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Genetic Engineering of Horticultural Crops
Genetic Engineering of Horticultural Crops
Genetic Engineering of Horticultural Crops
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Genetic Engineering of Horticultural Crops

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Genetic Engineering of Horticultural Crops provides key insights into commercialized crops, their improved productivity, disease and pest resistance, and enhanced nutritional or medicinal benefits. It includes insights into key technologies, such as marker traits identification and genetic traits transfer for increased productivity, examining the latest transgenic advances in a variety of crops and providing foundational information that can be applied to new areas of study. As modern biotechnology has helped to increase crop productivity by introducing novel gene(s) with high quality disease resistance and increased drought tolerance, this is an ideal resource for researchers and industry professionals.

  • Provides examples of current technologies and methodologies, addressing abiotic and biotic stresses, pest resistance and yield improvement
  • Presents protocols on plant genetic engineering in a variety of wide-use crops
  • Includes biosafety rule regulation of genetically modified crops in the USA and third world countries
LanguageEnglish
Release dateJan 8, 2018
ISBN9780128104408
Genetic Engineering of Horticultural Crops

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    Genetic Engineering of Horticultural Crops - Gyana Ranjan Rout

    Genetic Engineering of Horticultural Crops

    Editors

    Gyana Ranjan Rout

    Department of Agricultural Biotechnology, Orissa University of Agriculture and Technology, Bhubaneswar, India

    K.V. Peter

    World Noni Research Foundation, Perungudi, Chennai, India

    Table of Contents

    Cover image

    Title page

    Copyright

    List of Contributors

    Foreword

    Preface

    Chapter 1. Status of Horticultural Crops: Identifying the Need for Transgenic Traits

    1. Introduction

    2. Need for Genetically Engineered Crops in the Present Scenario

    3. Transgenic Research in Major Horticulture Crops

    4. Benefits of Commercialization of Transgenic Horticultural Crops in Developing Countries

    5. Future Prospectives and Conclusion

    Chapter 2. Genetic Engineering of Horticultural Crops: Present and Future

    1. Introduction

    2. Traditional Breeding Technology

    3. Molecular Breeding Technology

    4. Transgenic Technology

    5. Biofarming

    6. Genome Editing Technology

    7. Future Challenges

    Chapter 3. Gene Silencing in Horticultural Transgenic Crops

    1. Introduction

    2. Post-Transcriptional Gene Silencing

    3. Mechanism of Gene Silencing

    4. Application of RNAi in Transgenic Plants

    5. Conclusion

    Chapter 4. Transgenic Research in Fruit Crops

    1. Introduction

    2. Genetic Improvement of Fruit Crops Through Genetic Engineering

    3. Future Prospects

    Chapter 5. Genetic Engineering of Temperate Fruit Crops

    1. Introduction

    2. Genetic Improvement of Temperate Fruits

    3. Genetic Engineering in Temperate Fruits

    4. Gene Silencing in Temperate Fruit Crops

    5. Future Prospects

    Chapter 6. Transgenic Research in Floricultural Crops

    1. Introduction

    2. Genetic Transformation

    3. Target Traits for Genetic Modification

    4. Gene Isolation and Current Gene Availability

    5. Genetic Modification: Status by Species and Traits

    6. Transformation of Floricultural Crops

    7. The Regulation of Genetically Modified Floricultural Plants

    8. Overview of Regulations Worldwide

    9. Cost of Regulation

    10. International Trade Constraints

    11. New Breeding Technologies

    12. Status of Commercialization of Genetically Modified Floricultural Plants

    13. Future Prospects

    Chapter 7. Genetic Engineering in Papaya

    1. Introduction

    2. Genetic Transformation Study in Papaya

    3. Marker-Free Technology

    4. Advantages of Each Strategy

    5. Achievements in Transgenic Papaya in Hawaii

    6. Future Prospects

    Chapter 8. Transgenic Research in Vegetable Crops With Special Reference to Brinjal

    1. Introduction

    2. Shoot, Stem, and Fruit Borer in Brinjal

    3. Efficacy of Transgenic Bt-Brinjal

    4. Flow of Transgene: Is It a Real Risk?

    5. Present Scenario of Transgenic Brinjal in India and Its Commercialization Controversy

    6. Probable Strategy to Combat the Biosafety Issue and Extranuclear Transformation

    7. Conclusion

    Chapter 9. Genetic Engineering of Oil Palm

    1. Introduction

    2. Origin and Distribution

    3. Taxonomy

    4. Genetics and Breeding

    5. Genetic Engineering

    6. Future Perspectives and Conclusions

    Chapter 10. Genetic Improvement of Vegetables Using Transgenic Technology

    1. Introduction

    2. National and International Status of Vegetable Production

    3. Constraints in Vegetable Production

    4. Need of the Transgenic Vegetable

    5. History and Background of Development of Transgenic Vegetables

    6. Factors Involved in Selecting the Target Crop and Trait

    7. Genetic Transformation

    8. Traits Involved in the Development of Genetically Modified Crops

    9. Global Status of Transgenic Vegetables

    10. Ethical and Biosafety Issues, Risks, and Concerns

    11. Future Prospects

    Chapter 11. Transgenic Research in Tuber and Root Crops: A Review

    1. Introduction

    2. Background of Genetic Engineering in Tuber and Root Crops

    3. Genetic Engineering of Potato

    4. Genetic Engineering in Sweet Potato

    5. Genetic Engineering of Taro [Colocasia esculenta (L.) Schott]

    6. Genetic Engineering of Ginger and Turmeric

    7. Biosafety Issues Involved in Transgenic Tuber and Root Crops

    8. Conclusion and Future Prospects

    Chapter 12. Genetic Engineering in Medicinal and Aromatic Plants

    1. Introduction

    2. Plant Secondary Metabolites and Their Role

    3. Molecular Elucidation of Plant Secondary Metabolites

    4. Genetic Transformation Systems

    5. Functional Genomics Approaches for Engineering of Secondary Metabolic Pathways

    6. Genetic Engineering for Abiotic Stress resistance

    7. Future Prospects

    Chapter 13. Biosafety and Bioregulatory Mechanisms for Transgenic Crops

    1. Introduction

    2. Defining Biotechnology, Biosafety, and Risks

    3. Risks Associated With GM Crops and Its Analysis

    4. Bioregulatory Mechanisms for GM Crops in India

    5. Bioregulatory Mechanisms for GM Crops in the European Union

    6. Bioregulatory Mechanisms for GM Crops in United States of America

    7. Bioregulatory Mechanisms for GM Crops in Canada

    8. Conclusion

    Chapter 14. Critical Evaluation of the Benefits and Risks of Genetically Modified Horticultural Crops

    1. Introduction

    2. Needs-Based Transgenic Research in Horticultural Crops

    3. Crop Productivity With Special Reference to Abiotic and Biotic Stresses

    4. Contribution of Biotech Crops to Food Security, Sustainability, and Climate Change

    5. Benefits of Genetically Engineered Crops

    6. Role of Marker-Free Technology in Transgenic Crops

    7. Risk Associated in Transgenic Crops: Agricultural, Ecological, and Social Issues

    8. Biosafety Regulation in Transgenic Crops

    9. Future Prospects

    Chapter 15. Transgenic Development for Biotic and Abiotic Stress Management in Horticultural Crops

    1. Introduction

    2. Transgenic Research in Biotic Stress Management in Horticultural Crops

    3. Transgenic Research in Abiotic Stress Management in Horticultural Crops

    4. Role of Marker-Free Transgenic Technology

    5. Genome Editing Technology in Horticultural Crop Improvement

    6. Future Prospects

    Chapter 16. Transgenic Research in Spices

    1. Introduction

    2. Comparative Genomics and Gene Tagging

    3. Cloning and Isolation of Genes

    4. Genetic Transformation

    5. Regulatory Issues

    6. Future Perspectives

    Chapter 17. Genomics of Cucurbits

    1. Introduction

    2. Genome-Wide Characterization Studies in Cucurbits

    3. Application of Omics Technologies for Molecular Breeding of Cucurbits

    4. Future Prospect of Cucurbits Genomics

    Index

    Copyright

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    List of Contributors

    B. Kalyana Babu,     ICAR-Indian Institute of Oil Palm Research, Pedavegi, India

    Kantipudi Nirmal Babu,     ICAR-Indian Institute of Spices Research, Kozhikode, India

    S. Backiyarani,     ICAR-National Research Centre for Banana, Tiruchirappalli, India

    Mehmet C. Baloglu,     Kastamonu University, Kastamonu, Turkey

    Rajib Bandopadhyay,     The University of Burdwan, Burdwan, India

    Aparna Banerjee,     The University of Burdwan, Burdwan, India

    Kaipa H. Bindu,     ICAR-Indian Institute of Horticultural Research, Bangalore, India

    Raju Biswas,     The University of Burdwan, Burdwan, India

    Stephen F. Chandler,     RMIT University, Bundoora, VIC, Australia

    Sharda Choudhary,     ICAR-National Research Centre on Seed Spices, Ajmer, India

    Rajarshi K. Gaur,     Mody University of Science and Technology, Sikar, India

    Sirhindi Geetika,     Punjabi University, Patiala, India

    Urmi Halder,     The University of Burdwan, Burdwan, India

    Kaur Harpreet,     Punjabi University, Patiala, India

    Devendra Jain,     MPUAT, Udaipur, India

    Cissin Jose,     ICAR-Indian Institute of Spices Research, Kozhikode, India

    K. Kalaiponmani,     ICAR-National Research Centre for Banana, Tiruchirappalli, India

    Yasin J. Khan,     ICAR-National Bureau of Plant Genetic Resources, New Delhi, India

    Satyendra M.P. Khurana,     Amity University Haryana, Gurgaon, India

    Prathapani Naveen Kumar,     ICAR-Indian Institute of Oil Palm Research, Pedavegi, India

    Deepu Mathew,     Kerala Agricultural University, Thrissur, India

    Ravi K. Mathur,     ICAR-Indian Institute of Oil Palm Research, Pedavegi, India

    Mintu R. Meena,     ICAR-Sugarcane Breeding Institute Regional Centre, Karnal, India

    Hidayatullah Mir,     Bihar Agricultural University, Sabour, India

    Jutti B. Mythili,     ICAR-Indian Institute of Horticultural Research, Bangalore, India

    Jogi Nanjundan,     ICAR-Indian Agricultural Research Institute Regional Station, Wellington, India

    Dogra Neha,     Punjabi University, Patiala, India

    Nehanjali Parmar,     Dr. Y.S. Parmar University of Horticulture & Forestry, Nauni, India

    Vishwa Bandhu Patel,     Bihar Agricultural University, Sabour, India

    K.V. Peter,     World Noni Research Foundation, Chennai, India

    Sunil Prajapati,     Jawaharlal Nehru Krishi Vishwa Vidyalaya, Jabalpur, India

    Rohini M. Radhika,     ICAR-Indian Institute of Horticultural Research, Bangalore, India

    Devarajan Ramajayam,     ICAR-Indian Institute of Oil Palm Research, Pedavegi, India

    Gyana Ranjan Rout,     Orissa University of Agriculture and Technology, Bhubaneswar, India

    Mushtaq Ruqia,     Punjabi University, Patiala, India

    Kailash C. Samal,     Orissa University of Agriculture and Technology, Bhubaneswar, India

    M.S. Saraswathi,     ICAR-National Research Centre for Banana, Tiruchirappalli, India

    Deepika Sharma,     ICAR-Directorate of Rapeseed-Mustard Research, Bharatpur, India

    Radheshyam Sharma,     ICAR-National Research Centre on Seed Spices, Ajmer, India

    Kaushik Shruti,     Punjabi University, Patiala, India

    Huaijun Si,     Gansu Agricultural University, Lanzhou, People’s Republic of China

    Kunwar H. Singh,     ICAR-Directorate of Rapeseed-Mustard Research, Bharatpur, India

    Lal Singh,     ICAR-Directorate of Rapeseed-Mustard Research, Bharatpur, India

    Sheesh P. Singh,     J.V. (PG) College, Baraut, India

    Yogendra Singh,     Jawaharlal Nehru Krishi Vishwa Vidyalaya, Jabalpur, India

    Erinjery Jose Suraby,     ICAR-Indian Institute of Spices Research, Kozhikode, India

    Yoshikazu Tanaka,     Suntory Global Innovation Center Ltd., Kyoto, Japan

    Xun Tang,     Gansu Agricultural University, Lanzhou, People’s Republic of China

    Ajay K. Thakur,     ICAR-Directorate of Rapeseed-Mustard Research, Bharatpur, India

    S. Uma,     ICAR-National Research Centre for Banana, Tiruchirappalli, India

    Anuradha Upadhyay,     ICAR-National Research Centre for Grapes, Pune, India

    Arvind K. Verma,     ICAR-National Research Centre on Seed Spices, Ajmer, India

    Rakesh K. Verma,     Mody University of Science and Technology, Sikar, India

    Li Wang,     Gansu Agricultural University, Lanzhou, People’s Republic of China

    Yikai Wen,     Gansu Agricultural University, Lanzhou, People’s Republic of China

    Jiangwei Yang,     Gansu Agricultural University, Lanzhou, People’s Republic of China

    Ning Zhang,     Gansu Agricultural University, Lanzhou, People’s Republic of China

    Xiangyan Zhou,     Gansu Agricultural University, Lanzhou, People’s Republic of China

    Foreword

    Malnutrition, hidden hunger, and resultant stunting are rampant in India, despite self-sufficiency in food grain production. Past achievements in agricultural production transformed India from a ship to mouth existence to the present farm to ship for export position, which has given it national honor and pride. Appropriate public policy, use of science and technology, enlightened farmers, a well-organized public distribution system, and ever-growing demand for food grains has resulted in the much applauded Green Revolution. India is presently promoting a Nutrition Revolution through home gardens and multiple cropping systems. Despite all efforts, productivity of horticultural crops—fruits, vegetables, tubers, plantation crops, medicinal and aromatic plants, mushrooms, and bamboos—is one of the lowest, except for a few crops. Both biotic and abiotic factors limit production and the problem of deficit is further accelerated by huge harvest and postharvest losses. Genetic engineering is a scientific technology used in developed and many developing countries to increase production, reduce losses, and improve quality. The present edited book Genetic Engineering of Horticultural Crops, with 17 chapters authored by 58 scientists from 30 research institutes from India, China, Japan, and Turkey, reveals scientific advances made in this modern science of biotechnology and molecular biology. Contributions from different experts on various issues relating to genetic engineering of horticultural crops are of high value and worth reading. I congratulate all the contributing authors for their time, patience, and commitment to science. I appreciate the efforts made by Professors Gyana Ranjan Rout and K.V. Peter for coordinating and compiling this important publication. I am sure the researchers, academia, students, and others will take advantage of this important and informative publication.

    Preface

    It gives me great pleasure in writing this Preface for Genetic Engineering of Horticultural Crop, edited by Prof. G.R. Rout (Professor and Head, Department of Agricultural Biotechnology, College of Agriculture, Orissa University of Agriculture and Technology, Bhubaneswar, India) and Dr. K.V. Peter (Director, World Noni Research Foundation, Perungudi, Chennai, India). I found the chapters in this book carefully designed to focus on the various transgenic research activities and tools, which address crop productivity through the introduction of novel gene(s) with high nutrient qualities, resistance to pests and diseases, and improvement of stress tolerance. The book also discusses insights into key technologies, such as markers for trait(s) identification and transfer of these (various) genetic traits toward improving crop productivity. It also examines the latest trends in transgenic advances in a variety of crops, thus providing foundational information for future crop improvement research programs.

    Genetic Engineering of Horticultural Crops covers a complete range of topics on the status of horticultural crop research, including identifying the need for transgenic traits, gene silencing in horticultural crops, transgenic research in fruit crops, spices, medicinal and aromatic plants, vegetables (with special reference to brinjal), root and tuber crops, nuts, and floriculture crops, transgenic development for biotic and abiotic stress management in horticultural crops, and, lastly, biosafety and bioregulatory mechanisms in transgenic crops. I am pleased to note that the latest developments in molecular biology research, including the role of marker-free technologies such as genome editing, have also been discussed in this book, which makes it not only an up-to-date book, but also a much-needed book in this area, covering topics that are essential for the development of transgenic (horticultural) crops. I am sure this book will become a great companion to researchers, postgraduate students, and plant biotechnologists, providing insights into the various gene transfer technologies relevant for sustainable agriculture and food production systems.

    I congratulate Academic Press, an imprint of Elsevier, United States, for the excellence and great outlook of the book.

    BSc (UKM, Malaysia), MSc and PhD (London, UK)

    Senior Vice President (Agriculture Biotechnology)

    Chapter 1

    Status of Horticultural Crops

    Identifying the Need for Transgenic Traits

    Yogendra Singh, and Sunil Prajapati     Jawaharlal Nehru Krishi Vishwa Vidyalaya, Jabalpur, India

    Abstract

    For five decades conventional plant breeding techniques have made considerable progress in the development of improved varieties including horticultural crops. However, it has not been able to keep pace with the increasing demand for vegetables and fruits in developing countries. Therefore an instant need is felt to incorporate modern recombinant DNA technology to speed up crop improvement programs. Recombinant DNA technology has offered wonderful scope and potential to conventional methods of crop improvement, crop protection, and crop quality management. Transgenic technology in horticultural crops is helpful in increasing crop production and productivity, managing abiotic and biotic stress, enhancing nutritional value, and increasing shelf-life. This is why the global land area occupied by genetically modified crops has been continuously increasing during the last two decades, and reached a record 181.5  million ha globally by 2014. There are some issues related to biosafety and intellectual property rights in transgenic crops, which should be addressed by experts so the real benefits of this can reach out to others.

    Keywords

    Abiotic stress management; Flavr Savr; Floriculture; Genetically modified crops; Nutritional value; Papaya ringspot virus; Shelf-life

    1. Introduction

    Although conventional plant breeding techniques have made considerable progress in the development of improved varieties, it has not been able to keep pace with the increasing demand for vegetables and fruits in developing countries. Therefore an instant need is felt to incorporate transgenic technology to speed up crop improvement programs. Three major constituents are involved in crop improvement programs, i.e., in vitro culture protocol development, marker-assisted breeding, and genetic engineering. Agricultural biotechnology has offered wonderful scope and potential to conventional methods for crop improvement, crop protection, crop quality management, and other horticultural traits. Biotechnology explores various opportunities in fruit production by providing new genotypes for breeding purposes, a supply of healthy and disease-free planting material, improvement in fruit quality, enhanced shelf-life, availability of biopesticides, biofertilizers, etc. The expression of undesirable genes can be blocked by the application of antisense gene technology and RNA-interference (RNAi) technology. Eventually, biotechnological interventions that could increase the efficiency of horticultural crop improvement are essential to generate plants with several desirable traits.

    1.1. Basic Concept of Genetically Modified Crops

    Genetically modified (GM) crops are products of the introduction of one or more characterized genes in a crop plant using recombinant DNA technology (RDT). The inserted gene is known as a transgene and the plants containing transgenes are often called GM crops or transgenic crops. GM crops are a possible solution for the widely discussed current problems of food and nutritional security. In contrast, conventionally bred cultivars result from artificial selection, random mutagenesis, or intra- or interspecific (rarely intergeneric) hybridization leading to the transfer of a number of uncharacterized genes from the same or a related alien species. Therefore the major difference between GM crops and conventionally bred cultivars is in the technology for transfer, and in the nature of genes transferred. Despite this debate among the proponents and opponents of GM crops, the global land area occupied by GM crops has been continuously increasing during the last 20  years, reaching a record 181.5  million ha globally in 2014, at an annual growth rate of between 3% and 4%, up 6.3  million ha from 175.2  million ha in 2013. The year 2014 was the 19th year of commercialization, when growth continued after a remarkable 18 consecutive years of increases; notably 12 of the 18  years were double-digit growth rates. In other words, biotech crops are the fastest adopted crop technology in the world.

    The first commercially grown GM vegetable crop was tomato (called Flavr Savr), modified to ripen without softening by the Californian company Calgene, which took the initiative to obtain approval for its release in 1994. Currently, a number of food crops such as soybean, corn, cotton, Hawaiian papaya, potatoes, rapeseed (canola), sugarcane, sugar beet, field corn, as well as sweet corn and rice have been genetically modified to enhance their yield and durability, etc. Scientists are also working on oil-yielding crops and medicinal plants for the cosmetics industry, crops with altered nutritional value, and even crops that produce pharmaceutical drugs.

    The basis of RDT is a key set of enzymes and techniques that allow DNA to be manipulated and modified precisely (Fig. 1.1). The fundamentals of RDT include:

    1. Cutting of DNA with sequence-specific bacterial endonuclease (restriction endonuclease) to generate defined DNA fragments and using the enzyme DNA ligase to join them;

    2. Separating nucleic acid on the basis of size by gel electrophoresis;

    3. Detecting specific sequences in complex mixtures by nucleic acid hybridization;

    4. Introducing DNA into cells;

    5. Amplification of specific DNA molecules either by molecular cloning or using polymerase chain reaction.

    Molecular cloning is an in vivo technique for producing large quantities of a particular DNA molecule (recombinant DNA molecule). The cloning process involves:

    1. Introduction of the recombinant vector into a suitable host cell;

    2. Selective propagation of cells containing the vector;

    3. Extraction and purification of the cloned DNA molecule (recombinant DNA molecule).

    1.2. Production of Genetically Modified Crops With Special Reference to Horticultural Plants

    With the rapid advances in biotechnology, a number of GM crops or transgenic crops carrying novel traits have been developed and released for commercial agriculture production. Commercial cultivation of transgenic crops started in the early 1990s. Herbicide tolerance and insect resistance are the main traits that are currently under commercial cultivation, and the main crops are: soybean, maize, canola, and cotton. A new and rigorous comprehensive global metaanalysis of 147 published biotech crop studies over the last 20  years confirmed the significant and multiple benefits that biotech crops have generated from 1995 to 2014. Millions of farmers, both large and small, worldwide have concluded that the returns from planting biotech crops are high, hence repeat planting is virtually 100%; good returns on their investment is the critical test applied by demanding farmers when judging the performance of any technology. Out of 28 countries involved in commercial cultivation of biotech crops, 20 were developing and only eight were industrial countries (Table 1.1, Fig. 1.2). More than half the world’s population, ∼60% or ∼4  billion people, live in the 28 countries planting biotech crops. Bangladesh, one of the smaller and poorest countries in the world, approved and commercialized Bt brinjal in record time in 2014. Vietnam and Indonesia moved toward planting their first biotech crops in 2015 for a total of nine biotech countries in Asia. The choice of GM crops varies among the developing countries, with insect-resistant cotton being the most important commercially produced transgenic crop in Asian and African countries, while herbicide-resistant soybean followed by insect-resistant corn are predominant in the Latin American continent. In most cases, these GM technologies are proprietary, developed by the private sector, and released for commercial production through licensing agreements. The cultivation and commercial production of GM crops are capital intensive because of the high costs of seed and technology. Nevertheless, their cultivation has increased, mainly because of the benefits accrued from lower labor and production costs, reduction in the use of chemical inputs, and improved economic gain.

    Figure 1.1  Basic steps of recombinant DNA technology.

    Research on GM potato, cucumber, carrot, eggplant, sweet corn, and other vegetables in many countries of the world is aimed at resistance to viruses, bacteria, fungi, and insects, at tolerance to herbicides, at improvement of economic properties, prolongation of the consumption time, improvement of nutrition values, and seedlessness of fruits. The development of transgenic fruit cultivars is in progress. Papaya resistant to papaya mosaic virus is grown in the United States and China (James, 2011). Biotech grapevine resistant to viral, bacterial, and fungal diseases with abiotic stress tolerance and health benefits was developed in South Africa. Biotech banana, apple, pear, and strawberry cultivars are under development. The result of the international research conducted over the past 20  years is the development of HoneySweet plum highly resistant to plum pox virus (PPV). GM HoneySweet plum resistant to PPV was deregulated in the United States in 2010. Plums (Prunus domestica) are an important source of vitamins, minerals, and phytonutrients and contain specific compounds that support good digestive function and bone health. Sharka disease is the most devastating disease of plum and is responsible for the reduction or loss of plum production in many areas of Europe (Cambra et al., 2006).

    Table 1.1

    a 19 biotech mega-countries growing 50,000  ha, or more, of biotech crops.

    b Rounded off to the nearest hundred thousand.

    Courtesy: James, C., 2014. Global Status of Commercialized Biotech/GM Crops. The International Service for the Acquisition of Agri-biotech Applications (ISAAA), summary available at: http://www.isaaa.org/.

    Figure 1.2  Global map of biotech crops countries and mega-countries in 2014.

    Courtesy: James, C., 2014. Global Status of Commercialized Biotech/GM Crops. The International Service for the Acquisition of Agri-biotech Applications (ISAAA), summary available at: http://www.isaaa.org/.

    Papaya was genetically modified to resist the ringspot virus. Sun Up is a transgenic red-fleshed Sunset papaya cultivar that is homozygous for the coat protein (CP) gene papaya ringspot virus (PRSV); Rainbow is a yellow-fleshed F1 hybrid developed by crossing Sun Up and transgenic yellow-fleshed Kapoho (Gonsalves, 2004). In the early 1990s The New York Times stated that Hawaii’s papaya industry was facing disaster because of the deadly PRSV. Its single-handed savior was a breed engineered to be resistant to the virus. Without it, the state’s papaya industry would have collapsed. Today, 80% of Hawaiian papaya is genetically engineered, and there is still no conventional or organic method to control ringspot virus (Ronald and McWilliams, 2010). The GM cultivar was approved in 1998. In China, a transgenic PRSV-resistant papaya was developed by South China Agricultural University and was first approved for commercial planting in 2006; as of 2012, 95% of the papaya grown in Guangdong province and 40% of the papaya grown in Hainan province were genetically modified. The NewLeaf potato, brought to market by Monsanto in the late 1990s, was developed for the fast food market. It was withdrawn in 2001 after retailers rejected it and food processors faced export problems. As of 2005, about 13% of the zucchini (a form of squash) grown in the United States was genetically modified to resist three viruses; that strain is also grown in Canada. In 2011, Badische Aniline and Soda Company (BASF) requested the European Food Safety Authority’s approval for cultivation and marketing of its Fortuna potato as feed and food. The potato was made resistant to late blight by adding resistant genes blb1 and blb2 that originate from the Mexican wild potato (Solanum bulbocastanum). In February 2013, BASF withdrew its application. In 2013, the United States Department of Agriculture (USDA) approved the import of a GM pineapple that is pink in color and that overexpresses a gene derived from tangerines and suppresses other genes, increasing production of lycopene. The plant’s flowering cycle was changed to provide more uniform growth and quality. The fruit does not have the ability to propagate and persist in the environment once it has been harvested. According to Del Monte’s submission, the pineapples are commercially grown in a monoculture that prevents seed production because the plant’s flowers are not exposed to compatible pollen sources. Importation into Hawaii is banned for plant sanitation reasons (Perkowski, 2013). In 2014, the USDA approved a GM potato developed by J.R. Simplot Company that contained 10 genetic modifications that prevented bruising and produced less acrylamide when fried. The modifications eliminate specific proteins from the potatoes, via RNA interference, rather than introducing novel proteins. In February 2015, Arctic Apples were approved by the USDA, becoming the first GM apple approved for sale in the United States. Gene silencing is used to reduce the expression of polyphenol oxidase, thus preventing the fruit from browning.

    1.3. International and National Status

    There has been a consistent increase in global areas planted to transgenic crops from 1996 to 2014, which was assisted by 18  million farmers in 28 countries. About 181.5  million ha was planted (James, 2014) to transgenic crops in 2014 with high market value, such as herbicide-tolerant soybean, maize, cotton, and canola; insect-resistant maize, cotton, potato, and rice; and virus-resistant squash and papaya. With genetic engineering, more than one trait can be incorporated or stacked into a plant. Transgenic crops with combined traits are also available commercially. These include herbicide-tolerant and insect-resistant maize, soybean, and cotton. Notably, Bangladesh approved Bt brinjal for the first time on October 30, 2013, and in record time less than 100  days after approval small farmers commercialized Bt brinjal on January 22, 2014. Innate potato, another food crop, was approved in the United States in November 2014. It has lower levels of acrylamide, a potential carcinogen in humans, and suffers less wastage from bruising; potato is the fourth most important food staple in the world. A safer product and decreased wastage in a vegetatively propagated and perishable crop can contribute to higher productivity and food security. Also in November 2014, a new biotech alfalfa (event KK179) with up to 22% less lignin, which leads to higher digestibility and productivity, was approved for planting in the United States. The first biotech drought-tolerant maize, planted in the United States in 2013 on 50,000  ha, increased over fivefold to 275,000  ha in 2014 reflecting high acceptance by US farmers. Importantly, a new 2014 comprehensive global metaanalysis on 147 published biotech crop studies over the last 20  years worldwide confirmed the significant and multiple benefits that biotech crops have generated from 1995 to 2014; on average GM technology adoption has reduced chemical pesticide use by 37%, increased crop yields by 22%, and increased farmer profits by 68%. These findings corroborate earlier and consistent results from other annual global studies, which estimated increases in crop productivity valued at US$133.3  billion for the period 1996–2013.

    Biotech crops were grown commercially in all six continents of the world. Of the 28 countries planting biotech crops in 2015, 19 countries planted 50,000  ha or more to biotech crops (Table 1.2). These megacountries include the United States, Brazil, Argentina, India, Canada, China, Paraguay, Pakistan, South Africa, Uruguay, Bolivia, Philippines, Australia, Burkina Faso, Myanmar, Mexico, Spain, Colombia, and Sudan. Bangladesh approved a biotech crop (Bt brinjal) for planting the first time on October 30, 2013, and it was recorded that less than 100  days after the approval, commercialization was initiated on January 22, 2014 when 20 very small farmers planted their first crop of Bt brinjal; a total of 120 farmers planted 12  ha of Bt brinjal in 2014. This approval by Bangladesh is important in that it serves as an exemplary model for other small, poor countries. Also, very importantly, Bangladesh has broken the impasse experienced in trying to gain approval for commercialization of Bt brinjal in both India and the Philippines.

    Table 1.2

    a 19 biotech mega-countries growing 50,000  ha, or more, of biotech crops.

    Courtesy: James, C., 2014. Global Status of Commercialized Biotech/GM Crops. The International Service for the Acquisition of Agri-biotech Applications (ISAAA), summary available at: http://www.isaaa.org/.

    Figure 1.3  Global area of biotech crops million hectares (1996–2014).

    Courtesy: James, C., 2014. Global Status of Commercialized Biotech/GM Crops. The International Service for the Acquisition of Agri-biotech Applications (ISAAA), summary available at: http://www.isaaa.org/.

    Remarkably, in 2014 global biotech crop hectarage continued to grow for the 19th consecutive year of commercialization; 18  million farmers in 28 countries planted more than 181  million ha in 2014, up from 175  million in 27 countries in 2013 (Figs. 1.3 and 1.4). Innate potato, another food crop, which was approved for the commercial cultivation in the United States in November 2014. It has lower levels of acrylamide, a potential carcinogen in humans, and suffers less wastage from bruising; potato is the fourth most important food staple in the world. A safer product and decreased wastage in a vegetatively propagated and perishable crop, can contribute to higher productivity and food security. Also in November 2014, a new biotech alfalfa (event KK179) with up to 22% less lignin, which leads to higher digestibility and productivity, was approved for planting in the United States. Importantly, a new 2014 comprehensive global metaanalysis, on 147 published biotech crop studies over the last 20  years worldwide confirmed the significant and multiple benefits that biotech crops have generated from 1995 to 2014; on average GM technology adoption has reduced chemical pesticide use by 37%, increased crop yields by 22%, and increased farmer profits by 68%. These findings corroborate earlier and consistent results from other annual global studies which estimated increases in crop productivity valued at US$133.3  billion for the period 1996–2013.

    Figure 1.4  Currently grown genetically modified crops in the United States.

    Traits for which they are modified, and percent of total acreage of the crop that is planted to GM varieties. DT, drought tolerant; HT, herbicide tolerant; IR, insect resistant; VR, virus resistant. Courtesy: Byrne, P., 2014. Genetically Modified (GM) Crops: Techniques and Applications. Colorado State University.

    2. Need for Genetically Engineered Crops in the Present Scenario

    2.1. Crop Production and Productivity

    The efficacy of transgenic plant varieties in increasing production and lowering production costs is already demonstrable. In 1996 and 1997, the cultivation of virus-, insect-, and herbicide-resistant plants accounted for a 5%–10% increase in yield as well as for savings on herbicides of up to 40% and on insecticides of between $60 and $120 (US dollars) per acre (James, 1998). However, these increases in productivity, impressive as they are, probably have a limited impact on the global food supply because the products currently available on the market are suitable only for large mechanized farms practicing intensive agriculture. In fact, most of the transgenic crops that have been produced to date, especially by the private sector, are aimed either at reducing production costs in agricultural areas that already have high productivity levels or at increasing the value of the final product (e.g., improving the oil quality of seed crops). In a global sense, a more effective strategy to ensure sufficient levels of food production would be to increase productivity in developing countries, especially in areas of subsistence farming, where an increase in food production is urgently needed and where crop yields are significantly lower than those obtained in other areas of the world. In developing countries in the tropics and subtropics, crop losses caused by pests, diseases, and poor soils are made worse by climatic conditions that favor insect pests and disease vectors and by the lack of economic resources to purchase high-quality seeds, insecticides, and fertilizers. In addition to low productivity levels, postharvest losses in tropical areas are very high because of the favorable climate for fungal and insect infestation and the lack of appropriate storage facilities. Despite efforts to prevent pre- and postharvest crop losses, pests destroy over half of all world crop production. Postharvest loss caused by insects, the majority of which occurs in the developing world, is estimated to be 15% of the world’s production. It is possible that many of these problems could be alleviated by plant biotechnology. A major advantage of plant biotechnology is that it often generates strategies for crop improvement that can be applied to many different crops. Genetically engineered virus resistance, insect resistance, and delayed ripening are good examples of strategies that could potentially benefit a diversity of crops. Insect-resistant plant varieties, using the dendotoxin of Bacillus thuringiensis, have been produced for several important plant species, including tomato, potato, walnut, and maize; these are already under commercial production. It is envisaged that these strategies can be used for many other crops important for tropical regions and other regions in the developing world. Genetically engineered delayed ripening, although only tested on a commercial scale for tomato, has an enormous potential application for tropical fruit crops, which suffer severe losses in developing countries because they ripen rapidly and because there is a lack of appropriate storage conditions and efficient transport systems for them to reach the final consumer.

    2.2. Abiotic Stress Management

    High temperature, high light intensity, humidity, drought, frost, and salinity are the major abiotic stresses that reduce the yield and quality of fruit by affecting the vegetative and reproductive stages of growth and development. Nevertheless, abiotic stresses remain the greatest constraint to crop production. Research on genetic modification of various horticultural crops for improved abiotic stress tolerance has been explored with transformed tomato plants with a DNA cassette containing an Arabidopsis C repeat/dehydration-responsive element binding factor 1 (CBF1) complementary DNA (cDNA) and a nos terminator, driven by a cauliflower mosaic virus 35S promoter (Tsai-Hung et al., 2002). These transgenic tomato plants were more resistant to deficit water stress than the wild-type plants. Plants when exposed to abiotic stress conditions produce several pathogenesis-related proteins to compensate the effect of stress conditions. Among these proteins, osmotin is one of the important ones released during abiotic stress conditions. Husaini and Abdin (2008) reported the overexpressed tobacco osmotin gene in strawberry (Fragaria x ananassa Duch.) and found that the transgenic strawberry plants exhibited tolerance to salt stress. Also Subramanyam et al. (2011) expressed the tobacco osmotin gene in Capsicum annuum and the transgenic chilli plants exhibited improved salt tolerance. Cheng et al. (2009) developed transgenic tomato plants expressing the yeast SAMDC gene, which improved the efficiency of CO2 assimilation and protected the plants from high-temperature stress (38°C) as compared to the wild-type plants. A bacterial mannitol-1-phosphate dehydrogenase (mtlD) gene driven by the constitutive cauliflower mosaic virus (CaMV) 35S promoter was transferred into tomato plants in an attempt to improve abiotic stress tolerance (Khare et al., 2010). Drought (polyethylene glycol in medium) and salinity (sodium chloride in medium) tolerance tests revealed that transgenic lines exhibited a higher tolerance for abiotic stresses than nontransformed plants.

    2.3. Biotic Stress Management

    Pathogens cause losses in 10%–16% of the global harvest (Chakraborty and Newton, 2011). This loss can be minimized by using transgenic technology. In traditional agriculture, only individuals of the same species or eventually closely related species can be crossbred. If in this naturally available gene pool, resistance to biotic stress does not exist, traditional breeders cannot create resistance or introgress this trait into new varieties. Therefore it is necessary to search for alternative sources of genes in other completely unrelated plant species or in microbial organisms. Besides, traditional methods are resource- and time-consuming and germplasm dependent (Roy et al., 2011). Also using chemical spray may have adverse effects on human health and the environment, including beneficial organisms, and may lead to the development of chemical-resistant insects and weeds (Wahab, 2009). Plant genetic engineering has been made possible during the last three decades. Currently, transgenic plants with herbicide, insect pest, and virus disease resistance are cultivated in more than 175.2  million ha globally (James, 2013), while in 1996 only 1.7  million ha of land were under transgenic crops. Out of the 29 countries currently contributing to the cultivation of transgenic plants, 21 are developing countries and eight are industrial. During the 1996–2014 period, cumulative economic benefits from transgenic plants were high in developing countries at US$47.9  billion compared to US$59  billion generated by industrial countries.

    The major achievements in biotic stress management are as follows.

    2.3.1. Pest Resistance

    The bioinsecticidal δ-endotoxin gene (Bt gene) isolated from B. thuringiensis is currently in use to make plants resistant to insect pests. Progress in engineering insect resistance in horticultural plants has been attained by the use of insect control protein genes of B. thuringiensis. Insect resistance was first reported in tomato using Bt genes in 1987. Transgenic Bt tomato plants exhibited resistance against Spodoptera litura and Heliothis virescens (Fischhoff et al., 1987). Fruit trees such as persimmon transgenic for the cry I gene were found resistant to Plodia interpunctata and Monema flavescens (Tao et al., 1997). Potato varieties engineered for resistance to Colorado potato beetle were in commercial production for several years and were technically and agronomically successful, allowing significant reductions in insecticide use (Shelton et al., 2002). Chakrabarty et al. (2002) transformed cauliflower var. Pusa Snowball K-1 with a synthetic cry IAb gene and the transgenic plants indicated the effectiveness of the transgene against infestation by diamondback moth (Plutella xylostella) larvae during insect bioassays. Paul et al. (2005) developed transgenic cabbage (Brassica oleracea var. capitata) with a synthetic fusion gene of B. thuringiensis encoding a translational fusion product of cry1B and cry1Ab δ-endotoxins and found the transgenic plants resistant to P. xylostella. Transgenic technology has also been found to deliver resistance against various nematodes. Roderick et al. (2012) developed transgenic plantain (Musa sp.) cv. Gonja manjaya plants expressing a maize cystatin gene that inhibits the digestive cysteine proteinases and a synthetic peptide that disrupts nematode chemoreception. The best level of resistance exhibited by the transgenic plants against the major pest species Radopholus similis was 84% for the cystatin, 66% for the peptide, and 70% for the dual defense.

    2.3.2. Disease Resistance

    One of the major constraints limiting the production of fruit crops is diseases caused by several fungi, bacteria, and viruses. Conventional breeding seems to have limited application because of nonavailability of resistant gene(s) in gene pools of a particular crop. Genetic engineering of disease resistance in crops has become popular and valuable in terms of cost and efficacy. In fruit crops, the CP-mediated approach to engineer virus resistance has been in application to introduce resistance against diseases such as PPV, citrus tristeza virus, and grape fan leaf virus, etc. Papaya is grown in many tropical countries, but its cultivation is being threatened by PRSV, a disease that is considerably lowering its yield. Using biotechnological interventions, the CP gene of the virus has been transferred to papaya to confer PRSV resistance. Since 1998, GM papayas have been cultivated in Hawaii, USA, which had shown considerable resistance to PRSV. PRSV-resistant transgenic papaya varieties Sun Up and Rainbow have now occupied >80% shelf space in the US market. Also transgenic papaya plants with the mutated replicase (RP) gene from PRSV showed high resistance or immunity against PRSV in the field (Xiangdong et al., 2007). Praveen et al. (2010) developed transgenic plants of tomato with an AC4 gene–RNAi construct and the transgenic plants were found to show the suppression of tomato leaf curl virus activity. Yu et al. (2010) transformed commercial watermelon cultivars with an untranslatable chimeric construct containing truncated zucchini yellow mosaic virus CP and PRSV WCP genes.

    HoneySweet plum is highly resistant to PPV, the most devastating disease of plums and other stone fruits. HoneySweet was deregulated in the United States in 2010. HoneySweet (aka C5) has been evaluated for 11  years (2002–12) in a regulated field trial in the Czech Republic for resistance to PPV, prune dwarf virus (PDV), and apple chlorotic leaf spot virus (ACLSV), all of them being serious diseases of plum. Even under the high and permanent infection pressure produced through grafting, PPV has been detected only in HoneySweet trees in several leaves and fruits situated close to the point of inoculum grafting. The lack of infection spread in HoneySweet demonstrates its high level of PPV resistance. Coinfections of PPV with PDV and/or ACLSV had practically no influence on the quantity and quality of HoneySweet fruits, which are large, sweet, and of a high eating quality. In many respects, they are superior to the fruits of the well-known cultivar Stanley. Many fruit growers and fruit tree nurseries in the Czech Republic are supportive of the deregulation of HoneySweet plum to help improve plum production and control the spread of PPV.

    RNAi technology is being used quite successfully in controlling various bacterial and viral diseases in plants by switching off the expression of certain endogenous genes. Transgenic tomato plants expressing hairpin RNA (hpRNA) constructs against Agrobacterium iaaM and ipt oncogenes were found to be resistant to crown gall disease (Escobar et al., 2001). Using an hpRNA gene silencing strategy, transgenic poinsettia plants resistant to poinsettia mosaic virus have been developed by Clarke et al. (2008). RNAi technology has been found to impart resistance to various bacterial plant diseases. For imparting bacterial and fungal resistance, various genes such as chitinase, glucanase, attacin, osmotin, cercopin, defensing, etc. are being transferred into various horticultural crops globally. The HcrVf2 gene from a wild apple conferred scab resistance to a transgenic cultivated variety of apple by Belfanti (2004). Faize et al. (2004) developed transgenic apple plants with a wheat puroindoline-b (pin B) gene under a CaMV35S promoter and observed that the expression of pin-b gene reduced scab susceptibility in transgenic apple plants. Girhepuje and Shinde (2011) developed transgenic tomato plants expressing a wheat endochitinase gene and during disease screening the transgenic plants exhibited enhanced resistance to Fusarium oxysporum. Rivera-Domínguez et al. (2011) carried out genetic transformation of mango (Mangifera indica) cv. Ataulfo embryos with the defensin J1 gene.

    2.3.3. Herbicide Resistance

    The herbicide glyphosate is a potent inhibitor of the enzyme 5-enolpyruvylshikimate-3-phosphate synthase (EPSP) in higher plants. Shah et al. (1986) developed a cDNA clone encoding EPSP synthase from a cDNA library of a glyphosate-tolerant Petunia hybrida cell line (MP4-G) that overproduces the enzyme. This cell line was shown to overproduce EPSP synthase messenger RNA as a result of a 20-fold amplification of the gene. A chimeric EPSP synthase gene was constructed with the use of the cauliflower mosaic virus 35S promoter to attain high-level expression of EPSP synthase and introduced into petunia cells. Transformed petunia cells as well as regenerated transgenic plants were tolerant to glyphosate. Transgenic pineapple plants transformed with the bar gene for bialaphos resistance were developed by Sripaoraya et al. (2006) and evaluated for tolerance to the herbicide Basta. Seven months after transfer to the field, plants were found tolerant to 1600  mL/rai of Basta X (stock concentration 15% w/v glufosinate ammonium), this being twice the dose recommended for field application of the herbicide. Transgenic plants tolerant to glufosinate ammonium should facilitate more effective weed control in pineapple plantations without damage to the crop.

    2.4. Nutritional Value

    One of the main claims of the biotechnology industry since GM crops were first commercialized in the United States in 1996 has been that a second generation of genetic modification will bring real consumer benefits, for example, by improving the nutritional value of foods. First-generation GM crops are aimed at farmers by inserting genes for herbicide tolerance and insect resistance, which were sold by the biotechnology companies as a way to cut inputs and labor (which is disputed). Consumers in the United Kingdom, the rest of the European Union, and around the world rejected them. Their main use is therefore in animal feed and, more recently, in biofuels, neither of which are obvious or labeled at the point of sale. The biotech industry now hopes to boost their market with second-generation nutritionally enhanced GM crops, which it is claimed will alleviate malnutrition and improve health. Many biotech proponents go even further and claim that consumers will be able to obtain foods specifically developed to prevent common diet-related diseases.

    2.5. Shelf-Life and Qualitative Traits

    A distinction is made in the genetic manipulation of plants between input traits and output traits. Input traits involve changing the agricultural characteristics of plants, offering the farmer technical advantages in cultivation. These include traits that affect the growth of the plant, such as herbicide or insect resistance or tolerance to drought, cold, or lack of nutrients. Output traits are the qualitative or quantitative improvement of characteristics relating to the condition of plants or the substances they contain. For example, attempts are being made to use gene technology to give plants and parts of plants a longer shelf-life once they have been harvested (Flavr Savr tomato). It was observed that excessive softening is the main factor limiting fruit shelf-life and storage. Other goals are to achieve a higher vitamin or protein content. Output traits aim to provide advantages that are of personal benefit to the end consumer, and offer improved processing quality to companies that carry out the further processing of the products.

    Transgenic plants modified in the expression of cell wall-modifying enzymes have been used to investigate the role of particular activities in fruit softening during ripening. Fruit ripening has been modified by altering the activity of cell wall enzymes such as polygalacturonases that are involved in tissue softening and deterioration. The biosynthesis of ethylene (the fruit-ripening hormone) has also been blocked in several ways to delay fruit ripening. Calgene Inc., USA (1994), developed the first commercialized transgenic plant, a long shelf-life tomato (Flavr Savr) by the suppression of the polygalacturonase (PG) gene by antisense strategy (Fig. 1.5). The Flavr Savr tomatoes have improved flavor and total soluble solids, in addition to enhanced shelf-life. However, this Flavr Savr variety was withdrawn from the market 3 years later because of its disease susceptibility and lack of productivity. Later on, other tomato varieties with increased shelf-life were developed through antisense RNA inhibition of 1-aminocyclopropane-1-carboxylate (ACC) synthase or ACC oxidase and two ethylene precursors. Delayed leaf senescence has been achieved in tobacco and petunia by manipulation of cytokinin synthesis (Clark et al., 2003). Researchers at Horticultural Research International, UK, have identified the genes that control the taste, smell, and color of strawberries. As a result, it would now be possible to create super strawberries that will taste sweeter using transgenic approaches. Nambeesan et al. (2010) expressed a yeast spermidine synthase (ySpdSyn) gene under constitutive (CaMV35S) and fruit-ripening-specific (E8) promoters in Solanum lycopersicum (tomato). The ySpdSyn transgenic fruits had a longer shelf-life, reduced shriveling, and delayed decay symptom development in comparison with the wild-type fruits. Crop maturity indicated by the percentage of ripening fruits on the vine was delayed in a CaMV35S-ySpdSyn genotype, with fruits accumulating higher levels of the antioxidant lycopene. Notably, whole-plant senescence in the transgenic plants was also delayed compared with wild-type plants. Zhang et al. (2011) developed transgenic tomato plants by silencing the expression of the mitochondrial APX gene by an RNAi mechanism and observed increased vitamin C content in the transgenic tomato fruits.

    Figure 1.5  Making of Flavr Savr tomato.

    Courtesy: http://www.google.com/Images.

    3. Transgenic Research in Major Horticulture Crops

    3.1. Fruit Crops

    Fruits are one of the major sources of vitamins, essential nutrients, antioxidants, and fibers in the human diet. During the last two to three decades, genetic engineering methods based on the use of transgenes have been successfully adopted to improve fruit plants and focused mainly on enhanced tolerance to biotic and abiotic stresses, increased fruit yield, improved postharvest shelf-life of fruit, reduced generation time, and production of fruit with higher nutritional value. However, the development of transgenic fruit plants and their commercialization have been delayed by many regulatory and social hurdles. Nowadays, new genetic engineering approaches, i.e., cisgenesis or intragenesis, receive increasing interest for the genetic modification of plants. The absence of selectable marker genes in the end product and the introduced gene(s) derived from the same plant or plants sexually compatible with the target crop should increase consumers’ acceptance.

    3.2. Vegetable Crops

    Several companies are working on tomatoes that can be vine ripened and shipped without bruising. Others are trying to improve tomatoes that are processed for catsup, soups, pastes, or sauces by genetically engineering them to contain more solids, be thicker, and to contain more lycopene, which provides the red color. These transgenic tomatoes will have higher viscosity (thickness and texture), higher soluble solids, better taste, improved color, and higher vitamin content. The objectives for fresh market tomatoes include enhancing overall flavor, sweetness, color, and health attributes. Genetic engineering is also being used to develop potatoes with more starch and less water to prevent damage when they are mechanically harvested. A potato with less water content may absorb less oil when it is fried, producing healthier french fries or potato chips.

    3.3. Floricultural Crops

    The global flower industry thrives on novelty. Genetic engineering is providing a valuable means of expanding the floriculture gene pool, therefore promoting the generation of new commercial varieties. The commercialization of genetically engineered flowers is currently confined to novel coloring. In general, engineered traits are valuable to either the consumer or the producer. At present only consumer traits appear able to provide a return capable of supporting what is still a relatively expensive molecular breeding tool. The biosynthesis of floral pigments, particularly anthocyanins, has been elucidated in great detail in model flowers such as petunia. This knowledge is now being applied to an understanding of a wide range of other flowers and provides a means of targeting color modification in these species. The expression of genes transferred across genera is not always predictable and so requires considerable trial and error to arrive at stable phenotypes of commercial interest. The manipulation of metabolic pathways, often requiring introduction of multiple genes, can also be problematic. This is a reflection of the complexity of interactions within and between cells at a gene and gene product level. An understanding of gene function is an essential step in engineering novel traits. The production of novel flower color has been the first success story in floriculture genetic engineering. Other traits that have received attention include floral scent, floral and plant morphology, senescence of flowers both on the plant and postharvest, and disease resistance.

    3.4. Medicinal and Aromatic Plants

    Biotechnology offers promising tools for the creation of novel crop varieties with improved nutritional value, resistance to herbicides, pests, diseases, pollutants, and adverse climatic conditions. Using RDT, it is now possible to manipulate the levels and composition of active pharmaceuticals and essential oils in medicinal and aromatic plants. However, to accomplish this task, it is first crucial to understand the biochemical pathways and the pattern of expression of the genes responsible for the synthesis of specific natural products. Novel pathways can be introduced into target plants to confer them with novel traits, and, in parallel, endogenous genes can be turned off using the proper DNA constructs. Thus it is possible to manipulate biosynthetic pathways to cause either the accumulation of valuable metabolites or to prevent their degradation. In addition, the organ specificity or temporal restrictions for the production of natural products can be overcome using genetic engineering. The way transgenic plants are obtained and how the implementation of molecular biology methodologies has improved crops and plant products is described. Exciting possibilities to use transgenic plants for the production of industrial enzymes and other materials, and different examples illustrating methods to either boost the levels of pharmacologically active compounds or change essential oil composition in medicinal and aromatic plants, using RDT are discussed. The genetic engineering is a new emerging opportunity for the full exploitation of the biosynthetic potential of aromatic and medicinal plants.

    4. Benefits of Commercialization of Transgenic Horticultural Crops in Developing Countries

    4.1. Increased Production and Productivity

    In 2014, the largest review yet concluded that GM crops’ effects on farming were positive. The study found that herbicide-tolerant crops have lower production costs, while for insect-resistant crops the reduced pesticide use was offset by higher seed prices, leaving overall production costs about the same. The yields increased 9% for herbicide tolerance and 25% for insect-resistant varieties. Farmers who adopted GM crops made 69% higher profits than those who did not. GM crops help farmers in developing countries, increasing yields by 14%. The researchers considered a number of studies that were not peer reviewed, and a few that did not report sample sizes. They attempted to correct for publication bias by considering sources beyond academic journals. The large data set allowed the study to control for potentially confounding variables such as fertilizer use.

    4.2. Enhanced Crop Protection

    The herbicide tolerance transgene confers tolerance to a specific herbicide. This trait allows farmers to apply an herbicide that acts on a wide range of weeds while not affecting the modified crop. Herbicide tolerance is currently the most commonly used GM trait worldwide. Herbicide-tolerant crops are mainly grown in developed countries with the primary aim of reducing applications of herbicides. The trait has also been achieved using other methods, particularly mutation breeding and gene transfer from wild relatives. In insect/pest resistance a transgene produces toxins to specific insects that feed on the crop. Such genes have been widely used and are already leading to substantial reductions in the use of pesticides and insecticides. Insect-resistant potato and maize varieties are being grown in both developed and developing countries. The biotech grapevine which is resistant to viral, bacterial, fungal diseases as well as abiotic stress tolerances were developed in South Africa. The other biotech crops like banana, apple, pear, and strawberry cultivars are under the development. The results of the international research done over the past 20 years is the development of HoneySweet plum highly resistant to PPV. Examples of crops in which these traits are being introduced include coffee, bananas, cassava, potato, sweet potato, beans, papaya, squash, and melon. In some cases, the transgenes used are genes that occur naturally in the same species. Abiotic stress resistance is the ability of some plants to survive in harsh climatic or soil conditions and is sometimes associated with specific groups of genes. These genes can be isolated and introduced into crops. Such applications promise to be particularly valuable for developing countries, where abiotic stresses such as drought, heat, frost, and acidic or salty soils are common. Transgenic research on crops such as coffee, potato, brassica, and tomato varieties are currently in different stages of development.

    4.3. Improvements in Shelf-Life

    Transgenic horticultural crops providing direct benefits to the

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