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Plant Biotechnology and Agriculture Prospects for the 21st
Century
E d i t e d b y
Arie Altman Robert H. Smith Institute of Plant Sciences and
Genetics in Agriculture Hebrew University of Jerusalem Rehovot,
Israel
Paul Michael Hasegawa Bruno C. Moser Distinguished Professor
Horticulture and Landscape Architecture Department Purdue
University West Lafayette, Indiana, USA
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Contents
Contributors xxi
Foreword xxv Roger N. Beachy
Preface xxvii Me Altman and Paul Michael Hasegawa
Introduction to plant biotechnology 2011: Basic aspects and
agricultural implications xxix Arie Altman and Paul Michael
Hasegawa
Section A Introduction to basic procedures in plant
biotechnology 1 1 Genetics and genomics of crop domestication 3
J.S. (Pat) Heslop-Harrison, Trude Schwarzacher Plants and
Domestication 3
Scope 3 Domesticated crops 3 Weeds 4 Invasive species 4 Model
species and crop sciences 5
Understanding Domestication Processes 5 Evidence of relatives
and processes of early domestication 5 Genes of domestication 6
Genetic variation and domestication 6 Genetic control related to
diversity and speciation 6 Domestication of maize 7 Domestication
of legumes 7 Yield traits 8
Hybrid Species and New Polyploids in Domestication 8
Post-Domestication Selection 8
Modifications in crop characteristics 8 New Domestication 9
Domesticated species 9 Lost crops 9 Trees and biofuels 9
Genetics and breeding for new uses: Ecosystem services 10
Features of Domesticated Genomes 11 Superdomestication 14
Acknowledgments 16
2 The scope of things to come: New paradigms in biotechnology 19
Maheshi Dassanayake, Dong-Ha Oh, Dae-Jin Yun, Ray A. Bressan, John
M. Cheeseman, J. Hans Bohnert Introduction 19
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Progress Enabled by Next-Generation DNA Sequencing 20 Mapping of
comprehensive, genome-wide, treatment-specific transcript profiles
23 Current next-gen sequencing 23 Behold the third generation
23
The Elephant in the Laboratory: Data Handling 24 From Sequences
to Comparative Genomics 24
Transcriptome profiling 25 Broadening the Genomics Toolbox:
Proteins and Metabolites 26
Proteomics advances 26 Metabolomics highlights 26
Genomics Unlimited: Getting Beyond Mere Genes 27 Into the
Future: Genomics-Based Biotechnology and Agriculture 28
From models to crops, from labs to fields 28 Genetic resources
from extremophile species 29 Exploring "unknown unknowns" 29 The
importance of stress "tolerance" engineering 29
Acknowledgments 30
3 Protein targeting: strategic planning for optimizing protein
products through plant biotechnology 35 Elizabeth Hood, Carole
Cramer, Giuliana Medrano, Jianfeng Xu Introduction: Strategic
Decisions about How to Express an Output Trait 35 Approaches 37
Routing proteins to the endomembrane system 37 Accumulating
proteins in the ER 38 Accumulating proteins in ER-derived protein
bodies 39 Accumulating proteins in the vacuole or vacuolar protein
bodies 39 Accumulating proteins in the apoplast 40 Accumulating
proteins in the chloroplast 40 Accumulating proteins on the surface
of oil bodies 41
Seed-Based Expression Systems 41 Leaf Systems 44
Stable versus transient leaf expression systems 44 Protein
bodies in leaves 47
Hairy Root Cultures 47 Advantages of the hairy root culture
system 48 Recombinant proteins expressed with hairy root cultures
48 Hairy root cultures in bioreactors and scale-up 48
Summary and Conclusions 50
4 Proteomics and its application in plant biotechnology 55
Sylvain Bischof, Jonas Grossmann, Wilhelm Gruissem Introduction 55
Mass Spectrometry-Based Proteomics 56
Sample preparation prior to mass spectrometry 56 Mass
spectrometry 57
- Spectra assignment for peptide and protein identification 58
Quantitative proteomics 58 Post-translational modifications 58
Proteomics in Plant Biotechnology 59 What has been achieved so
far in crop proteomics? 59 Arabidopsis thaliana as plant model
organism 59 Crops and other economically relevant plant species 60
Future applications and perspectives 61
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5 Plant metabolomics: Applications and opportunities for
agricultural biotechnology 67 Diane M. Beckles, Ute Roessner
Introduction 67 Metabolite Networks: The Basics 68 Metabolomics:
Technologies for Analyses 69
Analytical platforms 70 Data analysis and interpretation 71
Metabolomics: Applications in Agricultural Biotechnology 73
Metabolite profiling to test substantial equivalence 73
Phytochemical diversity, phenotyping, and classification.... 74
Postharvest quality of horticultural crops 74 Stress responses 74
Functional genomics 75 Breeding and metabolite quantitative trait
loci 75
Metabolomics: Challenges and Future Perspectives 76 From model
organisms to crop plants 76 Compartmentation of plant metabolism 76
High-resolution sampling 76 Primary and secondary metabolism pose
different challenges 76 Identifying the metabolome 77 Measurements
of metabolic flux 77
Outlook 78 Acknowledgments 78
6 Plant genome sequencing: Models for developing synteny maps
and association mapping.... 83 Delphine Fleury, Ute Baumann, Peter
Langridge Introduction 83 Genome Sequencing 84
Strategies for plant genome sequencing 84 High-throughput
sequencing technologies 86 Single molecule and real-time sequencing
86 Assembly and alignment programs 86 Genome browsers 87
Models for Developing Syntenic Maps 88 Definitions 88
Intraspecies comparison 88 Cytogenetics for interspecies comparison
89 Sequence comparison 89 Macro- versus micro-synteny 89 Nature of
the differences 89 Applications of syntenic maps 91 Tools and
limitations 91
Association Mapping 91 Definitions 91 Population size and
structure 92 Markers and marker density .( 93
Implications 94
7 Agrobacterium-rr\ed\a\ed, plant genetic transformation 99 Yoel
Shiboleth, Tzvi Tzfira Introduction 99 The Genetic Transformation
Process 99 Agrobacterium as a Tool for Plant Transformation 104
Novel and Specialized Vectors for Plant Transformation 106
Manipulating the Plant Genome to Improve and Control Transformation
108
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Using Novel Selection Methods and Restriction Enzymes to Control
T-DNA Integration 109 Conclusions and Future Prospects 110
Acknowledgments 111
8 Biolistic and other non-Agrobacterium technologies of plant
transformation 117 Trade K. Matsumoto, Dennis Gonsalves
Introduction 117 Other Non-Agrobacterium Transformation 117
Electrophoretic transfection 117 Electroporation 118
Bioactive-beads-mediated gene transfer 118 Microinjection 118
Pollen-tube pathway 119 Silica carbide whisker-mediated
transformation 119
Biolistic Transformation 120 The invention 120 Electric
discharge particle acceleration 120 Current status of the
"invention" hardware 121
Advantages of Biolistic Transformation 121 Implications of
Biolistics in Agricultural Biotechnology 122
Application of biolistics in agriculture crops 122 Papaya: A
case study of biolistic transformation 122
9 Plant tissue culture for biotechnology 131 Prakash P. Kumar,
Chiang Shiong Loh Introduction 131 Plant Tissue Culture Technology
131
The basic laboratory setup 131 Preparation of tissue for
culturing 132 Nutrient media 132 Types of culture 133 Environmental
aspects of tissue culture 133 Modes of regeneration 134
Implications for Agricultural Biotechnology 134 Haploid tissue
culture 135 Somatic embryogenesis 135 Artificial seeds 135 In vitro
flowering 136
Future Perspectives 136 Acknowledgments 136
Section B Breeding biotechnologies 139 10 Somatic (asexual)
procedures (haploids, protoplasts, cell selection) and their
applications 141 Tanya Tapingkae, Zul Zulkarnain, Masayo
Kawaguchi, Takashi Ikeda, Acram Taji General Introduction 141
Somatic Embryogenesis 141
Introduction 141 Patterns of somatic embryogenesis 142 Factors
affecting somatic embryo induction 142 Plant maturation 143 Plant
regeneration 144
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Gene expression during somatic embryogenesis 144 Mass
propagation and somaclonal variation 144
Haploid Technology 144 Introduction 144 Cytological basis
underlying haploids plants induction 145 Factors affecting the
induction of microspore embryos 146 Haploid induction via ovary and
ovule cultures 147
Protoplast and Somatic Hybridization 148 Introduction 148 Types
of somatic hybrids 148 Protoplast fusion methods 148 Selection of
somatic hybrids 150 Identification of somatic hybrids 150 Factors
affecting regeneration of hybrid plants 151
Screening and Development of Stress-Resistant Plants Using in
vitro Selection Techniques 151 Introduction 151 General methods of
screening and breeding using in vitro selection techniques 151
Biotic stress resistance 152 Abiotic stress tolerance 152 Future
perspective of screening and breeding using in vitro selection
techniques 155
Conclusions and Future Directions 155 Acknowledgments 155
11 Marker-assisted selection in plant breeding 163 Giora
Ben-Ari, Uri Lavi Background 163
The concept of marker-assisted selection 163 Historical review
164
Plant Traits, DNA Markers, Technologies, and Applications 164
Genes controlling important traits 164 DNA markers 165 Modern
genotyping technologies 168 Identification of genes controlling
commercially important traits 170 Application of DNA markers to
breeding 173 MAS in breeding programs 174
Discussion 176 Bottlenecks and difficulties in the application
of MAS 176 Future prospects of application of genetic variations to
breeding 177
Acknowledgment 178
12 Male sterility and hybrid seed production 185 Sally Mackenzie
Introduction 185 Male Gametogenesis 185
Pollen mitosis 1 185 Pollen mitosis II 186
Male Sterility Mutants Elucidate Anther Development 187 Hormonal
Influences on Male Reproduction in Plants 187
Gibberellic acid 187 GA regulates jasmonic acid biosynthesis 188
Brassinosteroids 188 Auxins 189
Cytoplasmic Male Sterility Systems in Agriculture 189 Plant
mitochondrial mutations 189 Fertility restoration 189 Stability of
the CMS trait 190
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Contents V
Male Sterility: Metabolic and Evolutionary Implications 190 CMS
is a naturally found condition 190 Organelle metabolism influences
pollen development 190
Genetic Engineering of Male Sterility 191 Implementation of Male
Sterility in Agricultural Systems 191
13 Advances in identifying and exploiting natural genetic
variation 195 Christian S. Hardtke, Kaisa Nieminen Natural Genetic
Variation in Crop Breeding: From Prehistory to the Green Revolution
195 The Genetic Limits of Evolving Domesticated Crops 196
; Tapping the natural genetic variation present in wild
ancestors 196 Natural Genetic Variation in Arabidopsis 197 QTL
Analyses in Arabidopsis 197
Novel Arabidopsis genes isolated through the natural variation
approach 198 What to Expect: Intraspecific Variation in Gene
Structure and Content 198
| Structural genome variation: Higher than expected? 198 QTL
Analysis and Sequence Variation in Crops 199
Domestication genes of maize 199 Examples from rice 199 Examples
from other cereals 200
Toward Prediction of Variation in Molecular Function: Why Model
Organisms are here to Stay 200 Crucial support from model organism
candidate genes 200 Model systems as references to characterize
allele activities 201
Beyond Simple Traits: Epigenetics, Heterosis, Genetic
Incompatibility, and Trade-offs 201 I Incompatibility between
natural accessions 201 \ Trade-offs between different beneficial
traits 202
Extending the Toolbox: Genome-wide Association Mapping 202 The
Route to Effectively Exploit Natural Variation for Plant
Biotechnology 202
14 From epigenetics to epigenomics and their implications in
plant breeding 207 I Athanasios Tsaftaris, Aliki Kapazoglou, Nikos
Darzentas
Mechanisms of Epigenetic Inheritance and their Interactions 207
Introduction 207 Epigenetic mechanisms and their interactions
208
From Epigenetics to Epigenomics 212 Deciphering epigenomes: A
matter of scale and complexity 212 Epigenomic methods and the type
of data collected 212 Epigenomic resources 213 Transposable
elements on the emerging epigenomic landscape(s) 216 An
illustrative and practical example of data and resources
integration 217
Epigenetic Phenomena and their Implications in Plant Breeding
217 Epigenetic controls during vegetative development and the role
of the environment 217 Epigenetic control of flowering 219
Endosperm development and parental imprinting 220
Conclusions and Prospects 222 Acknowledgments 222 Abbreviations
222
Section C Plant germplasm 227 15 An engineering view to
micropropagation and generation of true to type and
pathogen-free plants 229 Eli Khayat Preface 229
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Shoot Multiplication Through Meristem Culture 229 Stage 0:
disinfection and start of axenic culture 230 Stage I: Initiation of
culture 230 Stage II: Multiplication 230 Stage III: Elongation and
promotion of shoots and roots development 231 Stage IV:
Acclimatization and hardening 231
Automation 231 Energy and Lights 232 Photoautotrophic Cultures
232 Micropropagation in Liquid Media 233 Plant—Microbe Interaction
During in vitro and ex vitro Stages of Micropropagation 233
Inoculation with Beneficial Microorganisms 234 Elimination of
Viruses by in vitro Techniques 238 Concluding Remarks 238
Acknowledgments 238
16 Regulation of apomixis 243 Peggy Ozias-Akins, Joann A. Conner
Introduction 243 Overview of Ovule Development During Sexual
Reproduction 244 Overview of Ovule Development During Apomictic
Reproduction 244 Germline Specification 244 Apomeiosis 246
Megagametogenesis 247 Gamete Specification 247 Parthenogenesis 248
Endosperm Development 250 Chromatin Modification and Epigenetic
Regulation 251 Conclusions and Future Prospects for Apomixis in
Crops 251
17 Germplasm collection, storage, and conservation 255 Florent
Engelmann Introduction 255
Strategies for conserving plant biodiversity 255 Ex situ
conservation technologies 256
Applications of Biotechnologies for Conservation 257 In vitro
collecting 257 Slow growth storage 258 Cryopreservation 259
Conclusions 264
Section D Controlling plant response to the environment: Abiotic
and biotic stress 269
18 Integrating genomics and genetics to accelerate development
of drought and salinity tolerant crops 271 Zvi Peleg, Harkamat
Walia, Eduardo Blumwald Impact of Abiotic Stresses on Crop Plant
Productivity 271 Water Deficit: A Major Abiotic Stress Factor 272
Salinity 272 Plant Responses to Abiotic Stress 272 Breeding for
Drought and Salinity Tolerance: "The Conventional Approach" 273
Germplasm resources for drought and salinity tolerance 274
Genetic dissection of plant responses to abiotic stress 274
Introducing new technologies for abiotic stress breeding 275
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Engineering-Tolerant Crop Plants: The Transgenic Approach 275
Genes for osmoregulation 275 Dehydration-responsive element 278 NAC
proteins 279 Genes for ionic balance 279 Genes for redox regulation
279 Other transcription factors 280
Hormones and Abiotic Stress 280 Challenges and Prospects 280
Acknowledgments 281
19 Molecular responses to extreme temperatures 287 Rafael
Cataia, Aurora Diaz, Julio Salinas Introduction I. 287 Plant
Response to Low Temperature 287
Low temperature perception 288 Transducing the low-temperature
signal 289
Cross-talk between Plant Responses to Extreme Temperatures 297
The membrane as a node in the perception of temperature
oscillations 298 Transducing the signals initiated by temperature
variations 298
Conclusions 300 Acknowledgments 301
20 Biotechnological approaches for phytoremediation 309 Om
Parkash Dhankher, Elizabeth A. H. Piion-Smits, Richard B. Meagher,
Sharon Doty Introduction 309
Overview of results from biotechnological approaches for
different pollutants 311 Organic pollutants 317
Future Prospects 323 Acknowledgments 323
21 Biotechnological strategies for engineering plants with
durable resistance to fungal and bacterial pathogens 329 Dor
Salomon, Guido Sessa Introduction 329 Choosing the Target gene for
Transgenic Expression 330
Plant immune receptors mediating pathogen recognition 330
Elicitors of plant immunity 331 Plant genes involved in signaling
networks of plant immunity 332 Antimicrobial genes 334 Genes
targeting pathogen virulence determinants 335
How Many Transgenes Should be Expressed in a Single Plant for
Efficient Disease Control? 335 When and Where Should the
Transgene(s) be Expressed? 336
Pathogen-responsive and tissue-specific promoters 337
Pathogen-responsive elements and synthetic promoters 338
Conclusions and Prospects 339 Acknowledgments 339
22 Controlling plant response to the environment: Viral diseases
343 Munir Mawassi, Abed Gera Introduction 343 Phytosanitation and
Quarantine Regulation 344 Transmission of Plant Viruses 344
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Cultural Strategies of Virus Control 344 Management of
soil-borne viruses 344 Management of airborne viruses 345
Resistance to Virus Transmission by Insects 345 Pathogen-derived
resistance 345 RNA-mediated resistance 346
Application of the PDR Concept for Developing Transgenic Virus
Resistance to Horticultural Crops 346 RNA silencing-based
applications for developing virus resistant plants 347 PDR
stability and suppression of RNA silencing 348
Assessment of Risks Associated with Transgenic Virus Resistance
in Plants 348 Conclusion 349
23 Insects, nematodes, and other pests 353 Philip R. Watkins,
Joseph E. Huesing, Venu Margam, Larry L. Murdock, T.J. V. Higgins
Introduction — Genetically Modified Crops for Insect Resistance
353
History of B. thuringiensis ; 353 Cry proteins 354
Commercially Available Insect Protected Crops 354 Bt maize 354
Bt cotton 356 Discontinued Bt crops 357
Bt Crops Under Development 357 Bt brinjal 357 Bt rice 358 Other
Bt crops 358
Impact of Bt 359 Benefits of Bt crops 359 Concerns about Bt
crops 359 Improving Bt 360
Cowpea Trypsin Inhibitor 360 Novel Insecticidal Protection
361
VIP genes 361 Microorganism-derived toxins 361 Plant-derived
toxins 361 Secondary metabolites 362 Other toxins 363 RNAi 363
Nematode-Resistant Crops 364 Recombinant Insecticides 364
Conclusion 364
Section E Biotechnology for improvement of yield and quality
traits 371
24 Growth control of root architecture 373 Christopher N. Topp,
Philip N. Benfey Introduction to Root System Architecture 373
Genetic and Developmental Aspects of Root Growth 373
Stereotypical organization of root tissues 374 Architectural
possibilities 374 Signaling 375 Systems biology concept of cell
identity 376
Plant-Environment Interactions 376 Environmental sensing and
root exudation 376 Microbial interactions 377 Architectural
responses to nutrient availability 378
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Crop Root Systems 379 Types of root systems 379 Embryonic and
post-embryonic root systems 379 Evolutionary strategies and
trade-offs 380
Approaches to Study Root Architecture 380 Quantitative analysis
380 High-throughput sequencing 381 Phenomics 381
Concluding Remarks 382
25 Control of flowering 387 Alon Samach Introduction 387
A plant's perspective 387 A farmer's perspective 387
Proteins Controlling Flowering Time 388 Florigen and FLOWERING
LOCUS T (FT) 388 Transcription factors regulating FT 389 Proteins
parallel or downstream of FT. 390
Processes Affecting Flowering Time Proteins 391 Histone
modifications 391 Gibberellin 392 MicroRNAs 393 The circadian clock
394 Regulated proteolysis 394 Sugars 394
Developmental Decisions on Timing of Flowering 395 Juvenility
395 Seasonality 395 Reproductive cycles and alternate bearing
397
Summary 398 Acknowledgment 398
26 Fruit development and ripening: A molecular perspective 405
Avtar K. Handa, Martfn-Ernesto Tiznado-Hernandez, Autar K. Mattoo
Fruit Classification 405 Fruit Development 406
Fruit shape, size, and mass 406 Fruit Ripening 409
Ripening mutations 409 Nutritional mutations 411 Shelf life
mutations 412
Ethylene and Fruit Ripening 413 Ethylene biosynthesis 413
Ethylene perception and signal transduction 414 Genetic
intervention in ethylene biosynthesis and perception 416
Fruit Texture 417 Cell wall depolymerizing enzymes 417 Expansins
418 Protein glycosylation 418
Future Perspectives 418
27 Potential application of biotechnology to maintain fresh
produce postharvest quality and reduce losses during storage 425
Amnon Lers Introduction 425
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Contents :
Ethylene Biosynthesis or Perception and Its Relation to
Postharvest Quality of Fresh Produce 426 Senescence in Postharvest
of Leafy Vegetables and Flowers 427
Background 427 Senescence regulatory genes . 427
Senescence-associated hormone biosynthesis or perception 428
Oxidative stress involvement in senescence 429 Chlorophyll
degradation 429
Abscission of Fruits, Flowers, and Leaves During Postharvest 429
Background 429 Development of the dedicated AZ tissue 430
Regulatory genes involved in abscission control or mediating
hormonal signal transduction 430 Genes involved in actual execution
of cell separation in the later stage of abscission 431 Ethylene
and abscission 431 Regulated manipulation of abscission 431
Reducing Postharvest Chilling Sensitivity 431 Background ., 431
Membrane structure and chilling sensitivity 432 Oxidative stress
and chilling sensitivity or tolerance 433 Regulation of
low-temperature responses 433 Molecules with protective functions
during cold stress 434
Affecting Postharvest Texture and Appearance Qualities 435
Background 435 Softening and cell wall hydrolysis 435 Softening and
turgor i. 435 Tissue lignifications 435
Implications for Plant and Agricultural Biotechnology 436
28 Engineering the biosynthesis of low molecular weight
metabolites for quality traits (essential nutrients,
health-promoting phytochemicals, volatiles, and aroma compounds)
443 Fumihiko Sato, Kenji Matsui General Introduction 443 Lessons
from Essential Nutrients 444
Essential amino acids : 444 Fatty acids 446 Vitamins 446
Improvement of the bioavailability of minerals through metabolic
engineering 449 Multigene transfer for improved food quality
449
General Strategy for the Engineering of Secondary Metabolites
with Nutritional Value 449 Identification of biosynthetic genes 449
Identification of transcription factors and engineering through
integrated "omics" 450 Modulation of organelle development 450
Quality Improvement of Plants as Functional or Medicinal Food
451 Resveratrol 451 Anthocyanins and flavonoids 452 Catechins and
proanthocyanidins 452 Sesamins 452
Beloved Metabolites: Plant Volatiles 452 Biochemistry of plant
volatile secondary metabolites 453 Flavor compounds in fruits 454
Scent/aroma of flowers 455 Volatile organic chemicals in vegetative
organs of plants 455
Perspectives 456 Conclusion 458 Acknowledgments 458
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Section F Plants as factories for industrial products,
pharmaceuticals, biomaterials, and bioenergy 463
29 Vaccines, antibodies, and pharmaceutical proteins 465 Yuri Y.
Gleba, Anatoli Giritch Introduction 465 Expression Technologies:
Nuclear Transformation 466 Expression Technologies: Plastid
Transformation 469 Expression Technologies: Transient Expression
Systems 469
"Full virus" vectors 470 Magnifection 470 Derisking the new
manufacturing process 471
Plant-Made Pharmaceuticals: A Unique Selling Proposition? 471
Plant-Based Manufacturing, Post-Translational Modifications, and
Plant-Specific Sugars 472 Plant-Based Manufacturing and Downstream
Issues 473 Plant-Based Expression Systems: Advantages and
Limitations 474
Nuclear transformation 475 Plastid transformation 475 Transient
expression 476
Conclusions and Outlook 476 Acknowledgments 476
30 Plants as factories for bioplastics and other novel
biomaterials 481 Jan B. van Beilen, Yves Poirier Introduction 481
Major Natural Plant Biopolymers 482
Starch 482 Cellulose 482 Rubber 483 Proteins 485
Novel Polymers Produced in Transgenic Plants 485 A role for
transgenic crops in the production of biopolymers? 485 Which
biopolymers should be targeted for production in transgenic crops?
486 Which crops should be targeted? 487 Fibrous proteins 487
Cyanophycin 488 Polyhydroxyalkanoate 489
Conclusion and Prospects 491
31 Bioenergy from plants and plant residues 495 Blake A. Simmons
Introduction 495 Biochemical Conversion 497
Comminution 498 Pre-treatment 499 Saccharification 500 Fuel
synthesis 500
Thermochemical Conversion 501 Pyrolysis 501 Gasification 502
Concluding Remarks 503 Acknowledgment 503
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Section G Commercial, legal, sociological, and public aspects of
agricultural plant biotechnologies 507
32 Containing and mitigating transgene flow from crops to weeds,
to wild species, and to crops 509 Jonathan Gressel Introduction:
Does Transgene Flow Matter? 509
Transgene flow: To what ecosystem? 510 Thresholds matter 511
Gene containment and/or mitigation is often necessary 511
Methods of Containment 511 Containment by targeting genes to a
cytoplasmic genome 512 Male sterility 512 Rendering crops asexual
513 Genetic use restriction technologies alias "terminator" 513
Chemically induced promoters for containment 513 Recoverable block
of function 514 Repressible seed-lethal technologies 514
Trans-splicing to prevent movement. 514 A genetic chaperon to
prevent promiscuous transgene flow from wheat to its wild and weedy
relatives 515 Transiently transgenic crops 515
Mitigating Transgene Flow 516 Demonstration of transgenic
mitigation 516 Will transgenic mitigation traits adversely affect
wild relatives of the crop? Models that suggests that mitigation is
deleterious 517
Traits that can be Used in Tandem Transgenic Mitigation
Constructs 518 Morphological traits and genes for mitigation 518
Chemical mitigation of transgene flow 519 Special cases where
transgenic mitigation is needed 520
Concluding Remarks 521
33 Intellectual property rights of biotechnologically improved
plants 525 Antoine Harfouche, Richard Meilan, Kannan Grant, Vincent
K. Shier Introduction: Capitalizing on Research and Development in
Agricultural Biotechnology with Intellectual Property Protection
525 Intellectual Property Protection of Biotechnologically Improved
Plants 526
International intellectual property protection agreements 526
Types of intellectual property protection in plant biotechnology
528
Freedom-to-Operate in Agricultural Biotechnology: The Road from
a Research Idea to Commercialization of a Biotechnologically
Improved Plant Product 532 Technology Transfer as a Means to
Facilitate the Development of Biotechnology-Based Agriculture 534
Conclusion and Future Needs 536 Acknowledgments 537
34 Regulatory issues of biotechnologically improved plants 541
Elizabeth £ Hood, Deborah Vicuna Requesens, Kellye A. Eversole
Introduction 541 Commercializing an Agricultural Biotechnology
Product 542 The Regulatory Framework 543
The U.S. Coordinated Framework. 545 Perspectives 547
Specialty crops regulatory assistance: A new paradigm 547
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Standardization 548 Conclusions 548
35 Prospects for increased food production and poverty
alleviation: What plant biotechnology can practically deliver and
what it cannot 551 Martina Newell McGloughlin Introduction 551
Progress to Date 552 The Next Generation 554 Barriers to
Introduction 557
36 Crop biotechnology in developing countries 563 Hugo De Groote
Introduction 563 Agriculture and Food in Developing Countries: The
Needs 564
Feeding a growing world population 564 Undernutrition and
poverty 564 Technology 565
Current State of GM Crops 565 Geographic distribution 565 Crops,
traits, and farmers 565 Future and trends 565
Economic Impact of Transgenic Crops in Developing Countries 567
Main effects of current GM crops 567 Empirical evidence of farm
level benefits 567 Effect of GM crops on poverty and inequality 568
Combined effects on farmer income 568 Macro level impacts 569
Health Impact 569 Safety concerns 569 Nutritional benefits of
biofortification 570 Nutritional impact of GM biofortification 570
Reduced exposure to toxins, pesticides, and anti-nutrients 571
The Environment 571 Consumer Acceptance of GM Food 572
Regional differences 572 Factors influencing acceptance 572
Regulatory Systems 573 Importance of regulatory systems 573
Regional differences 573 Economics of regulation 573 The way
forward. 574
Conclusions 574 Acknowledgments 574
Index 577
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