Table Of ContentHERBICIDE-
RESISTANT
CROPS
Agricultural, Environmental,
Economic, Regulatory, and
Technical Aspects
Edited by
Stephen O. Duke
Boca Raton London New York
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______________________________ PREFACE
I have had an interest in herbicide-resistant crops since the idea of using biotechnology to produce
such crops began to be discussed around 1980. My interest intensified in 1990, when I was asked to chair
a panel to write a report on this topic for the Council for Agricultural Science and Technology. Since
then, I have co-authored two more reviews that deal exclusively with the topic and have attended various
workshops, symposia, seminars, and panels on herbicide-resistant crops all over the United States. In
1994, I visited with biotechnologists, regulatory officials, and scientists assessing risk who are dealing
with herbicide-resistant crops in Denmark. In 1995, I attended a workshop on herbicide-resistant crops
in Australia and the First International Conference on Herbicide-Resistant Crops and Weeds in Spain.
I have watched the scientific and regulatory interest in herbicide-resistant crops become intense
throughout most of the developed world.
The amount of literature on herbicide-resistant crops has grown from a trickle fifteen years ago to
a flood. Yet, no book that deals exclusively with this topic has been produced. This book was conceived
to fill the void. Moreover, my intention was to produce a book that would cover most, if not all,
significant aspects of the topic, from the technology used to produce these crops to various potential
impacts of their use, as perceived by authors with different points of view. This book should be a good
reference for regulatory officials, biotechnologists, economists, agriculturists, and environmentalists
with an interest in herbicide-resistant crops.
Herbicide-resistant crops have been a matter of hot debate over the past decade, a time when they
have had no significant impact on agriculture. At the time of the printing of this book, the first significant
impacts of these products of biotechnology will begin to be felt in the United States. Over the next
several years comparing the predictions and conclusions of the various authors in this book with the
actual use, performance, and impact of herbicide-resistant crops should be an interesting endeavor.
I thank the authors who contributed their time, effort, and expertise to this volume. All of the authors
have had a deep involvement in herbicide-resistant crops, and most of them have published substantially
in the areas that they cover. Many of the chapters in this book are the most up-to-date and detailed
coverage of the specific topic available.
As with most edited books, the final list of chapters and authors differs somewhat from the one with
which I started. It was difficult to find authorities in some areas who could produce a chapter in a
reasonable period. I feel very fortunate to have accomplished this, despite having some of my initial
authors drop out after some time. This put the new authors under considerable pressure to generate
manuscripts in a limited time period. I especially thank those authors who came to my rescue to keep
the book well balanced in terms of context.
Stephen 0. Duke
_________________________ THE EDITOR
Stephen 0. Duke, Ph.D., is Director of the Southern Weed Science Laboratory, Agricultural
Research Service, United States Department of Agriculture at Stoneville, Mississippi.
Dr. Duke received a B.S. in biology from Henderson State College (Arkadelphia, Arkansas) in
1966, a M.S. in botany from the University of Arkansas (Fayetteville) in 1968, and a Ph.D. in botany
from Duke University (Durham, North Carolina) in 1975.
He is a member of numerous scientific societies and organizations in which he has held many
offices and assignments and has won numerous awards. These include: the American Association for
the Advancement of Science (Fellow); the American Society for Plant Physiologists (President of the
Southern Section); the Weed Science Society of America (President-Elect, Fellow, Scientist of the
Year), the Southern Weed Science Society (President, Scientist of the Year), the American Society for
Photobiology, the American Chemical Society (Vice Chair of Natural Products Subdivision), and the
Council for Agricultural Science and Technology (Board of Directors, Chair of Editorial Committee).
Among other awards, he has won the USDA Edminster Award, the USDA-ARS Mid-South Area
Scientist of the Year Award, and the Ciba-Geigy Agricultural Recognition Award.
Dr. Duke has published more than 100 refereed research journal articles and more than 50 invited
reviews. He is co-author of Physiology of Herbicide Action and editor of four other volumes. He has
presented well over 100 invited lectures at national and international meetings and has organized
numerous symposia for many different organizations. His current research interests include the use of
natural products as herbicides and the interaction of biological weed control agents with pesticides.
____________ CONTRIBUTORS
Greg Baldwin, M.S. Stephen 0. Duke, Ph.D.
Stoneville Pedigreed Seed Company Agricultural Research Service
Stoneville, Mississippi United States Department Agriculture
Stoneville, Mississippi
Gerard G. Barry, Ph.D.
The Agriculture Group
Donald N. Duvick, Ph.D.
Monsanto
Department of Agronomy
St. Louis, Missouri
Iowa State University
Ames, Iowa
Newell F. Bascomb, Ph.D.
American Cyanamid Company
William E. Dyer, Ph.D.
Princeton, New Jersey
Department of Plant, Soil, and
Environmental Sciences
Peter Boger, Ph.D.
Montana State University
Department of Plant Physiology
Bozeman, Montana
University of Konstanz
Konstanz, Germany
Charlotte V. Eberlein, Ph.D.
Department of Plant, Soil, and
Margaret R. Bolick, Ph.D.
Entomological Sciences
Nebraska State Museum
University of Idaho
University of Nebraska
Aberdeen, Idaho
Lincoln, Nebraska
David E. Eichholtz
Orvin C. Burnside, Ph.D.
Department of Agronomy The Agriculture Group
and Plant Genetics Monsanto
University of Minnesota St. Louis, Missouri
St. Paul, Minnesota
Graeme Evans, Ph.D.
Harold D. Coble, Ph.D. Bureau of Resource Sciences
Crop Science Department Department of Primary Industries
North Carolina State University and Energy
Raleigh, North Carolina Canberra, Australia
Bruce Coulombe, Ph.D. Robert T. Fraley, Ph.D.
Stoneville Pedigreed Seed Company The Agriculture Group
Stoneville, Mississippi Monsanto
St. Louis, Missouri
Xavier Delannay, Ph.D.
The Agriculture Group Roy L. Fuchs, Ph.D.
Monsanto The Agriculture Group
St. Louis, Missouri Monsanto
St. Louis, Missouri
Michael J. Donnelly-V anderloo
Department of Crop Science Rebecca J. Goldburg, Ph.D.
University of Guelph Environmental Defense Fund
Guelph, Ontario, Canada New York, New York
Jonathan B. Gressel, Ph.D. Danny Llewellyn, Ph.D.
Department of Plant Genetics C.S.I.R.O.
Weizmann Institute of Sciences Division of Plant Industry
Rehovot, Israel Canberra, Australia
J. Christopher Hall, Ph.D. Carol Mallory-Smith, Ph.D.
Depanment of Environmental Biology Department of Crop and Soil Science
University of Guelph
Oregon State University
Guelph, Ontario, Canadn
Corvallis, Oregon
F. Dan Hess, Ph.D. C. Jeffry Mauvais, Ph.D.
Sandoz Agro, Inc. Stine-Haskell Research Center
Palo Alto, California DuPont Agricultural Products
Wilmington, Delaware
D. Douglas Hopkins, J.D.
Environmental Defense Fund George D. McLean, Ph.D.
New York, New York
Department of Primary Industries and Energy
Bureau of Resource Sciences
Catherine M. Houck, Ph.D.
Canberra, Australia
Department of Molecular Biology
Calgene, Inc.
Norihiko Misawa, Ph.D.
Davis, California
Kirwin Brewery Company
Central Laboratories for Key Technology
David J. Hume, Ph.D.
Yokohama, Japan
Department of Crop Science
University of Guelph
Guelph, Ontario, Canada David W. Monks, Ph.D.
Honicultural Science Department
Kathleen H. Keeler, Ph.D. Nonh Carolina State University
School of Biological Sciences Raleigh, Nonh Carolina
University of Nebraska
Lincoln, Nebraska Thomas B. Moorman, Ph.D.
United States Depanment of Agriculture
Kyle E. Keller, Ph.D. Agricultural Research Service
United States Depanment of Agriculture National Soil Tilth Laboratory
Agricultural Research Service Ames, Iowa
National Soil Tilth Laboratory
Ames, Iowa Stephen R. Padgette, Ph.D.
The Agriculture Group
John A. Kiser, M.S.
Monsanto
Calgene, Inc.
St. Louis, Missouri
Davis, California
Diane B. Re
Ganesh M. Kishore, Ph.D.
Department of Regulatory Affairs
The Agriculture Group
Monsanto
Monsanto
St. Louis, Missouri
St. Louis, Missouri
Stephen G. Rogers, Ph.D.
David Last, Ph.D.
C.S.I.R.O. Depanment of Regulatory Affairs
Division of Plant Industry Monsanto
Canberra, Australia St. Louis, Missouri
Leonard L. Saari, Ph.D. Loren W. Tauer, Ph.D.
Stine-Haskell Research Center Department of Agricultural, Resource,
DuPont Agricultural Products and Managerial Economics
Newark, Delaware Cornell University
Ithaca, New York
Gerhard Sandmann, Ph.D.
Biosynthesis Group Donald C. Thill, Ph.D.
Department of Botany Department of Plant, Soil, and
J. W. Goethe University Entomological Sciences
Frankfurt, Germany University of Idaho
Moscow, Idaho
Frank S. Serdy, Ph.D.
Department of Regulatory Affairs Charles E. Turner, Ph.D.
Monsanto United States Department of Agriculture
St. Louis, Missouri Agricultural Research Service
Western Regional Research Center
Dale L. Shaner, Ph.D.
Albany, California
American Cyanamid Co.
Princeton, New Jersey
Indra K. Vasil, Ph.D.
Department of Horticultural Sciences
Timothy D. Sherman, Ph.D.
University of Florida
Department of Biological Sciences
Gainesville, Florida
University of Southern Alabama
Mobile, Alabama
Kevin C. Vaughn, Ph.D.
United States Department of Agriculture
Wendy Smith, Ph.D.
Agriculture Research Service
American Cyanamid Co.
Stoneville, Mississippi
Princeton, New Jersey
Stephen Waters, Ph.D.
David A. Somers, Ph.D.
Monsanto
Department of Agronomy and
Europe S.A.
Plant Genetics
Brussels, Belgium
University of Minnesota
St. Paul, Minnesota
John W. Wilcut, Ph.D.
David M. Stalker, Ph.D. Department of Crop Science
Calgene Inc. North Carolina State University
Davis, California Raleigh, North Carolina
Terry B. Stone Alan C. York, Ph.D.
Department of Regulatory Affairs Department of Crop Science
Monsanto North Carolina State University
St. Louis, Missouri Raleigh, North Carolina
___________________________ CONTENTS
Chapter 1 Herbicide-Resistant Crops-Background and Perspectives ....................................... 1
Stephen 0. Duke
Part I The Technology
Chapter 2 Mechanisms of Action and Resistance to Herbicides ............................................ 13
Timothy D. Sherman, Kevin C. Vaughn, and Stephen 0. Duke
Chapter 3 Techniques for Producing Herbicide-Resistant Crops ........................................... 37
William E. Dyer
Chapter 4 New Weed Control Opportunities: Development of Soybeans with a
Roundup Ready"' Gene ......................................................................................................... 53
Stephen R. PadgeUe, Diane B. Re, Gerard F. Barry, David E. Eichholtz,
Xavier Delannay, Roy L. Fuchs, Ganesh M. Kishore, and Robert T. Fraley
Chapter 5 Phosphinothricin-Resistant Crops ........................................................................... 85
lndra K. Vasil
Chapter 6 Cotton Weed Control Using the BXN System ....................................................... 93
David M. Stalker, John A. Kiser, Greg Baldwin, Bruce Coulombe, and
Catherine M. Houck
Chapter 7 Triazine-Resistant Crops: The Agronomic Impact and
Physiological Consequences of Chloroplast Mutation ......................................... 107
J. Christopher Hall, Michael J. Donnelly-Vanderloo, and David J. Hume
Chapter 8 Sulfonylurea Herbicide-Resistant Crops ............................................................... 127
Leonard L. Saari and C. Jeffry Mauvais
Chapter 9 Imidazolinone-Resistant Crops: Selection, Characterization, and
Management .......................................................................................................... 143
Dale L. Shaner, Newell F. Bascomb, and Wendy Smith
Chapter 10 Genetic Engineering of Crops for Tolerance to 2,4-D ......................................... 159
Danny Llewellyn and David Last
Chapter 11 Aryloxyphenoxypropionate- and Cyclohexanedione-Resistant Crops ................. 175
David A. Somers
Chapter 12 Steps Towards Genetic Engineering of Crops Resistant Against
Bleaching Herbicides ............................................................................................ 189
Gerhard Sandmann, Norihiko Misawa, and Peter Boger
Chapter 13 Possible Pleiotropic Effects in Herbicide-Resistant Crops .................................. 201
Carol Mallory-Smith and Charloue V. Eberlein
Description:Edited by a recognized leader in the field, Herbicide-Resistant Crops is the first book to cover all of the issues related to the controversial topic of herbicide-resistant crops. It provides extensive discussions of the modern biotechnological methods that have been used to develop such crops, and
