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PRINCIPLES OF PLANT AND ANlmC EjQFONTROL Control of â -~ * Plant-Parasitic Nematodes SUBCOMMITTEE ON NEMATODES COMMITTEE ON PLANT AND ANIMAL PESTS AGRICULTURAL BOARD NATIONAL RESEARCH COUNCIL Publication 1696 NATIONAL ACADEMY OF SCIENCES WASHINGTON, D.C. 1968
This report is one of a series on principles of controlling pests and diseases of plants and animals. The following volumes are in the series: Volume 1 Plant-Disease Development and Control Volume 2 Weed Control Volume 3 Insect-Pest Management and Control Volume 4 Control of Plant-Parasitic Nematodes Volume 5 The Vertebrates That Are Pests: Problems and Control Volume 6 Effects of Pesticides on Fruit and Vegetable Physiology The reports were prepared by six subcommittees working under the direction of the Com- mittee on Plant and Animal Pests. The following organizations sponsored this work: Agricultural Research Service, U.S. Department of Agriculture Agency for International Development, U.S. Department of State National Agricultural Chemicals Association Rockefeller Foundation Bureau of Sport Fisheries and Wildlife, U.S. Department of the Interior Available from PRINTING AND PUBLISHING OFFICE NATIONAL ACADEMY OF SCIENCES 2101 CONSTITUTION AVENUE WASHINGTON, D.C. 20418 Library of Congress Catalog Card Number 68-60085
Foreword The objective of the project on Plant and Animal Pest Control was to outline, for each of the several classes of pests, the principles of control where these are established; to call attention to effective procedures where true principles are not yet established; and to indicate areas of research that appear to war- rant early attention. The reports are not intended to be textbooks in the usual sense, nor encyclopedias, but are intended to deal with basic problems, the principles involved in controlling pests, and the criteria that should be considered in conducting research and in evaluating published information. Specific instances of control practices are cited only to illustrate principles and procedures. It is hoped that these reports will be useful to researchers at all levels, to pest-control agencies, to administrators seeking guidance on priorities for application of resources, and to general field workers in the United States and elsewhere. The National Academy of Sciences selected a committee of outstanding scientists to represent the diverse aspects of the problem and assigned to them responsibility for carrying out the project. To assist that committee, six subcommittees of specialists were appointed. Appropriate members of the parent committee were assigned as liaison members of the subcommittees, and in due time all reports were reviewed by the parent committee. Some seventy scientists have collaborated over a four-year period to pro- duce this series. Many others have contributed, to a lesser degree, in preparing statements and in reviewing and commenting on drafts of individual sections. Final responsibility for the content of these volumes rests with the parent committee. The Agricultural Board, under whose direction the Committee on Plant and Animal Pests operated, has reviewed and approved each manuscript.
Committee Members CHARLES E. PALM, Cornell University, Chairman WALTER w. DYKSTRA, Fish and Wildlife Service, U.S. Department of the Interior GEORGE R. FERGUSON, Geigy Agricultural Chemicals ROY HANSBERRY, Shell Development Company WAYLAND J. HAYES, JR., Communicable Disease Center, U.S. Department of Health, Education, and Welfare LLOYD W. HAZLETON, Hazleton Laboratories, Inc. JAMES G. HORSFALL, Connecticut Agricultural Experiment Station E. F. KNIPLING, Agricultural Research Service, U.S. Department of Agriculture LYSLE D. LEACH, University of California, Davis ROY L. LOVVORN, North Carolina State University GUSTAVA.SWANSON, Colorado State University
Preface This volume considers the principles involved in control of plant-parasitic nematodes and points to research avenues that might lead to improved methods in the future. The biology of the organisms themselves and the en- vironmental stresses to which they are subjected are included because this information is crucial to progress toward their control. We hope the material will prove useful to scientists trained in areas other than nematology who find themselves confronted with nematode-control problems as well as to students newly moving into this area of study and research. Specific nematocides are cited only in support of general statements relative to control principles, and trade names are used only when there is no widely recognized common name or when specific reference is made to a given commercial product. The first time such a common or trade name appears in a chapter, the principal active ingredient is indicated. The abbreviations DBCP, EDB, and 1,3-D are used to designate the compounds 1,2-dibromo-3- chloropropane, ethylene dibromide, and 1,3-dichloropropene, respectively. In preparing this material, sections were developed by individual Sub- committee members. The whole was then reviewed and integrated by the group collectively. Selected references are provided in literature citations following each chapter. The views of fellow workers were solicited and in- corporated in the discussions, and we are greatly indebted to S. D. Van Gundy,
A. M. French, R. Mankau, and J. F. Spears, who served as consultants in this endeavor. SUBCOMMITTEE ON NEMATODES W. F. MAI, Cornell University, Chairman E. J. CAIRNS, Auburn University L. R.KRUSBERG, University of Maryland B. F. LOWNSBERY, University of California, Davis C. W. McBETH, Shell Development Company D. J. RASKI, University of California, Davis J. N. SASSER, University of North Carolina IVAN J. THOMASON, University of California, Riverside
Contents PART I INTRODUCTION Chapter 1 THE SCIENCE OF NEMATOLOGY 3 Chapter 2 NEMATODES AND THEIR IMPORTANCE TO MAN 6 PLANT-PARASITIC NEMATODES IMPORTANT TO MAN 6 PRINCIPAL CHARACTERISTICS OF NEMATODES 8 Life Cycle/11 Soil Inhabitants / 11 Ectoparasitic Species; Endoparasitic Species Aboveground Parasites / 12 Bud and Leaf Nematodes; Seedgall Nematodes; Stem Nematode Significance of Life Habits /13 Geographic Distribution / 13 BIBLIOGRAPHY 14 PART II FACTORS INFLUENCING NEMATODE CONTROL Chapter 3 ECOLOGICAL RELATIONSHIPS VERTICAL DISTRIBUTION OF NEMATODES NEMATODE SURVIVAL 17 17 18 vii
NEMATODE POPULATIONS 18 THE SOIL ENVIRONMENT 20 Temperature / 21 Moisture / 21 Soil Texture and Structure / 22 Soil Solution / 23 CLIMATE 23 THE PLANT ENVIRONMENT 24 The Rhizosphere / 24 BIBLIOGRAPHY 26 Chapter 4 THE PHYSIOLOGY OF NEMATODES IN RELATION TO CONTROL 27 CHEMICAL COMPOSITION 27 METABOLISM 28 Respiration / 28 Temperature / 29 Moisture / 30 DORMANCY AND LONGEVITY 31 HATCHING, MOLTING, GROWTH, AND SEX DETERMINATION 32 NUTRITION AND LABORATORY CULTURING 33 CONCLUSIONS 34 BIBLIOGRAPHY 34 Chapter 5 PATHOGENIC RELATIONSHIPS 36 NEMATODE DISEASE SYMPTOMS AND DISEASE DIAGNOSIS 36 Top Symptoms and Signs; Root Symptoms and Signs Experimental Determination of Pathogenicity / 38 HISTOPATHOLOGY OF NEMATODE-PARASITIZED PLANTS 45 Gall Formation; Lesion Formation; Inhibition of Apical Growth Physiology of Diseased Tissues / 58 CONCLUSIONS 61 BIBLIOGRAPHY 61 viii
Chapter 6 NEMATODE INTERRELATIONSHIPS WITH OTHER PLANT-DISEASE ORGANISMS 63 FUNGI AND BACTERIA 63 NEMATODES 65 VIRUSES 65 PRACTICAL APPLICATIONS 66 FUTURE RESEARCH NEEDS 67 BIBLIOGRAPHY 68 Chapter 7 CONSIDERATIONS BASIC TO NEMATODE CONTROL 69 NEMATODES-PRIMARILY A SOIL PROBLEM 69 NEMATODE INFESTATIONS IN THE FIELD 70 HAZARDS OF MONOCULTURE 70 CONTROL METHODS AND LOCAL SITUATIONS 70 IDENTIFICATION OF NEMATODES AND DEGREES OF INFESTATIONS 72 COLLECTION OF SOIL AND PLANT SAMPLES 73 EXTRACTION OF NEMATODES FROM SOIL AND PLANTS 73 Mechanical Isolation / 73 Separation by Specific Gravity, Size, and Shape / 73 Wet Seiving of Soil; Elutriation Separation by Nematode Movement / 77 POPULATION LEVELS AND PREDICTION OF PLANT DAMAGE 82 BIBLIOGRAPHY 82 PART III BASIC PRINCIPLES OF CONTROL Chapter 8 PREVENTION OF SPREAD 85 MEANS OF DISSEMINATION 85 Soil and Plant Tissue / 85 Machinery, Reusable Containers, and Fertilizer / 86 Animals / 87 Water / 87 Wind / 88 NATURAL BARRIERS 88 ix
PRACTICES TO RESTRICT SPREAD Sanitation / 89 Nematocidal Treatments / 89 Certified Plant Materials / 90 Quarantines and Regulations / 90 BIBLIOGRAPHY 89 93 Chapter 9 REDUCTION OF NEMATODE POPULATIONS THROUGH LAND-MANAGEMENT AND CULTURAL PRACTICES 94 FALLOW 95 FLOODING 95 COVER CROPS 96 CROP ROTATION 97 TIME OF PLANTING 97 ORGANIC MANURING 101 REMOVAL OR DESTRUCTION OF INFECTED PLANTS 101 TRAP AND ANTAGONISTIC CROPS 102 NUTRITION AND GENERAL CARE OF HOST 103 SANITATION AND USE OF NEMATODE-FREE PLANTING STOCK 104 FUTURE RESEARCH NEEDS 104 Chapter 10 BIOLOGICAL CONTROL OF NEMATODES 106 USE OF BIOTIC AGENTS TO CONTROL NEMATODES 106 Fungi / 106 Bacteria and Viruses / 108 Protozoans/ 109 Other Nematodes / 109 Other Invertebrates / 110 INFLUENCE OF ORGANIC MANURING ON NEMATODE CONTROL 111 PLANT-ROOT EXUDATES TOXIC TO NEMATODES 111 BIBLIOGRAPHY 112 Chapter 11 PLANT RESISTANCE DEVELOPMENT OF NEW VARIETIES NATURE OF RESISTANCE 113 115 117
INFLUENCE OF ENVIRONMENT 119 GENETIC BASIS FOR RESISTANCE 120 RESISTANCE TO COMPLEX DISEASES 120 RESISTANCE-BREAKING BIOTYPES 120 FUTURE RESEARCH 121 BIBLIOGRAPHY 121 Chapter 12 CONTROL BY PHYSICAL FACTORS 123 CONTROL BY HEAT-BASIC CONSIDERATIONS 123 DISINFESTATION BY HEAT 124 Dry Heat/125 Moist Heat/ 126 Hot Water/126 Steam/126 Steam plus Chemical / 127 Endogenic Heat / 127 DISINFECTION BY HEAT 128 CONTROL BY LOW TEMPERATURE 129 CONTROL BY ELECTRICITY 130 CONTROL BY IRRADIATION 130 CONTROL BY MISCELLANEOUS PHYSICAL FACTORS 131 Chapter 13 CONTROL BY CHEMICALS 132 CHARACTERISTICS OF NEMATOCIDES 132 Types/132 Phytotoxicity / 133 Residues in Plants and Soil / 134 FACTORS INFLUENCING NEMATODE KILL IN SOIL 134 Soil Type / 135 Soil Condition / 135 Soil Moisture and Temperature / 136 Application Depth / 136 Application Rate / 137 Sealing Soil Surface After Application / 138 APPLICATION METHODS 138 Formulations/ 138 Applicators/ 141 Injectors; Plow Application xi
TYPES OF TREATMENT 144 Pieplant Treatment / 144 Treatment at Planting Time / 147 Postplant Treatment / 147 Bare-Root Dip / 149 Seed Treatment / 149 BASIC STUDIES ON NEMATOCIDES 150 Movement to Site of Action in Soil /ISO Penetration into the Nematode / 151 Mode of Action of the Nematocide on the Nematode / 151 Halogenated Hydrocarbons; Organophosphates; Carbamates Selectivity of Nematocides / 153 Development of Resistance to Nematocides / 154 FUTURE RESEARCH NEEDS 155 BIBLIOGRAPHY 156 Chapter 14 EVALUATION AND SELECTION OF CONTROL MEASURES 157 INTEGRATION OF CONTROL MEASURES 157 EFFECTIVENESS AND ECONOMICS 158 LEVEL OF AGRICULTURAL DEVELOPMENT 161 Recognition of a Problem / 161 Growth of Soil Fumigation in the United States / 162 SPECIFIC METHODS OF EVALUATING CONTROL MEASURES 162 Nematode Control / 162 Economic Evaluation / 163 PART IV RESEARCH NEEDS Chapter 15 FUTURE RESEARCH NEEDS 167 Xll