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CRC REVIVALS CRC REVIVALS

Edited by Jack Altman

Pesticide Interactions in Crop Production
Pesticide Interactions in Crop
Production
Beneficial and Deleterious Effects

Edited by
Jack Altman

ISBN 978-1-315-89635-9

,!7IB3B5-ijgdfj!
www.crcpress.com
 PESTICIDE
INTERACTIONS
zn
•

CROP
PRODUCTION
Beneficial and Deleterious Effects

Edited by
Jack Altman, Ph.D.
Plant Pathology and Weed Science
Colorado State University
Fort Collins, Colorado

Boca Raton London New York

CRC Press
CRC Press is an imprint of the
Boca Raton AnnGroup,
Taylor & Francis Arbor London
an informa business Tokyo
First published 1993 by CRC Press
Taylor & Francis Group
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Boca Raton, FL 33487-2742

Reissued 2018 by CRC Press

© 1993 by CRC Press, Inc.

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Library of Congress Cataloging-in-Publication Data

Pesticide interactions in crop production : beneficial and deleterious

effects / Jack Altman.
  p. cm.
Includes bibliographical references and index.
ISBN 0-8493-6339-X
1. Pesticides. 2. Plants. Effect of pesticides on. 3. Crops-
-Physiology. 4. Pesticides—Environmental aspects. I. Altman,
Jack, 1924-
SB951.P432 1993
632’ .95—dc20 92-26913

A Library of Congress record exists under LC control number: 92026913

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 Dedication

This book is dedicated to my wife, Roslyn

 PREFACE
Modem pesticides are designed to act against crop pests while exerting minimum impact on non-
target organisms and the environment. However, some pesticides may affect the environment if they
are misused or persist. Priorities for pest control are constantly changing with increasing emphasis on
environmental issues. The pesticide industry is more sensitive than ever to such issues. The aim is to
find the proper balance between protecting crops and protecting the environment.
We depend on crops produced in fertile soils for almost all our food and much of our clothing, so
it behooves us to nourish and protect these soils by assessing the safety of pesticides applied to crops
and to soils. Harmful products should be rejected and only crop protection agents which can help
maintain pest-free crops and productive-state soils should be developed.
Crop protection agents (CPA) can improve the productivity of soil by controlling insects and diseases
that attack plants. Some of these agents (CPA) also conserve water and nutrients by eliminating weeds
that compete with crops for these items.
Soil contains substantial amounts of organic matter formed by organisms feeding on plant and
animal remains. The cultivated layer of soil normally contains approximately 25 thousand pounds of
organic matter per acre. By contrast, crop protection agents are usually applied at .01 to l.O pounds
per acre. Nevertheless, the process of decomposition and soil-pesticide dilution should continue to be
evaluated until it is clear how pesticides break down into basic non-toxic substances.
The aim of this book is to evaluate pesticide effects on plants by exploring the physical, chemical,
biological, and ecological interactions of pesticides that influence a crop, its environment, and the pests
that affect these crops.
The authors have compiled the most current information to describe the beneficial and deleterious
pesticide interactions in crop production. The successful completion of this book is due to the cooperation
and hard work of all contributing authors. I am also indebted to my wife who was highly cooperative
and supportive of my efforts, during the last three years, to make this book a reality.
It is my hope that this book will be of value to university scientists, students, and researchers at
Government agencies alike, that are interested in the role of pesticides in crop production.
We would appreciate any comments and suggestions to help improve any future editions.

Jack Altman
 THE EDITOR
Jack Altman, Ph.D., is Professor of Plant Pathology in the Department of Plant Pathology and
Weed Sciences in the College of Agriculture at Colorado State University. Dr. Altman received his
B.S. in 1954 and his Ph.D. in 1957 from Rutgers University. Dr. Altman has authored 46 scientific
publications and 22 semi-scientific articles. He has written two books and in 1987 he wrote 14 disease
fact sheets in English and Arabic while on assignment in Jordan.
He has been invited to lecture, develop cooperative research projects, and present symposiums
throughout the U.S., Europe, the Middle East, Asia, and Australia. He was a Senior U.S. Scientist
and Alexander von Humboldt awardee in West Germany from 1977 to 1978. While there, he worked
cooperatively with Dr. Fritz Schonbeck, Dean of Agriculture at Hannover University in Hannover,
Germany, and conducted research on the side effects of herbicides on plant diseases. He was invited
to return and continue his research in Hannover, Germany for an additional four months in 1985. In
1981, he was selected to the PRC (China) Distinguished Scholar Exchange Program by the National
Academy of Sciences. He lectured in Beijing and Harbin and worked cooperatively with Chinese
scientists for two months on a commune in Hulan county north of Harbin. Later, he lectured and
discussed soilborne disease research for an additional month at universities in Shanghai, Nanjing, Jinan,
and Tsingtao. In 1988, Dr. Altman was awarded a Reserve bank of Australia Senior Rural Agricultural
Fellowship and worked cooperatively with Dr. Albert Rovira, head of the CSIRO Soils Division in
Adelaide, South Australia, on research to evaluate the pesticide (herbicide) syndrome for Rhizoctonia,
Gaeumannomyces (take-all), and the cereal cyst nematode.
In 1989, he was the leader of an international Plant Pathology Delegation of 20 scientists to Northern
Europe to exchange the latest scientific information on plant pathology, plant protection, and biological
controls. He spent three weeks visiting scientists and research laboratories in Sweden, Russia, Yugo-
slavia, and Germany.
He is a member of the American Phytopathological Society, the Society of Nematology, Sigma
Xi, and the International Cooperation Committee for A.P.S. He chaired the A.P.S. Soil Microbiology
Committee in 1983 and 1984.
Dr. Altman has completed research on such complex issues as the use of conventional herbicides,
the evaluation of pesticide interaction involving diseases, and soil pesticide toxicity with various iso-
thiocyanates. His current research "Herbicide Plant-Pathogens Interaction in the Plant Disease Syn-
drome," was initiated 25 years ago.
 CONTRIBUTORS
Don V. Allemann, Ph.D. David J. Drahos, Ph.D.
Manager Director of Research
Insect Control Research SBP Technologies, Inc.
CIBA-Geigy Corporation Stone Mountain, Georgia
Greensboro, North Carolina
Stephen 0. Duke, Ph.D.
Jack Altman, Ph.D. Director Southern Weed Science Laboratory
Professor USDA, Agricultural Research Service
Plant Pathology and Weed Science Stoneville, Mississippi
Colorado State University
Fort Collins, Colorado David J. Eagle, M.S.A.
Pesticide Residues Unit
Michael Barrett, Ph.D. Agricultural Development and Advisory
Associate Professor Service
Department of Agronomy Cambridge, England
University of Kentucky
Lexington, Kentucky Philip L. Eberbach, Ph.D.
Lecturer
Albert B. Bassi, Jr., Ph.D. School of Agriculture
Phytopathology Research Specialist Charles Sturt University-Riverina
Department of Biological Research Wagga Wagga, Australia
Agriculture Division
CIBA-Geigy Corporation Susanne Elmholt, Ph.D.
Greensboro, North Carolina Scientist
Department of Soil Biology and Chemistry
Gerrit J. Bollen, Ph.D. Danish Institute of Plant and Soil Science
Senior Lecturer Research Centre Foulum
Wageningen Agricultural University Tjele, Denmark
Wageningen, Netherlands
Virginia A. Ferreira, M.S.
Allan J. Cessna, Ph.D. Mathematician
Research Scientist Great Plains Systems Research Unit
Environmental Chemistry and Application USDA, Agricultural Research Service
Agriculture Canada, Research Station Ft. Collins, Colorado
Regina, Saskatchewan, Canada
Ian G. Ferris, Ph.D.
Raghavan Charudattan, Ph.D.
Coordinator
Professor Herbicide Persistence Research
Plant Pathology Department New South Wales Department of Agriculture
University of Florida Tamworth, New South Wales, Australia
Gainesville, Florida

J. Dekker, Ph.D. Jens C. Frisvad, Ph.D.

Professor Associate Professor
Department of Phytopathology Department of Biotechnology
Agricultural University Technical University of Denmark
Wageningen, The Netherlands Lyngby, Denmark
Ellis Griffiths, Ph.D., D.Sc. Stephen M. Neate, Ph.D.
Professor Senior Research Scientist
Department of Agricultural Sciences Department of Soil Biology
University College of Wales CSIRO Division of Soils
Aberystwyth, Dyfed, Great Britain Glen Osmond, South Australia

Stephen 0. Guy, Ph.D. Juan A. Ocampo, Ph.D.

Assistant Extension Professor Department of Soil Microbiology
Plant, Soil, and Entomological Science Estacion Expermental del Zaidin
University of Idaho C.S.I.C.
Moscow, Idaho Granada, Spain

Bruce M. Haigh, B.E. David Pimentel, Ph.D.

Pesticide Chemistry Professor
New South Wales Department of Agriculture Entomology and Section of Ecology and
Tamworth, New South Wales, Australia Systematics
Cornell University
Dana Kelly Heiny, Ph.D. Ithaca, New York
Research Associate
Department of Plant Pathology Hans-Peter Piorr, Ph.D.
University of Arkansas Associate Professor
Fayfetteville, Arkansas Department of Organic Agriculture
University of Bonn
F. Dan Hess, Ph.D. Bonn, Germany
Director of Biological and Biochemical
Research P. Lawrence Pusey, Ph.D.
Sandoz Agro, Inc. Research Plant Pathologist
Palo Alto, California USDA, Agricultural Research Service
Southeastern Fruit and Tree Nut Research
Stanley J. Kostka, Ph.D. Laboratory
Director of Research Byron, Georgia
Aquatrols Corporation
Cherry Hill, New Jersey
John P. Quinn, Ph.D.
Charles C. Kupatt, Ph.D. Lecturer
Herbicide Research Specialist Department of Biology
Department of Biological Research Queen's University
CIBA-Geigy Corporation Belfast, North Ireland
Greensboro, North Carolina
Philip A. Roberts, Ph.D.
William K. Lauenroth, Ph.D. Professor
Professor Department of Nematology
Range Science Department University of California
Colorado State University Riverside, California
Fort Collins, Colorado

John Lydon, Ph.D. Albert D. Rovira, Ph.D.

Plant Physiologist Director
Weed Science Laboratory Cooperative Research Center for Soil and Land
USDA, Agricultural Research Service Management
Beltsville, Maryland Glen Osmond, South Australia
Snorre Rufelt, Ph.D. George E. Templeton, Ph.D.
Research Information Officer Distinguished Professor
Swedish University of Agricultural Sciences Department of Plant Pathology
Alnarp, Sweden University of Arkansas
Fayetteville, Arkansas
Fritz SchOnbeck, Ph.D.
Institut fur Pflanzenkrankheiten und Ulf Thrane, Ph.D.
Pflanzenschutz Senior Scientist
Universitiit Hannover Department of Biotechnology
Hannover, Germany Technical University of Denmark
Lyngby, Denmark
Ulrike Steiner, Ph.D.
Institut fur Pflanzenkrankheiten und Vilhelm Umaerus, Agr.Dr.
pflanzenschutz Professor and Head
Universitat Hannover Section Plant Pathology
Hannover, Germany Swedish University of Agricultural Sciences
Uppsala, Sweden
Donald R. Sumner, Ph.D. Neil D. Westcott, Ph.D.
Professor Research Scientist
Plant Pathology, Coastal Plan Expt. Stn. Cereals Protection
University of Georgia Agriculture Canada, Research Station
Tifton, Georgia Saskatoon, Saskatchewan, Canada

Jedrzej B. Szerszen, Ph.D. Charles L. Wilson, Ph.D.

Assistant Professor Research Plant Pathologist
Department of Botany and Plant Pathology USDA, Agricultural Research Service
Michigan State University Appalachian Fruit Research Station
East Lansing, Michigan Kearneysville, West Virginia

Michael E. Wisniewski, Ph.D.

Plant Physiologist
Appalachian Fruit Research Station
USDA, Agricultural Research Service
Kearneysville, West Virginia
When tillage begins, other arts follow. The farmers, therefore, are the founders
of human civilization.
-Daniel Webster
 TABLE OF CONTENTS
Introduction
Chapter I
Pesticide Interactions in Crop Production ......................................................... 3
Jack Altman

Physiological, Metabolic, and Morphologic Effects of Pesticides on Plants

Chapter 2
Herbicide Effects on Plant Structure, Physiology, and Biochemistry .............................. 13
F. Dan Hess
Allelopathy

Chapter 3
The Role of Pesticides on Host Allelopathy and Their Effects on Allelopathic
Compounds ................................................................. ...... 0 •••••••••••• 37
John Lydon and Stephen 0. Duke

Fate of Pesticides
Chapter 4
Fate of Pesticides Applied to Cereals Under Field Conditions .................................... 59
Allan J. Cessna and Neil D. Westcott

Chapter 5
Computer Simulation Modeling of Pesticide Fate ................................................ 87
Virginia Ferreira and William K. Lauenroth

Chapter 6
Interactions of Herbicides and Other Agrochemicals in Plants: Interactions in Mixtures
with other Herbicides and with Safeners, Fungicides, Insecticides, and Nematodes .............. 113
Michael Barrett
Persistence of Pesticides
Chapter 7
Herbicide Persistence and Movement in Australian Soils: Implications for Agriculture ........... 133
Ian G. Ferris and Bruce M. Haigh
Resistance to Pesticides
Chapter 8
The Fungicide Resistance Problem: Current Status and Role of Systemics . . . . . . . . . . . . . . . . . . . . . . . 163
J. Dekker

Influence of Pesticides on Soil Symbionts, Soil Microflora, and Soil Mycoflora

Chapter 9
The Effect of Herbicides and Fungicides on Legume-Rhizobium Symbiosis ...................... 183
Philip L. Eberbach

Chapter 10
Influence of Pesticides on VA Mycorrhiza ............................................ 0 ••••••••• 213
J. A. Ocampo
Chapter 11
The Influence of Fungicides on Soil Mycoflora with Special Attention to Tests of
Fungicide Effects on Soil-Borne Pathogens ..................................................... 227
Susanne Elmholt, J. C. Frisvad, and U. Thrane

Chapter 12
Interactions of the Herbicides Glyphosate and Glufosinate (Phosphinothricin) with the
Soil Microflora ................................................................................ 245
John P. Quinn
Iatrogenic Responses and Pesticide - Pathogen Interactions
Chapter 13
Iatrogenic Effects of Pesticides on Plant Disease - An Update and Overview ................... 269
Ellis Griffiths

Chapter 14
Mechanisms Involved in Non-Target Effects of Pesticides on Soil-Borne Pathogens .............. 281
Gerrit J. Bollen

Chapter 15
Interactions of Pesticides with Diseases of Vegetables .......................................... 303
Donald R. Sumner

Chapter 16
Pesticide-Pathogen Interactions in Plant Disease ................................................ 315
Jack Altman
Influence of Nematicides on Nematode Pathogens and their Host Plants
Chapter 17
Influence of Nematicides on Nematode Pathogens and their Host Plants ......................... 335
Philip A. Roberts
Crop Injury

Chapter 18
Agrochemical Damage to Crop Plants .......................................................... 355
David J. Eagle
Safeners

Chapter 19
The Role of Safeners in the Pesticide-Disease Interaction: Influence on the Disease
Syndrome ..................................................................................... 377
Jedrzej B. Szerszen

Biologicals to Replace Pesticides

Chapter 20
Economic Comparisons of Mycoherbicides to Conventional Herbicides .......................... 395
Dana Kelly Heiny and George E. Templeton

Chapter 21
Genetic Engineering of Microorganisms for Pest Control - Survival and Potential
Effectiveness of Such Microorganisms in Field Crop Systems ................................... 409
Stanley J. Kostka and David J. Drahos
Chapter 22
The Role of Pesticides in Altering Biocontrol Efficacy ......................................... 421
Raghavan Charudattan

Ecological Agriculture and Conventional and Biodynamic Cropping Systems

Chapter 23
Reducing Pesticide Use Through Alternative Agricultural Practices- Fungicides and
Herbicides ................................................................ .................... 435
David Pimentel

Chapter 24
Changes in Fanning Systems in the Scandinavian Countries Focused on Pesticide Use ........... 449
Vilhelm Umaerus and Snorre Rufelt

Chapter 25
Phytopathological Advantages and Risks of Organic Fanning Systems - Future
Perspectives ................................................................ ................... 461
H. P. Piorr

Management of Postharvest Diseases

Chapter 26
Management of Postharvest Diseases of Fruits and Vegetables- Strategies to Replace
Vanishing Fungicides ................................................................ .......... 477
P. Lawrence Pusey, Charles L. Wilson, and Michael E. Wisniewski

Induced Resistance

Chapter 27
Induced Resistance as a Means of Plant Disease Control. ....................................... 495
Ulrike Steiner and Fritz Schonbeck

Pesticide-Disease Interactions in No-Till Agriculture

Chapter 28
Pesticide Disease Interactions in Conservation Tillage Systems .................................. 515
Stephen M. Neate and Albert D. Rovira

Economic and Agronomic Factors Related to Pesticide Use

Chapter 29
Pesticide Induced Economic and Agronomic Responses in Cereal Crops ......................... 533
Stephen 0. Guy

Future Trends for Pest Control

Chapter 30
Future Methods for Controlling Weeds, Plant Diseases, and Insects ............................. 545
Charles C. Kupatt, Albert B. Bassi, Jr., and Don V. AUemann

Index ................................................................ ......................... 569

Introduction
 Chapter 1

PESTICIDE INTERACTIONS IN CROP PRODUCTION

Jack Altman

TABLE OF CONTENTS
I. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

II. Insect Resistance Affecting Other Pests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

III. Biocontrol of Resistant Insects ........................................................... 5

IV. Resistance of Weeds ................................................................. .... 5

V. Dimensions of the Resistance Problem and the Development of Biopesticides .............. 6

VI. Purpose of This Book . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

0-8493-6339-X/93/$0 .00 +$.50

© 1993 by CRC Press, Inc. 3
4 Pesticide Interactions in Crop Production

I. INTRODUCTION
Production of food to sustain the population of the earth is an engaging occupation filled with
numerous risks. These include pests and weather (i.e., floods or droughts). Crop losses from pests
average between 30 and 40% annually. New challenges to crop production continue to occur, resulting
in even greater crop losses in isolated areas.
One of the most recent pest problems includes an outbreak of a new strain of the sweet potato
whitefly (Bemisia tabaci) in the Imperial Valley of California in October and November 1991. U.S.
populations of the sweet potato whitefly, (also referred to as the poinsettia whitefly) have sequentially
developed resistance to current pesticides including organophosphates, carbamates, and pyrethroids.
The collapse of existing controls due to the emergence of resistant strains in the late 1980s and early
1990s in many areas of the southern U.S. has resulted in the development of a control crisis which
greatly threatens future produce production. 2 As a result of the whitefly infestation, the 1991 cantaloupe
crop in the Imperial Valley has been severely damaged, and the winter lettuce crop was also in danger
of becoming a complete loss. In addition, cole crops grown in the valley are also being threatened.
A recent novel approach for whitefly control involving a biological control organism, Delphastus
pusilla, has been offered for sale by a small number of suppliers. This insect is a small species of lady
beetle. Adults feed on whitefly eggs and the lady beetle larvae primarily eat whitefly nymphs. Little
work has been done with D. pusilla, but it has the ability to eat large numbers of whiteflies. Studies
suggest that it likely will do best where very high numbers of whiteflies are present, since the adult
beetles need to eat 100 or more whitefly eggs in order to maintain reproduction. 2 Several additional
efforts for whitefly control are currently being evaluated in the Imperial Valley, including a change in
cropping patterns to allow for a 2- or 3-week noncrop interval between crops. However, such a drastic
change wi.U have to be evaluated on a regional basis. 2

II. INSECT RESISTANCE AFFECTING OTHER PESTS

Other important insect pest species that have become resistant to current insecticides are the Colorado
potato beetle (Leptinotarsa decemlineata), the Western flower thrips (Franklinella occidentalis), the
tobacco budworm (Heliothis virescens), and the green peach aphid (Myzus persicae). Populations of
the Colorado potato beetle in the eastern U.S. and Europe have rapidly developed resistance to a wide
range of insecticides as each has been introduced, including the pesticides mentioned previously. Insect
resistance problems in the U.S. are moving westward, with serious problems recently developing in
the Red River Valley production area of North Dakota and Minnesota. There is also evidence that the
potato beetle is becoming resistant to some of the new beetle-active Bacillus thuringiensis (Bt) strains.
Strains of western flower thrips (Frankliniella occidentalis) resistant to most organophosphate and
pyrethroid insecticides caused this insect, and the tomato spotted wilt virus that it transmits, to greatly
increase in importance in the 1980s, emerging as the single most important species of greenhouse insect
in the U.S. during that time. 2
The history of cotton production in the U.S. and Latin America has been closely tied to an ability
to stay ahead of resistance development in the tobacco budworm (Heliothis virescens). By far the single
largest insecticide market, due to tobacco budworm resistance, has also greatly influenced development
and marketing of many other insecticides. At present, the tobacco budworm is highly resistant to most
organophosphate and pyrethroid insecticides, as well as all carbamate and chlorinated hydrocarbon
insecticides.
Highly insecticide-resistant strains of the green peach aphid (Myzus presicae) occur throughout the
U.S. on both vegetable crops (notable potatoes) and greenhouse crops. Since the removal of aldicarb
from the market, only one insecticide, methamidophos, a highly toxic organophosphate insecticide,
has provided reliable control. 2
Altman 5

III. BIOCONTROL OF RESISTANT INSECTS

In spite of these examples of insect resistance, genetically engineered biopesticides are becoming
available. Ecogen Inc. has been granted a U.S. patent for its FOIL® potato biopesticide. FOIL®, which
is based upon a genetically enhanced strain of Bacillus thurinigiensis is used to control Colorado potato
beetles and com borers on potato crops. These pests caused crop losses of an estimated $369 million
in 1990. The patent also covers a novel beetle-active Bt isolate discovered by Ecogen scientists which
is the key to developing the genetically enhanced Bt strain in FOIL®. FOIL® is the only Bt-based
bioinsecticide on the market thus far that controls two different types of insects: beetles (Coleoptera)
and caterpillars (Lepidoptera).
Also available this summer (1991) are two genetically engineered biopesticides developed by
Mycogen Corp. MVP kills the diamondback moth and other caterpillar pests that attack cabbage,
broccoli, lettuce, and other vegetables, while M-TRAK is targeted primarily on the potato beetle, which
attacks all solanaceous vegetables. Mycogen Corp. believes that MVP and M-TRAK will prove to be
more persistent that other biopesticides. 3 Although the Mycogen products incorporate the Bt toxin gene,
the gene is inserted in the Pseudomonas fluorescens bacterium, which can be mass produced. The
Pseudomonas cells are killed and encased in a tiny "biocapsule" that preserves the toxin until it is
ingested by the target pest. By utilizing killed bacteria cells, Mycogen has produced a pesticide that is
more environmentally benign and easier for EPA to register as a pesticide. EPA has also exempted
these products from residue tolerance checks.
Growers in some instances have also revived the practice of using insecticidal oils, and these appear
to be promising for sweet potato whitefly control. Growers are also looking at older insecticides such
as sodium flusaluminate for Colorado potato beetle control.

IV. RESISTANCE OF WEEDS3-7

The challenge for pest control is omnipresent. Weeds, as well as insects and plant pathogens,
readily develop resistance to pesticides, resulting in pest buildup and new challenges to agriculture to
try and control them. Weeds are developing resistance at an alarming rate, as evidenced by resistance
from the use of sulfonureas and carbamates. For example, Homer LeBaron stated at a 1990 workshop
on herbicide resistance, "Obviously, we knew before the sulfonylurea herbicides were on the market
that they had the potential for developing resistance partly because of the single site of action and partly
because of the site of action itself. The enzyme system seems to be quite variable within certain plants.
I would not say that all the newer ones (herbicides) might be less subject to resistance, but anything
that has a single site of action, regardless of where it is, is probably going to be subject to a certain
frequency of resistance. We are also beginning to see, after quite a few years of use, resistance to the
old herbicides we have been using over the years . . . " 4 •5 (See also Chapter 8 and 27 for additional
information or resistance.)
Another (cause) for concern is the potential for multiple resistance, although it has occurred only
in two places in the world, as far as is known ... But there is a potential for the development of a
weed situation, similar to that of the more prominent (resistant) insects, ... where nothing one de-
veloped will control them for more than a year or two, or sometimes not at all, so that chemical ceases
to be of any significance in control of certain insects. That is what is occurring in Lolium rigidum that
has developed multiple resistance (in Australia). Lebaron concludes "I hope we don't see that in this
country, but we have to be careful and watch for it because it could occur ... " 4
In a paper in the Journal of Sugar Beet Research, 5 Schweizer and Westra have reviewed the potential
for weeds to develop resistance to sugar beet herbicides and advocate "Preventive action against the
evolution of herbicide-resistant weeds.'' They state that ''Sixty eight percent of the 37 important problem
weeds in sugarbeets have developed biotypes that are resistant to one or more herbicide classes in
Africa, Australia, Europe, or North America. Eighteen of these weeds have biotypes that are resistant
to herbicides in North America, but only two of these weeds have biotypes that are resistant to sugarbeet
herbicides. Common lambsquarters is resistant to pyrazon in Switzerland and green foxtail is resistant
to trifluralin in Canada. Diclofop methyl, fluazifop, paraquat, pyrazon, and trifluralin are considered
6 Pesticide Interactions in Crop Production

TABLE 1
Losses of Potential Crop Production from Weeds, Insects, and
Diseases in Various Regions of the Worlds

Percentage of losses due to Loss in production as

Insect percentage of
Weeds pests Diseases potential crop value

North and Central 8.0 9.4 11.3 28.7

America
South America 7.8 10.0 15.2 33.0
Europe 6.8 5.1 13.1 12.9
Africa 15.7 13.0 12.9 41.6
Asia 11.3 20.7 11.3 43.3
Oceania 8.3 7.0 12.6 27.9
U.S.S.R. 10.1 10.5 9.1 29.7
Peoples Republic of China 9.7 12.3 11.8 33.8

• Adapted from 1976 National Science Foundation Report on World Crop Loss by Glass.

high risk sugarbeet herbicides. Presently, it seems unlikely that problem weeds in sugarbeets in North
America will develop resistance to sugarbeet herbicides or to other herbicides currently registered for
use in sugarbeet rotations because sugarbeet growers integrate weed management programs, including
crop and herbicide rotations and tillage." 5
Insects likewise develop resistance readily since they are capable of rapid reproduction of new
generations. Plant pathogens too are capable of becoming resistant to modem pesticides, such as oxamyl,
metalaxyl, and benlate. Examples are Cercospora leaf spot on peanuts and sugar beets, and Botrytis
resistance to oxamyl has also been documented recently.
Genetic engineering to develop transgenic plants shows promise as a means to control these resistant.
pests with reduced levels of pesticides or without pesticides. However, this development is progressing
far too slowly; thus, the need for pesticides is still a requirement for economic crop production. These
needs will continue since the rate of resistance by pests to pesticides continues to grow, making many
pesticides inefficient or useless. Annual crop losses from various pests are presented in Table 1 and
pesticide use in recent years is illustrated in Figure 1.
Pest resistance is a growing problem that demands both a research and a policy response. The
scientific response has been growing in recent years. However, the policy response has only recently
been noted. At the federal level, programs with 1990 Farm Bill have been authorized at USDA research
laboratories to specifically deal with pest resistance detection and monitoring. There is also an increased
emphasis on changing agricultural operations to deal with resistance, assuming that pest systems like
IPM are compatible with pest resistance management goals. 3 ·6 •7
Reauthorization of the Federal Insecticide, Fungicide, and Rodenticide Act (FIFRA) in 1991 presents
further opportunities to include pest resistance considerations in the regulatory process. This process
also provides an opportunity to better integrate resistance research advances into regulatory decisions
through better coordination with the USDA. 3
There has also been an increasingly constructive response to resistance problems by the private
sector. This needs to be encouraged, and a public-private partnership in this area is essential. During
any governmental policy discussions, the private sector needs to be involved, and their willingness to
voluntarily cooperate needs to be factored into any regulatory proposals.

V. DIMENSIONS OF THE RESISTANCE PROBLEM AND THE

DEVELOPMENT OF BIOPESTICIDES
Pest resistance to conventional control measures, using chemical pesticides, is expanding. The list
of pests which developed or are developing resistance since the 1940s now includes 107 species of
Altman 7

Million lbs.

1960 1965 1970 1975 1980 1985

64& M.lll. TOll! 177 M.U.. TOll! 1034 M.lla. TOll! 1603 M.lla. TOll! 1461 M.lla. TOll! 1239 MJ1IL TOll!

~ Herbicides II Insecticides l;:,j:j:;:::!l Fungicides

FIGURE 1. Changes in pesticide production in the U.S. in 25 years. (Adapted from Statistical Abstract of the U.S.,
1988, 108th ed., U.S. Department of Commerce, Bureau of Census [J.A.])

weeds, 150 species of plant pathogens, and 450 species of insects, according to the National Academy
of Sciences (NAS) report in 1986. G.P. Georghio from the University of California, Riverside, reported
in 1981 that these numbers for resistant weeds were 30, resistant plant pathogens 81 , and pesticide-
resistant insects 447. If the 1981 report is compared with the 1986 NAS report, the resistance problem
seems to be increasing. This pest resistance has resulted in increased pesticide use and/or increased
losses due to pests. 6 •7 As a result of resistance buildup, the rates at which new pesticides are being
developed is slowing. Although biotechnology bodes well for the future of pest control, it is contributing
to this slowdown of pesticide development. 4
Biopesticides have been available for some 20 years, but despite their obvious environmental
advantage, they haven't been as effective generally as conventional chemical treatments in controlling
insect pests. 6 As a consequence, they haven't been widely used by growers. Currently, biopesticides
have only about a 1% share of the $20 billion a year global pesticide market, 25% of which is in the
U.S. Companies such as Ecogen and Mycogen have reportedly produced a more effective second
generation of biopesticides by bringing biotechnology to bear on the traditional problems of a slow rate
of insect kill and rapid degradation in the environment. 3
These promising developments come at a time when a rising tide of producer and consumer concerns
about food, soil, and water contamination from agriculture chemicals has created a new interest in
biopesticides. In addition, biotechnology is poised to contribute to the development of improved bio-
products. The big question is whether biopesticides really can compete with chemicals. According to
one financial analyst, adoption of new pesticides by farmers is usually slow. Traditionally, they ex-
periment with a new product for 2 or 3 years on increasingly larger fields before switching over
completely. This analyst points out that for the farmer, performance is the most important factor in
choosing an insecticide, followed second by the relative cost, and third by environmental considerations. 3
The analyst believes that farmers will initially use biopesticides to eradicate pests that have become
resistant to chemical agents before making a bigger changeover. Farmers who are accustomed to quick
kills from chemicals must also be educated on how the slower acting biopesticides work. Biopesticides
"manage" rather than eradicate pests. In terms of effectiveness, individual biopesticides probably can
give some chemicals a run for their money. However, can the fledgling biopesticide industry compete
financially with the chemical giants? Many of the innovative new bioproducts are developed by small,
at times undercapitalized companies. R&D periods are long and costs are high for testing and registration.
8 Pesticide Interactions in Crop Production

To truly compete with chemicals, a well-financed biopesticide industry must be developed. Venture
capital, in particular, is needed so that companies can develop and market new and better products.
Thus far, the financial markets appear to be more interested in the biomedical product companies as
opposed to the lower profit agricultural biotech firms.
Widespread commercial availability of new biopesticides is not a current reality. 3 •7 With early work
concentrating on a limited number of biopesticides, such as Bacillus thuringensis and its endotoxin for
insect control and FOIL®, MVP, and M-TRAK being evaluated in 1991 for insect control in limited
areas (actually the biggest use of Bt in the U.S. is for control of forest insects including budworms
and gypsy moth) 2 and "Devine" and "Collego" used as weed control pathogens. 8 Other than these,
there may not be a broad array of pest control options available in the near future.

VI. PURPOSE OF THIS BOOK

This book, with contributions from an international array of scientists, attempts to provide some
insight into present and future challenges to crop production and to review how pesticides affect crops.
It will attempt to address the influence of pesticides on above-ground portions of plants, including
foliage and fruits, and the below-ground portions of plants to show how symbiotic relationships with
mycorrhiza and rhizobia play an integral part in crop production. Chapters by Ocampa and Eberbach
address this latter issue. It will also try to address crop production with alternative means that allow
for pesticide reduction. These points are addressed by Umaerus and Rufelt in a discussion of •'ecological
agriculture,'' in which Scandinavian countries are pointing the way toward crop production with reduced
levels of pesticides or without pesticides. Ecological agriculture embodies regulations to reduce pesticide
use by 50% in the next 5 years in Sweden, Denmark, and Norway and to develop farming methods
that "induce" biological pest control.
In addition to an ecological agriculture approach, another method for pest control is addressed by
the section on induced resistance by Seiner and Schonbeck. Viable crop production involves economics,
and this point is addressed by Heiny and Templeton8 and Guy. 9 Pesticide use in crop production is
necessary in current production methods of modern agriculture, and the physiological effects of pesticides
on plants are reviewed by Hess and by Lydon and Duke. Since the use of pesticides is inevitable, the
chapters by Altman, Sumner, and Griffiths discuss some of the problems and challenges that result
from their use.
Facing the inevitable, this book attempts to provide some insight into how pesticides affect crop
production. Pesticide injury and toxicity are reviewed by Eagle. The effect of pesticides on the internal
structure of a crop plant is addressed by Hess. This book provides some information on iatrogenic
(nontarget) responses. The book also addresses the persistence of chemicals and the resistance of pests
as well as how these factors affect crop production. Finally, the book attempts to discuss the economics
of pesticide or biopesticide use and the future trends for replacing pesticides with biocontrol agents
and/or genetic engineering to develop pest-resistant or -tolerant crops.

REFERENCES
1. Perring, T. M., Cooper, A., Kazmer, D. V., Shields, C., and Shields, J., New strain of sweet potato whitefly
invades California vegetables, Calif. Agric., 45, 6, 1991.
2. Cranshaw, W., Personal communication, 1991.
3. NBIAP. News Report. National Biological Impact Assessment Program. University of Arizona, Dept. of Plant
Pathology. Tuscon. August 1991.
4. LeBaron, H., Herbicide resistance: a workshop, "A Call for Industry Action", Weed Techno/., Vol. 4(1), Jan.-
March, Weed Science Society of America, Champaign, IL, 219, 1990.
5. Schweitzer, E. E. and Westra, P., Potential for weeds to develop resistance to sugar beet herbicides in North
America, J. Sugar Beet Res., 28, 1, 1991.
Altman 9

6. National Academy of Sciences, Pesticide Resistance, Strategies and Tactics for Management, National Academy
Press, Washington, DC, 1987.
7. Dover, M. and Croft, B., Getting Tough: Public Policy and the Management of Pesticide Resistance, World
Resources Institute, Study No. I, Washington, D.C., 1984.
8. Heiny, D. K. and Templeton, G. E., Economic comparisons of mycoherbicides to conventional herbicides, in
Pesticide Interactions in Crop Production: Beneficial and Deleterious Effects, Altman, J., Ed., CRC Press, Boca
Raton, FL, 1993.
9. Guy, S. 0., Pesticide induced economic and agronomic responses in cereal crops, in Pesticide Interactions in
Crop Production: Beneficial and Deleterious Effects, Altman, J., Ed., CRC Press, Boca Raton, FL, 1993.
Physiological, Metabolic, and Morphologic Effects
of Pesticides on Plants
 Chapter 2

HERBICIDE EFFECTS ON PLANT STRUCTURE, PHYSIOLOGY, AND

BIOCHEMISTRY
F. Dan Hess

TABLE OF CONTENTS
I. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

II. Inhibition of Plant Biosynthesis Reactions ............................................... 14

A. Inhibition of Carotenoid Biosynthesis ............................................ 14
B. Inhibition of Lipid Biosynthesis .................................................. 16
C. Inhibition of Essential Amino Acid Biosynthesis .................................. 18
I. Aromatic Amino Acids ................................................... 18
2. Branched Chain Amino Acids ............................................ 19

III. Disruption of Membrane Integrity of Plant Cells ......................................... 21

A. Inhibition of Photosystem II Electron Flow ....................................... 22
B. Interaction at Photosystem I ..................................................... 23
C. Accumulation of Protoporphyrin IX .............................................. 23
D. Inhibition of Glutamine Synthetase (GS) ......................................... 25

IV. Effects on Plant Growth and Development ............................................... 26

A. Effects on Cell Division ......................................................... 27
I. Inhibition of Cell Division . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . 27
2. Disruption of Cell Division ............................................... 28
B. Inhibition of Cell Wall Formation ................................................ 29
C. Auxin Agonists .................................................................. 30

V. Summary and Conclusions .............................................................. 30

References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

().8493-6339-X/93/$0.00 +$.50
<r> 1993 by CRC Press, Inc. 13
14 Pesticide Interactions in Crop Production

I. INTRODUCTION

In crops, interactions between different components present in an agroecosystem can lead to a

detrimental influence on the quality or quantity of the crop produced, whereas effects of individual
components have no measurable influence. One important interaction is the potential additive or syn-
ergistic effect between phytotoxicity caused by plant diseases and the phytotoxicity caused by pesti-
cides. '·3 Alone, the degree of phytotoxicity caused by a pesticide or a disease may not be measurable,
whereas when both components are present, interaction of the two leads to a significant loss of yield.
To better evaluate the basis for this interaction, the target site of the pesticide needs to be understood.
The only pesticides designed to have direct effects on plants are herbicides. In fact, optimally other
pesticide groups, e.g., insecticides, fungicides, and nematocides, do not have any measurable effect
on plants after application. Therefore, even though this book deals with all pesticides, this chapter will
be limited to the effect of herbicides on plants.
Herbicides that are currently in development or are commercially used interfere with a wide variety
of plant biochemical reactions. The purpose of this chapter will be to catalog herbicides as to how they
interact with structural, physiological, and biochemical events in the plant (their mode of action). The
majority of herbicide use is in crop production. In most instances the herbicide site of action is present
and functional in the crop, and the lack of crop injury is due to events such as herbicide inactivation
by degradation or conjugation.
Most herbicide action can be categorized into one of three general types: (I) interference with
biosynthesis of plant constituents, (2) disruption of plant cell membranes, or (3) inhibition of plant
growth and development. Although some herbicides can fit into more than one category, for the purpose
of this chapter, only the principle category will be chosen. Biosynthetic pathways in plants that are
inhibited by herbicides include carotenoid pigment biosynthesis, lipid biosynthesis, and essential amino
acid biosynthesis. Herbicide target sites within essential amino acid biosynthesis inhibit both aromatic
and branched chain amino acids. Herbicide target sites cause plant membrane disruption by three
different mechanisms that eventually induce lipid peroxidation: inhibiting electron flow in photosystem
IT, capturing electrons in photosystem I, and accumulation of the chlorophyll precursor, protoporphyrin
IX. Many herbicides that are applied prior to the emergence of the weed and crop inhibit growth by
interfering with cell division or cell enlargement after seed germination. In some instances, the bio-
chemical or biophysical target sites of these herbicides are known, and in others they are not. Even
though growth inhibition in itself does not cause lethality, if the weed does not emerge above the soil
surface, death will occur once the available food reserves in the seed are utilized. Other herbicides
influence growth and development by interfering with cell wall biosynthesis or by being plant hormone
agonists.

II. INHIBITION OF PLANT BIOSYNTHESIS REACTIONS

Many biosynthetic pathways produce substances that are essential for normal plant growth and
development. Inhibition of one of these processes can lead to major disruptions in normal growth and
development and, directly or indirectly, the overall effect is often lethal. Currently available commercial
herbicides are known to inhibit three important biosynthetic pathways: carotenoid pigments, lipids, and
essential amino acids. Although other herbicides have sites of action in biosynthetic pathways (e.g.,
diphenyl ether herbicides), their action on the plant is not directly related to the depletion of the product
and, thus, will be discussed elsewhere in this chapter.

A. INHIBITION OF CAROTENOID BIOSYNTHESIS

The most striking symptom resulting from treating plants with herbicides that inhibit carotenoid
biosynthesis is the totally white foliage produced following treatment. This is sometimes termed "albino
growth". Growth does continue for a time, but without production of green photosynthetic tissue to
sustain carbohydrate production during photosynthesis, growth cannot be maintained. After growth
ceases, necrosis begins to occur due to peroxidative reactions. Those herbicides that inhibit carotenoid
biosynthesis do not affect preexisting carotenoids. Thus, plant tissues formed before treatment do not
Hess 15

LIPID LIPID
RADICAL ____. PEROXIDATION
LIGHT

~ LIPID/

• 1 CHL i.s.c.
~

!
CHL 3CHL

OXYGEN ~ LIPID\

~ SINGLET
OXYGEN
~ •
LIPID
RADICAL

PS II

FIGURE 1. Chlorophyll (CHL) reactions that lead to membrane disruption by lipid peroxidation. The energy in singlet
chlorophyll ('CHL) is normally passed to the photosystem II reaction center, but can undergo intersystem crossing (i.s.c.)
to the more reactive triplet chlorophyll ('CHL) state. Triplet chlorophyll can pass its energy to carotenoids. If carotenoids
are not available or if triplet chlorophyll is present in abundance, triplet chlorophyll can initiate lipid peroxidation.

show typical albino symptoms. Carotenoid turnover does occur in these tissues; thus, chlorosis develops
in preexisting tissue as turnover proceeds.
Even though the new growth in treated plants is white, these herbicides do not inhibit chlorophyll
biosynthesis. In fact, plants treated with carotenoid biosynthesis inhibitors and then grown in very low
light (10 lux) will produce new growth that is green and contains up to 70% of the chlorophyll present
in nontreated plants. 4 The loss of chlorophyll is the result of destruction of chlorophyll by light
(photooxidation). One important role of carotenoids is to protect chlorophyll from photooxidation. After
chlorophyll is synthesized and becomes functional, some of the chlorophyll which has been electronically
excited by absorbing light photons is transformed from the singlet form to the longer lived (but more
reactive) triplet form 5 (Figure I). Normally the energy from this reactive form of chlorophyll is dissipated
through carotenoids. When carotenoids are not present, these triplet states undergo degrading reactions,
among which are membrane and chlorophyll destruction.
The biosynthesis pathway of carotenoids is shown in Figure 2. The first series of reactions forms
a 20-carbon intermediate, geranylgeranyl pyrophosphate (GGPP). Formation of GGPP occurs in the
general isoprenoid pathway. Two GGPP molecules combine to form the 40-carbon intermediate phy-
toene. Then, after a series of desaturation reactions (dehydrogenation), a cyclization occurs. After final
molecular modifications, the carotenoids become functional.
Herbicides that inhibit carotenoid biosynthesis react at several sites, although there appears to be
a few common sites that are susceptible to attack. The best studied herbicide site of action in carotenoid
biosynthesis is the inhibition of phytoene desaturation. The enzyme inhibited is phytoene desaturase.
Because inhibition occurs in an in vitro system (e.g., isolated daffodil chromoplasts), the mechanism
of action is proposed to be directly on the functioning enzyme. 6 Inhibition of this enzyme causes a
large accumulation of phytoene and phytofluene in treated plants and is the most commonly reported
proof for herbicide action at this site. Herbicides that have been shown to inhibit this reaction are
norflurazon, fluridone, difunon (EMD-IT 5914), flurochloridone (R-40244), cyclohexane diones (SC
0051), flurtamone (RE 40885), and others. Another site of action for carotenoid synthesis inhibitors
is zeta-carotene desaturation, which results in the accumulation of high zeta-carotene levels. Examples
of herbicides that block at this site are dichlormate, 6-methyl pyrimidines, methoxyphenone, and
16 Pesticide Interactions in Crop Production

MEVALONIC ACID PHYTOENE

t
YTOENE
SATURASE

2H
ISOPENTENYL
PYROPHOSPHATE (5 C)
PHYTOFLUENE

liPP
~2H
ISOMERASE

DIMETHYLALLYL

riPP
PYROPHOSPHATE (5 C)
ZETA CAROTENE

A CAROTENE
SATURASE

2H
GERANYL
PYROPHOSPHATE (10 C)
NEUROSPORENE

FARNESYL
PYROPHOSPHATE (15 C)
~2H
LYCOPENE

GERANYLGERANYL
PYROPHOSPHATE (20 C) ALPHA-CAROTENE BETA-CAROTENE

t
PHYTONE (40 C)
~
LUTEIN
~
ZEAXANTHIN

FIGURE 2. The carotenoid biosynthesis pathway. Most herbicides that inhibit carotenoid biosynthesis inhibit the phytoene
or the zeta-carotene desaturase enzymes.

dihydropyrones. A third site of action has been proposed for the carotenoid biosynthesis pathway. Duke
et al. 7 reported that treating plants with clomazone causes an accumulation of gossypol, a triterpenoid;
thus, they suggested that clomazone inhibition of carotenoid biosynthesis occurs prior to GGPP. Later
research suggested the specific site of action of clomazone was between isopentenyl pyrophosphate
(IPP) and GGPP (Figure 2) (IPP isomerase and prenyl transferase). 8 Inhibition at this site decreases
carotenoids, chlorophylls, and gibberellic acid. However, recent research9 did not find any in vitro
inhibition by clomazone of the plastid enzymes catalyzing the steps leading from IPP to GGPP (IPP
isomerase and prenyl transferase). A final identified site of action for carotenoid biosynthesis inhibitors
is at the cyclization step following lycopene synthesis. This results in an accumulation of lycopene.
The only herbicides shown to act at this site are the substituted triethylarnines. 10 For a detailed discussion
of carotenoid synthesis inhibitors, see the reviews by Ridley 10 and Sandmann and Boger. 11

B. INHIBITION OF LIPID BIOSYNTHESIS

During plant growth, constant supplies of new lipids are needed for production of new membranes
as cells divide and expand. Because of this, inhibiting lipid biosynthesis will rapidly translate into
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