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Discover the latest edition of this authoritative textbook on plant biotechnology and genetics
Plant biotechnology is a field of research and development in which scientific techniques are brought to bear on the creation and modification of new, beneficial plants and strains. Biotechnological techniques can be used to add nutritive value, increase resistance to diseases and pests, increase yields, and more. The production of biotech crops has increased over one hundred times since their introduction into commercial agriculture in 1996, making them the most rapidly-adopted crop category in the history of modern agriculture.
Plant Biotechnology and Genetics is the essential introduction to this thriving research subject. Beginning with an overview of basic plant biology and genetics, it then moves to the fundamental elements of biotechnology. Now fully updated to reflect the latest research advances and technological breakthroughs, it continues to be a must-own for readers interested in the future of food production and more.
Readers of the third edition of Plant Biotechnology and Genetics will also find:
Plant Biotechnology and Genetics is ideal for advanced undergraduate and masters students in plant biotechnology courses, as well as professionals seeking a helpful reference guide.
C. Neal Stewart Jr., PhD, holds the Racheff Chair of Excellence in Plant Molecular Genetics and is a Professor in the Department of Plant Sciences at the University of Tennessee, Knoxville. He also serves as co-director of the Center for Agricultural Synthetic Biology, which Stewart co-founded in 2018. In addition to the prior editions of Plant Biotechnology, he has written Weedy and Invasive Plant Genomics, Plant Transformation Technologies, and Research Ethics for Scientists: A Companion for Students, all published by Wiley.
List of Contributors xv
Preface xvii
1. The Impact of Biotechnology on Plant Agriculture 1Graham Brookes
1.0. Chapter Summary and Objectives 1
1.1. Introduction 1
1.2. Cultivation of Biotechnology (GM) Crops 2
1.3. Why Farmers Use Biotech Crops 4
1.4. GM Crop Trait Use on Production and Farming 6
1.5. How the Adoption of Plant Biotechnology has Impacted the Environment 7
1.6. Conclusions 13
References 15
2. Mendelian Genetics and Plant Reproduction 17Matthew D. Halfhill and Suzanne I. Warwick
2.0. Chapter Summary and Objectives 17
2.1. Genetics Overview 17
2.2. Mendelian Genetics 20
2.3. Mitosis and Meiosis 24
2.4. Plant Reproductive Biology 28
2.5. Conclusion 34
Life Box 2.1. Richard A. Dixon 35
Life Box 2.2. Michael L. Arnold 36
References 38
3. Plant Breeding 39Nicholas A. Tinker and Elroy R. Cober
3.0. Chapter Summary and Objectives 39
3.1. Introduction 40
3.2. Central Concepts in Plant Breeding 41
3.3. Objectives in Plant Breeding 52
3.4. Methods of Plant Breeding 53
3.5. Breeding Enhancements 64
3.6. Conclusions 71
Life Box 3.1. Gurdev Singh Khush 71
Life Box 3.2. P. Stephen Baenziger 73
References 75
4. Plant Development and Physiology 76Glenda E. Gillaspy and Catherine P. Freed
4.0. Chapter Summary and Objectives 76
4.1. Plant Anatomy and Morphology 77
4.2. Embryogenesis and Seed Germination 78
4.3. Meristems 84
4.4. Leaf Development 87
4.5. Flower Development 90
4.6. Hormone Physiology and Signal Transduction 93
4.7. Conclusions 99
Life Box 4.1. Natasha Raikhel 99
Life Box 4.2. Brenda S.J. Winkel 101
References 103
5. Tissue Culture: The Manipulation of Plant Development 105Vinitha Cardoza
5.0. Chapter Summary and Objectives 105
5.1. Introduction 105
5.2. History of Tissue Culture 106
5.3. Media and Culture Conditions 107
5.4. Sterile Technique 109
5.5. Culture Conditions and Vessels 111
5.6. Culture Types and Their Uses 113
5.7. Regeneration Methods of Plants in Culture 119
5.8. Rooting of Shoots 122
5.9. Acclimation 122
5.10. Automation in Plant Tissue Culture 123
5.11. Artificial Intelligence (AI) and Machine Learning in Plant Tissue Culture 123
5.12. Problems That Can Occur in Tissue Culture 123
5.13. Conclusions 124
Acknowledgments 124
Life Box 5.1. Vinitha Cardoza 125
Life Box 5.2. Raymond D. Shillito 126
References 129
6. Molecular Genetics of Gene Expression 133Maria Gallo and Alison K. Flynn
6.0. Chapter Summary and Objectives 133
6.1. The Gene 134
6.2. DNA Packaging into Eukaryotic Chromosomes 134
6.3. Transcription 135
6.4. Translation 144
6.5. Protein Postranslational Modification 150
Life Box 6.1. Maarten Chrispeels 150
Life Box 6.2. Hong S. Moon 152
References 153
7. Plant Systems Biology 155Wusheng Liu, Yongil Yang, and C. Neal Stewart, Jr.
7.0. Chapter Summary and Objectives 155
7.1. Introduction 156
7.2. Defining Plant Systems Biology 157
7.3. Properties of Plant Systems 158
7.4. A Framework of Plant Systems Biology 160
7.5. Disciplines and Enabling tools of Plant Systems Biology 162
7.6. Conclusions 179
Life Box 7.1. C. Robin Buell 180
Life Box 7.2. Joshua Yuan 182
References 183
8. Recombinant DNA, Vector Design, and Construction 185Stephen L. Gasior, David G.J. Mann, and Mark D. Curtis
8.0. Chapter Summary and Objectives 185
8.1. Plasmids are Unique Genetic Elements in Nature 186
8.2. DNA Vectors 189
8.3. Recombinant DNA Methods 195
8.4. Vector Design in Plant Research and Trait Development 206
8.5. Vectors for Targeted Genome Manipulations 213
8.6. Prospects 216
Life Box 8.1. David Mann 216
References 218
9. Genes and Traits of Interest 224Joanna H. Kud and Kenneth L. Korth
9.0. Chapter Summary and Objectives 224
9.1. Introduction 225
9.2. Identifying Genes of Interest Via Omics Technologies 225
9.3. Traits for Improved Crop Production Using Transgenics 228
9.4. Conclusion 245
Life Box 9.1. Tony Shelton 246
References 247
10. Promoters and Marker Genes 249Wusheng Liu, Debao Huang, C. Neal Stewart, Jr., and Brian Miki
10.0. Chapter Summary and Objectives 249
10.1. Introduction 250
10.2. Promoters 250
10.3. Marker Genes 259
10.4. Marker- Free Strategies 270
10.5. Conclusions 272
Life Box 10.1. Wusheng Liu 274
Life Box 10.2. Yunde Zhao 275
References 277
11. Transgenic Plant Production 282John J. Finer and Ning Zhang
11.0. Chapter Summary and Objectives 282
11.1. Overview of Plant Transformation 283
11.2. Agrobacterium Tumefaciens 286
11.3. Particle Bombardment 293
11.4. Other Methods of Transformation 297
11.5. The Rush to Publish 300
11.6. A Look to the Future 306
Life Box 11.1. John Finer 306
Life Box 11.2. Kan Wang 308
Life Box 11.3. Ted Klein 310
References 312
12. Analysis of Transgenic Plants 315C. Neal Stewart, Jr.
12.0. Chapter Summary and Objectives 315
12.1. Essential Elements of Transgenic Plant Analysis 316
12.2. Assays for Transgenicity, Insert Copy Number, and Segregation 317
12.3. Transgene Expression 323
12.4. Knockdown or Knockout Analysis Rather Than Overexpression Analysis 326
12.5. The Relationship Between Molecular Analyses and Phenotype 327
Life Box 12.1. Neal Stewart 327
Life Box 12.2. Nancy A. Reichert 329
References 331
13. Plastid Genetic Engineering 332Alessandro Occhialini and Scott C. Lenaghan
13.0. Chapter Summary and Objectives 332
13.1. Introduction 333
13.2. Plastid Biology and Molecular Genetics 334
13.3. Plastid Genetic Engineering History and Motivations 336
13.4. Plastome Engineering Versus Nuclear Genome Engineering 336
13.5. Key Components for Plastome Engineering of Plants 338
13.6. Plastome Transformation Vector Design 340
13.7. Beyond Transplastomics: The Use of Episomal Vectors for Minisynplastome and Minichromosome Approaches 347
13.8. Removing DNA from Plastids 349
13.9. The Future of Plastid Engineering 350
Life Box 13.1. Henry Daniell 351
Life Box 13.2. Pal Maliga 354
Life Box 13.3. Alessandro Occhialini 356
Life Box 13.4. Ralph Bock 357
References 358
14. CRISPR- Cas: Genome Editing from Small- Scale to High Throughput for Plant Biology and Biotechnology 366S.P. Avinash, Mirza J. Baig, and Kutubuddin A. Molla
14.0. Chapter Summary and Objectives 366
14.1. Introduction 367
14.2. Diverse CRISPR Approaches and Tools for Precisely Editing Genomes 371
14.3. Changing Gene Expression by CRISPR 374
14.4. CRISPR Screening for Large- Scale Functional Genomics 376
14.5. CRISPR- Enabled Crop Improvement 377
14.6. Commercialized Genome- Edited Crops 380
14.7. Conclusions 380
Life Box 14.1. Kutubuddin Molla 381
Life Box 14.2. Dan Voytas 382
Life Box 14.3. Yiping Qi 383
References 385
15. Regulations and Biosafety 390Alan Mchughen and Stuart J. Smyth
15.0. Chapter Summary and Objectives 390
15.1. Introduction 390
15.2. History of Genetic Engineering and its Regulation 392
15.3. Regulation of GE Plants 394
15.4. Regulatory Flaws and Invalid Assumptions 402
15.5. The State of Genome Editing Regulation 406
15.6. Conclusion 409
Life Box 14.1. Alan Mchughen 411
References 412
16. Field Testing of Transgenic Plants: Risk Assessment and Performance 415Detlef Bartsch, Achim Gathmann, Arti Sinha, and Christiane Saeglitz
16.0. Chapter Summary and Objectives 415
16.1. Introduction 416
16.2. Environmental Risk Assessment Process 416
16.3. An Example Risk Assessment: The Case of Bt Maize 418
16.4. Proof of Safety Versus Proof of Hazard 422
16.5. Modeling the Risk Effects on a Greater Scale 422
16.6. Proof of Benefits: Agronomic Performance 423
16.7. Conclusions 424
Life Box 18.1. Detlef Bartsch 426
References 427
17. Intellectual Property in Agricultural Biotechnology: Strategies for Open Access 429Gregory Graff, David Jefferson, Monica Alandete-Saez, Cecilia Chi-Ham, Sara Boettiger, and Alan B. Bennett
17.0. Chapter Summary and Objectives 429
17.1. Intellectual Property and Agricultural Biotechnology 430
17.2. The Relationship Between Intellectual Property and Agricultural Research 433
17.3. Patenting Plant Biotechnology: The Anti- Commons Problem 434
17.4. What Is Freedom to Operate? 438
17.5. Strategies for Open Access 441
17.6. Conclusions 443
Life Box 17.1. Alan Bennett 444
Life Box 17.2. Maud Hinchee 445
References 446
18. Why Transgenic Plants Are So Controversial 451Jennifer Trumbo and Douglas Powell
18.0. Chapter Summary and Objectives 451
18.1. Introduction 452
18.2. Perceptions of Risk 454
18.3. Responses of Fear 456
18.4. Feeding Fear: Case Studies 457
18.5. How Many Benefits Are Enough? 459
18.6. Continuing Debates 460
18.7. Business and Control 462
18.8. Conclusions 462
Life Box 18.1. Wayne Parrott 464
References 465
19. Plant Synthetic Biology 467Scott C. Lenaghan
19.0. Chapter Summary and Objectives 467
19.1. What is Synthetic Biology? 467
19.2. Design: Plant Synthetic Biology 470
19.3. Build: Components of Plant Synthetic Biology 474
19.4. Test: Components of Plant Synthetic Biology 485
19.5. Conclusion 485
Life Box 19.1. Nicola J. Patron 486
Life Box 19.2. Scott C. Lenaghan 487
References 488
Index 491
GRAHAM BROOKES
PG Economics Ltd, Dorchester, UK
Since the first stably transgenic plant produced in the early 1980s and the first commercialized transgenic plant in 1994, biotechnology has revolutionized plant agriculture. In the United States, more than 90% of the maize (corn), soybean, cotton, and canola crops are transgenic for insect resistance, herbicide resistance, or both. Plant biotechnology has been one of the most rapidly adopted technologies in the history of agriculture and continues to expand in much of the developed and developing world.
The technology of genetic modification (GM, also stands for "genetically modified"), which consists of genetic engineering and also known as genetic transformation, has now been utilized globally on a widespread commercial basis for more than a quarter of century; and in 2020, over 18 million farmers in 26 countries had planted 185.6 million hectares of crops using this technology. These milestones provide an opportunity to critically assess the impact of this technology on global agriculture. This chapter therefore examines specific global socioeconomic impacts on farm income and environmental impacts with respect to pesticide usage and greenhouse gas (GHG) emissions of the technology. Further details can be found in Brookes (2022a, b and c).
Although the first commercial GM crops were planted in 1994 (tomatoes), 1996 was the first year in which a significant area of crops containing GM traits were planted (1.66 million hectares). Since then, there has been a dramatic increase in plantings, and in 2020, the area planted to crops utilizing GM seed technology was 185.6 million hectares.
Almost all of the global GM crop area derives from soybean, maize (corn), cotton, and canola (Fig. 1.1). In 2020, GM soybean accounted for the largest share (50%) of total GM crop cultivation, followed by maize (33%), cotton (12%), and canola (5%). In terms of the share of total global plantings to these four crops accounted for by GM crops, GM traits accounted for a majority of soybean grown (72%) in 2020 (i.e., non-GM soybean accounted for 28% of global soybean acreage in 2020). For the other three main crops, the GM shares in 2020 of total crop production were 31% for maize, 76% for cotton, and 27% for canola (i.e., the majority of global plantings of maize and canola continued to be non-GM in 2020). The trend in plantings of GM crops (by crop) from 1996 to 2020 is shown in Figure 1.2. In terms of the type of biotechnology trait planted, Figure 1.3 shows that GM herbicide-tolerant soybeans dominate, accounting for 33.6% of the total, followed by herbicide-tolerant and insect-resistant (largely Bt) maize, herbicide-tolerant maize, insect-resistant soybeans (also containing herbicide-tolerant technology), and insect-resistant cotton with respective shares of 21, 20.9, 10.8, and 8.3%. It is worth noting that the total number of plantings by trait produces a higher global planted area (274.3 million hectares) than the global area by crop (185.6 million hectares) because of the planting of crops containing the stacked traits of herbicide tolerance and insect resistance (e.g., a single plant with two biotech traits). In total, 46% of global GM crop plantings in 2020 contained stacked traits, 44% contained only herbicide-tolerant (GM HT) traits, and 10% only insect-resistant (GM IR) traits. Finally, looking at where biotech crops have been grown, the United States had the largest share of global GM crop plantings in 2020 (36%: 66.8 million hectares), followed by Brazil (55.7 million hectares: 30% of the global total) and Argentina (12%: 22.3 million hectares). The other main countries planting GM crops in 2020 were India, Canada, and China (Fig. 1.4). In 2020, there were also additional GM crop plantings of papaya (187 hectares), squash (1000 hectares), alfalfa (1.26 million hectares), sugar beet (462,300 hectares), and potatoes (1780 hectares) in the United States, of papaya (9000 hectares) in China, of sugar beet (17,000 hectares) in Canada and 6309 hectares of insect resistant brinjal in Bangladesh.
Figure 1.1. Global GM crop plantings in 2020 by crop (base area: 185.6 million hectare).
(Source: ISAAA, Canola Council of Canada, CropLife Canada, USDA, CSIRO, ArgenBio.)
Figure 1.2. Global GM crop plantings by crop 1996-2020.
Figure 1.3. Global GM crop plantings by main trait and crop: 2020.
(Source: Various, including ISAAA, Canola Council of Canada, CropLife Canada, USDA, CSIRO, ArgenBio.)
Figure 1.4. Global GM crop plantings 2020 by country.
(Sources: ISAAA, Canola Council of Canada, CropLife Canada, USDA, CSIRO, ArgenBio, Various including ISAAA, Canola Council of Canada, CropLife Canada, USDA, CSIRO, ArgenBio, National Ministries of Agriculture (Mexico, Philippines, Spain, Vietnam), Grains South Africa.)
The primary driver of adoption among farmers (both large commercial and small-scale subsistence) has been the positive impact on farm income. GM technology has had a significant positive impact on farm income derived from a combination of enhanced productivity and efficiency gains (Figs. 1.5 and 1.6). In 2020, the direct global farm income benefit from GM crops was $18.8 billion. This is equivalent to having added 5.9% to the value of global production of the four main crops of soybeans, maize, canola, and cotton. Since 1996, farm incomes have increased by $261.3 billion.
Figure 1.5. Global GM crop farm income benefits 2020: baseline total $18.8 billion. Notes: Others = HT sugar beet, Virus resistant papaya and squash, drought tolerant maize and IR brinjal.
Figure 1.6. Cumulative global GM crop farm income benefits 1996-2020: baseline total $261.3 billion. Notes: Others = HT sugar beet, Virus resistant papaya and squash, drought tolerant maize and IR brinjal.
The largest gains in farm income in 2020 have arisen in the maize sector, largely from yield gains. The $3.7 billion additional income generated by GM insect resistant (GM IR) maize in 2020 has been equivalent to adding 6.3% to the value of the crop in the GM crop-growing countries, or adding, the equivalent of 2.8% to the $133 billion value of the global maize crop in 2020. Cumulatively since 1996, GM IR technology has added $67.8 billion to the income of global maize farmers.
Substantial gains have also arisen in the cotton sector through a combination of higher yields and lower costs. In 2020, cotton farm income levels in the GM adopting countries increased by $3.94 billion, and since 1996, the sector has benefited from an additional $73.11 billion. The 2020 income gains are equivalent to adding 12.1% to the value of the cotton crop in these countries or 12% to the $32.7 billion value of total global cotton production. This is a substantial increase in value added terms for two categories of cotton seed technology.
Significant increases to farm incomes have also resulted in the soybean and canola sectors. The GM herbicide tolerant (HT) technology in soybeans has boosted farm incomes by $5.64 billion in 2020, and since 1996 has delivered $74.65 billion of extra farm income. The adoption of "Intacta" soybeans (combining HT and IR traits) in South America since 2013 also provided $16 billion of additional farm income. In the canola sector (largely North American) an additional $8.2 billion has been generated (1996-2020).
Figures 1.7 and 1.8 summarize farm income impacts in key GM crop adopting countries. These highlight the important farm income benefit arising from GM HT soybeans in South America (Argentina, Bolivia, Brazil, Paraguay, and Uruguay), GM IR cotton in China and India, and the range GM crop adoption in the US. Figure 1.8 in particular (the increasing share of "other countries") also illustrates the growing level of farm income benefits being obtained in countries that were later adopters of GM crop technology such as Pakistan, the Philippines, and Colombia.
In terms of the division of the economic benefits obtained by farmers in developing countries relative to farmers in developed countries, in 2020, 55% of the farm income benefits have been earned by developing country farmers. The vast majority of these income gains for developing country farmers have been from GM IR cotton and GM HT soybeans. Over the 25 years, 1996-2020, the cumulative farm income gain derived by developing country farmers was 52% ($136.6 billion).
Figure 1.7. Cumulative global GM crop farm income benefits 1996-2020 by country: baseline total $261.3 billion
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