Market-Driven Plant Breeding for Practicing Breeders: Plant Breeding for Practicing Breeders 9789811954337, 9789811954344, 981195433X

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Table of contents :
Foreword 1
Foreword 2
Preface
Acknowledgments
Contents
About the Authors
1: Plant Breeding: Its Evolution and Recent Trends
1.1 Introduction
1.2 History and Evolution of Plant Breeding
1.3 Market Drivers for Plant Breeding Products
1.3.1 Advanced Technologies
1.3.2 Hybrid Seed Production
1.3.3 Decreasing Arable Land
1.3.4 Increasing Population
1.3.5 Globalization
1.3.6 Legal Framework
1.4 The Global Perspective on Plant Breeding Research in the Seed Business
1.5 The Indian Perspective on Plant Breeding Research in the Seed Business
1.5.1 Increasing Participation of the Private Sector in Plant Breeding
1.5.2 Trends in Plant Breeding Research in the Private Sector in India
1.5.2.1 Yield and Quality Parameters
1.5.2.2 Adaptability to Climatic Conditions
1.5.2.3 Commodity Specificity
1.5.2.4 Location
1.5.2.5 Postharvest Qualities
1.5.2.6 New Breeding Technologies (NBT)
1.5.2.7 Trait Novelty
1.6 Summing Up: The Need for this Book
References
2: Germplasm Management in Commercial Plant Breeding Programs
2.1 Introduction
2.2 Germplasm: Definition, and Scope in Breeding
2.3 Accessing Germplasm
2.3.1 Domestic and International Sources for Germplasm
2.3.1.1 GRIN and Germplasm Repositories, USA
2.3.1.2 Germplasm Bank at World Vegetable Centre, Taiwan
2.3.1.3 Germplasm Information, India
2.3.1.4 Other Sources of Germplasm
2.3.2 Accessibility and Quarantine
2.3.2.1 Accessibility
2.3.2.2 Quarantine
2.4 The National Biodiversity Act (NBA): Its Implications
2.5 Role of Consortiums in Providing Germplasm Access
2.5.1 Public/Private
2.5.2 Public/Public
2.5.3 Private/Private
2.6 Germplasm Collection and Curation Practices
2.6.1 Collection
2.6.2 Record Keeping
2.6.3 Curation
2.7 Germplasm Conservation, Evaluation, and Data Management
2.7.1 Stage 1: Seed Increase and Maintenance of Original Collections
2.7.2 Stage 2: Evaluation
2.7.3 Stage 3: Data Management
2.7.3.1 Manual Data Input and Analysis
2.7.3.2 Software-Based Data Input and Analysis
2.8 Germplasm Seed Inventory and Management
2.9 Legal Issues in Exploitation of Local/Indigenous Variety
2.9.1 Accessing Genetic Materials from the Public or Private Sector
2.9.1.1 Open Pollinated Varieties (OPVs)
2.9.1.2 Hybrids
2.10 Germplasm Improvement Projects: Case Studies of Impact
2.10.1 Self-pollinated Crops: Varietal Development
2.10.2 Self-pollinating Crops: Heterosis and Impact (as in Tomato)
2.10.3 Cross-Pollinated Crops
2.10.3.1 Seedless Watermelon
2.10.3.2 Improvement of Maize Germplasm
2.11 Pre-breeding
2.11.1 MLN Virus Resistance in Maize for Africa
2.11.2 Technology-Driven Breeding
2.12 Conclusions
References
3: Designing Plant Breeding Programs for Targeted Deliveries
3.1 Introduction
3.2 Plant Breeding in History
3.3 Designing Plant Breeding Programs Aligned with Organizational Objectives
3.3.1 Aligning Plant Breeding Programs with Commercial Strategies
3.3.2 Market Analysis to Predict Future Product Design Requirements
3.3.2.1 STEEP Analysis and Future Market Prediction
3.3.3 Designing a New Plant Variety Based on Predicted Market Needs
3.4 The Planning Process: Breeding Objectives and Processes
3.4.1 The New Variety Design
3.4.1.1 Planning a New Variety Design
3.4.1.2 Trait Profiling in New Variety Designing
3.4.2 Plant Breeding Process Mapping
3.4.2.1 Developing Progress Indicators
3.4.2.2 Focusing on Key Success Factors
3.4.3 Role of a Breeder in the Plant Breeding Process
3.5 Execution of a Planned Plant Breeding Program
3.5.1 Creating New Variability
3.5.1.1 Selection of Parental Lines
3.5.1.2 Choice of Mating Designs
3.5.1.3 Selection Methods
3.5.1.4 Selection Criteria
3.5.1.5 Selection Environment and Screening Methodology
3.5.2 Stabilizing the New Variation: Recombinant Plant to Recombinant Inbred Line
3.5.2.1 Nursery Management
3.5.2.2 Improving Breeding Process Efficiencies; Speed Breeding and Dihaploidy
3.5.2.3 Maintaining the New Recombinant Line
3.6 Hybrid Seed Production for Testing and Trialing
3.6.1 Hybrid Breeding
3.6.2 Crossing Block Management
3.6.3 Hybrid Testing
3.6.3.1 Stages of Testing-Initial, Advanced, and Multilocation
3.6.3.2 Data Generation and Management
3.6.3.3 Selection Environments
3.6.3.4 Hybrid Advancement Decisions
3.7 Conclusion
References
4: Inbred Line Development and Hybrid Breeding
4.1 Introduction
4.2 Why Hybrids?
4.3 Specifics of Commercial Plant Breeding
4.4 Inbred Line Development (ILD)
4.4.1 Doubled Haploidy (DH)
4.4.1.1 DH in Commercial Plant Breeding
4.4.1.2 Economic Impact of DH
4.4.1.3 Haploid Inducer Lines
4.4.1.4 Doubled-Haploid Breeding and In Vitro Mutagenesis in Brassicas
4.4.1.5 In vitro Androgenesis
4.4.1.6 DH in Cucurbits
4.4.1.7 DH Production Services
4.4.2 Fast Generation Cycling System (FGCS) Technology
4.4.2.1 Advantages of FGCS
4.5 Inbred Line Evaluation
4.6 Combining Ability (CA)
4.6.1 Relative Importance of Combining Ability
4.6.1.1 Early Generation Testing
4.6.2 Combining Ability Estimation Techniques
4.6.2.1 Diallel Mating Design
4.6.2.2 Line x Tester Design
4.6.2.3 North Carolina Design
4.7 Applications of the Concept of CA in Commercial Plant Breeding
4.7.1 CA for Yield
4.7.2 CA for Disease, Insect Pest, and Weed Resistance
4.7.3 CA and Germplasm Classification
4.7.4 CA and QTL Mapping
4.8 Breeding for Biotic and Abiotic Stress Resistances
4.8.1 Breeding for Abiotic Stress Resistances
4.8.1.1 Breeding for Drought Resistance
4.8.1.2 Breeding for Salinity Tolerance
4.8.1.3 Screening Techniques for Abiotic Stresses
4.8.2 Breeding for Biotic Stress Resistance
4.8.2.1 Disease Resistance Breeding
4.8.2.2 Breeding of ``Single Gene´´ Resistant Varieties
4.8.2.3 Alternative Strategies for Breeding Resistant Varieties
4.8.2.4 Challenges in the Breeding of Resistant Crops
4.8.2.5 Crop-specific Examples of Successes in Resistance Breeding
4.8.2.6 The Way Ahead
4.9 Pollination Control Mechanisms
4.9.1 Self-incompatibility (SI)
4.9.1.1 Applications of SI to Crop Production and Breeding
4.9.1.2 SI versus SC in Crop Production, Yield, and Quality
4.9.1.3 SI as an Alternative to Male Sterility for Developing Hybrids
4.9.1.4 SI in Hybrid Breeding Schemes
4.9.1.5 Achieving SC and Its Role in Hybrid Breeding
4.9.1.6 SI in Overcoming Interspecific Reproductive Barriers (IRBs)
4.9.1.7 Patented Applications of Plant SI Systems
4.9.1.8 Future Prospects of SI in Crop Breeding
4.10 Male Sterility
4.10.1 Male Sterility in Alliums
4.10.2 Male Sterility in Brassicas
4.10.3 Male Sterility in Carrot
4.10.4 Male Sterility in Maize
4.10.5 Male Sterility in Pearl Millet
4.10.6 Male Sterility in Peppers
4.10.7 Male Sterility in Sorghums
4.10.8 Male Sterility in Sunflower
4.10.9 Male Sterility in Rice
4.10.10 Future Prospects of Male Sterility
4.11 Breeding of Vegetable Crops: Success Stories
References
5: Use of Molecular Technologies in Plant Breeding
5.1 Overview of Marker-Assisted Breeding
5.1.1 Development of Markers
5.1.2 Molecular Marker Lab Resources
5.1.3 Plant Tissue Sampling Activity
5.1.4 Data Generation, Handling, and Interpretation
5.1.5 Marker Methodologies for Quantitative Traits
5.1.5.1 Linkage and Association Mapping Methods
5.1.6 Application of Markers in Plant Breeding
5.1.6.1 Diversity Analysis and Parent Selection
5.1.6.2 Heterotic Grouping
5.1.6.3 F1 or Hybridity Verification Tests
5.1.6.4 Marker-Assisted Selection (MAS)
5.1.6.5 Marker-Assisted Backcross Breeding (MABB)
5.1.6.6 Genomic Selection (GS)
5.1.6.7 Seed Purity Testing
5.1.6.8 DNA Fingerprinting for Varietal Identification
5.2 Transgenic Breeding
5.2.1 Creation of Transgenic Lines
5.2.1.1 Choosing the Gene of Interest
5.2.1.2 Gene Cloning
5.2.1.3 Plasmid Cassette Designing
5.2.1.4 Plant Cell Transformation
5.2.2 Event Selection
5.2.2.1 Criteria for Event Selection
5.2.2.2 Event Testing
5.2.2.3 Elite Line Development
5.2.2.4 Multi-stack of Events
5.2.2.5 Confined Field Trial (CFT) for Event Selection
5.2.3 Quality Control of GM Seed
5.2.4 GM Trait Regulations and the Registration Process
5.3 Genome Editing
5.3.1 CRISPR-Cas9 Method for Creating GE Plants
5.3.1.1 Applications
5.3.2 Regulations for Gene-Edited (GEd) Crop Varieties
5.4 Licensing of Traits
5.5 Introgression into Licensee Germplasm
5.6 Summary and Future Perspective
References
6: Product Development in Market-Driven Plant Breeding
6.1 Definition of Product Development
6.2 Key Elements of PD
6.2.1 PD Activities in the Seed Business
6.2.2 The Steps Involved in PD Activities
6.2.3 Seed and Logistics Requirements for Wide-Area Testing
6.2.4 Wide-Area Testing, Based on Mapping of Market Segments
6.2.5 Trial Calendar for Various Crops
6.2.6 Trait Dictionaries for Each Crop
6.2.7 Data Management and Conclusions
6.2.8 Operational Plan for Testing
6.2.9 Skills Development Plan for the Organization
6.2.10 Product Advancement Meetings
6.2.11 Registration of New Products: How and When
6.3 Preparing Operational Budgets for PD
6.4 Handover to Marketing and Sales
6.4.1 Role of PD
6.4.2 Role of Breeder
6.4.3 Role of Marketing
6.4.4 Role of Supply Chain
6.4.5 Role of Sales
6.5 Product Stewardship: GMO and Non-GMO
6.5.1 GMO Products
6.5.2 Non-GMO Products
6.6 Innovation in Product Development
6.7 Conclusion
References
7: Maintenance Breeding
7.1 Introduction
7.2 Monitoring of Hybrid/Product Performance
7.3 Maintenance Breeding of Parental Lines
7.3.1 Pollination Mechanisms
Box 7.1 Glossary of Abbreviated Terms Used in MB (Maintenance Breeding)
7.3.2 Timelines for Completion of the Maintenance Breeding Cycle
7.3.3 General Procedures of Maintenance Breeding in Annual Crops
7.3.4 Specific Deviations from General Practices in the Different Hybrid Systems of Annual Crops
7.3.5 Features of MB in Annual Crops Requiring Special Breeding Systems
7.3.5.1 Special Features of MB in Castor
7.3.5.2 Precautions in MB for Crops Requiring the CMS System
7.3.5.3 Special Features of MB in Crops Requiring the GMS System
7.3.5.4 Special Features of MB in Crops Requiring the TGMS System
7.3.6 General Procedure for MB in Biennial Crops
7.3.7 Specific Deviations from General Practice in Different Hybrid Systems of Biennial Crops
7.3.8 Special Features of MB in Some Breeding Systems of Biennial Crops
7.3.8.1 Precautions in MB in Brussels Sprout, Radish, and Cauliflower
7.3.8.2 Precautions in MB of Biennial Crops with the CMS System
7.3.8.3 Precautions in MB of Biennial Crops with the GMS System
7.3.8.4 Precaution in MB of Biennial Crops with the TGMS System
7.4 Conclusion
References
8: Seed Production Research
8.1 Introduction
8.2 SPR in Conventional Production Areas
8.2.1 Characteristics of Parents Determining Seed Production Potential
8.2.1.1 Seed Yield of the Female Parent
8.2.1.2 Agronomic and Seed Characteristics of the Female Parent
8.2.1.3 Agronomic Characteristics of the Male Line
8.2.1.4 Nicking
8.2.1.5 Seed Quality Features
8.2.2 Seed Production Research Strategy
8.2.2.1 Information on Seed Producibility of Hybrids
8.2.2.2 Data Generation and Recommendations
8.2.3 Practices at Sowing/Transplanting to Achieve Good Nicking at Flowering
8.2.3.1 Aged Seedling Transplanting
8.2.3.2 Seed Priming
8.2.3.3 Prediction and Adjustment of Synchronization
8.3 SPR in a New Area and/or New Season
8.3.1 Selection of New Area
8.3.2 Effect of Climatic Factors in Seed Production
8.3.2.1 Temperature
Box 8.1 Effect of High Temperature on Rice Seed Production
Box 8.2 Effect of High Temperature on Wheat Seed Production
Box 8.3 Effect of High Temperature on Maize Seed Production
Box 8.4 Effect of High Temperature on Cotton Seed Production
Box 8.5 Effect of High Temperature on Sorghum Seed Production
Box 8.6 Effect of High Temperature on Canola Seed Production
Box 8.7 Effect of High Temperature on Vegetable Seed Production
8.3.2.2 Precipitation
8.3.2.3 Photoperiod
8.3.2.4 Wind Velocity
8.3.2.5 Altitude
8.3.2.6 Seed Production Environment and Seed Quality
8.4 Sex Modifications Through Hormones and Chemicals in Vegetables
8.5 Breakdown of Incompatibility
8.6 Swapping of Parents
8.7 Pollen Preservation
8.7.1 Pollen Preservation Method
8.8 Agronomic Research in Seed Production Crops
8.9 Summing Up
References
9: Breeders´ Role in Supporting Allied Functions in the Seed Business
9.1 Introduction
9.2 Quality Seed Production
9.2.1 Parent Seed Production
9.2.2 Hybrid Seed/Certified Seed
9.2.3 New Product Development
9.2.4 Seed Quality Assurance
9.3 Stewardship
9.3.1 Assurance of Product Performance
9.4 Sales and Marketing
9.4.1 Competition Scanning
9.4.2 Third-Party Hybrid Evaluations
9.4.3 Product Registration
9.4.4 Market Development
9.4.5 Export of Seed
9.4.6 Consumer Complaint Redressal
9.5 Management
9.5.1 Germplasm Acquisition and Use
9.5.1.1 Germplasm Acquisition Under ITPGRFA and CBD
9.5.1.2 Germplasm Access Under the PBR/PVP System
9.5.1.3 Accessing Off-PVP Germplasm
9.5.1.4 Accessing Germplasm from the Open Source Seed Initiative (OSSI)
9.5.1.5 Accessing Germplasm from Research Collaborations
9.5.1.6 Accessing Germplasm via Licensing Agreements
9.5.1.7 Sharing Germplasm for Developing Joint Hybrids for Marketing
Box 9.1 SOP for Tracking the Existing Germplasm Presently Used in the Breeding Program
Box 9.2 SOP for Acquiring New Germplasm
9.5.2 Intellectual Property Rights (IPR) for the Hybrids/Parental Lines
9.5.2.1 Applying for PBR
9.5.3 Licensing
9.5.4 Business Planning
9.5.5 Mergers and Acquisitions
9.6 Summing Up
References
10: Regulatory Aspects of the Seed Business in Relation to Plant Breeding
10.1 Introduction
10.2 Regulations on the Seed Industry
10.2.1 Regulation of the Seed Industry in India
10.2.2 Some Important Laws Related to the Seed Business/Crop Breeding in India
10.2.3 Recognition of R&D Units
10.3 Regulations on Germplasm
10.3.1 The International Treaty on Plant Genetic Resources for Food and Agriculture (ITPGRFA)
Box 10.1 International Treaty on Plant Genetic Resources for Food and Agriculture (ITPGRFA)
Box 10.2 The International Treaty on Plant Genetic Resources for Food and Agriculture (ITPGRFA) and Consultative Group for Int...
Box 10.3 The ITPGRFA in India, with Reference to Agricultural Crops
Box 10.4 Convention on Biological Diversity (CBD) and the Nagoya Protocol
Box 10.5 The ITPGRFA and the Nagoya Protocol in India, with Reference to Agricultural Crops
Box 10.6 Benefit Sharing Under the Nagoya Protocol in India
10.3.2 Convention on Biological Diversity (CBD)
10.3.3 Intellectual Property Rights (IPR) Protection for Germplasm
10.3.3.1 Intellectual Property Rights (IPR) Protection for Germplasms in the USA
10.3.3.2 Intellectual Property Rights (IPR) Protection for Germplasm in India
10.3.4 Plant Breeder Rights (PBR)/Plant Variety Protection (PVP) System
10.3.5 Plant Breeders´ Rights (PBR) System in India
10.3.5.1 PBR for the Hybrids/Parental Lines
10.3.5.2 PBR for Essentially Derived Varieties (EDVs)
Box 10.7 Financial and Legal Implications of PVP in India
10.3.5.3 Applying for PBR in India
Box 10.8 Filing Process for PVP Registration in India
Box 10.9 The Data Compilation Process for PVP Registration in India
Box 10.10 Actions to be Taken for Filing and After Filing the PVP Application
Box 10.11 Vigilance on Violation of Rights and Defending Counterclaims
10.3.6 Registration of Germplasm in India
10.4 Germplasm Access
10.4.1 Germplasm Access Under the PBR/PVP System
10.4.2 Accessing Off-PVP Germplasm
10.4.3 Accessing Germplasm from the Open Source Seed Initiative (OSSI)
10.4.4 Accessing Germplasm from Research Collaborations
10.5 Crop Variety Registration
10.5.1 Crop Variety Registration in India
Box 10.12 Varietal Registration Regulation in Gujarat State, India
10.5.2 Release and Notification of Crop Cultivars in India
10.5.2.1 Evaluation
10.5.2.2 Identification
10.5.2.3 Release and Notification
10.5.3 OECD Scheme for the Varietal Certification of Seed
10.6 Import and Exports of Seed and Germplasm
10.6.1 Import/Export of Seed and Germplasm in/from India
Box 10.13 Regulations on Import of Seed/Germplasm in India
10.7 Regulations and the Registration Process for Genetically Modified (GM) Plants
10.7.1 GM Trait Regulations in India
10.7.1.1 Regulations on Initiating Research/Experimental Work on GM Crops
10.7.1.2 Safety Assessment Guidelines
10.7.1.3 GM Crops´ Release Procedures
10.7.1.4 Current Regulatory Scenario in India
10.7.2 GM Trait Regulations in the USA
10.7.3 GM Trait Regulations in European Union (EU) Countries
10.7.4 GM Trait Regulation in China and in Some Asian Countries
10.7.4.1 GM Trait Regulations in Bangladesh
10.7.4.2 GM Trait Regulations in Pakistan
10.7.4.3 GM Trait Regulations in Vietnam
10.7.4.4 GM Trait Regulations in the Philippines
10.7.4.5 GM Trait Regulation in Japan
10.7.4.6 GM Trait Regulation in Indonesia
10.7.5 GM Trait Regulation in Other Countries
10.7.5.1 GM Trait Regulation in Australia
10.7.5.2 GM Trait Regulation in New Zealand
10.7.5.3 GM Trait Regulation in Ethiopia
10.7.5.4 GM Trait Regulation in Brazil
10.7.5.5 GM Trait Regulation in Canada
10.8 Regulation of Genome Edited Crops
10.9 Licensing of Traits
10.10 Summing Up
References
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Aparna Tiwari · Surinder K. Tikoo · Sharan P. Angadi · Suresh B. Kadaru · Sadananda R. Ajanahalli M. J. Vasudeva Rao

Market-Driven Plant Breeding for Practicing Breeders

Market-Driven Plant Breeding for Practicing Breeders

Aparna Tiwari • Surinder K. Tikoo • Sharan P. Angadi • Suresh B. Kadaru • Sadananda R. Ajanahalli • M. J. Vasudeva Rao

Market-Driven Plant Breeding for Practicing Breeders

Aparna Tiwari Foundation for Advanced Training in Plant Breeding (ATPBR) Aurangabad, Maharashtra, India

Surinder K. Tikoo Tierra Agrotech Ltd Hyderabad, India

Sharan P. Angadi R&D Foundation for Advanced Training in Plant Breeding (ATPBR) Bangalore, India

Suresh B. Kadaru International Rice Research Institute Los Banos, Laguna, Philippines

Sadananda R. Ajanahalli Agreeva Technologies LLP Hyderabad, India

M. J. Vasudeva Rao GRSV Consulting Services Bangalore, India

ISBN 978-981-19-5433-7 ISBN 978-981-19-5434-4 https://doi.org/10.1007/978-981-19-5434-4

(eBook)

# The Editor(s) (if applicable) and The Author(s), under exclusive license to Springer Nature Singapore Pte Ltd. 2022 This work is subject to copyright. All rights are solely and exclusively licensed by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors, and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, expressed or implied, with respect to the material contained herein or for any errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. This Springer imprint is published by the registered company Springer Nature Singapore Pte Ltd. The registered company address is: 152 Beach Road, #21-01/04 Gateway East, Singapore 189721, Singapore

This Book was prepared under the aegis of ATPBR Foundation for Advanced Training in Plant Breeding

Foreword 1

Agriculture in India has been essentially a production-driven system for the past many decades, as it is in many developing countries. As we struggled to feed a growing population, the emphasis was producing more at affordable prices. Faced with shortages, the country had to focus on increasing production, and rightly so. But the situation has also been changing simultaneously. With the rapid increase in per capita income in India from Rs. 25,987 in 2004–2005 to Rs. 1,28,829 in 2020– 2021 at current prices (source: Ministry of Statistics & Programme Implementation GOI, World Bank and IMF World Economic Outlook: https://statisticstimes.com/ economy/country/india-gdp-per-capita.php), the purchasing power of the common man has increased considerably. This is expected to grow by 58% in the next 5 years. The recent 15-year period has witnessed consumer preferences leaning towards private markets. In many fields, modern digital technology and big data analytics have made it possible for businesses to offer tailor-made products to suit consumer needs in different market segments. Food cannot lag behind. The bottom of the pyramid still needs a large volume of food at affordable prices, thereby continuing the need for our agriculture to focus on higher yields and stress tolerance. However, the need for differentiated food products for the population with higher incomes has become markedly evident in the last 10–15 years, thereby propelling the need for market-driven agriculture, which meets the needs of different market segments. In this context, market-driven plant breeding is essential to support market-driven agriculture. I am very happy that this book is coming out at a crucial juncture in the vii

viii

Foreword 1

development of agricultural research, which is gearing itself to meet future challenges posed by food and nutrition security, climate change, changing food baskets, depletion of natural resources, stagnant yields, farm mechanization, digital transformation of agriculture, and other similar changes. The stupendous achievements of plant breeding in improving crops have been laudable. The world in general—and India in particular—has hugely benefited by the advances in plant breeding and plant biotechnology. But, if we observe closely, the most successful crop varieties are those which met either felt or latent needs in the market. While some varieties may have succeeded mainly due to marketing efforts, most of them have succeeded because they fitted well—by design or by accident— with the needs of the customer. Farmers are running a production system that is aimed at meeting the requirements of the consumers. In the value chain of a crop, the players include farmers, primary and secondary processors, traders, and the consumer. Each of them has his/her own expectations or requirements, which may be different from others. Understanding and incorporating these requirements for maximizing the synergistic impact of the germplasm is important. It is critical that breeders stay connected to the markets, which in their entirety include farmers and consumers, as well as the players in the value chain in between. If the breeders are developing seed varieties for export markets, the farmers, consumers, and value chain players in those countries become the customers. Similarly, if the breeding is aimed at improving crops to make them suitable for export purposes, either as agricultural produce or as processed food, breeders need to understand the requirements of those market segments and tailor the varieties accordingly. We cannot ignore the role of the society in acceptance of technologies, especially those which are used in agriculture and the manufacture of food. Hence, it is also a politically sensitive subject. Plant breeders and biotechnologists need to always keep in mind the social acceptance of their products. In this context, social scientists play an important role. It may be necessary for seed technology scientists to consult social scientists while developing technologies or products that are different from the traditional ones. It is important that breeders in both the public and the private sectors keep the market requirements in mind. In the private sector, the planning process takes care of some of this aspect, as the companies are closely connected to the markets of farmers. They are still not fully connected, however, to the ultimate consumers, food companies, and exporters. This gap needs to be bridged by forming collaborative breeding programs with the food industry. The seed industry should also participate in consumer surveys conducted to understand the likely future trends in consumption. This is a highly dynamic area and needs constant monitoring. As the CEO of a seed company, I always spent considerable time with breeding teams when they were setting up the objectives of their breeding programs. This is a crucial part of the preparation of our research teams to cater to the needs of the market. Breeders need to visit markets regularly and interact with farmers, processors, and exporters to understand the trends. Within the organization, whether

Foreword 1

ix

public or private, there should be an institutionalized process of connecting markets to the breeders. The formation of teams for each crop, which bring together marketing and seed production personnel with the breeders of that crop, helps in this process, as observed by me during my career in the seed industry. Functions should not operate in silos. Biotechnology has had—and will have—a huge impact on breeding strategies and possibilities. As human endeavor advances its efforts to find answers to evernew challenges that come up, there will be greater technological developments in the future. They will facilitate market-driven plant breeding in a more precise and quicker fashion. However, it is the plant breeder who needs to have that “eye” for the continuously changing market requirements, so that he/she can utilize the new technological tools for a well-directed research effort. Some of those aspects find detailed explanation in this book. I have no doubt that the book will be extremely useful for breeders involved in crop improvement programs. The authors have made a tremendous effort to incorporate significant knowledge, which makes the book a “must have” asset in your portfolio. This results from their vast experience in the field, and from their ability to convert all that knowledge into useful guidelines for breeders, business managers, policymakers, marketing professionals, and other stakeholders. This is the first book of its kind in this field. I am confident that many agri-input industry professionals will benefit from it. I congratulate the authors for bringing out this invaluable book. Federation of Seed Industry in India (FSII) New Delhi, India

Ram Kaundinya

Foreword 2

It is a privilege and pleasure to write the foreword for this book. I consider it futuristic in the present-day context, where plant breeding is becoming increasingly important to meet the demands for food and nutrition security for the ever-increasing global population, which is expected to reach nine billion by 2050. During my Master’s degree program in Plant Breeding (at the GB Pant University of Agriculture and Technology, Pantnagar, Uttarakhand, India) in the early 1970s, I considered plant breeding to be the same in public sector institutions and in private sector seed companies. However, over the years, after working with private sector seed companies (during my career with ICRISAT, the International Crops Research Institute for the Semi-Arid Tropics, Hyderabad, India), my perception has changed. The discussions and deliberations for the formation of the ICRISAT-Private Sector Hybrid Parents Research Consortia (HPRC) were highly educative. I was able to understand the importance and the need for commercial or market-driven plant breeding to be different from the public sector plant breeding efforts. Broadly speaking, plant breeding can be defined as “fast-track evolution of economically important plant species to develop improved varieties, which produce better quality and quantity of the produce for food, feed, and fodder to enhance quality of life in human beings.” Working in an International public sector research and development (R&D) institute, (i.e., ICRISAT), plant breeding was considered a public good activity that produces International Public Goods (IPGs), including early-generation and xi

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advanced-generation breeding materials, which could then be shared with both public sector institutions and private sector seed companies, for them to further develop and release the breeding materials as improved crop cultivars (varieties/ hybrids) to the farming community. The private sector translates public goods from publicly funded research institutions into private sector assets, which can earn profits for their companies. Plant breeding in the private sector is different from that in the public sector for the following considerations: (1) market-oriented approach; (2) profit-driven goals; (3) speed and cost-effectiveness of the breeding efforts; (4) farmer-centric, targeted trait development; and (5) clarity of end-user needs in the selection of traits. While efforts in the public sector aim at releasing a variety, the private sector plant breeders’ efforts continue to be invested on seed production and on placing the seed in the market at the right time and place, and at a price that is affordable to the targeted farmer. The private sector has an effective network of seed producers, as well as seed distributors, in the marketing system. That is how they succeed in marketing the varieties or hybrids that they produce. The private sector has also invested in strategic research and development as and when needed, including in biotechnology, so that they can always stay ahead of the competitors. The book on Market-Driven Plant Breeding has been well conceived, and the chapters in the book are well thought out; they cover the most critical aspects of commercial plant breeding programs. In the past four decades, we have seen large investments by the private sector seed companies not only into basic and strategic research, but also on practical, adaptive, and applied research. Many new disruptive technologies that include cutting-edge technologies are being deployed from both information technology and biotechnology spheres. These new-age research methods and technologies have tremendously helped both the speed and accuracy of plant breeding, and the return on investment has been very good. The ten chapters in the book cover a range of highly relevant aspects and issues, as they relate to market-oriented plant breeding. A major consideration in marketdriven plant breeding is the need to retain productive staff for a long time because plant breeding is a long-term investment, and it will need committed staff to ensure that the products are delivered on time to the farmer. Despite the efforts by the private sector seed companies to retain committed and high-performing plant breeders, a high attrition rate and poaching within the industry have been common. Many private sector seed companies are investing in continuous training and in upgrading the skills of their plant breeders. That effort must be strengthened further under the Foundation for Advanced Training of Plant Breeders (ATPBR). The effort should also ensure that, in addition to the staff in many private sector companies, ATPBR can also train the public sector plant breeders. This book can be an effective instrument in such efforts. ICRISAT Patancheru, India

C. L. Laxmipathi Gowda

Preface

Plant breeding, the science of altering heritable patterns of economically important traits of plants to increase their value, is broadly defined as purposeful manipulation of plant genotypes to create desirable phenotypes for the benefit of mankind. A science-driven, inventive process of genetically improving plant varieties culminates in cultivar development and crop improvement, via improvement of the seed. It encompasses the creation of multi-generation, genetically diverse populations on which human knowledge and selection skills are applied to generate genetically altered plants, with precise new combinations of desirable traits. Commercial-scale plant breeding is a multifaceted process in which new crop varieties are continuously developed to enhance their yield and agronomic performance over the prevalent varieties. Plant breeders exercise planned and extensive selection practices to identify top performing plants with desired trait combinations, while minimizing inadvertent unintended changes. In our current times, there is a pressing need to reorient plant breeding to create “smart” crop varieties that yield more with fewer inputs, which are also resilient to climate change. In addition, plant breeding helps to achieve food security and sustainability through the development of high-yielding, climate-resilient, nutritious, improved crop varieties; it thereby addresses the challenges posed by the needs of a growing global population, shrinking genetic diversity, and uncertain environmental conditions. Both public and private sector organizations have contributed significantly to the development of improved varieties, which have had a strong positive impact on the economic well-being of its stakeholders (farmers, vendors, seed producers, marketing personnel, researchers, etc.), as well as on employment, environmental protection, and biodiversity conservation. Over the last three to four decades, there has been a significant increase in plant breeding activities in the private sector across the globe. On the one hand, this reflects the integration of advanced technologies into conventional plant breeding processes, the need for intellectual property protection of new plant varieties, and the globalization of seed trade. On the other hand, it is a consequence of shrinking or stagnant budgets devoted to applied plant breeding in the public sector. Historically, the private sector has developed effective and efficient mechanisms to translate public sector research and innovations into commercially viable plant xiii

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breeding products, delivered through seeds and/or other planting materials. Further, more recent efforts in the private sector have been devoted to efficient utilization of technological tools, as well as wider geographic and commodity coverage, involving greater fiscal commitment towards strengthening R&D capabilities to produce appropriate innovations. Those efforts have made it a front runner in the acceleration of plant breeding programs, development of improved varieties, and dissemination of improved varieties through scalable, high-quality seed delivery systems. The outcomes sought and successes achieved by plant breeding in the future will be measured in terms of sustainable improvements in agricultural production of food, feed, fiber, fuel, and other plant products that meet the needs of society. Since plant breeding research in the private sector is driven by ROI (Return on Investment) and changing market preferences, plant breeders need to work very efficiently to deliver targeted outputs, under stricter timelines and clearly defined budgets, so that new varieties become available in the shortest possible time. To do so effectively, private sector plant breeding needs to respond quickly to changes in the market requirements and to spot new opportunities for improved plant varieties and hybrids as they occur. And despite the growing private sector involvement in plant breeding, presence of the public sector will continue to be crucial in the areas of plant breeding education, pre-breeding, new germplasm development, and conservation. Over the past 30–40 years, private sector investments in R&D have increased substantially, and so has the requirement of skilled plant breeders. To build the capability in human skills required for crop improvement in the future, a new generation of plant breeders must be trained. In India, the private sector engages a major part of the existing workforce of plant breeders, but it needs trained plant breeders in greater numbers to augment breeding capabilities, and thus meet the growing demand of food, as well as plant-derived products, for an ever-increasing population. The current availability of skilled breeders is inadequate to fully capitalize on the benefits from available plant genetic resources. To meet the growing need for plant breeders at regional, national, and global levels, it is essential both to train the existing workforce in the best practices and to develop a stream of highly skilled breeders to serve the growing needs of the industry. Plant breeders must be trained with an integrated set of capabilities to build and sustain effective regional, national, and global plant breeding capacities. Structured education and training are the traditional methods to identify and develop breeders. There is a need to update plant breeding education programs, both to meet the current needs of a rapidly expanding job market and to provide the requisite knowledge and skill for the future. The authors have attempted to elucidate the finer aspects of plant breeding practices, as carried out in a commercially driven seed business, based on their collective experience over a substantial period of working in both the public and private sectors, nationally and internationally. This book encompasses their accumulated understanding of a lifetime, as it relates to practical and successful plant breeding. The ten chapters in the book cover a range of topics from the evolution of plant breeding to recent trends, including contemporary regulations governing plant

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breeding. The range encompasses germplasm management; designing breeding programs for targeted deliveries; inbred line development and hybrid breeding; use of molecular technologies; product development; maintenance breeding; seed production research; and the role of breeders in related functions of the seed value chain. Each chapter contains detailed explanations, integrating knowledge and experience, for both potential and practicing plant breeders. Given the diverse experience of the authors, there is a diversity in approaches to their topics as well, and no attempt has been made to impose (or even arrive at by discussion) a consistency or uniformity of approach across the chapters. The book highlights plant breeding in a competitive market-driven environment, and it describes the crucial aspects of the seed value chain. We earnestly hope that this book will prove to be a valuable source of practical knowledge, and thus help readers to effectively accelerate their plant breeding efforts in a market-driven world. Aurangabad, Maharashtra, India Hyderabad, Telangana, India Bangalore, Karnataka, India Los Banos, Laguna, Philippines Hyderabad, India Bangalore, Karnataka, India

Aparna Tiwari Surinder K. Tikoo Sharan P. Angadi Suresh B. Kadaru Sadananda R. Ajanahalli M. J. Vasudeva Rao

Acknowledgments

Thinking about writing a book and converting that idea into a book are two different sides of a coin. Writing a book is an emotional journey, and this experience is challenging as well as rewarding. The authors were inspired by their brilliant friends, colleagues, and family members to deliver their best experience in the form of this book. The authors wish to record thanks to the following (mostly unseen) contributors. Dr. Aparna Tiwari would like to thank her parents Dr. Om Prakash Tiwari and Mrs. Archana Tiwari, as well as Mr. Ravi Datt Mishra and Mrs. Gita Mishra, for their continuous support and encouragement. She also would like to extend her thanks to her precious gems Master Varchas Mishra and Ms. Vanshika Mishra for their unspoken support, so that she should commit more hours to work. Finally, she wants to thank her husband, Dr. Arvind Mishra, for believing in her and for tolerating her incessant disappearances into her home office. Dr. Sadananda A. R. would like to acknowledge the continuous inspiration and support of his wife, Poornima Sadananda, and for her tolerating the loneliness from the long hours of his desk work. He also thanks his children, Abhijith and Vishwajith, and their better halves, Dr. Dwitiya Sawant and Dr. Malavika Prasad, for their continued inspiration in his penning of his lifelong experience of crop breeding. His special thanks also go to his 4-year-old, cute, granddaughter, Reeva Saima Sadananda, for kindling the childish wish of finding something new in every passing moment of life, by her virtual interaction on a daily basis. He is also very grateful to his beloved parents and all his brothers, for allowing him to pursue his chosen field of work and their constant encouragement. He is thankful to many ex-colleagues and managers of his more than 40-year professional career, for the exposure to different roles and responsibilities in different facets of a plant breeder’s profession. His special thanks also go to many of his ex-colleagues and friends, from both private seed companies and academic institutions, who took time to review the manuscripts and provided very useful suggestions. His special thanks also go to Dr. Sharan Angadi for inviting him to contribute to this book, and to Dr. Vasudev Rao, who, as a mentor of over 50 years, always encouraged him with his critical insights and inputs. Dr. Sharan Angadi would like to thank Aparna Tiwari for initiating the book idea, supporting the preparation of the manuscript, and pursuing each and every step with xvii

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the publishers, under the aegis of ATPBR. Thanks to M. J. Vasudeva Rao for his enthusiastic support in giving material shape to the concept, detailing the contents, keeping track of the progress, and providing valuable inputs, and to Suren Tikoo and Sadananda, for reading through the manuscript and suggesting improvements. Thanks to my wife Sunanda and daughter Sahana, for their constant support. Dr. Suresh Kadaru would like to thank his wife Sudha, daughter Tanmai, son Tanay, and his parents for their constant support and encouragement throughout his scientific career. He wants to admit that his scientific endeavor might not have been possible without Syngenta, and he extends his sincere gratitude for the learnings, opportunities, and encouragement it provided all these years. He also expresses his thanks to all the authors of this book for their help, patience, and constructive suggestions during the chapter preparation. Dr. Surinder K. Tikoo would like to thank his worthy parents, Late Mr. Prem Nath Tikoo and Mrs. Premrani Tikoo, for always supporting him to pursue his field of work; his wife Renu for a four plus decades of loving companionship and continuous inspiration to do his best; his son Dhiren, daughter-in-law Aarti, daughter Smita and grandson Aryaman, for being a source of joy and for their constant encouragement. Thanks are also due to his sisters, Vineeta, Ranjana, and Veena and brothers, Narender and Late Ravi, and their families and close friends and associates, for being always supportive. Finally, he would like to dedicate this effort to his youngest brother and joy of the family, Ravi, who was snatched away from the family due to Covid on 27 April 2021. Dr. Vasudeva Rao would like to express his heartfelt thanks to his family members, wife Champa, daughter Anupama, son Dr. Abhay, and grandchildren, Mihir, Neil, and Nia. It was their love, encouragement, and support which helped him to stay on course. He is especially thankful to the unstinted support from coauthors of this book, and close friends who helped him to coordinate the book successfully. He expresses his deepest gratitude to all his teachers and several wellknown plant breeders, with whom he worked across institutions in the public and private sectors, and who have inspired him to excel. The authors collectively express their deep sense of gratitude to Dr. D.R. Mohan Raj, who has long and significant experience in editing documents relating to international agricultural research, and who edited many of these books. He was a crucially important part of our efforts. In some instances, he converted raw drafts from relatively “inexperienced authors” into coherent contents, through his refined editorial skills; in other cases, he made valuable suggestions for rewriting or reorganizing materials, which would help the book reach a larger and diverse audience more effectively. We are deeply indebted to him for his untiring efforts and his disciplined approach, which set a high professional standard for our team in this publication endeavor. The authors are indebted to Mr. Ram Kaundinya (Director General, FSII) and Dr. C. L. Laxmipathi Gowda (Formerly DDG, ICRISAT, Hyderabad) for their keen insights on the topic and for writing the Forewords. We value their testimonial, as we hope the potential readers of this books will do as well.

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The authors would like to extend their thanks to the entire Springer team, who helped in the publishing of this book. Special thanks to Ms. Aakanksha Tyagi, Senior Editor (Books), Life Sciences, for all her patience and cooperation. Lastly, this book is dedicated to all those who strive to grow and help others to grow. Following authors have contributed specific chapters as mentioned against their names in the book: Chapter 1. Plant Breeding: Its Evolution and Recent Trends - Aparna Tiwari Chapter 2. Germplasm Management in Commercial Plant Breeding Programs Surinder K. Tikoo Chapter 3. Designing Plant Breeding Programs for Targeted Deliveries - M. J. Vasudeva Rao Chapter 4. Inbred Line Development and Hybrid Breeding - Sharan Angadi Chapter 5. Use of Molecular Technologies in Plant Breeding - Suresh B. Kadaru and Sadananda R. Ajanahalli Chapter 6. Product Development in Market-Driven Plant Breeding - Surinder K. Tikoo Chapter 7. Maintenance Breeding - Sadananda R. Ajanahalli Chapter 8 Seed Production Research - Sadananda R. Ajanahalli Chapter 9. Breeders’ Role in Supporting Allied Functions in the Seed Business Sadananda R. Ajanahalli Chapter 10. Regulatory Aspects of the Seed Business in Relation to Plant Breeding - Sadananda R. Ajanahalli —The Authors

Contents

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Plant Breeding: Its Evolution and Recent Trends . . . . . . . . . . . . . 1.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.2 History and Evolution of Plant Breeding . . . . . . . . . . . . . . . . 1.3 Market Drivers for Plant Breeding Products . . . . . . . . . . . . . 1.3.1 Advanced Technologies . . . . . . . . . . . . . . . . . . . . 1.3.2 Hybrid Seed Production . . . . . . . . . . . . . . . . . . . . 1.3.3 Decreasing Arable Land . . . . . . . . . . . . . . . . . . . . 1.3.4 Increasing Population . . . . . . . . . . . . . . . . . . . . . . 1.3.5 Globalization . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.3.6 Legal Framework . . . . . . . . . . . . . . . . . . . . . . . . . 1.4 The Global Perspective on Plant Breeding Research in the Seed Business . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.5 The Indian Perspective on Plant Breeding Research in the Seed Business . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.5.1 Increasing Participation of the Private Sector in Plant Breeding . . . . . . . . . . . . . . . . . . . . . . . . . 1.5.2 Trends in Plant Breeding Research in the Private Sector in India . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.6 Summing Up: The Need for this Book . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Germplasm Management in Commercial Plant Breeding Programs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.2 Germplasm: Definition, and Scope in Breeding . . . . . . . . . . . 2.3 Accessing Germplasm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.3.1 Domestic and International Sources for Germplasm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.3.2 Accessibility and Quarantine . . . . . . . . . . . . . . . . . 2.4 The National Biodiversity Act (NBA): Its Implications . . . . . 2.5 Role of Consortiums in Providing Germplasm Access . . . . . . 2.5.1 Public/Private . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.5.2 Public/Public . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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2.5.3 Private/Private . . . . . . . . . . . . . . . . . . . . . . . . . . . . Germplasm Collection and Curation Practices . . . . . . . . . . . . . 2.6.1 Collection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.6.2 Record Keeping . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.6.3 Curation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.7 Germplasm Conservation, Evaluation, and Data Management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.7.1 Stage 1: Seed Increase and Maintenance of Original Collections . . . . . . . . . . . . . . . . . . . . . . 2.7.2 Stage 2: Evaluation . . . . . . . . . . . . . . . . . . . . . . . . . 2.7.3 Stage 3: Data Management . . . . . . . . . . . . . . . . . . . 2.8 Germplasm Seed Inventory and Management . . . . . . . . . . . . . 2.9 Legal Issues in Exploitation of Local/Indigenous Variety . . . . . 2.9.1 Accessing Genetic Materials from the Public or Private Sector . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.10 Germplasm Improvement Projects: Case Studies of Impact . . . 2.10.1 Self-pollinated Crops: Varietal Development . . . . . . 2.10.2 Self-pollinating Crops: Heterosis and Impact (as in Tomato) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.10.3 Cross-Pollinated Crops . . . . . . . . . . . . . . . . . . . . . . 2.11 Pre-breeding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.11.1 MLN Virus Resistance in Maize for Africa . . . . . . . . 2.11.2 Technology-Driven Breeding . . . . . . . . . . . . . . . . . 2.12 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.6

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Designing Plant Breeding Programs for Targeted Deliveries . . . . . 3.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.2 Plant Breeding in History . . . . . . . . . . . . . . . . . . . . . . . . . . 3.3 Designing Plant Breeding Programs Aligned with Organizational Objectives . . . . . . . . . . . . . . . . . . . . . . . . . . 3.3.1 Aligning Plant Breeding Programs with Commercial Strategies . . . . . . . . . . . . . . . . . . . . . 3.3.2 Market Analysis to Predict Future Product Design Requirements . . . . . . . . . . . . . . . . . . . . . . 3.3.3 Designing a New Plant Variety Based on Predicted Market Needs . . . . . . . . . . . . . . . . . . 3.4 The Planning Process: Breeding Objectives and Processes . . . 3.4.1 The New Variety Design . . . . . . . . . . . . . . . . . . . . 3.4.2 Plant Breeding Process Mapping . . . . . . . . . . . . . . 3.4.3 Role of a Breeder in the Plant Breeding Process . . . 3.5 Execution of a Planned Plant Breeding Program . . . . . . . . . . 3.5.1 Creating New Variability . . . . . . . . . . . . . . . . . . . 3.5.2 Stabilizing the New Variation: Recombinant Plant to Recombinant Inbred Line . . . . . . . . . . . . .

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Hybrid Seed Production for Testing and Trialing . . . . . . . . . . 3.6.1 Hybrid Breeding . . . . . . . . . . . . . . . . . . . . . . . . . . 3.6.2 Crossing Block Management . . . . . . . . . . . . . . . . . 3.6.3 Hybrid Testing . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.7 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Inbred Line Development and Hybrid Breeding . . . . . . . . . . . . . . 4.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.2 Why Hybrids? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.3 Specifics of Commercial Plant Breeding . . . . . . . . . . . . . . . . 4.4 Inbred Line Development (ILD) . . . . . . . . . . . . . . . . . . . . . . 4.4.1 Doubled Haploidy (DH) . . . . . . . . . . . . . . . . . . . . 4.4.2 Fast Generation Cycling System (FGCS) Technology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.5 Inbred Line Evaluation . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.6 Combining Ability (CA) . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.6.1 Relative Importance of Combining Ability . . . . . . . 4.6.2 Combining Ability Estimation Techniques . . . . . . . 4.7 Applications of the Concept of CA in Commercial Plant Breeding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.7.1 CA for Yield . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.7.2 CA for Disease, Insect Pest, and Weed Resistance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.7.3 CA and Germplasm Classification . . . . . . . . . . . . . 4.7.4 CA and QTL Mapping . . . . . . . . . . . . . . . . . . . . . 4.8 Breeding for Biotic and Abiotic Stress Resistances . . . . . . . . 4.8.1 Breeding for Abiotic Stress Resistances . . . . . . . . . 4.8.2 Breeding for Biotic Stress Resistance . . . . . . . . . . . 4.9 Pollination Control Mechanisms . . . . . . . . . . . . . . . . . . . . . . 4.9.1 Self-incompatibility (SI) . . . . . . . . . . . . . . . . . . . . 4.10 Male Sterility . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.10.1 Male Sterility in Alliums . . . . . . . . . . . . . . . . . . . . 4.10.2 Male Sterility in Brassicas . . . . . . . . . . . . . . . . . . 4.10.3 Male Sterility in Carrot . . . . . . . . . . . . . . . . . . . . . 4.10.4 Male Sterility in Maize . . . . . . . . . . . . . . . . . . . . . 4.10.5 Male Sterility in Pearl Millet . . . . . . . . . . . . . . . . . 4.10.6 Male Sterility in Peppers . . . . . . . . . . . . . . . . . . . . 4.10.7 Male Sterility in Sorghums . . . . . . . . . . . . . . . . . . 4.10.8 Male Sterility in Sunflower . . . . . . . . . . . . . . . . . . 4.10.9 Male Sterility in Rice . . . . . . . . . . . . . . . . . . . . . . 4.10.10 Future Prospects of Male Sterility . . . . . . . . . . . . . 4.11 Breeding of Vegetable Crops: Success Stories . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Use of Molecular Technologies in Plant Breeding . . . . . . . . . . . . . . 5.1 Overview of Marker-Assisted Breeding . . . . . . . . . . . . . . . . . 5.1.1 Development of Markers . . . . . . . . . . . . . . . . . . . . . 5.1.2 Molecular Marker Lab Resources . . . . . . . . . . . . . . 5.1.3 Plant Tissue Sampling Activity . . . . . . . . . . . . . . . . 5.1.4 Data Generation, Handling, and Interpretation . . . . . 5.1.5 Marker Methodologies for Quantitative Traits . . . . . . 5.1.6 Application of Markers in Plant Breeding . . . . . . . . . 5.2 Transgenic Breeding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.2.1 Creation of Transgenic Lines . . . . . . . . . . . . . . . . . . 5.2.2 Event Selection . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.2.3 Quality Control of GM Seed . . . . . . . . . . . . . . . . . . 5.2.4 GM Trait Regulations and the Registration Process . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.3 Genome Editing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.3.1 CRISPR-Cas9 Method for Creating GE Plants . . . . . 5.3.2 Regulations for Gene-Edited (GEd) Crop Varieties . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.4 Licensing of Traits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.5 Introgression into Licensee Germplasm . . . . . . . . . . . . . . . . . . 5.6 Summary and Future Perspective . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

157 157 158 163 164 165 168 170 181 182 186 193

Product Development in Market-Driven Plant Breeding . . . . . . . . 6.1 Definition of Product Development . . . . . . . . . . . . . . . . . . . 6.2 Key Elements of PD . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.2.1 PD Activities in the Seed Business . . . . . . . . . . . . 6.2.2 The Steps Involved in PD Activities . . . . . . . . . . . 6.2.3 Seed and Logistics Requirements for Wide-Area Testing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.2.4 Wide-Area Testing, Based on Mapping of Market Segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.2.5 Trial Calendar for Various Crops . . . . . . . . . . . . . . 6.2.6 Trait Dictionaries for Each Crop . . . . . . . . . . . . . . 6.2.7 Data Management and Conclusions . . . . . . . . . . . . 6.2.8 Operational Plan for Testing . . . . . . . . . . . . . . . . . 6.2.9 Skills Development Plan for the Organization . . . . . 6.2.10 Product Advancement Meetings . . . . . . . . . . . . . . 6.2.11 Registration of New Products: How and When . . . . 6.3 Preparing Operational Budgets for PD . . . . . . . . . . . . . . . . . 6.4 Handover to Marketing and Sales . . . . . . . . . . . . . . . . . . . . . 6.4.1 Role of PD . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.4.2 Role of Breeder . . . . . . . . . . . . . . . . . . . . . . . . . .

205 205 206 206 210

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193 194 194 196 196 197 198 199

. 211 . . . . . . . . . . . .

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Contents

6.4.3 Role of Marketing . . . . . . . . . . . . . . . . . . . . . . . . 6.4.4 Role of Supply Chain . . . . . . . . . . . . . . . . . . . . . . 6.4.5 Role of Sales . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.5 Product Stewardship: GMO and Non-GMO . . . . . . . . . . . . . 6.5.1 GMO Products . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.5.2 Non-GMO Products . . . . . . . . . . . . . . . . . . . . . . . 6.6 Innovation in Product Development . . . . . . . . . . . . . . . . . . . 6.7 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

8

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230 231 231 232 232 233 234 236 237

Maintenance Breeding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.2 Monitoring of Hybrid/Product Performance . . . . . . . . . . . . . . 7.3 Maintenance Breeding of Parental Lines . . . . . . . . . . . . . . . . . 7.3.1 Pollination Mechanisms . . . . . . . . . . . . . . . . . . . . . 7.3.2 Timelines for Completion of the Maintenance Breeding Cycle . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.3.3 General Procedures of Maintenance Breeding in Annual Crops . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.3.4 Specific Deviations from General Practices in the Different Hybrid Systems of Annual Crops . . . 7.3.5 Features of MB in Annual Crops Requiring Special Breeding Systems . . . . . . . . . . . . . . . . . . . . 7.3.6 General Procedure for MB in Biennial Crops . . . . . . 7.3.7 Specific Deviations from General Practice in Different Hybrid Systems of Biennial Crops . . . . . . . 7.3.8 Special Features of MB in Some Breeding Systems of Biennial Crops . . . . . . . . . . . . . . . . . . . 7.4 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

239 239 242 243 244

Seed Production Research . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.2 SPR in Conventional Production Areas . . . . . . . . . . . . . . . . . 8.2.1 Characteristics of Parents Determining Seed Production Potential . . . . . . . . . . . . . . . . . . . . . . . 8.2.2 Seed Production Research Strategy . . . . . . . . . . . . 8.2.3 Practices at Sowing/Transplanting to Achieve Good Nicking at Flowering . . . . . . . . . . . . . . . . . . 8.3 SPR in a New Area and/or New Season . . . . . . . . . . . . . . . . 8.3.1 Selection of New Area . . . . . . . . . . . . . . . . . . . . . 8.3.2 Effect of Climatic Factors in Seed Production . . . . . 8.4 Sex Modifications Through Hormones and Chemicals in Vegetables . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.5 Breakdown of Incompatibility . . . . . . . . . . . . . . . . . . . . . . .

246 246 246 251 261 265 265 269 270

. 273 . 273 . 275 . 276 . 278 . . . .

285 286 288 291

. 298 . 298

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8.6 8.7

Swapping of Parents . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pollen Preservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.7.1 Pollen Preservation Method . . . . . . . . . . . . . . . . . . 8.8 Agronomic Research in Seed Production Crops . . . . . . . . . . . 8.9 Summing Up . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

10

Breeders’ Role in Supporting Allied Functions in the Seed Business . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9.2 Quality Seed Production . . . . . . . . . . . . . . . . . . . . . . . . . . . 9.2.1 Parent Seed Production . . . . . . . . . . . . . . . . . . . . . 9.2.2 Hybrid Seed/Certified Seed . . . . . . . . . . . . . . . . . . 9.2.3 New Product Development . . . . . . . . . . . . . . . . . . 9.2.4 Seed Quality Assurance . . . . . . . . . . . . . . . . . . . . 9.3 Stewardship . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9.3.1 Assurance of Product Performance . . . . . . . . . . . . . 9.4 Sales and Marketing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9.4.1 Competition Scanning . . . . . . . . . . . . . . . . . . . . . . 9.4.2 Third-Party Hybrid Evaluations . . . . . . . . . . . . . . . 9.4.3 Product Registration . . . . . . . . . . . . . . . . . . . . . . . 9.4.4 Market Development . . . . . . . . . . . . . . . . . . . . . . 9.4.5 Export of Seed . . . . . . . . . . . . . . . . . . . . . . . . . . . 9.4.6 Consumer Complaint Redressal . . . . . . . . . . . . . . . 9.5 Management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9.5.1 Germplasm Acquisition and Use . . . . . . . . . . . . . . 9.5.2 Intellectual Property Rights (IPR) for the Hybrids/ Parental Lines . . . . . . . . . . . . . . . . . . . . . . . . . . . 9.5.3 Licensing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9.5.4 Business Planning . . . . . . . . . . . . . . . . . . . . . . . . 9.5.5 Mergers and Acquisitions . . . . . . . . . . . . . . . . . . . 9.6 Summing Up . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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299 299 299 300 301 302

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305 305 306 307 308 309 309 310 311 311 312 312 313 313 314 314 315 315

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319 320 321 321 322 322

Regulatory Aspects of the Seed Business in Relation to Plant Breeding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10.2 Regulations on the Seed Industry . . . . . . . . . . . . . . . . . . . . . . 10.2.1 Regulation of the Seed Industry in India . . . . . . . . . . 10.2.2 Some Important Laws Related to the Seed Business/Crop Breeding in India . . . . . . . . . . . . . . . 10.2.3 Recognition of R&D Units . . . . . . . . . . . . . . . . . . . 10.3 Regulations on Germplasm . . . . . . . . . . . . . . . . . . . . . . . . . . 10.3.1 The International Treaty on Plant Genetic Resources for Food and Agriculture (ITPGRFA) . . . . . . . . . . .

323 323 324 325 325 327 328 329

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10.3.2 10.3.3

Convention on Biological Diversity (CBD) . . . . . . Intellectual Property Rights (IPR) Protection for Germplasm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10.3.4 Plant Breeder Rights (PBR)/Plant Variety Protection (PVP) System . . . . . . . . . . . . . . . . . . . . 10.3.5 Plant Breeders’ Rights (PBR) System in India . . . . 10.3.6 Registration of Germplasm in India . . . . . . . . . . . . 10.4 Germplasm Access . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10.4.1 Germplasm Access Under the PBR/PVP System . . . 10.4.2 Accessing Off-PVP Germplasm . . . . . . . . . . . . . . . 10.4.3 Accessing Germplasm from the Open Source Seed Initiative (OSSI) . . . . . . . . . . . . . . . . . . . . . . 10.4.4 Accessing Germplasm from Research Collaborations . . . . . . . . . . . . . . . . . . . . . . . . . . . 10.5 Crop Variety Registration . . . . . . . . . . . . . . . . . . . . . . . . . . 10.5.1 Crop Variety Registration in India . . . . . . . . . . . . . 10.5.2 Release and Notification of Crop Cultivars in India . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10.5.3 OECD Scheme for the Varietal Certification of Seed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10.6 Import and Exports of Seed and Germplasm . . . . . . . . . . . . . 10.6.1 Import/Export of Seed and Germplasm in/from India . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10.7 Regulations and the Registration Process for Genetically Modified (GM) Plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10.7.1 GM Trait Regulations in India . . . . . . . . . . . . . . . . 10.7.2 GM Trait Regulations in the USA . . . . . . . . . . . . . 10.7.3 GM Trait Regulations in European Union (EU) Countries . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10.7.4 GM Trait Regulation in China and in Some Asian Countries . . . . . . . . . . . . . . . . . . . . . . . . . . 10.7.5 GM Trait Regulation in Other Countries . . . . . . . . 10.8 Regulation of Genome Edited Crops . . . . . . . . . . . . . . . . . . . 10.9 Licensing of Traits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10.10 Summing Up . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. 336 . 336 . . . . . .

338 340 349 350 350 350

. 351 . 351 . 352 . 352 . 353 . 357 . 358 . 358 . 361 . 362 . 370 . 371 . . . . . .

372 378 379 382 382 383

About the Authors

Aparna Tiwari is cofounder and Chief Executive Officer at the Foundation for Advanced Training in Plant breeding (ATPBR), where she leads professional training, technology liaising, awareness sessions, and other capacity building activities. She also serves as a conference secretary and coordinates 5F Farming national conferences annually. She has worked previously in various positions with the FAO (Food and Agriculture Organization), Monsanto India Pvt. Ltd., ICRISAT (International Crops Research Institute for the Semi-Arid Tropics), and the MGM– Institute of Technology. She obtained her MSc and PhD from the Wageningen University, The Netherlands, and her Bachelor’s (BSc) from Banaras Hindu University. She has authored a book and published several research papers in peer-reviewed international journals and conferences. Surinder Kumar Tikoo currently Cofounder and Director Research—Breeding and Development, Tierra Seed Science Pvt. Ltd., Pune, has five decades of experience as a successful vegetable breeder, teacher, and research and development manager. He has held global leadership positions over long periods in two private sector organizations, as well as spent 20 years with the Indian Council of Agricultural Research (ICAR) as a vegetables breeder, teacher, and research guide. He has developed several nationally and internationally released varieties and hybrids of tomato and okra, some of which are market leaders in their respective segments. He has served on several advisory committees of the Government of India and various professional bodies. Dr. Tikoo, a PhD from Bangalore University, has published over 50 research and review papers. He also holds one patent. He has been conferred the Lifetime Achievement Award for Excellence in Translational Research, jointly by the Society of Plant Biochemistry and Biotechnology and Springer. Sharan Angadi is a plant geneticist and breeder who worked for 17 years in India, first with UAS, Bangalore, and then with the Indian Council for Agricultural Research (ICAR), contributing significantly to the genetic improvement of oilseed crops and horticultural crops. He then devoted 22 years to Pro Agro PGS and Nunhems Seeds, Netherlands, where he served as Head of Breeding for Asia Pacific and Global Coordinator for peppers, spanning several countries in Asia, Europe, and the Americas. He has particular experience in applying emerging technologies to xxix

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About the Authors

viable commercial plant breeding. Dr. Angadi has a PhD from IARI, New Delhi; an MSc from Tamil Nadu Agricultural University, Coimbatore; and a BSc from UAS, Bangalore. After retirement, he continues to provide expert R&D advice to seed companies in India and abroad, besides serving on various expert bodies. The Asia and Pacific Seed Association (APSA) honored him with The Most Influential Plant Breeding Researcher for Horticultural Crops Award for the year 2021. Suresh B. Kadaru has nearly 15 years of industry research experience in developing superior seed products for farmers, using modern breeding tools, technologies, and data analytics. He completed his BSc (Agri.) at Acharya NG Ranga Agriculture University, Rajendranagar, and his MSc (Biotechnology) from Tamil Nadu Agricultural University, Coimbatore, India. After completing his PhD in Agronomy and Applied Statistics from Louisiana State University, Baton Rouge, he worked as a Marker Data Analyst in Syngenta, USA, pioneering the application of molecular markers in plant breeding. Then relocating to India, he has provided functional leadership and served on the regional Seeds Product Development Leadership Team, which defines R&D strategies for maize and rice crops in the Asia-Pacific region. In addition to his publications, he has a joint patent and considerable training experience to his credit. He has received several internal awards in Syngenta for his markers assisted breeding work on rice and corn hybrids. Sadananda R. Ajanahalli an expert in agricultural research and technology management, has more than 40 years’ experience in the seed industry and in international and national public research institutions, including the International Maize and Wheat Improvement Center (CIMMYT), with whom he was associated as Maize Seed Systems Specialist for Asia. His various roles and responsibilities have included plant breeding, product development, and seed system management. After retiring from CIMMYT, he has pursued consultancy activities through the firm, AGREEVA. He has managed R&D in multinational and Indian seed companies for over 17 years and facilitated the development and deployment of Bt cotton hybrids (BGI & BGII) and maize hybrids in the shortest possible time. He has developed 47 hybrids and varieties in different field crops and served on various industrial bodies for biodiversity, variety protection, and biotechnology. Dr. Sadananda holds a PhD in Genetics from IARI, New Delhi. M. J. Vasudeva Rao (PhD in Genetics, IARI; MSc in Plant Breeding, UAS, Bangalore) has spent 15 years as a plant breeder in the public sector, and then nearly 35 years in the private seed sector. While at ICRISAT, he won the Hari Om Ashram Award from ICAR for his work on Striga resistance in sorghum; he also did pioneering work on early-maturing groundnut. His responsibilities in multinational and national seed companies in India ranged from Breeding and Research Management to include supply chain, sales, marketing, and business opportunities. He last served as President, Agricultural Technology, at Metahelix Life Sciences Private Ltd. (a Tata enterprise). His contributions to plant breeding in developing successful new varieties and hybrids, new selection technologies, and in mentoring plant

About the Authors

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breeders have won him the Lifetime Achievement Award in Plant Breeding by the ATPBR (Foundation for Advanced Training in Plant Breeding). He has published extensively and served on many committees to promote the science of Plant Breeding.

1

Plant Breeding: Its Evolution and Recent Trends

Abstract

Plant breeding is the process of improving crops. It continuously addresses the evolving needs of consumers by introducing new genetic diversity through new products. The principles of plant breeding and tools of biotechnologies are being effectively utilized by breeders to improve crop performance; they do so by developing varieties with better nutritional content and agronomic traits (e.g., yield, disease resistance, etc.). Breeders work in close association with farmers as well as consumers to understand their preferences and priorities, and accordingly design and execute comprehensive breeding strategies to maximize revenue opportunities of all stakeholders through the new improved varieties. A successful breeder requires technical knowledge and acquires extensive experience over time. But breeding successes are often confined to a handful of experts, who know how to translate breeding challenges into opportunities, and how to take ownership of efforts to develop an improved variety, by minimizing total cost without compromising on productivity and quality of produce. With the increasing need for food from a growing human population, it has now become indispensable for us all to learn from the best commercial plant breeding practices. This chapter aims to emphasize the need to train breeders with the knowledge and wisdom acquired from experienced breeders in a commercial setting.

1.1

Introduction

The world population has been growing continuously, necessitating a corresponding rise in the demand for food. By 2050, we may need to feed about 2.3 billion more mouths than we do today (FAO 2009). Much of the population growth is predicted to occur in urban areas, while urbanization already exerts a slowing effect on # The Author(s), under exclusive license to Springer Nature Singapore Pte Ltd. 2022 A. Tiwari et al., Market-Driven Plant Breeding for Practicing Breeders, https://doi.org/10.1007/978-981-19-5434-4_1

1

2

1 Plant Breeding: Its Evolution and Recent Trends

agriculture. While, on the one hand, food consumption per capita increases tremendously, on the other hand, skilled labor in rural areas is becoming scarce. This eventually poses a serious threat to food security. There are several other factors affecting food security, such as the following: 1. Currently over 50,000 edible plant species are available in the world, of which, only 30 species are grown, providing 95% of human dietary energy. Rice, wheat, maize, and potato are the four crops that provide more than 60% of the world’s food energy intake. This dependence on a few selected crop species would be a serious threat for food security. 2. The global climate is undergoing severe changes, with rising incidences of unpredictable storms, shrinking the available area of arable farmland. Increases in global temperature have been disrupting the normal agricultural production patterns. This intensifies the difficulties of maintaining genetic resistances against pests, diseases, and weeds among crops. 3. There is a continuously growing demand for nutritious and quality food, mainly due to economic progress in the developing countries, such as China, India, Indonesia, Brazil, and Mexico. In modern times, there are numerous questions related to food security, which breeders and agricultural experts need to address. For example, 1. Can the existing food systems suffice to feed the growing global population, which is projected to cross 9 billion by 2050? 2. Can agriculture generate more livelihood opportunities, which, in turn, can ease the mass migration to cities? 3. Can breeders achieve the expected increases in the rates of production? 4. Can all stakeholders of the agriculture system (i.e., public sector, private sector, farmers, consumers, government bodies) work in unison to promote the sustainable development of agriculture? The answer to all these questions is affirmative, in that sustainable production can be achieved by using climate-smart farming techniques in efforts led by competent breeders, who aim to stabilize the food security via the development of crop varieties that are tolerant to pests, diseases, drought, waterlogging, and salinity (FAO 2013). Over the past few decades, advanced technologies, and introduction of enhanced seed varieties, with traits such as higher yield, improved nutritional quality, reduced crop damage, and disease resistance, have encouraged farmers to shift from the usage of farm-saved seeds to the use of value-added, innovative, commercial hybrid seeds. This shift has been possible with the advancements in plant breeding. Plant breeding is an effective tool to make genetic changes in plants to achieve the desirable phenotypes. Plant breeding helps to develop improved varieties for resistance to biotic and abiotic stresses, improve crop productivity, and thus help stabilize the market and reduce price volatility, contribute to reduction of greenhouse gas emission, and help improve farmers’ incomes. These are achieved, however, by

1.2 History and Evolution of Plant Breeding

3

skilled expert plant breeders, using innovative tools and exploiting the available pool of germplasm. Today’s innovations in plant breeding use sophisticated science and technology that includes speed breeding, genomic selection, gene mapping, precision phenotyping, mutations, and marker-assisted breeding (MAB). Though plant breeders play a very critical role in crop improvement, their current set of skills and competencies may not be adequate to meet the current and future demands for improved varieties. This chapter provides an overview of the plant breeding industry and elaborates the importance of plant breeding research to meet the needs of growing populations. It attempts to cover the evolution of plant breeding, along with the market forces that will drive the future direction of plant breeding research, which indicates the growing prominence of the private sector in plant breeding research to meet the trends of the future market. The importance of plant breeding is increasing and so is the need for competent plant breeders.

1.2

History and Evolution of Plant Breeding

Plant breeding has a very long history. If selection is considered as the beginning of crop improvement, then plant breeding began with the start of agriculture in human civilization. Every single person of those times was the first or primitive plant breeder. In the early years of plant breeding, variability was exploited by natural selection, which is still followed by farmers who use farm-saved seeds from selected plants for sowing. Over the years, this selection process has evolved into a very precise and efficient scientific body of knowledge, i.e., the science of plant breeding. Agriculture is a continuous evolutionary process and so is plant breeding (see Table 1.1), where a few methods are basic and evergreen, to be always applied in any breeding program (such as introduction, selection, hybridization, mutation, etc.). A few others are still in the process of improvement and adaptation (such as genetically modified organisms, or GMO, and genomic selection), while some are in the fully mature phase (e.g., marker-assisted selection, or MAS, mapping). Experts have classified the history of plant breeding into four stages or eras: Pre-Mendelian, Mendelian, Post-Mendelian, and Modern (see Table 1.1 and Fig. 1.1; see also Flannery 1973; Palladino 1993; Diamond 2002; Davies 2003). Toward the latter part of the seventeenth century, Marcello Malpighi and Nehemiah Grew contributed substantially to early understandings of plant anatomy and reproduction, which marked the beginning of scientific forays into plant breeding. Marcello Malpighi was the first scientist to describe the plant structures and the generation of seeds in 1675, which was elaborated further to identify the parts involved in the germination and growth of young plants. The discovery of the laws of inheritance by Mendel toward the end of the nineteenth century accelerated the innovations in crop improvement. The scientific basis of these laws allowed the creation of combinations of beneficial traits among parent plants more precisely and efficiently than before. Later, the discovery of DNA structure in 1953 by Watson and Crick increased the knowledge of genetic material

4

1 Plant Breeding: Its Evolution and Recent Trends

Table 1.1 The four stages or eras in the history of plant breeding “Pre-Mendelian” - before 1900 • 9000 BC - First evidence of domescaon • 1717 - first arficial hybrid Fairchild's mule by crossing Carnaon and Dianthu (Sweet william variety) • 1801 - Lamarck's theory of inheritence of acquired characters • 1859 - Darwin's "Origin of Species" published - has details on inbreeding, sterility etc. • 1866 - Mendel's Laws of Inheritance - "recombinaon", "segregaon", "dominance"

“Mendelian” - 1900 - 1920 • 1899 - Terms “allelomorph”, “homozygote”, “heterozygote”, “F1” and “F2” (Bateson) • 1900 - deVries' Mutaon Theory • 1902 - Pure line theory (Johannsen) • 1903 - Embryo culture (Hannig) • 1906 - Term "Genecs" (Bateson) • 1906 - Mulple Factor Hypothesis (George Yule)

“Post Mendelian” - 1921 - 1950 • 1928 - Mutagenic effects of X-rays in barley • 1929 - 10 chromosomes in maize idenfied • 1931 - Cytological proof of crossing-over • 1933 - Cytoplasmic male sterility in maize discovered (Rhoades) • 1944 - DNA idenfied as the hereditary material • 1945 - Recurrent selecon method(Hull) • 1946 - Reciprocal (half-sib) recurrent selecon • 1950 - Ac-Ds system of transposable elements in maize (McClintock)

“Modern” - 1951 to present • 1952 - Use of mul lines in oats (Janssen) • 1953 - method of developing mullines in Wheat outlined (Borlaug) • 1953 - Structure of DNA (Watson and Crick) • 1956 - G for gene hypothesis for host-parasite resistance (Flor) • 1964 - Developed high yielding semi dwarf variees of wheat which resulted in green revoluon (Borlaug) • 1965 - First applied Single Seed Descent (SSD) method in oats (Graphius) • 1968 - Developed the concept of crop ideotype in wheat (Donald) • 1970 - World’s first coon hybrid for commercial culvaon in India (Patel) • 1970 - rDNA technology (Berg, Cohen and Bayer) • 1976 - World’s first rice hybrid (CMS based) for commercial culvaon in China • 1983 - First GE transgenic plant – tobacco - in USA (Fraley et al.) • 1987 - World’s transgenic coon plant in USA (Monsanto) • 1991 - World’s first pigeon pea hybrid (ICPH 8) (GMS based) for commercial culvaon in India • 1994 - “FlavrSavr” tomato, first GM food produced for the market (Calgene, California) • 1994 - GMO, Insect resistant coon • 1995 - BT corn developed • 1996 - Glyphosate Resistant Crop® soyabean • 1997 - Terminator gene allowing germinaon of seed for one generaon only (Monsanto, USA) • 1998 - Idenficaon of traitor gene, which responds to specific brand of ferlizers and inseccides • 2000 - Marker Assisted Selecon Barley resistant to yellow dwarf virus • 2004 - Glyphosate Resistant Crop® wheat developed • 2010 - Genomic selecon • 2012 - CRISPR-Cas9 (Gene eding) • 2018 - Speed breeding

substantially. Targeting the DNA to restructure plant anatomy through mutation breeding became the center of research in plant breeding around 1960. In 1983, with the development of recombinant DNA or GM (Genetic Modification) technology, modern plant breeding techniques, such as genomic selection, became a reality. During the same period, the “Green Revolution” (from 1960 to 1980) was noted bringing about remarkable increases in the production of wheat and rice. The establishment of CIMMYT (International Maize and Wheat Improvement Centre) in 1966 and IRRI (International Rice Research Institute) in 1960 contributed

Fig. 1.1 Evolution of plant breeding over the millennia (source: ISF 2008)

1.2 History and Evolution of Plant Breeding 5

6

1 Plant Breeding: Its Evolution and Recent Trends

significantly to the development of improved varieties of wheat and rice, respectively, and provided the impetus for the success of the green revolution. International regulatory bodies, such as the International Seed Testing Association (ISTA) (1924), the International Plant Protection Convention (IPPC) (1951), the OECD Seed Schemes (1953), and the International Union for the Protection of New Varieties of Plants (UPOV) (1961), were established, which set the ground rules for plant breeding. Crossbreeding techniques were further improved through DNA analytical methods to achieve marker-assisted selection (MAS). This technique involves selection of plants carrying a DNA segment called the “marker,” linked to a specific trait of interest, which enables the expression of a desirable phenotype. With further advancement and availability of an array of molecular markers, complemented by the dense molecular genetic maps, MAS enables the identification of specific plant phenotypes governed by major genes, as well as quantitative trait loci (QTL). This method has been more effective than traditional cross-breeding techniques. Further evolution of technologies led to new breeding techniques (NBTs), which involve the introduction of sequence-specific nuclease (SSN) complexes, which induce a gene mutation in a plant genome, or delete the gene or regulatory sequences from the genome when introduced temporarily. A few examples of NBTs are cisgenesis, intragenesis, sequence-specific nuclease technology, oligo-directed mutagenesis, RNA-dependent DNA methylation, CRISPR/Cas (mutation breeding techniques), and reverse breeding. These techniques of crop improvement paved the way for a debate on the associated risks and the technical aspects of plant breeding. And thus, the NBTs came under increased evaluation from a regulatory perspective. The recent introduction of speed breeding as another powerful tool to accelerate crop breeding research has helped to achieve up to six generations per year in spring wheat, barley, pea, and four generations in canola, instead of the 2–3 generations of these crops per year under normal conditions. Further integration of this technology with other modern crop breeding technologies, such as high-throughput genotyping, genome editing, and genomic selection, is expected to further accelerate the process of crop improvement. Plant breeding has thus evolved from simple selection for basic needs by primitive men and women to multifaceted advancements in technology. It has been transformed from a simple food production activity to an ever-evolving, globally interconnected, industrial activity, aimed at feeding the numerous needs of millions of people. Consequently, the role of a plant breeder has also largely transformed from being a sole decision-maker to the role of an all-round personality to lead and contribute through a multidisciplinary team. The tools kit which a breeder uses today for crop improvement would have been unthinkable some 50 years ago. Today, the use of tech-oriented tools is imperative, together with conventional tools, to develop improved genetic material, which allows farmers to deliver higher yields and better quality than before. With appropriate, modern, and sophisticated management, a breeder can do his work more efficiently and economically than at any other time in history (Godwin et al. 2019).

1.3 Market Drivers for Plant Breeding Products

1.3

7

Market Drivers for Plant Breeding Products

Plant breeding plays a significant role in food production, biodiversity conservation, and the resilience and sustainability of farming. It can mitigate the effects of population growth, climate change, and other social and physical challenges, and thus support the growth of national economies by driving the global seed market. During the last four decades, plant breeding businesses have evolved in size and level of performance. Based on geography and its constraints, researchers have worked toward overcoming local challenges and started developing varieties that focus on quality and nutritional aspects. Because of a large incidence of mergers and acquisitions of agrochemical (including pesticides, fungicides, or plant protection chemicals) companies and seed industries, the plant breeding industry is expanding continuously (Prifti 2015). The merger of Dow AgroSciences (USA) with DuPont (USA), Monsanto (USA) with Bayer (Germany), and Syngenta (Switzerland) with ChemChina (China) provide some recent instances of large-scale mergers and acquisitions. These consolidations have led to changes in the structure, competition, investment trends, and supply-demand dynamics in the industry, as well as increased the concentration of plant breeding and biotechnological companies. The Indian seed market, which reached a value of US$ 4.9 billion in 2020, is fragmented by crop types, varieties, and local needs, and is projected to grow further (IMARC 2020a). The several factors that drive plant breeding research in this context are depicted in Fig. 1.2.

Fig. 1.2 Market drivers for plant breeding research

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1.3.1

1 Plant Breeding: Its Evolution and Recent Trends

Advanced Technologies

The complementation of biotechnology to plant breeding has revolutionized the process of variety development, by reducing the length/duration and cost for the release of an improved variety. The driving force behind innovation and technology investment is the need to deliver better value to the producers and consumers. The integration of advanced technology facilitates cheaper and faster mapping of the genetic codes of plants, and in doing so, helps solve the major agricultural challenges quicker than was possible earlier. The continuously increasing demand for improved crop varieties and hybrid seeds, and the need to reduce pre-harvest crop losses, are the main factors that have encouraged the deployment of advanced plant breeding techniques. These technologies improve the effectiveness of the breeding program, enhance the precision, reduce the time taken to develop improved varieties, and help to discover more information about the functions of the genes of interest. In India, about 92% of the cotton crop area is under transgenic crop production (i.e., Bt cotton), which is a clear indication that farmers are increasingly adopting transgenic seeds for both yield and non-yield traits (Barwale et al. 2004). Industries are investing in a huge way in biotechnology to strengthen their biotech products pipeline. The main reasons for this are as follows: 1. Biotech crops have led to a reduction in the use of insecticides; there has been more than 50% reduction in the use of insecticides in India and China after the introduction of Bt cotton (Naveen et al. 2017). 2. Increased farmers’ income: Because of enhanced crop yields, there is a significant rise in farmers’ incomes per hectare (i.e., 250 US dollars in India, 220 US dollars in China, 117 US dollars in South Africa, and 135 US dollars in the Philippines). For Bt cotton, increase in yield was 10% in China and 31% in India (Martin 2020). 3. An almost four-fold growth in the biotech seed business has been reported in India between 1979 and 2018. This accelerated introduction of biotech crops is due to growing demand for animal feed, shrinking per capita farmland, and growing population (Biotech Seeds 2020; ISF 2008; Knight et al. 2007; Medgadget report 2020; Paroda 2018; Martin 2020). 4. There is a need to develop varieties with unique traits, such as insect resistance, herbicide tolerance, resistance to environmental stresses, as well as value-added traits (nutrient-enhanced seed, pharmaceuticals, biofuel, etc.). A substantial reduction in the cost of genomic solutions has also enabled their use in the accelerated growth of plant breeding. Over a longer period, the cost of genomics and other advanced breeding techniques is expected to be lower than the cost of conventional breeding, owing to the availability of skilled labor, adequate infrastructure, and other favorable factors.

1.3 Market Drivers for Plant Breeding Products

1.3.2

9

Hybrid Seed Production

Hybrid seed production is another important driver in the growth and development of plant breeding research. Farmers have been shifting from traditional farm-saved seeds to hybrid seeds, owing to the advantages conferred by the hybrid seeds, such as high uniformity, better performance because of heterosis, or hybrid vigor. Hybrids are advantageous for seed companies as well, as they can protect their intellectual property rights (IPR) better than inbred varieties. Currently, the application of hybridization is prominent and well established in the case of cotton (>90%), corn (>60%), and sunflower (80%), but limited in the case of rice and wheat (