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Copyright © 2011. Michigan State University Press. All rights reserved.
Environmental Safety of Genetically Engineered Crops, Michigan State University Press, 2011. ProQuest Ebook Central,
E n v i ro n m e n ta l S a f e t y o f
G e n e t i c a l l y E n g i n ee r e d C r o p s
Copyright © 2011. Michigan State University Press. All rights reserved.
Environmental Safety of Genetically Engineered Crops, Michigan State University Press, 2011. ProQuest Ebook Central,
Copyright © 2011. Michigan State University Press. All rights reserved.
Environmental Safety of Genetically Engineered Crops, Michigan State University Press, 2011. ProQuest Ebook Central,
E n v i ro n m e n ta l
Safety of
G e n e t i c a l ly
E n g i n ee r e d
Crops
Environmental Safety of Genetically Engineered Crops, Michigan State University Press, 2011. ProQuest Ebook Central,
Copyright © 2011 by Michigan State University
i The paper used in this publication meets the minimum requirements of ANSI/NISO Z39.48-1992 (R 1997)
(Permanence of Paper).
17 16 15 14 13 12 11 1 2 3 4 5 6 7 8 9 10
Environmental safety of genetically engineered crops / edited by Rebecca Grumet . . . [et al].
p. cm.
Includes bibliographical references and index.
ISBN 978-1-61186-008-5 (cloth : alk. paper)
1. Crops—Genetic engineering—Environmental aspects. 2. Crops—Genetic engineering—Safety measures.
3. Crops—Genetic engineering—Government policy. I. Grumet, Rebecca.
SB123.57.E68 2010
631.5'233—dc22 2010052158
G Michigan State University Press is a member of the Green Press Initiative and is committed to developing
and encouraging ecologically responsible publishing practices. For more information about the Green
Press Initiative and the use of recycled paper in book publishing, please visit www.greenpressinitiative.org.
Copyright © 2011. Michigan State University Press. All rights reserved.
Visit Michigan State University Press on the World Wide Web at:
www.msupress.msu.edu
Environmental Safety of Genetically Engineered Crops, Michigan State University Press, 2011. ProQuest Ebook Central,
This book is dedicated to the Father of Green Revolution, Dr. Norman
E. Borlaug (1914–2 009), a visionary scientist and inspirational leader
who dedicated his life to the development and use of new science and tech-
nology for the benefit of farmers and consumers around the world.
Copyright © 2011. Michigan State University Press. All rights reserved.
Environmental Safety of Genetically Engineered Crops, Michigan State University Press, 2011. ProQuest Ebook Central,
Copyright © 2011. Michigan State University Press. All rights reserved.
Environmental Safety of Genetically Engineered Crops, Michigan State University Press, 2011. ProQuest Ebook Central,
Contents
Foreword by M. S. Swaminathan ix
Preface xi
Environmental Safety of Genetically Engineered Crops, Michigan State University Press, 2011. ProQuest Ebook Central,
viii
| Contents
Contributors 225
Index 231
Environmental Safety of Genetically Engineered Crops, Michigan State University Press, 2011. ProQuest Ebook Central,
Foreword
M. S. Swaminathan
from population growth, biodiversity loss, and climate change. Therefore, every country
should equip itself with the capacity to assess and manage environmental risks. This book will
be of invaluable help in this task. We owe the authors deep gratitude for their comprehensive
and authoritative treatment of a difficult and often emotionally charged subject. All biotech-
nology regulatory authorities should derive benefit from this publication.
ix
Environmental Safety of Genetically Engineered Crops, Michigan State University Press, 2011. ProQuest Ebook Central,
Copyright © 2011. Michigan State University Press. All rights reserved.
Environmental Safety of Genetically Engineered Crops, Michigan State University Press, 2011. ProQuest Ebook Central,
Preface
individual farmers’ choices. While science cannot stand alone in determining appropriate
policies and decisions about use of new technologies, discussions of policy must be rooted in
accurate information. This is especially important with respect to questions of biosafety. As
a starting point in assessing validity, we can ask, Is there scientific consensus at the highest
levels of available expertise? How far does the consensus extend? Where is there divergence of
opinion, and what are the gaps in our knowledge?
More than one hundred U.S., European, and international scientific societies from around
the world have addressed the relative safety of genetically engineered and conventionally bred
crops. All have reached the conclusion that properly regulated genetic engineering of crop
plants does not pose new risks to human health or the environment relative to conventional
plant breeding. GE crops are not inherently dangerous. This conclusion is cogently sum-
marized by the 2005 report of the International Council for Science (ICSU—http.iscu.org),
headquartered in Paris, and representing 111 national academies of science and 29 scientific
xi
Environmental Safety of Genetically Engineered Crops, Michigan State University Press, 2011. ProQuest Ebook Central,
xii
| Preface
unions. The ICSU’s report was based on 50 independent scientific enquiries carried out by
different groups in different parts of the world to investigate the food and environmental
safety of GE crops. With respect to food safety, the conclusion was that “currently available
genetically modified foods are safe to eat. This view is shared by several intergovernmental
agencies, including the FAO/WHO Codex Alimentarius Commission on Food Safety, which
has 162 member countries.” However, the report also states that “although currently available
GM foods are considered safe to eat, this does not guarantee that no risks will be encoun-
tered as more foods are developed with novel characteristics. Ongoing evaluation of emerging
agricultural products is required to ensure that new foods coming to market are safe for con-
sumers. Food safety evaluation must be undertaken on a case-by-case basis.”
With respect to environmental safety, the appropriateness of specific technologies depends
on the current agricultural system and practices, and the surrounding natural environment.
The ICSU report concluded: “The effect of genetic technologies may be either positive or
negative. They may either accelerate the environmentally damaging effects of agriculture, or
they may contribute to more sustainable agricultural practices and the conservation of natural
resources. It is a matter of application and choice.” Thus, it is not GE technologies per se,
but, like virtually any technology, the manner in which they are employed that can be safe
or unsafe. Scientific assessments must be coupled with the governmental and institutional
frameworks and capacity to interpret and make nationally and locally appropriate choices of
technologies.
It is from this perspective—that the safety of genetically engineered crops is a matter of
application and choice—that this book is written. We describe the basis for the develop-
ment and use of genetically engineered crops and examine the possible impacts that such
technologies may have on the environment. To make these determinations, specific genes
and traits must be evaluated within the context of the environment into which they will be
introduced, the manner in which they will be used, and the current agricultural system and
attendant environmental impacts of the technology. Our intent is to provide the reader with a
framework by which to address these issues within the context of the varying geographies and
circumstances in which the adoption of GE crop technology is being considered.
Copyright © 2011. Michigan State University Press. All rights reserved.
Environmental Safety of Genetically Engineered Crops, Michigan State University Press, 2011. ProQuest Ebook Central,
Part 1
Introduction to Environmental
Biosafety in Relation to Genetically
Engineered Crops
Copyright © 2011. Michigan State University Press. All rights reserved.
Environmental Safety of Genetically Engineered Crops, Michigan State University Press, 2011. ProQuest Ebook Central,
Copyright © 2011. Michigan State University Press. All rights reserved.
Environmental Safety of Genetically Engineered Crops, Michigan State University Press, 2011. ProQuest Ebook Central,
Production of Genetically
Engineered Crops, Relationship to
Conventional Plant Breeding, and
Implications for Safety Assessment
Rebecca Grumet
Environmental Safety of Genetically Engineered Crops, Michigan State University Press, 2011. ProQuest Ebook Central,
4
| Rebecca Grumet
it is not surprising that the genome (total collection of genes) of a typical plant comprises
20,000–40,000 genes (Chrispeels and Sadava 2003). It has been analogized that if all of the
information contained in the genes of a rice plant (estimated 37,500 genes) were to be written
out as ATGCs, it would fill 40 books, each with 1,000 pages (Lemaux 2006). Any typical gene
would take up less than one page.
Importantly, the language of DNA (the ability to encode the amino acid sequence of
proteins) is the same regardless of the type of organism it came from. This is a critical fac-
tor underlying genetic engineering techniques, which makes it possible to move genes from
one organism to another. Genetic engineering depends on two types of technologies primar-
ily developed in the 1970s and 1980s, recombinant DNA technology and transformation
technologies. Recombinant DNA technology refers to a collection of techniques used to take
genes from different places and put them together in new combinations (i.e., recombinant).
Transformation refers to moving the recombined gene into a new organism. The resultant
organism is referred to as transformed, genetically modified, genetically engineered, or transgenic.
The process of producing a GE crop can be broadly divided into five general steps: obtain
and engineer the desired gene (recombinant DNA technology); introduce the gene into indi-
vidual cells/chromosomes (transformation); regenerate the transformed cell into a whole plant
(tissue culture technology); verify the presence and expression of the introduced gene and
desired new trait (lab, greenhouse, and field studies); and incorporate the new trait into a
high-performing variety (conventional breeding) (figure 1). Each of these steps will be dis-
cussed briefly to give a general overview of the process. More complete descriptions of the
relevant technologies can be found in several texts and reviews, such as Chrispeels and Sadava
(2003); Sharma et al. (2005); Thomson (2006); Watson et al. (1992).
The length of time and sophistication of technologies needed to obtain the gene of inter-
est varies widely, depending on the state of background knowledge. If the gene of interest is
closely related to one that has already been cloned from another organism, or if the protein the
gene codes for has already been isolated and characterized, obtaining the gene can be relatively
easy, and relies primarily on standard recombinant DNA techniques. These include cloning
(making copies of the gene of interest so it can be worked with in the laboratory), and cutting
and pasting to precisely assemble the intended pieces of DNA (Watson et al. 1992). If the
gene has never been cloned before, and little or nothing is known about the protein it encodes,
initial cloning can be more difficult, and may require more sophisticated genomic technolo-
gies developed in the last two decades that can examine thousands of genes at the same time.
Desirable genes about which little is known might include a gene coding for resistance to a
specific disease, or one coding for enzymes to produce an herbal medicinal compound.
Once the gene of interest is in hand, modifications may be needed to engineer the gene
Environmental Safety of Genetically Engineered Crops, Michigan State University Press, 2011. ProQuest Ebook Central,
Production, Breeding, and Safety
| 5
Figure 1. Steps to Produce a GE Crop: (1) obtain and engineer the desired gene (recombinant DNA technology); (2) introduce
the gene into individual cells/chromosomes (transformation); (3) regenerate the transformed cell into a whole plant (tissue
culture technology); (4) verify the presence and expression of the introduced gene and desired new trait (lab, greenhouse
and field studies); (5) and incorporate the new trait into a high performing variety (conventional breeding).
Copyright © 2011. Michigan State University Press. All rights reserved.
into a form for expression in the new plant. Generally the most important modification is
adding an appropriate promoter, a piece of that tells the cell when, where, and how
much of the gene should be expressed. or example, certain genes may need to be expressed
DNA
everywhere, others only in the seeds, others only in response to drought.
F
Step 2: ransfer the Engineered Gene into Cells of the Desired Plant
T
ntroducing the gene into cells (and ultimately chromosomes) of the desired plant requires
transformation technology. he two most widely used methods are Agrobacterium-mediated
I
transformation and particle bombardment ( hrispeels and Sadava 2003; Sharma et al. 2005).
T
C
Environmental Safety of Genetically Engineered Crops, Michigan State University Press, 2011. ProQuest Ebook Central,
6
| Rebecca Grumet
Step 3: Regenerate Cells with the New Gene into the Whole Plant
In the first stage of transformation, the gene of interest becomes incorporated into a single cell.
It is then necessary to culture the cell to become a whole plant. This process, called regeneration,
is accomplished by placing the treated plant tissue onto a culture medium that allows the cells to
divide and differentiate to form plant parts. The necessary media conditions must be developed
for each species of interest. The regenerated plant is ultimately transferred to soil and used to
produce progeny. Tissue culture techniques are also used for a variety of other purposes, such as
propagating species that are not typically grown from seeds (i.e., clonally propagated species), or
to produce disease-free sources of planting materials (Lindsey and Jones 1990).
In any transformation process, only a tiny fraction of a percent of the treated cells acquire
the gene of interest. One of the challenges is to be able to separate the few transformed cells
from the thousands of nontransformed cells. This is most often accomplished by introduc-
ing two genes, the desired gene (e.g., for insect resistance), and a second gene that serves as a
marker to find the transformed cells (selectable marker gene) (Sharma et al. 2005). The select-
able marker allows the transformed cells to grow in conditions that inhibit nontransformed
cells, such as in the presence of antibiotic or herbicide resistance. In this way, only the cells
that have become transformed are able to grow and divide and ultimately regenerate.
Copyright © 2011. Michigan State University Press. All rights reserved.
The first step after regenerating a putatively transgenic plant is to verify presence and expres-
sion of the gene. Presence of the gene is determined using DNA detection techniques such as
PCR or Southern analysis. Stable inheritance is verified by transfer to subsequent generations.
Expression of the gene can be verified by testing for transcription to RNA or production of
the desired protein. Once it is determined that the gene is present, stably inherited, and prop-
erly expressed, the plants can be tested for expression of the desired trait, (e.g., virus resistance,
salt stress resistance) in greenhouse and confined field trials. At this stage, if the desired trait
has been obtained, proof of concept has been established, demonstrating that the introduced
gene can be potentially useful.
Environmental Safety of Genetically Engineered Crops, Michigan State University Press, 2011. ProQuest Ebook Central,
Production, Breeding, and Safety
| 7
For many, if not most crops, the plant varieties that perform well in tissue culture for trans-
formation and regeneration are not the best varieties for performance and productivity in the
field. Furthermore, even if a high-performing cultivar can be successfully transformed, no one
variety can be used everywhere. Varieties must be bred for the specific conditions in which
they will be grown with respect to temperatures, availability of water, length of growing sea-
son, day length, prevalent diseases, and so forth. As an example, seed company catalogs offer
hundreds of “Roundup Ready” (glyphosate resistant) soybean varieties for sale in the United
States. For genes/traits being introduced into new locations, the gene will need to be trans-
ferred into locally adapted varieties. Varieties also vary depending on the use of the crop, for
example, fresh market versus processing, food versus feed, specific processing characteristics
(e.g., oil or starch qualities), again necessitating incorporation of the gene of interest into the
types appropriate for the intended use.
Controlled crosses (pollinations) are made between plants of the genetically engineered
variety and the desired recipient variety, and backcrossed as needed to obtain the appropri-
ate combination of traits. The resultant progeny are tested in small-plot field trials to verify
performance. The most promising lines are moved to more extensive multiyear, multilocation
trials to be sure the crop and the new trait perform well over a range of conditions that may
be encountered by the new variety.
in the area, thin seed coats, and loss of the inner fibrous pod layer which in the wild expels
beans from the pod (Kaplan et al. 1973). These modifications demonstrate that prehistoric
plant breeders were able to make impressive gains in productivity and agricultural and culi-
nary quality.
Selection
The primary tool available to the earliest plant breeders was selection, that is, choosing the
best types for further propagation. Selection can occur both intentionally and inadvertently.
Examples of intentional selection include selection for increased size, better flavor, or ease of
cooking. Inadvertent selection plays a predominant role in local adaptation. Humans move
crops from one location to another. In this process, the environmental conditions change, and
Environmental Safety of Genetically Engineered Crops, Michigan State University Press, 2011. ProQuest Ebook Central,
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