pathogenesis Related Proteins) in Infected
Host Cell 184
Defense Through Genetically Engineered Disease-Resistant Plants 184
Defense with Pathogen-Derived Genes 185
Defense Through RNA Silencing by Pathogen Derived Genes 186
Selected References 187
Questions 191

7. Genetics of Plant Pathogen Interaction 192

Genetics of Host–Parasite Interaction 192
Resistance and Susceptibility 192
The Common Antigen Hypothesis 195
The Phytoalexin-Induced Hypothesis 196
Vertical and Horizontal Resistance 196
Mutation 196
Heterokaryosis 197
Parasexual Recombination 199
Adaptation 199
Saltation 200
Cytoplasmic Variation 200
Sexual Recombination 200
Transformation 201
Transduction 201
Physiological Specialization 202
Adaptation of Fungi to Different Hosts 205
Apparent Resistance 207
 Contents xvii

Genetics of Virulence in Pathogens and of Resistance in Host Plants 208

Selected References 212
Questions 215

8. Effect of Environmental Factors and Nutrition on Disease Development 216

Effect of Temperature on the Development of a Disease 216
Humidity and Moisture 218
Soil pH 220
Soil Texture 220
Effect of Light 221
Effect of Oxygen and Carbon Dioxide Concentration 221
Effect of Wind/Air 222
Effect of Herbicides/Fungicides 222
Effect of Air Pollution 222
Influence of Nutrition on Disease Development 223
Role of Biotic Environment 226
Role of Environmental Factors in Epiphytotics 226
Selected References 227
Questions 228

9. Survival and Dispersal of Plant Pathogens 230

Survival of Plant Pathogens 230
Direct Transmission 233
Indirect Transmission 234
Overwintering of Pathogens and Their Role in Epidemics 243
Survival Strategies Adopted by Plant Pathogens 244
Selected References 244
Question 246

10. Plant Disease Epidemiology and Plant Disease Forecasting 247

Introduction 247
Some Important Epiphytotics of the Past and Present 249
Epiphytotic Growth and Analysis 250
Computer Simulation of Epidemics 250
System Approach in Epidemiology 251
Methods Used in Plant Disease Forecasting 254
Examples of Plant Disease Forecasting Systems 260
Selected References 261
Questions 264

11. Management of Plant Diseases 266

Cultural Methods 266
Chemical Methods 281
Disease Control by Immunizing the Host 304
Control Through Use of Transgenic Plants 305
xviii Contents

Breeding for Disease Resistance 306

Innovative Methods of Plant Disease Control 320
Selected References 322
Questions 328

12. Plant Diseases Caused By Fungi—Characteristics and Classification of

Plant Pathogenic Fungi 330
General Features of Fungi 330
Classification of Fungi 332
Selected References 348
Questions 349

13. Rots, Damping Offs, Downy Mildews and White Rusts 350
Damping Off Fungal Pathogens, Downy Mildews and White Rusts 350
Key to the Families of the Order Peronosporales 351
Key to the Important Genera of Peronosporaceae 351
Key to the Important Genera of Pythiaceae 351
Damping Off Disease of Seedlings 352
Damping Off of Vegetables Seedlings 352
Fruit Rot of Cucurbits 356
Stem or Foot Rot of Papaya (Carica papaya) 357
Rhizome Rot of Ginger (Zingiber officinale) 358
Late Blight of Potato (Solanum tuberosum) 359
Blight of Colocasia (Colocasia esculenta) 366
“Koleroga” or “Mahali” of Areca Palms (Areca catechu) 369
Bud Rot of Palms 370
Root Rot, Crown Rot, Foot Rot, Gummosis and Brown Rots of Citrus Fruits 372
Leaf Rot and Foot Rot of “Pan” (Piper betle) 374
Collar Rot of Apple 378
Downy Milew of Peas (Pisum sativum) 379
Downy Mildew of Crucifers 380
Downy Mildew of Cucurbits 382
Downy Mildew of Grapes (Vitis vinifera) 383
Downy Mildew of Bajra (Pennisetum typhoides) 385
Downy Mildew or Leaf Shredding Disease of Sorghum 388
White Rust of Crucifers 390
Some Other Diseases Caused By Oomycota 392
Selected References 393
Questions 403
 Contents xix

14. Powdery Mildews 404

Powdery Mildews 404
Powdery Mildew of Wheat (Triticum spp.) and Barley 407
Powdery Mildew of Peas (Pisum sativum) 410
Powdery Mildew Of Grapevine 413
Powdery Mildew of Cucurbits 415
Powdery Mildew of Apple 418
Powdery Mildew of Mango 419
Selected References 420
Questions 424

15. Smuts and Bunts 425

Modes of Infection 427
Loose Smut of Wheat 429
Flag Smut of Wheat 432
Hill Bunts or European Bunts or Stinking Smuts of Wheat 434
Karnal Bunt of Wheat 437
False Smut of Paddy 440
Bunt of Rice (Syn. Black Smut, Kernel Smut, Rice Smut) 442
Leaf Smut of Rice 443
Covered Smut, Kernel Smut, Short Smut or Grain Smut of Jowar (Sorghum vulgare) 445
Loose Smut of Sorghum 447
Head Smut of Sorghum and Maize 448
Long Smut of Sorghum 450
Smut Of Bajra (Pennisetum typhoides) or Pearl Millet 452
Common Smut of Maize 453
Whip Smut of Sugarcane 456
Covered Smut of Barley (Hordeum vulgare) 458
Loose Smut of Barley 460
Loose and Covered Smuts of Oats (Avena sativa) 460
Selected References 462
Questions 466

16. Rusts 467

Plant Rusts 467
Life Cycles 468
Heteroecism and Autoecism 469
Rusts of Wheat 474
Black Stem Rust of Wheat 474
Brown Rust of Wheat 481
Yellow Rust of Wheat 483
Rust of Linseed (Linum usitatissimum) 485
Rust of Beans (Phaseolus vulgaris) 488
Rust of Pea (Pisum sativum and Lens esculenta) 489
Rust of Gram (Cicer arietinum) 490
xx Contents

Rust of Coffee 492

Rust of Groundnut 494
Selected References 495
Questions 498

17. Wilts, Root Rots and Head Blight 499

Vascular Wilts 499
Wilt of Pigeon Pea (Cajanus cajan) 505
Wilt of Cotton (Gossypium arboreum and G. herbaceum) 509
Wilt of Linseed (Linum usitatissimum) 511
Fusarium Head Blight of Wheat 512
Panama Disease of Banana 515
Dutch Elm Disease 517
Root Rots 520
Rhizoctonia Root Rot of Cotton 520
Root and Stem Rot of Jute 522
Black Scurf of Potato 523
Stem Rot of Rice 524
Sheath Blight of Rice (Oryza sativa L.) 526
Bakanae or Foot Rot Disease of Rice 528
Selected References 531
Questions 537

18. Leaf Spots, Leaf Blights and Anthracnoses 538

Leaf Spots and Leaf Blights 538
Leaf Spot of Turmeric (Curcuma longa) 539
Brown Spot of Maize 540
Early Blight of Potato 542
Leaf Blight of Wheat 544
Stripe Disease of Barley 546
Brown Leaf Spot (Helminthosporiosis) Disease of Rice 548
Blast Disease of Rice 551
Tikka Disease of Groundnuts 557
Anthracnoses 560
Red-Rot of Sugarcane 560
Ripe Fruit-Rot and Die-Back of Chillies 564
Mango Anthracnose 566
Anthracnose of Grapes 567
Ascochyta Blight of Chickpea (Cicer arietinum L.) 569
Selected References 571
Questions 577

19. Galls and Abnormal Growths 578

Clubroot Disease of Cabbage (Brassica oleracea var. capitata) 578
Wart Disease of Potato (Solanum tuberosum L.) 581
 Contents xxi

Stem Gall of Coriander (Coriandrum sativum) 583

Leaf Curl of Peach (Prunus persica) 586
Ergot of Bajra (Pearl Millet) 587
Malformation of Mango 591
Selected References 596
Questions 600

20. Post-Harvest Diseases of Fruits, Vegetables and Post-Harvest Decay of

Grains and Legumes 601
Bacterial Rots 602
Fungi 603
Non-Pathogenic Diseases 605
Mode of Infection and Factors Influencing Post-Harvest Diseases 607
Control of Post-Harvest Diseases 608
Post-Harvest Decay of Grains and Legume Seeds 617
Methods of Prevention 619
Selected References 621
Questions 623

21. Root Diseases 624

Root Exudates and Soil-Borne Pathogens 625
Associative and Antagonistic Effects 626
Predisposing Factors 628
Survival 629
Dispersal of Soil-Borne Pathogens 629
Control of Root Diseases 630
Selected References 634
Questions 637

22. Seed-Borne Diseases 638

Seed-Borne Bacterial Diseases 638
Seed-Borne Viral Diseases 639
Seed-Borne Fungal Diseases 640
Seed-Borne Nematode Diseases 645
Pathological Effects of Seed-Borne Diseases 646
Quarantine Programmes for Seed Borne and Seed Stocks 648
Seed Treatment 649
Selected References 652
Questions 655

23. Plant Diseases Caused by Mollicutes: Phytoplasmas and Spiroplasmas 656

Sandal Spike 659
Sesamum Phyllody 661
Grassy Shoot Disease (GSD) of Sugarcane 662
Little Leaf of Brinjal (Solanum melongena) 664
xxii Contents

Coconut (Root) Wilt 665

Selected References 668
Questions 674

24. Bacteria and Bacterial Diseases 675

Structure, Reproduction and Classification of Bacteria 675
Growth and Reproduction in Bacteria 680
Classification of Bacteria (Procaryotes) 685
Plant Pathogenic Bacteria 692
Symptoms of Bacterial Plant Diseases 694
Bacterial Plant Diseases 695
Bacterial Blight of Paddy 695
Bacterial Leaf Streak 700
Citrus Canker 702
Bacterial Blight, Angular Leaf Spot and Black Arm of Cotton 704
Black Rot of Crucifers 708
Wilt and Brown Rot of Potato 709
Yellow Ear Rot or Tundu Disease of Wheat 713
Bacterial Soft Rot and Blackleg of Potato 715
Fire Blight of Apples and Pears 717
Crown Gall of Stone Fruits 718
Stalk Rot of Maize 720
Common Scab of Potato 721
Ratoon Stunting Disease (RSD) of Sugarcane 723
Citrus Greening Disease (CGD) 724
Angular Leaf Spot and Wildfire of Tobacco 726
Selected References 727
Questions 734

25. Viruses, Viroids and Their Diseases 736

Plant Virsues and Plant Viral Diseases 736
Symptoms of Viral Infections in Plants 739
Transmission of Plant Viruses 743
Control of Viral Diseases 747
Structure of Plant Viruses 749
Classification of Plant Viruses 750
Virus Purification 753
Viroids 754
Physical and Chemical Properties of Plant Viruses 755
Infections, Multiplication and Movement of Viruses Within Plants 756
Tobacco and Tomato Mosaic 757
Viral Diseases of Potato 761
Leaf Curl of Tomato 764
Mosaic Diseases of Tomato 765
Viral Diseases of Papaya 766
 Contents xxiii

Yellow Vein Mosaic of Bhindi (Abelmoschus esculentus) 767

Viral Diseases of Sugarcane 768
Bunchy Disease of Banana 769
Bean Common Mosaic 770
Tristeza of Citrus 770
Citrus Exocortis (Viroid Disease) 772
Cadang-Cadang Disease of Coconut (Viroid Disease) 773
Potato Spindle Tuber (Viroid Disease) 774
Selected References 775
Questions 779

26. Diseases Caused by Nematodes and Flagellates 780

Diseases Caused by Plant Parasitic Nematodes 780
Isolation of Nematodes 782
Symptoms Caused by Nematodes 784
Mechanism of Nematode Injury to Plants 785
Interrelationship between Nematodes and Other Plant Pathogens 787
Ecology and Spread 788
Characteristics of Plant Pathogenic Nematodes 789
General Methods of Control of Nematode Diseases 792
Root Knot of Vegetables 798
Molya Disease of Barley and Wheat 800
Citrus Nematode Disease 801
Ear-Cockle of Wheat 802
Soybean Cyst Nematode 804
Plant Diseases Caused by Flagellate Protozoa 805
Phloem Necrosis of Coffee 806
Hartrot of Coconut Palms 806
Selected References 807
Questions 809

27. Diseases Due to Angiospermic Parasites 810

Parasitic Flowering Plants 810
Dissemination of Angiosperm Parasites 811
Control of Angiosperm Parasites 813
Loranthus or “Banda” or Giant Mistletoe 813
Boomrape (Orobanche spp.) 814
Striga spp. (Witchweed) 815
Cuscuta 816
Selected References 820
Questions 821

28. Non–Infectious or Abiotic Diseases 822

Non–Infectious or Abiotic Diseases of Plants 822
Nutritional Deficiencies 823
xxiv Contents

Tip Burn of Paddy or Pansukh of Rice 826

Blossom End Rot of Tomato 826
Heat Canker of Linseed 827
Black Heart of Potato 827
Mango Necrosis or Black Tip 828
Khaira Disease of Rice (Zinc Deficiency Disease of Rice) 829
Zinc Deficiency Disease of Citrus 829
Selected References 830
Questions 831

29. Application of Biotechnology in Plant Pathology 832

The Use of Tissue Culture Techniques of Importance to Plant Pathology 832
Selection for Disease Resistance 838
Recombinat-DNA Technology 838
Use of Monoclonal Antibodies in Plant Pathology 839
Somaclonal Variants as a Source of Disease Resistance 841
Innovative Tools for Detection and Diagnosis of Plant Pathogenic Microbes 842
Selected References 846
Questions 848

30. Phytopathological Techniques in Plant Pathology 850

Isolation of Pathogens 850
Requirements for Isolation of Pathogens 850
Maintenance and Preservation of Microbial Cultures 857
Plant Disease Assessment Methods 861
Selected References 864
Questions 866

Index 867
 Introduction 1

Nature and Concept of Plant Diseases

It is imperative to define and discuss the meanings of certain terms commonly used in phytopathology
(phyton- plant; pathos- suffering; logos- knowledge) before we discuss the basic principles and concept
of plant pathology. It is essential to include these definitions and terms in an introductory chapter, since
an understanding of these terms is required before the reader can be made aware of the details of the
processes involved in plant diseases.
Phytopathology is the study of the diseases of plants and covers the entire field of biological and sci-
entific activity that is concerned with the understanding of this complex phenomenon. Phytopathology
is thus the study of the nature, development, and control of plant diseases.
Disease, being a complex phenomenon, is difficult to define in a few words. We have to keep in mind
what Locke said in Human Understanding: “Though definitions will serve to explain the names of
substances as they stand for our ideas, yet they leave them not without great imperfection as they stand
for things.” According to modern conception, disease is an interaction among the host, parasite, and
environment. A simple dictionary meaning of disease is: any departure from health, presenting marked
symptoms, malady, illness, disorder. Diseased plants are distinguished by changes in their morphologi-
cal structures or physiological processes, which are brought about by unfavourable environment, or by
parasitic agencies. Several definitions of diseases in plants have been proposed. Some examples are:
“a series of harmful physiological processes caused by continuous irritation of the plant by a primary
agent”; “a harmful deviation from the normal functioning of physiological processes”; “a continuous
impairment of metabolism”. Stakman and Harrar (1957) defined plant disease as “a physiological dis-
order or structural abnormality that is harmful to the plant or to any of its part or products that reduces
the economic value”.
Agrios (1997) has defined disease in plants as a series of invisible and visible responses of plant cells
and tissues to a pathogenic microorganism or environmental factor that results in adverse changes in the
form, function, or integrity of the plant and may lead to partial impairment or death of the plant or its
parts. However, A Guide to the Use of the Terms in Plant Pathology (1973) prepared by the Federation
of British Plant Pathologists recommends that the use of the term should be restricted to malfunc-
tions caused by plant pathogenic organisms or viruses and those caused by other factors (the so called
non-infectious diseases) should be termed as “disorders”. According to Horsfall and Cowling (1977a),
2 Plant Pathology

disease is a malfunctioning process that is caused by continuous irritation. Of course this process may
result in some suffering, and this produces symptoms. This conception of the disease is accepted by the
Committee of Terminology of the American Phytopathological Society and by their counterpart, Com-
mittee of the British Mycological Society.

Simple Interest Disease

This denotes a disease, the increase of which is mathematically analogous to simple interest in money.
There is only one generation of disease in the course of one epidemic.
A simple interest disease develops from a common source of inoculum, that is, the capital is constant,
and often there is one generation of infection in a season. In the increase of simple interest disease, the
loge [1/(1-x)] is plotted against time (where x = the proportion of diseased tissue), thus allowing for the
fact that with increasing infection the number of plants susceptible to infection decreases (Fig. 1.1).
Soil-borne diseases such as vascular wilts and soil-borne smuts, which infect seedlings and subsequent-
ly sporulate in the inflorescence of the mature plants, provided that the pathogen does not spread from
the smutted heads, are examples of simple interest diseases.

0.50

0.25
x

0.0

0.50
x
1
1

0.25
Loge

0.0
5 15 25 4 14 24 3
July Aug. Sept.

Figure 1.1 The progress of an epidemic of wilt of cotton caused by Fusarium oxysporum f. vasinfectum. In the upper
graph x is plotted against time; the lower part plots Loge [1(/1-x)] against time. (Redrawn from JE Van der
Plank. Plant Disease Epidemics and Control, Academic Press, New York and London, 1963, p.34.)

Compound Interest Disease

This denotes a disease, the increase of which is mathematically analogous to compound interest money.
There are several or many generations of the pathogen in the life of the crop, that is, the capital is in-
creased by the amount of interest. In plotting the amount of disease against time, the transformation
loge (x/1-x) is used, where x represents the proportion of infected susceptible tissue or of infected plants
if the pathogen develops in a systemic manner (Fig.1.2). Late blight of potato, stem rust of wheat, or
powdery mildew of wheat are examples of compound interest diseases.
If we know that a particular disease is of the simple interest type, suitable control measures can be ap-
plied. For example, sanitation affects the simple interest disease more than the compound interest type.
 Introduction 3

Figure 1.2 Progress of stem rust of wheat caused by Puccinia graminis tritici. (Redrawn from JE Van der Plank. Plant
Disease Epidemics and Control, Academic Press, New York and London, 1963, p.32.)

If the disease is of the former type, sanitation, including the eradication of the inoculum of the pathogen
from planting material and sites, should be insisted upon. If it is of the compound interest type, and the
epidemic spreads rapidly and lasts long, the infection rate should be reduced by a timely application
of fungicides. The stress should shift from sanitation to reducing the infection rate in proportion to the
increase in the rate of spread and duration of the epidemic.

Cause of Diseases
When a parasitic microorganism enters into the causal complex of a disease, it is commonly considered
to be the cause of the disease. However, we know that sometimes a variation in the environment also
makes a plant liable to infection. The microorganism is thus not the sole causal factor. After a microor-
ganism has successfully infected a susceptible plant, the subsequent interaction between the plant and
the microorganism (which results in a disease) is also subject to environmental influences. Thus, strictly
speaking, it is not correct to refer to a microorganism as the cause of a given disease since this implies
that it is the sole cause. The term ‘causal organism’ is proper since it implies that the organism is a part
of the causal complex. Most causal organisms are parasites. A parasite is not synonymous with the
pathogen. In some cases, organisms are parasitic without becoming causal factors in the disease. Many
parasitic establishments do not become pathogenic if the parasite compensates for the damage done by
its presence. The mycorrhizal fungus is certainly parasitic on the roots of trees but it is not pathogenic.
The root nodule bacterium (Rhizobium leguminosarum) is another example. In other cases, the byprod-
ucts of a strict saprophyte are an important part of the causal complex. Leaf tip injury to oats may be due
to the absorption of Griseofulvin from soil. Milo root rot is caused by a powerful toxin produced by the
fungus (Periconia circinata) growing in the root zone.
Thrower (1996) modified a definition offered by the British Mycological Society in 1950 and defined
a parasite as “an organism or virus existing in an intimate association with another living organism from
which it derives an essential part of the materials for its existence”. In a literal sense, a pathogen is an
4 Plant Pathology

agent that causes damage. According to A Guide to the Use of Terms in Plant Pathology (1973) prepared
by the Federation of British Plant Pathologists, a pathogen is an organism or virus capable of causing
disease in a particular host or range of hosts. Most, but not all pathogens, are also parasites in that they
derive the materials they need for existence from a living plant (the host or the suscept) as distinct from
saprophytes, which derive these materials from dead organic matter.

Pathotype
This is a subdivision of a species distinguished by the common characters of pathogenicity, particularly
in relation to the range of hosts.

Serotype
A serotype is the population of a pathogen (usually a bacterium or virus) in which all individuals possess
a given character of serology in common (Robinson, 1969). On the basis of serological tests, differ-
ences between apparently similar organisms or viruses may be found and the subdivisions thus formed
are called serotypes or strains. For example, only one serotype was designated amongst 45 isolates of
Xanthomonas oryzae pv. oryzae on the basis of agglutination reactions and gel diffusion tests (Addy
and Dhal, 1977).

Physiotype
It is a population of pathogens in which all individuals have a particular character of physiology (but not
pathogenicity) in common (Robinson, 1969).

Pathogenicity
It is the quality or characteristic of a pathogen capable of causing disease.

Pathogenesis
It is the sequence of progress in disease development from the initial contact between a pathogen and
its host to the completion of the syndrome. Horsfall and Dimond (1960) liken the relationship between
the microorganisms and higher plants to a situation in which your mother-in-law lives with you. If she
contributes to the harmonious running of the household, your relation is symbiotic; if you support her
and she does not do her share of work, she is a parasite and you are the host; if she stirs up trouble be-
tween you and your wife, she is a pathogen. Her actions, your reactions, and her reactions constitute
pathogenesis.

Host
An organism that harbours or supports the activities of a parasite is known as the host.
Whetzel (1929) termed the diseased plant suscept, but suscept may actually mean the plant that is
susceptible or prone to disease.
 Introduction 5

Inoculum
Inoculum is the infectious material that can cause disease, and it is that portion of individual pathogens
that is brought into contact with the host.
Inoculum Potential The term inoculum potential describes the degree of infectivity. Garrett (1956)
defined inoculum potential as “the energy of a fungal parasite available for infection of a host at the
surface of the host organ to be affected”.
Primary Inoculum The overwintering or oversummering stage may be in or on the seed of the host, in
the perennial wild host, in debris of infected plant of the previous season, in dormant spores, sclerotia or
other organs in the soil, or in the form of an inhabitant of the soil flora. Whatever the source, it causes
primary infection. It may serve as an infective agent and thus becomes the primary inoculum, or it may
multiply as a saprophyte (or as a parasite on the perennial host) to produce spores which become the pri-
mary inoculum. The rust fungus oversummers in India in the uredinial (uredial) stage on summer wheat
in the hills, from where the primary inoculum is carried by air currents to the next crop grown at the foot
of the hills and then to the plains. When the host becomes diseased, the organism commonly produces
another crop of spores or infective bodies which are known as the secondary inoculum and cause sec-
ondary infection. This last cycle may be repeated several times. The uredinial (uredial) pustules formed
in the wheat crop as a result of primary infection become a source of secondary infection to other plants
as the urediospores are blown down to these leaves and plants.
Penetration This is the first step in the contact of the inoculum with the host. Penetration may take
place in one of a number of ways characteristic of the organism and are sometimes influenced by the
environment. The methods of penetration have been discussed in Chapter 3. There is a clear distinction
between penetration and infection. Penetration refers to the initial invasion of the host by an organism.
We may have many cases of penetration without infection, for example, in the case of apple scab fungus
(Venturia inaequalis) conidia, the infection hyphae penetrate readily whether or not the host variety and
fungal strain are compatible; but in the resistant host variety, the fungus is not able to develop further
and cause infection due to the influence of the vital activities of the cell.
Infection It implies the establishment of the pathogen inside the host following penetration in which a
parasitic relationship between the two organisms is established.
Incubation Period There may be a long period between penetration, infection, and the appearance of
the disease, but ordinarily one follows the other. This interval of time between the penetration of the host
by the pathogen and the first appearance of the symptoms on the host is known as the incubation period.
In the case of black stem rust of wheat, the pathogen Puccinia graminis tritici enters the host (wheat) a
day or so after the urediniospores (urediospores) have germinated, but the actual symptoms of the dis-
ease in the form of rust pustules develops only after a week or so at 20°C. In the case of Sphacelotheca
sorghi (covered smut of Sorghum), the very young seedling is infected from soil-borne or seed-borne
spores, but no visible symptoms appear until the infected plant produce smutted flowers. In such cases,
the incubation period is technically several months, although increased respiration of the diseased plant
may occur much sooner. In case of the pathogen Mycosphaerella musicola, the incubation period is
about three or four weeks.
Disease Development It refers to a series of events which occur between the time of infection and
complete development of the disease.
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 "Tehän lähdette nyt, eikö niin, Brannon?" sanoi hän hymyillen.
"Sanokaa
Bettylle — —"

Samassa hän kiljahti, kun Brannon liikahti. Hän tunsi Brannonin

käden olkapäätään vasten ja hän horjahti siitä voimakkaasta
sysäyksestä, jonka Brannon hänelle antoi, syöksyi sivuttain ja kaatui
maahan selälleen noin kahdentoista jalan päähän Brannonista.

Kaatuessaan ei hän hetkeksikään ollut kadottanut Brannonia

näkyvistään. Jännittynein lihaksin oli hän koettanut lievittää
kaatumistaan ja oli tullut maahan hitaasti, joten Brannonin jokainen
liike jäi ainaiseksi hänen sieluunsa kuin valokuva.

Työntäessään Josephinen syrjään, hypähti Brannon muutaman

askeleen taaksepäin ja hänen oikea kätensä oli laskeutunut
vyötäisten kohdalle. Käsi kirposi siitä jälleen taaksepäin, nousi
ylöspäin ja syöksi tulta ja savua. Kuusipiippuisen kumahteluun
sekoittui rihlan kimakka, ilkeä-ääninen paukahtelu Denverin pyssystä
kiven takaa, savun puskiessa suorana viivana harjanteelta. Brannon
lähetti yhden kuulan sinnekin, käsitellen pistooliaan aivan kuin hän
olisi heittänyt jotakin harjanteelle päin. Ase laski taas ja asettui
vaakasuoraan ja sen suusta syöksyi yhtämittainen tuli Lattimeria
kohti, joka oli saanut aseensa esille ja koetti tähdätä Brannoniin.

Lattimerin ase laukesi, mutta hänen voimattomaksi ammuttu

kätensä ei enää jaksanut ohjata sitä ja se kirposi hänen
hervottomista sormistaan ja putosi kolahtamatta pehmeään santaan.

Brannonin liikkeet olivat olleet niin nopeat, että Josephine oli

varma siitä, että Denver ja Lattimer olivat kaatuneet yhtaikaa. Niin
hänen jännittynyt ja kauhistunut sielunsa ainakin tulkitsi asian.

Lattimerin horjahdellessa eteenpäin outo ihmetyksen ilme

lasimaisissa silmissään ja hitaasti lyyhistyen polvilleen, vieri Denver
hervottomana rinnettä alas. Hän liukui hiljaa aivan pehmeään
santaan asti ja jäi siihen makaamaan oudossa rykelmässä. Lattimer
taas näytti ikäänkuin valinneen itselleen määrätyn kuolinpaikan,
vaipui siihen hitaasti ja oikaisihe aivan kuin makuulle ainakin.

Syntyi aavemainen, syvä hiljaisuus. Brannon seisoi synkkänä

katsellen vuoroon kumpaakin kaatunutta vihollistaan.
Kumpainenkaan ei liikahtanut ja Josephine käsitti, että kaikki oli ohi.

Brannon ei sanonut sanaakaan eikä edes katsahtanutkaan

Josephineen.
Josephine nousi päästäkseen tappopaikalta pois ja meni
ruohokentän
toiseen päähän. Siinä lysähti hän maahan ja itki näkemiään kauhuja.
Mutta hän tunsi sisässään, että oikeus oli tapahtunut.

Hänen oli täytynyt pyörtyä, sillä kun hän epämääräisen ajan

perästä aukaisi silmänsä, näki hän Brannonin seisovan lähellään
olevalla harjanteella.

Oli tullut hämärä. Maa näytti harmaalta ja kuolleelta ja taivas oli

kalpean sininen ja siellä täällä tuikahti tähti. Josephine nousi.

Brannon kuuli hänen liikkeensä ja tuli hänen luokseen, katsoen

häntä suoraan silmiin.

"Tuolla tulee joku" sanoi hän. "Pohjoisesta. Jotenkin varmasti se

on Cole Meeder ja joku hänen miehistään. Mitä aiotte sanoa heille?
Aiotteko kertoa heille saman valheen kuin minullekin, että menitte
omasta ehdostanne Lattimerin kanssa?"

"Tiesittekö sen"? sanoi hän hämmästyneenä. "Tiesittekö, että

valehtelin?"

Brannon nauroi hiljaa.

"Denver ja Lattimer unohtivat yhden asian", sanoi hän. "Se oli se,
että minulla on silmät. Sieltä, missä olin vähän aikaisemmin kuin te
juoksitte vastaani, saatoin nähdä vuorenharjanteen. Näin Denverin
kiipeevän sinne ja pidin silmällä mihin hän kätkeytyi. Sitäpaitsi,
vaikka en olisi nähnytkään häntä, niin —" Hän pysähtyi, pani
molemmat kätensä Josephinen olkapäille ja loi häneen katseen,
jossa oli moitetta ja samalla ihailua.

"Te ette valehtele vakuuttavasti, Jo", sanoi hän, "te ette ole kylliksi
harjaantunut siihen. Mutta te teitte suurenmoisen työn —
suurenmoisen työn!"

Josephine itki hiljaa.

Vähän myöhemmin talutti Brannon hänet vuoren harjanteelle,

jossa he varjojen tummetessa vartoivat kun Cole Meeder ja Starin
miehet ratsastivat heidän luokseen.

Neljäsneljättä luku.

Kun Josephine myöhemmin eräänä päivänä istui Triangle L:n

kuistilla mukavassa kiikkutuolissa, katseli hän miettiväisenä avaralle
alangolle.

Rauhan palattua saattoi hän antaa ajatustensa viipyä menneissä

eräänlaisella tyytyväisyydellä. Sillä elleivät hänen kokemuksensa
ehkä olleet antaneet hänelle muuta, niin olivat ne kuitenkin
opettaneet hänelle sen seikan, ettei hän yhdellä iskulla pystynyt
muuttamaan luontoaan siinä määrässä, että se vastaansanomatta
sopeutuisi mihin uuteen ympäristöön tahansa.

Hän sulki silmänsä ja muistutteli mielessään tapahtumain kulkua

alusta alkaen, kun hän ensi kerran katsoi Brannonia silmiin rautatien
varrella, aina siihen hetkeen saakka kun hän kolme päivää sitten
taasen katsoi niihin heidän seisoessaan vuoren harjanteella
odotellen Starin miehiä. Hän näki, etteivät elämän ohjesäännöt,
sellaisina kuin hän oli ne oppinut tuntemaan, olleet niin joustavat,
että niitä olisi voinut ulottaa poikki mantereen tähän erämaahan. Hän
tunsi, että Brannon edusti Länttä samoinkuin hän itse oli Idän
edustaja. Hän oli kuullut ihmisten sanovan, että ihmisluonto on
kaikkialla samanlainen ja vaikkei hän pystynytkään väittämään sitä
vääräksi, oli hän vakuutettu siitä, ettei sitä ollut tarpeeksi perusteltu.
Siihen olisi hänen mielestään pitänyt lisätä jotakin, joka koski sitä
ympäristöä jossa kukin eli.

Brannon esimerkiksi, oli elänyt koko ikänsä Lännessä. Josephine

muisteli Bettyn sanoneen, että hän polveutui Booneista, joka taas
viittasi siihen, että missä aavat erämaat levisivät, siellä oli myös
Brannon kotonaan. Olisiko hän sama Brannon jos hänet aivan äkkiä
sijoitettaisiin keskelle sivistynyttä yhteiskuntaa?

Josephine arveli, ettei hän olisi sama. Mahdollisesti olisi hän yhtä
taipumaton ja jyrkkä, mutta Josephinen kotiseudun ankarien lakien
alaisena murtuisi hänen terästahtonsa ja hänen täytyisi hillitä
väkivaltaisia taipumuksiaan. Laki ei siellä olisi hänen kädessään,
vaan koko kansan käsissä, joka sitä vain valitsemainsa edustajain
kautta toteutti. Sen vuoksi hän ei olisi sama Brannon siellä.

Ja joka tapauksessa, olipa hän Lännen tai Idän mies, pelkäisi

Josephine häntä. Alussa taisteli hän Brannonia vastaan luullen
vihaavansa häntä. Nyt hän älysi, että hän oli taistellut ainoastaan sitä
runollista lumousta vastaan, joka häntä ympäröi. Sen jälkeen kun
hän oli nähnyt hänen tappavan Lattimerin ja Denverin, oli tämä
lumous hävinnyt. Josephine ei rakastanut häntä, hän ei edes pitänyt
hänestä, mutta hän ihaili hänen alkuperäistä sielullista voimaansa ja
hänen rautaista itsehillintäänsä. "Teräs" Brannon! Josephine
muistaisi hänet aina, se oli varma!

Hän näki nyt Brannonin tulevan ratsastaen häntä kohti

pohjoisesta, aavan tasangon poikki. Hän katseli häntä kun hän
rauhallisen kepeästi ratsasti mustallaan. Hän tunsi taas tuon
lumouksen. Mutta hän pudisti hymyillen päätään muistellessaan
erästä erikoista Idän miestä, joka olisi sangen karsaasti katsonut
nahkahousuja ja kannuksia.

Brannon tuli perille, pysäytti hevosensa kuistin nurkalla ja kääntyi

istumaan poikkipuolin satulassa. Hänen hymynsä oli leveä,
ystävällinen — ei mitään muuta.

"Tehän olette aikeissa jättää meidät", sanoi hän. "Kuulin sen juuri
eräältä miehistä. Tulin sanomaan hyvästiä teille."

"Kaksikymmentä mailia!" sanoi Josephine, viitaten matkaan, jonka

Brannon sitä varten oli ratsastanut. Hän oli kuullut Bettyn kertovan
siitä.
 "Niin", sanoi Brannon, katsoen häntä vakavasti, "ettehän tahtone
kieltää minulta sitä iloa?"

Josephine nauroi ystävällisesti moitteelle hänen äänessään.

"Pelkään, että tulette hyvinkin iloiseksi kun lähden", sanoi hän

tarkaten Brannonin ilmettä.

"Enhän toki" sanoi hän, silmät leimuten, "te olette ollut erittäin
mielenkiintoinen."

"Brannon", sanoi Josephine — sillä kun hän nyt oli lähdössä pois,
tahtoi hän olla varma siitä, ettei Brannonille jäänyt pahaa mieltä, sillä
hän muisti sen illan kun Brannon oli pannut kätensä hänen päänsä
päälle miltei hyväillen, "oletteko varma, että minä olen mielestänne
ollut ainoastaan mielenkiintoinen?"

"Luulen, että siinä on kaikki, Miss Hamilton", vastasi Brannon

katsoen häntä vakavasti silmiin. "Ymmärrättehän, että kun te
vihaatte —"

"Vihasin", oikaisi Josephine.

"Kun te vihasitte minua", korjasi Brannon hymyillen, "niin ei minulla

ollut rohkeutta —"

"Brannon", keskeytti Josephine, "eikö se ole Betty?"

Brannonin silmät säihkyivät. Hän punastui, mutta hänen äänensä

oli luja.

"Niin luulen", sanoi hän. Ja silloin näki Josephine ensikerran

Brannonin luonteen ystävällisen, inhimillisen puolen. Puna levisi
koko hänen kasvoilleen, hänen silmänsä elivät ja niissä oli
poikamaisen hämmästynyt ilme.

"Jos hän huolii minusta", lisäsi hän sitten. Josephinen silmät

kertoivat suloista tietoa ja hän hymyili tyytyväisenä.

"Miksi ette kysy häneltä, Brannon?" sanoi Josephine ystävällisesti.

"Aion tehdä sen", sanoi Brannon vakavasti. "Olen ajatellut sitä jo

kaksi vuotta, mutta näettehän, ettei Betty juuri kehoita siihen."

"Betty ei juokse miesten perässä, Brannon."

"Se on totta!" sanoi Brannon, ihaillen Bettyn aitoutta naisena.

"Betty on tyttöjen tyttö, eikö niin, Miss Hamilton?"

"Jos minä olisin mies, niin pitäisin häntä suuressa määrässä

tavoiteltavan arvoisena, Brannon", sanoi Josephine hymyillen. "Ja",
jatkoi hän kuivasti, "minä olisin hyvin levoton siitä, että joku toinen
mies ennättäisi pyytää häntä ennen minua. Teillä on toivoa,
Brannon."

Brannon oli kiitollinen ja poikamaisen iloinen. Hän loi Josephineen

sellaisen silmäyksen, että se sai hänen sydämen lyömään
kiivaammin ja antoi hänen aavistaa, että hän olisi ehkä ollut hyvinkin
surullinen täytyessään jättää Brannonille hyvästit, jos kohtalo olisi
toisin määrännyt.

Mutta mielenliikutus meni pian ohi. Seuraavassa tuokiossa pudisti

hän
Brannonin kättä, sillä hän oli astunut satulasta sanomaan
jäähyväiset.
 "Te kai lähdette viiden aikaan?" sanoi Brannon.

"Lin Murray valjastaa paraikaa hevosia", vastasi Josephine

"Lähden kymmenen minuutin kuluessa."

"Betty saattaa teitä Willetiin saakka?"

"Niin."

"Hyvä", sanoi Brannon vakavasti. "Sitten sanon jäähyväiset vasta

kaupungissa. Lin Murrayhin ei aina ole luottamista."

Hän jätti Josephinen kuistille ja meni karja-aitaukselle, missä

Murray valjasti hevosia.

Kymmenen minuutin kuluttua vierivät ajoneuvot kuistin eteen.

Brannon istui yksin ajajan paikalla. Hänen kasvonsa olivat
ilmeettömät.

Josephine oli mennyt sisään ottamaan tavaroitaan ja kun Betty tuli

ulos, oli Josephine aivan hänen kintereillään.

Betty oli jo puolivälissä rappuja, ennenkuin hän huomasi

Brannonin, joka istui ajajan istuimella välinpitämättömän näköisenä.

Betty pysähtyi ja punastui.

"Brannon", sanoi hän, "Missä Murray on? Käskin häntä tulemaan

mukaan
Willetiin!"

"Murray on sairas", vastasi Brannon rauhallisesti, katsomatta

Bettyyn. "Eikö ollutkin onni, että satuin tulemaan kotiin äsken?" lisäsi
hän katsoen Bettyyn, jonka posket tulivat yhä punaisemmiksi. "Sinun
olisi täytynyt ajaa kotiin yksin."

Betty olisi yhtä hyvin voinut olla yksinkin paluumatkansa

alkutaipaleen, sillä niin vähän tiesi hän Brannonin läsnäolosta.
Brannonin keskustelu rajoittui nimittäin vain yksitavuisiin
maanitteluihin, hänen ohjatessaan ravakasti juoksevia hevosiaan
pimeän hämyssä.

Mutta Betty ei puhunut enempää itsekään. Heillä oli kummallakin

omat huolensa, jotka painoivat heidän sydämiään. He olivat
pahoillaan siitä, että nuori Idän nainen, jolla oli ollut aatteita, oli tullut
Suureen länteen, ja nähnyt siellä aatteittensa murskautuvan
kokonaan, tullessaan kosketuksiin sen ihmisluokan kanssa, jonka
tapaa kaikissa yhteiskunnissa, nimittäin lainrikkojien.

"Hänen ei olisi pitänyt sekaantua asioihin sillä tapaa", sanoi

Brannon äkkiä, pukien ajatuksiaan sanoiksi.

"Niin", sanoi Betty, ikäänkuin olisi jatkanut keskenjäänyttä

keskustelua. "Siinä hän teki erehdyksen."

"Toisen erehdyksen teki hän vihatessaan minua", sanoi Brannon.

"Minä koetin —"

"Ei hän sitä tehnyt, Brannon."

"Mitä ei tehnyt?"

"Ei hän vihannut sinua. Eikä koskaan tule sitä tekemäänkään. Kun
sinä puhuit hänen kanssaan kuistilla juuri ennen lähtöä, niin hän
sanoi —"
 "Kuuntelitko sinä?" syytti Brannon. Hän taivutti itseään Bettyn
puoleen nähdäkseen hänen kasvojaan hämärässä ja huomasi, että
ne olivat tulipunaiset.

"Minä — minä en voinut sille mitään, Brannon. Minä olin näet juuri
tulossa ulos ja kun minun piti — piti —"

Hän pysähtyi, uteliaana tietämään miksi Brannon huokaisi niin

syvään.

"Miksi sinä noin teit, Brannon?" kysyi hän.

"Helpotuksesta", vastasi Brannon. "Olen kaksi vuotta koettanut

sanoa sinulle sen, minkä kerroin hänelle. Luulen, ettei minulla
koskaan olisi ollut rohkeutta siihen.

"Mutta sinun täytyy, Brannon", kuiskasi Betty. "Minä en koskaan

tyytyisi siihen tietoon, että sinä olet sanonut sen jollekin toiselle
naiselle, vaikka minä olisin sen itse kuullutkin —"

Ohjaajan varman käden puutteessa pysähtyivät hevoset vihdoin

tien vieressä kasvavan tuoreen heinän ääreen. Sillä aikaa kertoi
mies, jonka rohkeus oli hankkinut hänelle nimen "Teräs" tulevalle
elämänkumppanilleen tuon ikivanhan, tarinan, joka on aina yhtä
uusi, aina sama, tänään ja huomenna ja joka ei tiedä mitään
pohjoisesta eikä etelästä, ei idästä eikä lännestä.
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