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Pesticide Resistance STRATEGIES AND TACTICS POR MANAGEMENT Committee on Strategies for the Management of Pesticide Resistant Pest Populations Board on Agriculture National Research Council NATIONAL ACADEMY PRESS Washington, D.C. 1986

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NATIONAL ACADEMY PRESS 2101 CONSTITUTION AVENUE, NW WASHINGTON, DC 20418 NOTICE: The project that is the subject of this report was approved by the Governing Board of the National Research Council, whose members are drawn from the councils of the National Academy of Sciences, the National Academy of Engineering, and the Institute of Medicine. The members of the committee responsible for the report were chosen for their special competences and with regard for appropriate balance. This report has been reviewed by a group other than the authors according to procedures approved by a Report Review Committee consisting of members of the National Academy of Sciences, the National Academy of Engineering, and the Institute of Medicine. The National Research Council was established by the National Academy of Sciences in 1916 to associate the broad community of science and technology with the Academy's purposes of furthering knowledge and of advising the federal government. The Council operates in accordance with general policies determined by the Academy under the authority of its congressional charter of 1863, which establishes the Academy as a private, nonprofit, self-governing membership corporation. The Council has become the principal operating agency of both the National Academy of Sciences and the National Academy of Engineering in the conduct of their services to the government, the public, and the scientific and engineering communities. It is administered jointly by both Academies and the Institute of Medicine. The National Academy of Engineering and the Institute of Medicine were established in 1964 and 1970, respectively, under the charter of the National Academy of Sciences. This project was supported under agreements between the following agencies and the National Academy of Sciences: Grant No. DAN-1406-G-SS-3076-00 from the U.S. Agency for International Development; Grants No. 59-32R6-2-132 and 59-3159-4-33 from the U.S. Department of Agricul- ture; and Contract No. CR-810761-01 from the U.S. Environmental Protection Agency. Support from the following corporate sponsors is also gratefully acknowledged: American Cyanamid Com- pany; Ciba-Geigy Cotporation; E. I. du Pont de Nemours & Company; FMC Corporation; ICI Americas, Inc.; Mobay Chemical Corporation; Monsanto Agricultural Products Company; NOR- AM Chemical Company; Rohm and Haas Company; Sandoz, Inc.; and Union Carbide Agricultural Products Company, Inc. Library of Congress Cataloging-in-Publication Data Main entry under title: Pesticide resistance. Contains papers from a symposium held in Washington, Nov. 27-29, 1984. Includes index. 1. Pesticide resistance Congresses. I. National Research Council (U.S.). Committee on Strategies for the Management of Pesticide Resistant Pest Populations. SB957.M36 1985 363.7'8 85-25919 ISBN 0-309-03627-5 Printed in the United States of America

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COMMITTEE ON STRATEGIES FOR THE MANAGEMENT OF PESTICIDE RESISTANT PEST POPULATIONS EDWARD H. GLASS (Chairman), New York State Agricultural Experiment Station, Cornell University PERRY L. ADKISSON, Texas A&M University GERALD A. CARLSON, North Carolina State University BRIAN A. CROFT, Oregon State University DONALD E. DAVIS, Auburn University JOSEPH W. ECKERT, University of California GEORGE P. GEORGHIOU, University of California, Riverside WILLIAM B. JACKSON, Bowling Green State University HOMER M. LeBARON, Ciba-Geigy Corporation BRUCE R. LEVIN, University of Massachusetts FREDERICK W. PLAPP, JR., Texas A&M University RICHARD T. ROUSH, Mississippi State University HUGH D. SISLER, University of Maryland Staff ELINOR C. CRUZE, Project Officer GERALDINE WILLIAMS, Secretary HALCYON YORKS, Secretary VANESSA LEWIS, Secretary ~

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BOARD ON AGRICULTURE WILLIAM L. BROWN (Chairman), Pioneer Hi-Bred International, Inc. JOHN A. PING (Vice Chairman), Inter-American Development Bank PERRY L. ADKISSON, Texas A&M University C. EUGENE ALLEN, University of Minnesota LAWRENCE BOGORAD, Harvard University ERIC L. ELLWOOD, North Carolina State University JOSEPH P. FONTENOT, Virginia Polytechnic Institute and State University RALPH W. F. HARDY, Cornell University and BioTechnica International, Inc. ROGER L. MITCHELL, University of Missouri CHARLES C. MUSCOPLAT, Molecular Genetics, Inc. ELDOR A. PAUL, University of California, Berkeley VERNON W. RUTTAN, University of Minnesota JAMES G. TEER, Welder Wildlife Foundation JAN VAN SCHILFGAARDE, U.S. Department of Agriculture/Agricultural Research Service VIRGINIA WALBOT, Stanford University CHARLES M. BENBROOK, Executive Director IV

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Contents PREFACE . EXECUTIVE SUMMARY . . . 1X ~ 1. INTRODUCTION ................... THE MAGNITUDE OF THE RESISTANCE PROBLEM George P. Georghiou ............... 2. GENETIC, BIOCHEMICAL, AND PHYSIOLOGICAL MECHANISMS OF RESISTANCE TO PESTICIDES MODES AND GENETICS OF HERBICIDE RESISTANCE IN PLANTS Jonathan Gressel ................... GENETICS AND BIOCHEMISTRY OF INSECTICIDE RESISTANCE IN ARTHROPODS: PROSPECTS FOR THE FUTURE Frederick W. Plapp, Jr. ................... RESISTANCE TO 4-HYDROXYCOUMARIN ANTICOAGULANTS IN RODENTS Alan D. MacNicoll ...................... PLANT PATHOGENS S. G. Georgopoulos ................... CHEMICAL STRATEGIES FOR RESISTANCE MANAGEMENT Bruce D. Hammock and David M. Soderlund ..... BIOTECHNOLOGY IN PESTICIDE RESISTANCE DEVELOPMENT Ralph W. F. Hardy .................... v 11 14 45 54 74 87 100 .111 .130

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Vl 3. POPULATION BIOLOGY OF PESTICIDE RESISTANCE: BRIDGING THE GAP BETWEEN THEORY AND PRACTICAL APPLICATIONS ................ CONTENTS .143 FACTORS INFLUENCING THE EVOEUTION OF RESISTANCE George P. Georghiou and Charles E. Taylor 157 POPULATION DYNAMICS AND THE RATE OF EVOLUTION OF PESTICIDE RESISTANCE Robert M. May and Andrew P. Dobson COMPUTER SIMULATION AS A TOOK FOR PESTICIDE RESISTANCE MANAGEMENT Bruce E. Tabashnik PEEIOTROPY AND THE EVOEUTION OF GENETIC SYSTEMS CONFERRING RESISTANCE TO PESTICIDES Marcy K. Uyenoyama QUANTITATIVE GENETIC MODELS AND THE EVOEUTION OF PESTICIDE RESISTANCE Sara Via MANAGING RESISTANCE TO RODENTICIDES ]. H. Greaves RESPONSE OF PLANT PATHOGENS TO FUNGICIDES M. S. Wolfe and ]. A. Barrett ........ . EXPERIMENTAL POPULATION GENETICS AND ECOLOGICAL STUDIES OF PESTICIDE RESISTANCE IN INSECTS AND MITES Richard T. Roush and Brian A. Croft ..... 4. DETECTION, MONITORING, AND RISK ASSESSMENT PREDICTION OR RESISTANCE RISK ASSESSMENT Johannes Keiding .............. DETECTION AND MONITORING OF RESISTANT FORMS: AN OVERVIEW K. ]. Brent . . 5. TACTICS FOR PREVENTION AND MANAGEMENT RESISTANCE IN WEEDS Fred W. Slife ..................... .170 .194 .207 .222 .236 .245 .257 .271 .279 .298 .313 .327 PREVENTING OR MANAGING RESISTANCE IN ARTHROPODS John R. Leeper, Richard T. Roush, and Harold T. Reynolds 335

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CONTENTS PREVENTING AND MANAGING FUNGICIDE RESISTANCE Johan DeEker ........................ CASE HISTORIES OF ANTICOAGULANT RESISTANCE William B. Jackson and A. Daniel Ashton .... . . 6. IMPLEMENTING MANAGEMENT OF RESISTANCE TO PESTICIDES . . . ~ . . . . . . . . . . . . . . . . . . . . . . . ACTIONS AND PROPOSED POLICIES FOR RESISTANCE MANAGEMENT BY AGRICULTURAL CHEMICAL MANUFACTURERS Charles ]. Delp ............... PESTICIDE RESISTANCE MANAGEMENT: AN EX- REGUEATOR S VIEW Edwin L. Johnson .................... THE ROLE OF REGULATORY AGENCIES IN DEALING WITH PESTICIDE RESISTANCE Lyndon S. Hawkins ................ THE ROLE OF COOPERATIVE EXTENSION AND AGRICULTURAL CONSULTANTS IN PESTICIDE RESISTANCE MANAGEMENT Raymond E. Frisbie, Patrick WeddIe, and Timothy I. Dennehy .................... INTEGRATION OF POLICY FOR RESISTANCE MANAGEMENT . . V11 .347 .355 .371 .388 . . .393 .403 410 Michael I. Dover and Brian A. Croft . . . . . . . . . . . . 422 ECONOMIC ISSUES IN PUBLIC AND PRIVATE APPROACHES TO PRESERVING PEST SUSCEPTIBILITY John A. Miranowski and Gerald A. Carison .... GLOSSARY INDEX .. 436 .449 .453

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Preface THE BRIGHT FUTURE projected for crop protection and public health as a result of the introduction of synthetic organic pesticides is now open to serious question because of an alarming increase in the number of instances of resistance in insects, plant pathogens, and vertebrates, and to a lesser extent in weeds. There are no longer available any effective pesticides against some major crop pests, such as the Colorado potato beetle on Long Island and the diamondback moth on cruciferous crops in much of the tropical world. Likewise, the malaria eradication programs of many countries are in disarray, in large part because vector mosquitoes are no longer adequately controlled with available insecticides. The incidence of malaria is resurging at an alarming rate. Because of the costs of bringing new pesticides to market, there are fewer new pesticides, and those produced are targeted only for major crops and pests. Resistance to pesticides, which first involved only insecticides, now exists for fungicides, bactericides, rodenticides, nemati- cides, and herbicides. Concern for the resistance problem has been expressed by the pesticide industry, farmers, crop protection scientists and practitioners, and govern- ment agencies. During the past 25 years there have been several symposia on the subject, and considerable research has been conducted on the genetic, biochemical, and physiological bases for resistance. As a result, much has been learned about the phenomenon; however, few methods have been de- veloped to date for preventing or delaying the onset of resistance to pesticides, other than eliminating or minimizing their use. In the past, problems have been overcome by the substitution of new pesticides. This procedure is IX

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x PREFACE threatened, because the rate of introduction of new pesticides has slowed dramatically during the last few years. New technologies and information have been developed in recent years that appear to have promise for application in finding ways to avoid or at least delay development of resistance. Thus, a new study was initiated, under the aegis of the Board on Agriculture. The evolutionary process by which organisms develop strains resistant to chemicals is universal throughout the extensive range of organisms in which the problem now exists. It was decided, therefore, to enlist the assistance of basic scientists in evolution, population genetics, modeling, and biochem- istry. It was also decided to make the study inclusive across pest classes and involve international experts from academia, government, and industry. In- asmuch as the application of solutions will have to take place in the field or wherever pests are found, we also enlisted crop protection practitioners. Finally, because resistance management systems may involve economics, regulations, and policy, representatives from these fields were recruited. The objectives of this study were to (1) identify promising strategies to avoid or delay the development of pesticide-resistant strains of pest species, as well as manage established resistant pest populations; (2) establish research priorities to develop these strategies and new approaches not currently in use; (3) stimulate pertinent research, not only in those disciplines concerned with resistance of pests affecting plants and animals, but in related fields as well; and (4) analyze the impact of changes in policy that will be needed to implement these strategies. To.accomplish these objectives, the committee organized a conference held in Washington, D.C., November 27-29, 1984. The conference consisted of a two-day symposium at which invited papers were presented, followed by a one-day workshop attended by the committee, symposium speakers, and additional scientists who were asked to participate. The conference was designed to produce this volume, which integrates a report prepared by the Committee on Strategies for the Management of Pesticide Resistant Pest Populations and the symposium papers themselves. The report is based on the committee's deliberations, the symposium papers, and the workshop discussions, while the papers represent the ideas of the individual authors. A group of papers follows each relevant section of the report. A glossary is included to communicate as broadly as possible among the disciplines and backgrounds of the many interests concerned with man- agement of resistance to pesticides. We hope this book will prove useful to many people, especially those involved in pest control, whether in industry, academia, government, applied pest management, or decision making. We are grateful to our many scientific colleagues who have given gen- erously of their knowledge and time to this study. Special thanks and ap

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an preciadon are extended to Drs. Raymond E. Fhsbie' Timothy Dennehy, and A. Daniel Ashton far their contributions. We also recognize and ~precia~ the fine support of Dr. Elinor C. Craze' stag officer far this study, and over Bag of the Bond on Agrkultum. Edward H. Cash CA~m~" Committee on S~a~gies far He Management of Pesticide Resistant Fast Populations

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