Molecular Interventions for Developing Climate-Smart Crops: A Forage Perspective: A forage perspective 981991857X, 9789819918577

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Rajesh Kumar Singhal Shahid Ahmed Saurabh Pandey Subhash Chand   Editors

Molecular Interventions for Developing Climate-Smart Crops: A Forage Perspective

Molecular Interventions for Developing Climate-Smart Crops: A Forage Perspective

Rajesh Kumar Singhal • Shahid Ahmed • Saurabh Pandey • Subhash Chand Editors

Molecular Interventions for Developing Climate-Smart Crops: A Forage Perspective

Editors Rajesh Kumar Singhal ICAR-Indian Grassland and Fodder Research Institute Jhansi, India Saurabh Pandey Department of Agriculture Guru Nanak Dev University Amritsar, India

Shahid Ahmed ICAR-Indian Grassland and Fodder Research Institute Jhansi, India Subhash Chand All India Coordinated Research Project on Forage Crops and Utilisation ICAR-Indian Grassland and Forage Research Jhansi, India

ISBN 978-981-99-1857-7 ISBN 978-981-99-1858-4 https://doi.org/10.1007/978-981-99-1858-4

(eBook)

# The Editor(s) (if applicable) and The Author(s), under exclusive license to Springer Nature Singapore Pte Ltd. 2023 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

Foreword 1

Forage crops are more resilient to biotic and abiotic stresses in comparison to other agricultural crops under unprecedented climate change regime. However, these crops are considered as “orphan” crops as they are mainly cultivated in less productive, uncultivable, and unmanaged lands of dryland areas where economically important crops cannot be cultivated. Modern problems need modern solutions, and forage crops serve as a valuable resource due to their diverse range of agriculturally important traits such as tolerance to various biotic and abiotic stresses as well as high-quality attributes. The important genetic resources carrying the desirable traits could be used directly or indirectly in the breeding programs to improve the existing high-yielding varieties that lack few traits through various modern breeding tools such as Marker Assisted Selection, Marker Assisted Backcross Breeding, and genome editing. The edited book Molecular Interventions for Developing Climate-Smart Crops: A Forage Perspective covers various aspects of molecular breeding, plant physiology, and biochemistry in forage crops that could play a vital role in developing climatesmart forage crops. The chapters highlight the existing genetic resources of grasses and their conservation methods, unconventional breeding tools such as allele mining to unlock the allelic variants for climate resilience, genetic improvement of treebased forage species, and strategies for developing forage cultivars tolerant to biotic and abiotic factors.

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Foreword 1

The editors deserve special compliments for this excellent book on recent molecular advancements in forage crops that have been neglected for many years in developing nations including India. I am sure that this book will be helpful to readers from different disciplines for their academic and research purposes including students, teachers, plant breeders, and research scholars due to its comprehensive subject coverage and in-depth analysis of recently available literature. I congratulate all the editors for this important publication and convey my best wishes for their future endeavors. Indian Council of Agricultural Research, Krishi Bhavan New Delhi, India

D. K. Yadava

Foreword 2

The unprecedented human population, urbanization, and infrastructure developments exert huge pressure on cultivated land, water, environment, and other resources. In India, forage crops are mainly cultivated in dryland areas, forests, community grazing lands, bunds, and unproductive lands where food crops cannot be grown. These crops are more resilient to biotic and abiotic stresses and are more input-use efficient; therefore, fewer resources are required for their production. However, they are reservoirs of various agriculturally important traits and can be used as donors for trait-specific breeding programs. The edited book Molecular Interventions for Developing Climate-Smart Crops: A Forage Perspective is well compiled with recent developments in forage research including allele mining to unravel the undisclosed alleles in forage genetic resources and other molecular interventions. The content of each chapter is nicely presented in easy-to-understand language with suitable examples. The contributions of the authors are praiseworthy. They conducted comprehensive exercises in collecting the content of each chapter and presented them in a simplified manner. The editors of the book also deserve special recognition for their excellent compilation of all the topics related to forage crops grown under challenging climatic conditions vis-à-vis up-to-date review of the subject. I am very sure that this book will be very useful for students, researchers as well as teachers of various disciplines and other stakeholders.

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Foreword 2

I once again congratulate all the authors for their efforts and hard work. ICAR-Indian Grassland and Fodder Research Institute Jhansi, Uttar Pradesh, India

Amaresh Chandra

Foreword 3

Forage crops are as essential to livestock as cereals are to human beings for their feed and nutritional security, being the main source of green and nutritious fodder to the livestock. However, these crops are mostly allocated to poor farming lands due to competition from more remunerative cash crops and grain crops, particularly in developing nations including India. Genetic improvement program of fodder crops mostly focused on higher forage yield, with comparatively less emphasis on quality traits as well as disease and pest tolerance. Feeding of nutritious green grasses along with appropriate portions of legumes can increase livestock productivity. The edited book Molecular Interventions for Developing Climate-Smart Crops: A Forage Perspective covers recent progresses in forage research including allele mining to untie the undisclosed alleles in forage genetic resources, biochemical and physiological changes after application of man-made substances against biotic and abiotic factors, for better fodder productivity and nutritional quality. Each chapter is nicely presented in easy-to-understand language with suitable examples. The authors have done a comprehensive exercise in collecting the content of each chapter and in interpreting every topic in a very simple way. The contribution of the editors of the book also deserves special appreciation for the excellent compilation of the topics. I am very sure that this book will be very useful for students, researchers as well as teachers of various disciplines.

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Foreword 3

I once again congratulate the editors for their hard work in bringing up this publication and best wishes for their future endeavors. AICRP on Forage Crops and Utilization, ICAR-Indian Grassland and Fodder Research Institute Jhansi, Uttar Pradesh, India

A. K. Roy

Preface

The UNESCO describes grassland as “land covered with herbaceous plants with less than 10% tree and shrub cover” and grasslands are among the largest ecosystems globally representing 40.5% coverage of the terrestrial area. Besides, forage crops are predominantly cultivated in the dry land areas where abiotic constraints are more common and persist throughout the season and, therefore, considered as “Orphan crops.” However, these crops are reservoir of many important but unexploited traits that could be game changer for increasing and sustaining productivity of food crops and livestock. Therefore, to meet the increasing demand of global food and feed, we must include innovative techniques with conventional approaches to develop climate-smart forage crops that can tolerate biotic and abiotic stresses. The chapters included in Molecular Interventions for Developing Climate-Smart Crops: A Forage Perspective provide sound scientific view by the leading scientists working on forage crops nationally and internationally. The book covers in-depth understanding of forage genetic resources, particularly tropical grasses, constraints in forage breeding, allele mining to discover important but undisclosed allelic variations, trees as forage and their genetic improvement, effect of biotic and abiotic factors and molecular interventions to reduce their adverse impact, impact of nanopriming on forage crops, agroforestry as fodder crops, synergy between livestock and fodder crops, use of crop waste as livestock feed, and interventions of agricultural engineering to solve present-day problems. We are enough confidant that this book will improve the understanding and knowledge of undergraduate and postgraduate students, researchers, and scientific staffs working on forage crops globally. Each chapter has been written in such a way that it would meet the scientific information related to forage improvement, production, and protection of the end user. Finally, we thank the editors and publishing managers of Springer and contributing authors for their consistent technical support, guidance, and patience throughout the process of drafting and publishing this book. Jhansi, India Jhansi, India Amritsar, India Jhansi, India

Rajesh Kumar Singhal Shahid Ahmed Saurabh Pandey Subhash Chand xi

Introduction

Climate change is related to long-term shifts in environmental parameters particularly temperature and weather patterns. Agricultural crops are extremely vulnerable to climate, and even slight changes in climatic factors such as temperature and soil water status cause tremendous loss in yield production. However, forage crops can tolerate climatic variations in a better way than food crops under field conditions and play a significant role in global food security and environmental sustainability. In addition, forage crops are repository of many valuable traits, including tolerance against biotic and abiotic stresses that could be used as a donor source for improving the trait of interest in agricultural crops. Research programs on forage crops are limited globally that not only increase the gap between fodder demand and supply but also affect the feed and nutritional security of livestock with respect to animal health, reproduction, and productivity. Besides, forage crops are resilient to climatic challenges as they are grown and mostly cultivated in unproductive soils and harsh conditions. By considering the above challenges, this book Molecular Interventions for Developing Climate-Smart Crops: A Forage Perspective encompasses different aspects related to plant physiology, biochemistry, and molecular breeding of forage crops that could play a crucial role in developing climate-smart forage crops. Chapter 1 covers the status of genetic resources of tropical grasses and their conservation methods, constraints in breeding methods and seed production and various approaches to overcome them. Chapter 2 emphasizes existing availability of forage genetic resources, their linkage with livestock, and environmental sustainability. These genetic resources harbor a wealth of undisclosed allelic variants for climate resilience, and allele mining would be rewarding to unravel the superior alleles present in genetic resources for deployment in forage breeding programs. Chapter 3 mainly focuses on different breeding approaches for genetic improvement of trees that are mostly being used as forage. These tree-based forage crops will not only provide additional fodder to livestock but also be beneficial for sustaining ecological balance under challenging climatic conditions. Chapter 4 highlights the impact of different biotic and abiotic factors on forage crop production, association of different alkaloids with forage crops and their mitigation practices using breeding and molecular approaches. Chapter 5 covers the effect of nano-priming, from seed germination to final seed yield, on maize. Nano-priming accelerated signaling xiii

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Introduction

pathways trigger hormone secretion and reactive oxygen species biosynthesis and improve disease resistance. In addition, Chap. 6 highlights the biochemical and physiological role of oxidative stress and antioxidant defense in forage crops ameliorating abiotic stresses. Chapter 7 underlines the different fodder cropping systems that ameliorate the adverse effect of salinity in soils under challenging climatic conditions. Chapter 8 highlights the various environmental stresses that hamper forage production globally and provides different approaches to increase or sustain forage production under challenging climatic conditions. Chapter 9 covers the potentials, challenges, and opportunities of agroforestry under climate change regimes. Agroforestry systems are bio-diverse and are linked to desertification and climate change mitigation in a variety of ways. Chapter 10 mainly focuses on the impact of climate change on forage crops, livestock keeping and health management and synergies between each other. Traditional and nontraditional livestock feeds are also discussed for feed and nutritional security of livestock under challenging periods. Nowadays, crop burning is an emerging issue in developing countries like India that adversely affects humans, livestock, and environmental health. These modern problems need modern solutions, and advancements in agricultural equipment can solve them. Therefore, Chap. 11 enlists the sustainable use of paddy straw as a livestock feed that will not only provide additional fodder to livestock but also be a climate-resilient approach to crop residue burning. Chapter 12 covers agricultural engineering interventions including artificial intelligence and hydroponic system that not only increase fodder production but also fodder quality. Modern agricultural machines can help solve the problem of crop residue burning issue and use crop residues for conservation agriculture. Chapter 13 highlights the realistic and pragmatic extension strategies for promoting forage crops and their climate-resilient cultivation technologies among the stakeholders particularly dairy farmers. We believe that this book will enrich the scientific knowledge of readers, including plant breeders, agronomists, biochemists, and plant pathologists about the pre-existing genetic variability in forage crops, modern tools to generate variability and their utilization in breeding programs to sustain food-feed-based systems.

Contents

Part I 1

2

3

4

5

6

Forage Crop Improvement

Genetic and Genomic Resources of Range Grasses: Status and Future Prospects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Vikas C. Tyagi, Tejveer Singh, Nilamani Dikshit, Sultan Singh, Maneet Rana, Rahul Kaldate, Prabhu Govindaswamy, Hanamant M. Halli, Avijit Ghosh, Rajesh Kumar Singhal, and Manjanagouda S. Sannagoudar Forage Genetic Resources and Scope for Allele Mining of Abiotic Stress Tolerance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brijesh K. Mehta, Surendra Kumar Meena, Nilamani Dikshit, P. Shashikumara, Anup Kumar, Praveen Kumar, Mahendra Singh, Gaurendra Gupta, and Shahid Ahmed Breeding for Developing Higher Productive Tree-Based Forage Under Stress Environments . . . . . . . . . . . . . . . . . . . . . . . . . Hirdayesh Anuragi, Srijan Ambati, Rajesh Kumar Singhal, Sukumar Taria, Alka Bharati, Kunasekaran Rajarajan, Arun Kumar Handa, and Ayyanadar Arunachalam

3

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Impact of Climate Change on Forage Crop Production with Special Emphasis on Diseases and Mitigation Strategies Through Breeding and Molecular Approaches . . . . . . . . . . . . . . . . Namburi Karunakar Reddy, Gaurav Rakhonde, Pooja Purushotham, Pooja S. Patel, and Shalaka Ahale

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Effect of Nano-Priming on Maize Under Normal and Stressful Environment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sananda Mondal, Bandana Bose, and Debasish Panda

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Oxidative Stress and Antioxidant Defense in Mitigating Abiotic Stresses in Forage Crops: A Physiological and Biochemical Perspective . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109 Meenakshi Goyal, Archana Kumari, Ankita Kumari, Himanshu Sharma, Pashupat Vasmatkar, and Namrata Gupta xv

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7

Contents

Forage Cropping Under Climate Smart Farming: A Promising Tool to Ameliorate Salinity Threat in Soils . . . . . . . . . 137 Eetela Sathyanarayana, B. Prem Kumar, Rupesh Tirunagari, G. Keerthana, Vilakar Kayitha, J. Bharghavi, S. Saranya, M. Rajashekhar, B. Rajashekhar, K. Charan Teja, and Saideep Thallapally

Part II

Sustainable Forage Production

8

Forage Cultivation Under Challenging Environment . . . . . . . . . . . 149 Jasmina Milenković, Mirjana Petrović, Snežana Andjelković, and Debasis Mitra

9

Potentials and Opportunities of Agroforestry Under Climate Change Scenario . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161 Manjanagouda S. Sannagoudar, G. K. Prajwal Kumar, Vanitha Khandibagur, Avijit Ghosh, Amit K. Singh, G. A. Rajanna, Hanamant M. Halli, V. K. Wasnik, B. R. Praveen, and R. T. Chethan Babu

10

Climate Change Impact on Forage Characteristics: An Appraisal for Livestock Production . . . . . . . . . . . . . . . . . . . . . . 183 Pooja Tamboli, Amit Kumar Chaurasiya, Deepak Upadhyay, and Anup Kumar

11

Sustainable Use of Paddy Straw as Livestock Feed: A Climate Resilient Approach to Crop Residue Burning . . . . . . . . . . . . . . . . . 197 B. R. Praveen, Manjanagouda S. Sannagoudar, R. T. Chethan Babu, G. A. Rajanna, Magan Singh, Sandeep Kumar, Rakesh Kumar, and V. K. Wasnik

12

Engineering Interventions for Climate-Resilient Forage Production . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215 Amit Kumar Patil, Naseeb Singh, Partha Sarathi Singha, Monika Satankar, Sheshrao Kautkar, S. K. Singh, and P. K. Pathak

13

Promotion of Improved Forage Crop Production Technologies: Constraints and Strategies with Special Reference to Climate Change . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229 Ashish Kumar Gupta, M. L. Sharma, M. A. Khan, and P. K. Pandey

Editors and Contributors

About the Editors Rajesh Kumar Singhal currently works as a scientist in the Division of Crop Improvement, ICAR-Indian Grassland and Fodder Research Institute, Jhansi (U. P), India. He has received his PhD from IAS, BHU, India. He has 8 years of experience in different areas of Plant Physiology aspects such as drought stress, seed priming, artificial night light pollution, and multiple abiotic stresses related to different crops. Currently, he is working on fodder oat improvement, oat modelling for climate change, and abiotic stress in forage crops. He has published more than 25 research and review papers in reputed journals. He also published more than 15 chapters with many publishers. Shahid Ahmed is a principal scientist and head of the Crop Improvement Division at ICAR-Indian Grassland and Forage Research Institute, Jhansi, India. In the last 15 years, he has been working on breeding and crop improvement in forage crops. He has a lot of experience in biofortification, genetic diversity, marker selection, disease resistance, and nutrient management in forage crops and develops a number of technologies. He published 32 research articles, 6 books, 3 chapters, 1 conference paper, 2 technical reports, and handled a number of projects related to forage crop improvements. He published research articles in very high-rated journals. Saurabh Pandey currently works as an Assistant Professor in the Department of Agriculture at Guru Nanak Dev University, Amritsar, Punjab, India. He was conferred a PhD (Plant Virology) from the National Institute of Plant Genome Research, New Delhi. He has 8 years of experience in different areas of Plant Biotechnology such as plant stress biology, plant-virus interactions, abiotic stresses, bio-fortification, molecular marker studies, and metabolism. He has published about half a dozen research papers in various journals of international repute. Dr. Pandey has authored many book chapters about molecular markers, genomics, plant breeding, and plant molecular virology, catering to the needs of modern agriculture research. He has been an active member of the Indian Virological Society, the Indian Academy of Horticultural Sciences, and the Society of Plant Biochemistry and

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Editors and Contributors

Biotechnology. Dr. Pandey is the recipient of the Gold Medal and Best PG thesis award for M.Sc. Thesis and various other awards for his research work. Subhash Chand is pursuing his PhD from the Division of Genetics, ICAR-Indian Agricultural Research Institute, New Delhi (India). He is posted at ICAR-Indian Grassland and Fodder Research Institute, Jhansi. He has 3 years of experience in forage and fodder crops including oat, berseem, lucerne, guinea grass, BN Hybrids, etc. while working in AICRP on forage crops and utilization. He has published more than ten research papers in reputed international and national journals. He has published more than ten book chapters in different books. He has published five books related to fodder crops published by AICRP on forage crops and utilization. His area of specialization is genetics and plant breeding, quantitative genetics, disease resistance breeding, and molecular breeding. He was also awarded NTS, JRF, and SRF during his academic period by the ICAR.

Contributors Shalaka Ahale Department of Plant Pathology, Punjab Agricultural University, Ludhiana, Punjab, India Shahid Ahmed ICAR-Indian Grassland and Fodder Research Institute, Jhansi, India Srijan Ambati Agriculture College Warangal, Professor Jayashankar Telangana State Agricultural University, Warangal, Telangana, India Snežana Andjelković Institute for Forage Crops Kruševac, Kruševac, Republic of Serbia Hirdayesh Anuragi ICAR-Central Agroforestry Research Institute, Jhansi, India Ayyanadar Arunachalam ICAR-Central Agroforestry Research Institute, Jhansi, India R. T. Chethan Babu Agronomy Section, ICAR - National Dairy Research Institute, Karnal, Haryana, India Alka Bharati ICAR-Central Agroforestry Research Institute, Jhansi, India J. Bharghavi Department of Crop Physiology, Agricultural College, Warangal, PJTSAU, Warangal, India Bandana Bose Department of Plant Physiology, Institute of Agricultural Sciences, Varanasi, Uttar Pradesh, India Amit Kumar Chaurasiya Department of Animal Nutrition, NDVSU, Jabalpur, MP, India

Editors and Contributors

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Nilamani Dikshit ICAR-Indian Grassland and Fodder Research Institute, Jhansi, India Avijit Ghosh ICAR - Indian Grassland and Fodder Research Institute, Jhansi, India Prabhu Govindaswamy ICAR - Indian Agriculture Research Institute, Jhansi, India ICAR - Indian Grassland and Fodder Research Institute, Jhansi, India Meenakshi Goyal Department of Plant Breeding and Genetics, Punjab Agricultural University, Ludhiana, Punjab, India Ashish Kumar Gupta Rani Lakshmi Bai Central Agricultural University, Jhansi, Uttar Pradesh, India Gaurendra Gupta ICAR-Indian Grassland and Fodder Research Institute, Jhansi, India Namrata Gupta Department of Botany, Punjab Agricultural University, Ludhiana, Punjab, India Hanamant M. Halli ICAR - National Institute of Abiotic Stress Management, Baramati, MH, India ICAR - Indian Grassland and Fodder Research Institute, Jhansi, India Arun Kumar Handa ICAR-Central Agroforestry Research Institute, Jhansi, India Rahul Kaldate ICAR-Indian Grassland and Fodder Research Institute, Jhansi, India Sheshrao Kautkar ICAR - Central Institute for Research on Cotton Technology (CIRCOT), Mumbai, India Vilakar Kayitha Keladi Shivappa Nayaka University of Agricultural and Horticultural Sciences (KSNUAHS), Shimoga, India G. Keerthana Department of Soil Science and Agricultural Chemistry, College of Agricultural Sciences, JKKM, Erode, India M. A. Khan Indira Gandhi Agricultural University, Raipur, Chhattisgarh, India Vanitha Khandibagur University of Horticultural Sciences, Bagalkot, India Anup Kumar ICAR-Indian Grassland and Fodder Research Institute, Jhansi, India Plant Animal Relationship Division, ICAR-IGFRI, Jhansi, UP, India B. Prem Kumar Division of Soil Science and Agricultural Chemistry, ICAR-IARI, New Delhi, India G. K. Prajwal Kumar Karnataka State Department of Agriculture, Bangalore, India

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Editors and Contributors

Praveen Kumar ICAR - Indian Grassland and Fodder Research Institute, Jhansi, India Rakesh Kumar Agronomy Section, ICAR - National Dairy Research Institute, Karnal, Haryana, India Sandeep Kumar Agronomy Section, ICAR - National Dairy Research Institute, Karnal, Haryana, India Ankita Kumari National Dairy Research Institute, Karnal, Haryana, India Archana Kumari Department of Biochemistry, Punjab Agricultural University, Ludhiana, Punjab, India Surendra Kumar Meena ICAR-Indian Grassland and Fodder Research Institute, Jhansi, India Brijesh K. Mehta ICAR-Indian Grassland and Fodder Research Institute, Jhansi, India Jasmina Milenković Institute for Forage Crops Kruševac, Kruševac, Republic of Serbia Debasis Mitra Department of Microbiology, Raiganj University, Raiganj, West Bengal, India Sananda Mondal Department of Crop Physiology, Institute of Agriculture, VisvaBharati, Sriniketan, West Bengal, India Debasish Panda Department of Crop Physiology, Institute of Agriculture, VisvaBharati, Sriniketan, West Bengal, India P. K. Pandey Indira Gandhi Agricultural University, Raipur, Chhattisgarh, India Pooja S. Patel Department of Plant Pathology, University of Agricultural Sciences, GKVK, Bangalore, Karnataka, India P. K. Pathak ICAR - Indian Grassland and Fodder Research Institute, Jhansi, India Amit Kumar Patil ICAR - Indian Grassland and Fodder Research Institute, Jhansi, India Mirjana Petrović Institute for Forage Crops Kruševac, Kruševac, Republic of Serbia B. R. Praveen Agronomy Section, ICAR - National Dairy Research Institute, Karnal, Haryana, India Pooja Purushotham Department of Plant Pathology, University of Agricultural Sciences, GKVK, Bangalore, Karnataka, India G. A. Rajanna ICAR - Directorate of Groundnut Research, Ananthapur, AP, India

Editors and Contributors

xxi

Kunasekaran Rajarajan ICAR-Central Agroforestry Research Institute, Jhansi, India B. Rajashekhar KVK, Palem, PJTSAU, Hyderabad, India M. Rajashekhar Agricultural Entomology, PJTSAU, Hyderabad, India Gaurav Rakhonde Department of Plant Pathology, University of Agricultural Sciences, GKVK, Bangalore, Karnataka, India Maneet Rana ICAR-Indian Grassland and Fodder Research Institute, Jhansi, India Namburi Karunakar Reddy Department of Plant Pathology, University of Agricultural Sciences, GKVK, Bangalore, Karnataka, India Manjanagouda S. Sannagoudar ICAR - Indian Grassland and Fodder Research Institute, Jhansi, India ICAR - Indian Institute of Seed Science, Regional Station, Bengaluru, India S. Saranya GD Goenka University, Gurugram, Haryana, India Monika Satankar IARI, New Delhi, India Eetela Sathyanarayana Department of Soil Science and Agricultural Chemistry, Agricultural College, Palem, PJTSAU, Hyderabad, India Himanshu Sharma Department of Biochemistry, Punjab Agricultural University, Ludhiana, Punjab, India M. L. Sharma Indira Gandhi Agricultural University, Raipur, Chhattisgarh, India P. Shashikumara ICAR-Indian Grassland and Fodder Research Institute, Jhansi, India Amit K. Singh ICAR - Indian Grassland and Fodder Research Institute, Jhansi, India Magan Singh Agronomy Section, ICAR-National Dairy Research Institute, Karnal, Haryana, India Mahendra Singh ICAR-Indian Grassland and Fodder Research Institute, Jhansi, India Naseeb Singh ICAR - Research Complex for NEH Region, Umiam, India S. K. Singh ICAR - Indian Grassland and Fodder Research Institute, Jhansi, India Sultan Singh ICAR-Indian Grassland and Fodder Research Institute, Jhansi, India Tejveer Singh ICAR-Indian Grassland and Fodder Research Institute, Jhansi, India Partha Sarathi Singha Assam University, Silchar, Assam, India Rajesh Kumar Singhal Division of Crop Improvement, ICAR-Indian Grassland and Forage Research, Jhansi, India

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Editors and Contributors

Pooja Tamboli Plant Animal Relationship Division, ICAR-IGFRI, Jhansi, UP, India Sukumar Taria ICAR-Central Agroforestry Research Institute, Jhansi, India K. Charan Teja Department of Agronomy, Agricultural College, Palem, PJTSAU, Hyderabad, India Saideep Thallapally Forest College and Research Institute, Mulugu, Hyderabad, Telangana, India Rupesh Tirunagari Division of Soil Science and Agricultural Chemistry, ICARIARI, New Delhi, India Vikas C. Tyagi ICAR-Indian Grassland and Fodder Research Institute, Jhansi, India Department of Soil and Crop Sciences, Texas A&M University, College Station, TX, USA Deepak Upadhyay Plant Animal Relationship Division, ICAR-IGFRI, Jhansi, UP, India Pashupat Vasmatkar Department of Biochemistry, Punjab Agricultural University, Ludhiana, Punjab, India V. K. Wasnik ICAR - Indian Grassland and Fodder Research Institute, Jhansi, India

Abbreviations

ACC ADF AF AI APX ARMS ATP CA CAM CAT CCS Chl CHS CP CSA DHAR DSS EC ECH EE F3H FAO G× E GDP GHG GoI GR GS GVA ha IAA ICE ILRI

1-aminocyclopropane-1-carboxylic acid Acid-detergent fiber Agroforestry Artificial intelligence Ascorbate peroxidase Amplification refractory mutation system Adenosine triphosphate Conservation agriculture Crassulacean acid metabolism Catalase Combined charging system Chlorophyll Chalcone synthase Crude protein Climate smart agriculture Dehydroascorbate reductase Decision support system Electrical conductivity Evaporative cool hydroponic Ether extract Flavanone 3-hydroxylase Food and Agriculture Organization Genotype × environment Gross domestic production Greenhouse gases Government of India Glutathione reductase (GR) Genomic selection Gross value added Hectare Indole-3-acetic acid Internal combustion engine International Livestock Research Institute xxiii

xxiv

IPCC IVDMD LTS MAS MDA MDHAR Mha MS MSI NAFPs NAGS NDF NGS OFT PAL PCD PCR PM POX PPO PS RAPD ROS RWC SALT SAR SMART SOC SOD SPM SSR STS TAL TCHQD TDN TILLING TOT TRGA UHPLC UV rays VOCs WSC

Abbreviations

Intergovernmental Panel on Climate Change In vitro dry matter digestibility Long-term salt stress Marker-assisted selection Malondialdehyde Monodehydroascorbate reductase Million hectare Mass spectrometry Membrane stability index Non-timber forest products National active germplasm site Neutral detergent fiber New generation sequencing On farm testing Phenylalanine ammonia-lyases Programmed cell death Polymerase chain reaction Particulate matter Peroxidase Polyphenol oxidase Photosystem Random amplified polymorphic DNA Reactive oxygen species Relative water content Sloping Agricultural Land Technology Sodium adsorption ratio Single-molecule real-time Soil organic carbon Superoxide dismutase Suspended particulate matter Simple sequence repeat Short-term salt stress Tyrosine ammonia-lyases Tetra chlorox hydroquinone dehalogenase Total digestible nutrients Targeting induced local lesions in genomes Transfer of technology Total reported geographic area Ultra-high performance liquid chromatography Ultraviolet rays Volatile organic compounds Water soluble carbohydrate

Part I Forage Crop Improvement

1

Genetic and Genomic Resources of Range Grasses: Status and Future Prospects Vikas C. Tyagi, Tejveer Singh, Nilamani Dikshit, Sultan Singh, Maneet Rana, Rahul Kaldate, Prabhu Govindaswamy, Hanamant M. Halli, Avijit Ghosh, Rajesh Kumar Singhal, and Manjanagouda S. Sannagoudar

1.1

Introduction

Grasses play a key role in human life, provide food for human beings, and feed for livestock. They have a crucial role in terms of food production and in the delivery of ecosystem services such as water supplies, biodiversity, and carbon sequestration (Ravagnani et al. 2012). Poaceae is considered the fifth most species-rich Angiosperm family (Davis et al. 2009) after Asteraceae, Leguminosae, Orchidaceae, and Rubiaceae. According to the latest estimate, Poaceae comprises 11,554 species under 759 genera (The plant-list-2013: version 1.1). In India, it is the largest family and is represented by 1334 species belonging to 261 genera (Karthikeyan 2005). This works out to be about 14% of the total grass species of the world. With a cosmopolitan distribution found in virtually all non-marine habitats, and especially prevalent in sub-arid habitats, grasses account for roughly 25% of global terrestrial primary production (Still et al. 2003) and form the crown constituent of grasslands, the most extensive biome on the planet, occupying an estimated 31–43% of the worlds land surface (World Resources 2000). Grasses rank as the top plant kingdom, having been in cultivation for over 10,000 years (Kilian et al. 2007), they form the V. C. Tyagi (✉) ICAR-Indian Grassland and Fodder Research Institute, Jhansi, India Department of Soil and Crop Sciences, Texas A & M University, College Station, TX, USA T. Singh · N. Dikshit · S. Singh · M. Rana · R. Kaldate · P. Govindaswamy · H. M. Halli · A. Ghosh ICAR-Indian Grassland and Fodder Research Institute, Jhansi, India R. K. Singhal Division of Crop Improvement, ICAR-Indian Grassland and Forage Research, Jhansi, India M. S. Sannagoudar ICAR-Indian Grassland and Fodder Research Institute, Jhansi, India ICAR-Indian Institute of Seed Science, Regional Station, Bengaluru, India # The Author(s), under exclusive license to Springer Nature Singapore Pte Ltd. 2023 R. K. Singhal et al. (eds.), Molecular Interventions for Developing Climate-Smart Crops: A Forage Perspective, https://doi.org/10.1007/978-981-99-1858-4_1

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most important food crop on Earth today, providing roughly 80% of the annual global food with domestic animals also being raised on diets partly or wholly of grasses. Grasslands provide fodder for cattle and the progress of the dairy industry mainly depends on grasses and grassland of a region. Grasses are found on all continents and in all climatic zones. They grow from sea level to the highest elevation. They grow in marshes, deserts, woodland, sand, rocks, and almost in all types of soils. The grasses are also called pioneer species. Range grasses primarily grow as natural vegetation in the rangeland and provide grazing and forage for livestock and wildlife. These grasses differ widely with respect to ecology, elevation range, genomic resources, usage, nutritional composition, and yield. These forage grasses are used as cut fodder or grazed pastures and also as harvested seed crops (generally from dual-purpose food and feed crops) which may be given directly to livestock after preservation as fermented green matter (silage and haylage) or as dried product (hay, pellets, and cube concentrates). In India, the livelihood of nearly 70% population in rural areas is dependent on farming. Further, the livestock sector is an integral component of agriculture and plays a key role in improving the economy and providing employment, especially in rural areas. It contributes to nearly 4.11% to the national GDP through milk, meat, wool as well as farmyard manure. India’s geographical area is 328.72 Mha, which supports 20% of the world’s livestock. The country is holding about 56.7% of the world’s buffaloes, 12.5% cattle, 20.4% small ruminants, 2.4% camel, 1.4% equine, 1.5% pigs, and 3.1% poultry (IGFRI Vision 2050). As per the 19th livestock census 2012, in India, the total livestock population in the country is 512.05 million. The important source of fodder supply is mainly from crop residues, cultivated fodder, and fodder from common property resources like forests, permanent pastures, and grazing lands. Presently, the country faces a net deficit of 35.6% green fodder, 10.95% dry crop residues, and 44.0% concentrate feed ingredients (IGFRI Vision 2050). The main reasons for the low productivity are the non-availability of sufficient and high-quality fodder and feed including grazing facilities (Roy 1993). The projected livestock population estimates in India are based on the 10 and 11th fiveyear plan document, by the Government of India, which shows the increasing rate of livestock population. Domestic animals are about 500 million and are projected to rise at a rate of 1.23% in the near future (Datta 2013). So, it is important to maintain sustainability in livestock feed and fodder but presently the nation is facing shortfalls in livestock feed and fodder. In India, more than 50% of animals are dependent on 12 Mha grazing lands (Roy and Singh 2013). There are two types of grass species, i.e., C3 and C4 species depending on their primary carbohydrates synthesized at the beginning of the photosynthesis (Carlier et al. 2009) are found in these grasslands including some clover and other legumes, soft and hardy herbs, and shrubs. Important grass species of Indian rangelands/ grasslands, common name, distribution, and origin are listed in Table 1.1. Further, grass species are also considered to be a suitable forage plant in a rangeland/ grassland in comparison to legumes due to their wider adaptability, rejuvenation capability, continuous vegetative growth, rapid ground coverage, an excellent source

Cenchrus setigerus Vahl

Chloris gayana Kunth Chrysopogon fulvus (Spreng.) Chiov. Cymbopogon citratus (DC.) Stapf

5

6

8

7

Cenchrus ciliaris L.

4

Bird wood grass, Dhaman grass, KalaDhaman Chloris, Rhodes grass Dhwalu, Gusia, Pandhri Kusal, and Karehull Lemongrass or oil grass

Ceylon sheep Grass, Congo signal grass Buffelgrass, Anjan Grass

West Indies, Sri Lanka, Java, Guatemala, Brazil, Mexico, Congo, Tanzania, India, Thailand, Bangladesh, Madagascar, Guinea, China, and the Philippines

Asia and East Africa and throughout India

South Africa, USA, India

Throughout Africa, Madagascar and eastwards to Burma and Ceylon African, Arabian country, Australia, and South America

Karnataka, Tamil Nadu, and southern states Aravalli Hills in Rajasthan, Central Plateau as well as lower range of Himalaya UP, Bihar, Manipur, Meghalaya, West Bengal

Rajasthan, Punjab, Haryana, Gujarat, Western UP, and Tamil Nadu Punjab, Rajasthan, Haryana, Gujarat

Coastal regions, i.e., Odisha, Sundarbans, Goa, Kerala

Brachiaria brizantha (A. Rich.) Stapf

3

Malaysia

(continued)

Tropical and subtropical regions

Sub-Saharan Africa

Africa, Arabia, the Middle East, and India

Africa, Arabia, the Middle East, and India

Tropical Sub-Saharan Africa

Asia

Bothriochloa pertusa (L.) A. Camus

2

Origin Asia

Accepted name Bothriochloa bladhii (Retz.) S.T. Blake

S. no. 1

Distribution (India) Throughout tropical to subtropical parts from Punjab to West Bengal Uttar Pradesh, Northern and Southern states

Table 1.1 Important grass species and their geographical distribution Distribution (globally) India, Australia, Africa, Pacific Island, Nepal, Tropical Asia, China, and Tropical Africa Kenya, Uganda, southern and north-eastern tropical Africa, Southeast Asia, Caribbean archipelago, Pakistan, Sri Lanka, Southeast Asia, Indochina, Thailand, and Indonesia Tropical Africa, Australia

Genetic and Genomic Resources of Range Grasses: Status and Future Prospects

Common name Forest blue grass, Bada Phulwa, Fulkara Indian blue grass, Phulwa

1 5

Heteropogon contortus (L.) P. Beauv. ex Roem. & Schult. Iseilema laxum Hack.

11

Panicum antidotale Retz.

Panicum maximum Jacq.

Paspalum notatum F.

Pennisetum pedicellatum Trin.

13

14

15

16

12

10

Accepted name Cymbopogon pendulus (Nees ex Steud.) W. Watson Dichanthium annulatum (Forssk.) Stapf

S. no. 9

Table 1.1 (continued)

Deenanath grass, Kaysuwa, Barn

Blue panic, Giant panic, Bansi Guinea grass, green panic, Gini grass Bahia grass

Lampa, Kusal, Kusali, Kusli, Lamb, Lap, Musel Musiyal or Machuri, Moshi

Common name Lemongrass, barbed wire grass, citronella grass Marvel grass, Kale Ghass

Brazil and distributed in Mexico, Texas South America, and India North tropical Africa

Tropical Africa

Australia, arid and semiarid regions of Afghanistan and Persia

Bihar, West Bengal, Haryana, Punjab, MP, and UP

West Bengal, Bihar

Assam, Punjab, Rajasthan, Kerala

North Himalaya to Cape Comorian and in the grassland of the east to west and whole of south Maharashtra, Gujarat, Kerala, Tamil Nadu, Andhra Pradesh, Madhya Pradesh, and Uttar Pradesh Assam, Punjab, Rajasthan, Kerala

Tropics and subtropics

Ceylon, Mauritius, Peninsular India, and Sri Lanka

Throughout India

Distribution (India) UP, Bihar, Manipur, Meghalaya, West Bengal

Burma, Africa, Australia, and southern USA

Distribution (globally) China, Nepal, India, Bangladesh, Myanmar, Vietnam

West Africa

Central America

Indian subcontinent

Indian subcontinent

Southern Asia

Africa, the Arabian Peninsula, the Indian Subcontinent, and southeastern Asia Tropical and subtropical part of the world

Origin Southeast Asia

6 V. C. Tyagi et al.

Setaria sphacelata (Schumach.) Stapf and C.E. Hubb. ex Moss Chrysopogon zizanioides (L.) Roberty

18

19

Sehima nervosum (Rottler) Stapf

17

Khas, Vetiver

Rat’s tail grass, White grass, Sen, Poona, Suekai Setaria, Golden millet, Nandi

India, Myanmar, and Sri Lanka

Tropical Africa, Australia, South Asia, America

Central East Africa and Sudan, Southeast Asia, Australia

Indian subcontinent

Tropical and subtropical Africa

Indian subcontinent

Madhya Pradesh, Uttar Pradesh, Gujarat, Karnataka, Maharashtra, Andhra Pradesh, Tamil Nadu Assam, Nagaland, Meghalaya, Uttar Pradesh, Madhya Pradesh, Odisha, Punjab Rajasthan, Uttar Pradesh, Assam, Karnataka, Tamil Nadu, Kerala, and Andhra Pradesh

1 Genetic and Genomic Resources of Range Grasses: Status and Future Prospects 7

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of carbohydrates, good candidates for soil conservation, improve soil physical properties, enrich soil carbon, highly productive and nutritional characteristics (Trivedi 2002a, b). There has been considerable progress in the knowledge of the different aspects of range grasses particularly the distribution, ecology, genetic resources, breeding, genomic resources, nutritive value, species suitability to different ecosystems, etc. Hence, an attempt has been made in this review to understand the important grass species and their geographical distribution, genetic resources of tropical grasses, grass cover and distribution of important grasses, forage grass genetic resources conservation and status, forage breeding, problems in forage crop breeding, ecotypic selection, hybridization, identification and diversification of sexual lines in tropical grasses, intraspecific hybridization, genomic resources, linkage maps, molecular markers closely associated with desirable forage breeding traits for use in markerassisted selection, the nutritive value of tropical range grasses, hay, and silage of grasses, matching nutritive value with the animal requirement, biomass yield and productivity, suitable pasture species for dry lands in different rainfall (mm) and their yield and future research.

1.2

Genetic Resources of Tropical Grasses

Rangelands are integral parts of the ecosystems that have many values beyond their use as forage, including watershed protection, ethnomedicinal, natural, and beauty. The maintenance of enormous genetic diversity is mandatory for broadening the genetic base of the present and future forage improvement programs. Extensive collection, proper evaluation, in-depth study of genetic attributes, and cataloging of germplasm are prerequisites for its efficient utilization. There are about 11,000–12,000 species of grasses divided among about 750–770 genera in the world (Kellogg 2015; Soreng et al. 2017). Out of these, about 600 species of grasses are currently used for grazing and livestock feeding in the world. Indian subcontinent is one of the world’s mega centers of crop origin and crop plant diversity, possesses rich genetic and species diversity in native forage grasses (1256 spp.), as about one-third of them, chiefly belonging to the tribes, namely Andropogoneae, Paniceae, and Eragrostideae, are of forage importance (ICAR 2009). Further, the distribution of grasses is primarily governed by climatic factors, chiefly by latitudinal influence followed by altitude and topography, particularly the soil moisture relationship. Dabadghao and Shankarnarayan (1973) prepared a comprehensive document entitled “The Grass Covers of India” dealing with grasses and grasslands of the entire territorial expanse of the country. Their study concluded that aridity of the soil coupled with abiotic (mainly fire) and biotic interferences (grazing and looping) favors the development of grasslands, whereas moist soils without these disturbances provide conditions for woodlands and forests. Five types of grass covers were identified for the entire country based on perennial and annual grasses, herbaceous, and tree species including information on environmental conditions of these regions and successional trends in relation to changes in soil humus and moisture.

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Genetic and Genomic Resources of Range Grasses: Status and Future Prospects

1.3

9

Forage Grass Genetic Resources Conservation and Status

Forage grasses are difficult to conserve but forage genetic resources are usually stored in seed gene banks. Grass seeds can be safely stored in situ as well as ex situ. There has been little research on seed storage of many kinds of forage grasses and as a result, there is limited information about the storage behavior of seeds of many species. Ex situ conservation of forages usually involves the storage of seed in gene banks in standard temperatures of 0–4 °C for active collections and -18 °C for base collections, both at low seed moisture (3–7%). However, some seeds of forage grass species have been found to survive more than five years of cold storage (at 5% moisture content and 8 °C) without substantial loss of viability and can be stored in seed gene banks. Seeds stored in gene banks for long-term conservation are better protected from external climatic or environmental factors. Many forage grasses have short-lived seeds, i.e., seeds remain viable for a very few years, e.g., Guinea grass (Harty et al. 1983) and signal grass (Whiteman and Mendra 1982); shy seeders, i.e., only a small percentage of the seeds have a caryopsis and will germinate, e.g., Pennisetum purpureum and Digitaria erianthum which need to be maintained in field gene banks. The main objective of in situ conservation is to maintain the environment which has led to the development of the distinctive properties of genetic resources. In situ conservation allows genetic evolution and specific environment to influence the further development of forage plants. By ex situ conservation, the genetic structure of the original seed sample is preserved. The durability of seed storage is influenced by seed condition and initial germination. Worldwide, there are nearly 1500 gene banks which are registered in the WIEWS (World Information and Early Warning System on PGR) database (http://apps3.fao. org/wiews/) with the holding of 7.1 million accessions belonging to 53,109 species, including major crops, minor or neglected crop species, as well as trees and wild plants. Out of the total germplasm stored, 651,024 accessions belong to forage crops (FAO. The State of ex situ Conservation: The Second Report on the State of the world’s PGRFA. 2010. p. 351. http://www.fao.org/docrep/013/i1500e/i1500e03. pdf). Conservation of Forage germplasm under the Consultative Group of International Agricultural Research (CGIAR) is a part of germplasm conservation activity coordinated on plant genetic resources. Among the international organizations, the major forage germplasm repository is CIAT Columbia, safeguarding one of the largest and most diverse tropical forage collections, with more than 700 different species from 75 countries. The collection’s main emphasis is on legumes (more than 21,000 accessions), although almost 1700 grasses are being conserved as well. More than half of the preserved germplasm was collected between 1977 and 1993 as part of 75 explorations. The ILRI gene bank conserves almost 19 thousand accessions of forages from over 1000 species, out of which 1820 accessions of grasses belong to 139 species. Similarly, CSIRO-Australia has conserved 2670 accessions of different forage grasses belonging to 252 species (Sandhu et al. 2015). The ICAR-National Bureau of Plant Genetic Resources (NBPGR) is the nodal agency for the characterization, evaluation, maintenance, conservation, documentation, and distribution of germplasm resources in India. Currently, a total of >8000

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accessions of cultivated and range grasses and legumes are being maintained at the long-term storage (LTS) module of the National Gene Bank at NBPGR, New Delhi. Likewise, ICAR-Indian Grassland and Fodder Research Institute (IGFRI) is a unique R & D organization in South Asia for sustainable agriculture through quality forage production for improved animal productivity. ICAR-IGFRI being the National Active Germplasm Sites (NAGS) on forages works with its three regional stations and All India Coordinated Research Project (AICRP) on forage crops with 18 coordinated centers. Besides conducting explorations, acquiring germplasm from various outside national and international agencies, followed by conservation in IGFRI’s Medium Term Storage, IGFRI now holds >9940 accessions representing >67 forage genera (Sahay et al. 2018).

1.4

Breeding System in Forage Grasses

The breeding system in tropical perennial forage grasses is quite different from the cultivated cereals. The temperate grasses represent all three types of modes of reproduction, i.e., sexual, asexual, and clonal. About 60% of tropical grasses are apomictic, in which identical true-to-type plants of the mother are produced. The advantage of this system is that hybrid vigor can be fixed if the phenomenon of apomixis is properly understood. Choice of the vigorous plants and the percentage of selection intensity have contributed astoundingly to genetic advance. Improvement of forage yield, quality, and adaptation has given primacies traditionally by phenotypic breeding. Breeding is the key to the future development of superior forage varieties. The selection of plants has involved vigorous nature and contributed greatly to genetic advance (Busey 1989). In all important tropical grasses, concentrated efforts have been made worldwide to develop varieties for different growing conditions including cut and carry and for pasture and grassland purposes.

1.4.1

Problems in Forage Crop Breeding

Breeding in forage crops involves the same principles of breeding for self or crosspollinated crops in sexually propagated crops. Dealing breeding with apomictic grasses should be taken into consideration while choosing the breeding methods. Some grasses are difficult to propagate as individual lines from seed, creating problems to maintain their identity due to their cross-pollination nature. Species with small floral parts make artificial hybridization and pollination control difficult. Diversity in the pollination of the different species, irregularities in fertilization, seed set, and perennial habit of crops in important forage species, etc. limit the extent to which they may be self-pollinated/cross-pollinated. This restricts the attempt for recombination breeding in the apomixis species causing problems in crossing and obtaining gene recombination. Similarly, low viability and seed setting in forage species are other important points that restrict the variability generated through seeds. Breeding lines may perform differently with different environments and

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11

systems of grazing management, along with taking long years to evaluate the perennial grasses for persistence and productiveness. Seventy percent of the species in the grass family are polyploids causing genome complexity (Poehlman 1987). The primary information on breeding behavior, methods of breeding, and evaluation procedures have been developed for many forage species. A large number of forage species increase the problem of assembling and maintaining adequate germplasm collections of each species. Breeding methods normally adopted in perennial forage grasses are as follows.

1.4.2

Ecotypic Selection

Ecotypic selection is the evaluation of a large number of accessions and the release of a superior one. Vegetative materials are increased from promising accessions tested in multi-locations over the years and if found superior in agronomic traits released as variety. Direct selection based on phenotypic traits in large germplasm and identification of superior accessions has given importance and assured 80–130% more productive in leaf, 26% increase in digestibility (Burton 1989; Burton et al. 1993). Most cultivars of tropical grasses available commercially are wild ecotypes selected from natural diversity (Hacker and Jank 1998).

1.4.3

Hybridization

Most tropical grasses are reproduced through apomictic mode. The apomictic mode of reproduction is the biggest bottleneck in recombination breeding in tropical grasses. Identification and generation of sexual lines in apomictic grasses may prove to be a new era of research direction. On the availability of appropriate sexual lines, the benefits of hybrid vigor can be fixed by taking the advantage of apomixis.

1.4.4

Identification and Diversification of Sexual Lines in Tropical Grasses

Some species such as guinea grass, buffelgrass, Paspalum, and Brachiaria are either obligate or facultative sexual lines. Buffelgrass is a mainly obligate apomictic species (Snyder et al. 1955), and the use of obligate sexual or apomictic genotypes with high levels of sexuality is the only alternative for conventional crosses (Burson et al. 2012; Hussey et al. 1993). Bashaw (1962) identified a source of sexuality, an off-type plant, product of a mutation that showed poor forage aptitude (Griffa et al. 2005), significant susceptibility to salt stress (Griffa 2010; Lanza Castelli et al. 2010), and little genetic contribution for apomictic F1s of agronomic interest (Griffa et al. 2005). Quiroga et al. (2013) further diversified two of them were identified as sexual genotypes. Both highly sexual genotypes could be used as female parents in crosses for obtaining improved cultivars of buffelgrass. Development of a synthetic

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autotetraploid sexual (2n = 4x = 36), B. ruziziensis genotype through colchicine doubling (Swenne et al. 1981) facilitated recombination between sexual plants and tetraploid apomictic Brachiaria pollen donors and enabled the establishment of breeding programs at the International Center for Tropical Agriculture (CIAT) and Empresa Brasileira de Pesquisa Agropecuária (EMBRAPA) (Brazilian Enterprise for Agricultural Research) in the late 1980s (Miles 2007). Sexual plants of guinea grass have already been found by Combes and Pernes (1970), Smith (1972), and Hanna et al. (1973), and their chromosome numbers were determined, as 2n = 16 (Combes and Pernes 1970) and 2n = 32 in five of 18 sexual plants (Hanna et al. 1973). Nakajima et al. (1979) isolated sexual plants from African collections or introductions, and their chromosome numbers were revealed to be 2n = 16, 18, and 32, diploid and tetraploid.

1.4.5

Intraspecific Hybridization

In Panicum maximum, increased foliage percentage, seed production, and rapid re-growth (Jank et al. 2001, 2004; Muir and Jank 2004; Resende et al. 2004) have been observed in F1 compared to parents. Many wide adaptable varieties of guinea grass (PGG-9, PGG-14, PGG-19, PGG-518, and CO-2) have been developed and released from PAU, Ludhiana, and TNAU, Coimbatore. The high forage quality and determined flowering cycle of B. ruziziensis, the yield, and resistance to spittlebug of B. brizantha, and the vigor and adaptation to acid, infertile soils of B. decumbens (Peters and Lascano 2003) have been observed.

1.4.6

Wide Hybridization (Interspecific and Intergeneric)

In tropical grasses, a large number of interspecific and intergeneric hybrids were developed both for genetic improvement programs and phylogenetic studies and the transfer of apomixes. Crossing between sexual and apomictic plants of B. ruziziensis directly or by chromosomal doubling with colchicine to make cross-compatible and to induce variability along with quality traits has been attempted in grasses (Lutts et al. 1994). The ploidy level resulted in the discovery of a sexual accession, which was artificially doubled to make crosses with tetraploid accessions feasible (Pinheiro et al. 2000). High seed producing cultivars of Pennisetum purpureum were obtained by crossing P. purpureum and P. glaucum (L.). The progenies from this cross are triploid and sterile, but once doubled by colchicine become hexaploid and fertile (Pereira and Lédo 2008). Ploidy series of grasses has been developed at ICARIGFRI, Jhansi, India, to decode the genome complexity of the majority of native grass species. ICAR-IGFRI has developed world largest ploidy series of Panicum maximum to understand the ploidy regulated phenomenon at the genetic level. Better fodder plant types in pearl millet generated through interspecific hybridization P. glaucum × P. squamulatum; P. glaucum × P. orientale at IGFRI, Jhansi. The resultant plant has shown higher biomass, better adaptability, better tolerance for

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Genetic and Genomic Resources of Range Grasses: Status and Future Prospects

13

moisture deficit, and improved forage attributes compared to other tropical grasses (IGFRI 2015). Novo et al. (2016) used a colchicine-induced sexual autotetraploid genotype of P. plicatulum Michx. to obtain interspecific hybrids using the apomictic species, P. oteroi, as the pollen donor. The hybrids showed less irregular meiotic behavior with fewer quadrivalents and more bivalents than either parent. Fertility among interspecific hybrids varied from complete sterility in some of them to seed productions in others that were approximately twice as much as for either parent. The great variability of seed set performance may well be a drastic genetic consequence of joining two homologous chromosome sets of P. plicatulum together with two homologous sets of P. oteroi that, in turn, have some homeology between them. Most hybrids reproduce by sexual means; thus, they could be used as female parents in backcrosses and in crosses with other species of the Plicatula group for interspecific gene transferring in breeding programs. In India and worldwide Bajra (P. glaucum) × Napier (P. purpureum) hybrids have been recognized as revolutionary due to their wide adaptation, wide biomass yield, better forage attributes, tolerance to abiotic stresses, and perennial nature.

1.5

Seed Production in Range Grasses

Seed is the primary and most essential input for enhancing crop yield. As per the estimation, only 25–30% of required quantity of quality seed is available for cultivated fodder crops and less than 10% in case of range grasses and legumes in India. Conversely, range grass seed production faces many challenges like lack of supply of good quality pasture seeds and a lack of knowledge on seed standards, production practices, and conservation practices (ASARECA 2007). Therefore, development of technologies to improve seed yield and quality will be a step forward toward quality forage production and livestock productivity.

1.5.1

Constraints in Seed Production

In spite of several interventions, seed production in tropical forage crops faces many more “problems and limitations” (Table 1.2). Since varietal development is the main focus in case of cultivated fodder production but seed production technology is not well documented, whereas in the case of range species due to a lack of large-scale production and the above-mentioned problems no specific seed production technology is accessible. Therefore, it is necessary to develop strategies for higher seed yields to obtain moderate yields of green forage as well as seed from the same crop by following appropriate management techniques (Geetha 2001).

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Table 1.2 Problems and limitations in seed productions Constraints 1. Indeterminate growth 2. Uneven maturity and seed shattering 3. Blank seed 4. Low density of ear-bearing tillers 5. Lodging

6. Lower harvest index 7. Lack of post-harvest technology 8. Degradation and destruction of natural grasslands/rangelands and biodiversity 9. Climate change/global warming related problems 10. Lack of knowledge among stakeholders regarding seed production

Limitations Major hindrance for large-scale cultivation on commercial basis and mechanization Makes impossible to realize the full potential of seed production and difficulty in harvesting Due to poor ovule to seed ratio and lead Only 30–50% tillers possess inflorescence at the time of peak flowering Common problem due to prolonged and vigorous vegetative growth, poor harvest index due to higher biomass production It is hardly 2–3% Leads to loss of final product and affect quality Reduces the area under cultivation Making the seed production risky and uncertain This results in poor quality and increased cost on production

Source: Sunil Kumar et al. (2012) and Trivedi (Trivedi 2002a, b)

Table 1.3 Principles of quality seed production 1 2 3 4 5 6 7 8 9 10 11 12

1.5.2

Selection of suitable crop/variety based on the weather, growing length, soil, and market availability Seedbed preparation to obtain proper crop establishment The optimum time for planting Selection of suitable plant geometry The optimum depth of sowing and seeding equipment Water management Weed management Nutrient management Management of pests and diseases Canopy management/defoliation Harvesting Burning/fall management

Principles of Range Grass Seed Production

In spite of high demand and lower seed yield for tropical range grasses little or no attention has been given to improving the performance (Mnene et al. 2017). Hence it is necessary to understand the principles and factors involved in seed multiplication. This helps to determine the correct management for quality seed production or to improve the current low seed yield per unit area (Table 1.3).

1

Genetic and Genomic Resources of Range Grasses: Status and Future Prospects

1.6

15

Genomic Resources in Range Grasses

Compared to dual-purpose crops, genomic resources are very meager in many of the range grasses. For the breeding of range grasses, unique problems and difficulties are there which are not observed in the cultivated crops, these include their polyploid and apomictic nature, non-synchronous flowering/anthesis and spikelet maturity, abscission of spikelets after maturity, self-incompatibility, small floral parts, poor ovule to seed ratio, low seed viability, poor seedling vigor, inadequate germplasm base, lack of disomic inheritance, and relatively short domestication history (Roy et al. 2019). However, over the last decade the genus Brachiaria, Cenchrus, Panicum, and Paspalum are well explored for the development of genomic resources. For a molecular breeding program, the availability and easy accessibility of genomic resources is a prerequisite. Genomic resources like restriction fragment length polymorphism (RFLPs), random amplified polymorphic DNA (RAPD), amplified fragment length polymorphisms (AFLPs), inter-simple sequence repeats (ISSR), and more efficient and informative simple sequence repeat (SSR) assays based on analysis of either expressed sequence tag (EST) or genomic DNA have been developed in range grasses (Cidade et al. 2013; Bishoyi et al. 2016; Abdi et al. 2019; Raja et al. 2019; Ribotta et al. 2019; Tegegn et al. 2019; Yadav et al. 2019) for use in diversity analysis, phylogenetic analysis, and for the identification of a taxonomic relationship among the members of a different genus of range grasses. The marker type along with the number and amplified alleles available in various range grasses has been summarized in Table 1.4. Further, in addition to conventional molecular marker technology, a large repertoire of potential molecular markers (5035 microsatellites (SSRs) and 3,46,456 single nucleotide polymorphisms (SNPs)) has been generated in Panicum maximum using de novo transcriptome assembly (Toledo-Silva et al. 2013). Such information will definitely help in marker-assisted selection-based breeding research in the near future. Recently a high throughput genotyping system in the form of a genotyping by sequencing (GBS) has also been utilized in Panicum maximum (Lara et al. 2019) for investigating the effect of the allele dosage. Genetic mapping and gene tapping in range grasses have not been attempted much either due to non-availability of mapping populations in apomictic species or due to poor genomic resources. The first and foremost step for mapping is the construction of linkage maps. Genetic linkage maps have become keystones in the basic genetic analysis as well as in applied plant breeding. Linkage maps have assisted in the identification of DNA markers linked to major genes of agronomic importance and have permitted identification of tightly linked DNA tags for use as diagnostic tools in plant breeding. In range grasses, linkage map has been successfully developed in Cenchrus ciliaris (Jessup et al. 2003; Yadav et al. 2012), Chloris gayana (Ubi et al. 2004), Panicum maximum (Ebina et al. 2005; Deo et al. 2019), and Paspalum notatum (Ortiz et al. 2001; Stein et al. 2007). The list of published maps along with cross-type, mapping population, markers mapped, no of linkage groups, distance (cM), and average marker density (cM) in various range grasses has been summarized in Table 1.5. Once a linkage map is developed, the next step is to

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Table 1.4 Genomic resources available in different range grasses S. no. 1 2 3

4

Range grass Bothriochloa bladhii (Retz.) S.T. Blake Bothriochloa pertusa (L.) A. Camus Brachiaria brizantha (A. Rich.) Stapf

Cenchrus ciliaris L.

Marker type –

Number –

Reference –

ctEST-SSR

16

RAPD

10 (107 allele)

SSR

13

SSR

15

SSR

93

ISSR

10 (441 allele)

SSR

39

SSR

23 (459 allele)

RAPD RFLP/STS RAPD

1 1 106

RAPD STS SCAR

2 1 6

RAPD SCAR RAPD

20 1 154 (1296 loci)

ISSR

SCAR

3 (27 reproducible bands) 1

AFLP

60

AFLP SCAR EST-SSR

180 8 116

AFLP SCAR

50 primer combinations (5713 bands) 4

Abdi et al. (2019) Ambiel et al. (2008) Jungmann et al. (2009) Vigna et al. (2011) Ferreira et al. (2016) Nitthaisong et al. (2016) Triviño et al. (2017) Tegegn et al. (2019) Lubbers et al. (1994) Gustine et al. (1996) Gustine et al. (1997) Roche et al. (1999) Dwivedi et al. (2007) Chandra and Dubey (2008) GutierrezOzuna et al. (2009) Kumar et al. (2010) KharratSouissi et al. (2011) Yadav et al. (2012) Abdi et al. (2019) Kumar and Saxena (2016)

(continued)

1

Genetic and Genomic Resources of Range Grasses: Status and Future Prospects

17

Table 1.4 (continued) S. no.

Range grass

Marker type AFLP

5

Cenchrus setigerus Vahl

RAPD AFLP SCAR

6

7

8

9

Chloris gayana Kunth

Chrysopogon fulvus (Spreng.) Chiov.

Cymbopogon citratus (DC.) Stapf

Cymbopogon pendulus (Nees ex Steud.) W. Watson

Number 8 primer combinations (565 bands) 154 (1296 loci)

ctEST-SSR

50 primer combinations (5713 bands) 3 115

AFLP

8

RAPD

14 (135)

AFLP

4 (227)

AFLP

3

AFLP

3 (237)

AFLP RFLP AFLP

SRAP ISSR RAPD

164 25 4 primer pair combinations (152 bands) 20 18 6

RAPD

13

CISP

6

EST-SSR

20 (151)

RAPD

60 (371)

PCR based functional markers RAPD

17 (119)

RAPD

60 (371)

5

Reference Yadav et al. (2019) Chandra and Dubey (2008) Kumar and Saxena (2016)

Abdi et al. (2019) Yadav et al. (2019) Pérez et al. (1999) Ubi et al. (2000) Ubi et al. (2001) Ubi et al. (2003) Ubi et al. (2004) Ribotta et al. (2013) Ribotta et al. (2019) Adams and Daffornli (1999) Adams et al. (1998) Chandra et al. (2013) Kumar et al. (2009) Khanuja et al. (2005) Kumar et al. (2007) Prasad and Shekhar (2013) Khanuja et al. (2005) (continued)

18

V. C. Tyagi et al.

Table 1.4 (continued) S. no.

Marker type EST-SSR

Number 20 (151)

SSR

13 (95)

RAPD ISSR RAPD

90 (785) 70 (579) 7 (55)

RAPD

25 (208)

RAPD

32 (307)

STS

14

ISSR RAPD

5 (61) 27 (269)

STS

14 (106 allele)

CISP

6

ctEST-SSR

17

Heteropogon contortus (L.) P. Beauv. ex Roem. & Schult.

RAPD

33

CISP

6

12 13

Iseilema laxum Hack. Panicum antidotale Retz.

– SRAP EST-SSR ctEST-SSR

– 826 (28 pairs) 25 17

14

Panicum maximum Jacq.

RAPD

16

AFLP RAPD SSR

56 41 (372 allele) 13 (190 allele)

EST-SSR

196

SSR

15 (63 allele)

10

11

Range grass

Dichanthium annulatum (Forssk.) Stapf

Reference Kumar et al. (2009) Kumar et al. (2007) Bishoyi et al. (2016) Chandra et al. (2004) Saxena and Chandra (2006) Chandra et al. (2006) Chandra and Dubey (2007) Saxena and Chandra (2010) Saxena and Chandra (2011) Chandra et al. (2013) Abdi et al. (2019) Carino and Daehler (1999) Chandra et al. (2013) – Huang et al. (2011) Abdi et al. (2019) Ebina and Nakagawa (2001) Ebina et al. (2005) Ebina et al. (2007) YamadaAkiyama et al. (2009) Chandra and Tiwari (2010) (continued)

1

Genetic and Genomic Resources of Range Grasses: Status and Future Prospects

19

Table 1.4 (continued) S. no.

15

Range grass

Paspalum notatum Flüggé

Marker type SSR

Number 7 degenerate primers

SSR

20

SSR

147

SSR SNP ctEST-SSR

5035 3,46,456 6

GBS

1223

RAPD

16

RFLP RAPD AFLP

71 16 5 primer combinations (62 bands) 343 (1700 bands) 30 primer combinations 85 2

RAPD AFLP RFLP RAPD RAPD AFLP AFLP

16

17 18

Pennisetum pedicellatum Trin.

Sehima nervosum (Rottler) Stapf Setaria sphacelata (Schumach.) Stapf and C.E. Hubb. ex Moss

ISSR

2 24 14 combinations (1342 bands) 91

SSR

30 (208)

ctEST-SSR

75

AFLP

8

CISP

6

ILP

26 (67)

SSR

147

SSR

159

Reference Tiwari and Chandra (2010) Sousa et al. (2011) Gupta et al. (2012) Toledo-Silva et al. (2013) Abdi et al. (2019) Lara et al. (2019) Ortiz et al. (1997) Ortiz et al. (2001)

Martínez et al. (2003)

Daurelio et al. (2004) Stein et al. (2004) Espinoza et al. (2006) Cidade et al. (2008) Cidade et al. (2013) Abdi et al. (2019) Yadav et al. (2019) Chandra et al. (2013) Gupta et al. (2011) Gupta et al. (2012) Pandey et al. (2013) (continued)

20

V. C. Tyagi et al.

Table 1.4 (continued) S. no.

19

Range grass

Chrysopogon zizanioides (L.) Roberty

Marker type eSSR

Number 40

miRNA based molecular marker SSR

66

RAPD

12 (158)

RAPD

13

RAPD RAPD

6 18 (220)

RAPD

7

RAPD ISSR HvSSR AFLP

18 (82) 10 (54) 36 14 (442)

SNP SSR CAPS RAPD

14 11 1 20 (152)

ISSR

15

8

Reference Kumari et al. (2013) Yadav et al. (2014) Randazzo et al. (2019) Ruanjaichon (1997) Adams et al. (1998) Adams (2000) Dong et al. (2003) Moosikapala and Te-Chato (2010) Singh et al. (2014) Celestino et al. (2015) Sigmon et al. (2017) Dhawana et al. (2018) Raja et al. (2019)

AFLP amplified fragment length polymorphism, CAPS cleavage amplified polymorphic sites, ctEST-SSR cross transferable EST-SSR, CISP conserved intron scanning primers, EST-SSR expressed sequence tags-simple sequence repeats, GBS genotyping by sequencing, HvSSR hypervariable SSR, ILP intron length polymorphic, ISSR inter-simple sequence repeats, miRNA microRNA, RAPD random amplified polymorphic DNA, RFLP restriction fragment length polymorphism, SCAR sequence characterized amplified region, SNP single nucleotide polymorphism, SSR simple sequence repeats, SRAP sequence-related amplified polymorphism, STS sequence tagged sites

use it in the mapping of traits of interest. In range grasses, one of the important traits of interest is the mapping of the apospory gene/locus. To this end, several workers attempted mapping of apospory locus in different range grasses viz. Brachiaria brizantha (Pessino et al. 1998), Cenchrus ciliaris (Yadav et al. 2012; Kumar and Saxena 2016), Panicum maximum (Ebina et al. 2005; Bluma-Marques et al. 2014), and Paspalum notatum (Martínez et al. 2003; Rebozzio et al. 2012) using RFLP, AFLP, SCAR, and single-dose markers (SDAFs) markers. Table 1.6 shows

Range grass Cenchrus ciliaris L.

Chloris gayana Kunth

Panicum maximum Jacq.

Paspalum notatum Flüggé

S. no. 1

2

3

4

Synthetic completely sexual tetraploid plant (Q4188) as

Q408410 × Tift9t (diploid genotypes)

Sexual, colchicine induced, autotetraploid “Noh PL1” × apomict “Natsukaze” Apomictic genotype of M. maximus cv. Mombaça and a sexual genotype, S10

Obligate apomictic (IG-963108) × sexual parents (IG-96443) Female “katambora” (Kat) × Male “Tochirakukei” (Toch)

Cross-type Sexual genotype (90C48507) × apomictic genotype (PI 409164)

Full-sib progeny of 136 F1 hybrids 126 F1 mapping population 113 Pseudo

71 Pseudo test-cross population

86 F2 mapping population F1 Pseudo test-cross population

Mapping population 87 F1 hybrids

Q4188 map consisted of

112

858 SNPs

Markers mapped Maternal map 322 RFLPs Paternal map 245 RFLPs Male parent42 AFLPs, 4 SCAR “Kat” genetic map-72 AFLP, 12 RFLP “Toch” genetic map-52 AFLP, 9 RFLP Parental map-360 simplex markers

Table 1.5 List of linkage map constructed in range grasses using DNA-based molecular markers

1703.5

39

10

1590.6

991

756.69

443.3

12

8

488.3

2757

42

14

Distance (cM) 3464

No of linkage groups 47

6.8

9

~1.13

4.7

9.0

7.8

11.3

Average marker density (cM) 10.8

(continued)

Ortiz et al. (2001)

Deo et al. (2019)

Ebina et al. (2005)

Yadav et al. (2012) Ubi et al. (2004)

Reference Jessup et al. (2003)

1 Genetic and Genomic Resources of Range Grasses: Status and Future Prospects 21

S. no.

Range grass

Table 1.5 (continued)

female parent × a natural aposporous individual (Q4117) as pollen donor

Cross-type test-cross F1 family

Mapping population

Q4117 map-183 SDAFs

233 single-dose markers (SDAFs)

Markers mapped

No of linkage groups 26 co-segregation groups 39 co-segregation groups 2265.7

Distance (cM)

12.4

Average marker density (cM)

Reference Stein et al. (2007)

22 V. C. Tyagi et al.

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Genetic and Genomic Resources of Range Grasses: Status and Future Prospects

23

Table 1.6 Molecular markers closely associated with desirable forage breeding traits for use in marker-assisted selection S. no. 1

2

Range grass Brachiaria brizantha (A. Rich.) Stapf Cenchrus ciliaris L.

Trait mapped Apospory

QTL/Gene Apo

Apomictic mode of reproduction

–

Apomixis

Apo

Sexual mode of reproduction Apomictic reproduction

Sexual locus

Apomictic reproductive behavior Aposporous apomixis

–

–

Apospory

Apomixis

3

Panicum maximum Jacq.

Apospory

Apo

Apo

qGM1and 2, qTDM3 and 4, qLDM5 and 6, qSDM7, qPLB8, and qRC9 and 10

Reference Pessino et al. (1998)

Apo-C270, Apo-C470, Apo-C730, Apo-C930 20G, 18G, and 19G 12FS, 4HS, and 12b

Kumar and Saxena (2016) Yadav et al. (2012)

UGT-197, P16R, Q8M, U12H, V4, and X18R UGT 197

Roche et al. (1999) Lubbers et al. (1994) Gustine et al. (1997) Dwivedi et al. (2007) Ebina et al. (2005)

J16–800 and M02–680 OPF08–600

Apo

Agronomic trait (GM, TDM, LDM, SDM, PLB, and RC)

Associated markers PAM52–5 and PAM49–13

Flanking: G7–428, A1–166 Co-segregating: A3–355, F2–258, E6–258, D5–139, E2–300, G2–131, G4–49, G4–88, G4–228 PM_16, PM_A01

S_14_29023868 and S_10_48091934 –

BlumaMarques et al. (2014) Deo et al. (2019)

(continued)

24

V. C. Tyagi et al.

Table 1.6 (continued) S. no.

Range grass

4

Paspalum notatum Flüggé

Trait mapped Nutritional trait (OM, CP, NDF, ADF, IVD, PL, CEL, and SIL) Apospory

QTL/Gene qOM, qCP, qNDF, qADF, qIVD, qPL, qCEL, and qSIL Apospory

Apospory

Associated markers –

BCU243–377, BCU259–1157, E36/M37–103, E36/M38–67, E32/M33–110, E33/ M32–340, C1069–4900, C1069–6230 SPNA1 and SPNA2

Reference

Martínez et al. (2003)

Rebozzio et al. (2012)

Apo aposporous apomixis locus

agronomically important traits mapped in various range grasses. Even with the number of QTLs that have already been mapped, very few markers have progressed to the MAS level in range grasses. However, recently Deo et al. (2019) mapped the apospory locus in Panicum maximum using the GBS approach and fine-mapped the apo locus at 1.13 cM distance with S_14_29023868 and S_10_48091934 flanking markers, which can be used for marker-assisted breeding in various range grasses including guinea grass.

1.7

Nutritive Value

The forage nutritive value is important in livestock nutrition as livestock production depends on the amount of nutrients in the forage (Schut et al. 2010). Total digestible nutrients (TDN), crude protein (CP), and metabolism energy (ME) are often used as a determinant of forage quality (Pinkerton 2005; White and Wight 1984). France et al. (2000) noted that the nutritional value of forage depends on the amount of proteins and digestible carbohydrates. The ash, lignin, cellulose, crude fiber, phosphorus, carotene, and some other plant chemical compounds are other determinants of forage quality. The dry matter digestible is also one of the main indexes for determining forage quality. The amount of nutrients accumulated in forage is influenced by several factors viz. species, age, environment, plant fraction, fertilization, etc. The availability of nutrients to animals depends on forage intake and the extent of digestibility. Evaluation of tropical range grasses revealed that their crude protein content varied in a narrow range between 7.23 and 8.85% adequate to meet the maintenance requirement of sheep and goats. The TDN contents of grasses

1

Genetic and Genomic Resources of Range Grasses: Status and Future Prospects

25

ranged between 36.12 (Vetiveria zizanioides) and 51.21% (Pennisetum TSH). The DE and ME contents of evaluated grasses ranged from 1.59 to 2.26 and 1.31 to 1.85 kcal/g DM, respectively. The DDM contents were as low as 47.70 and as high as 56.74% among the grasses. The RFV was lowest for Vetiveria zizanioides (54.50) and highest for Pennisetum TSH (79.77%).

1.8

Hay and Silage of Grasses

Grasses can be conserved as hay, silage, and haylage when available in excess to provide the quality fodder in lean periods where availability of forage is scanty and pasture growth is limited to sustain livestock production. Forage conservation should focus on minimizing losses, which start immediately after cutting. For good quality hay, grasses should be harvested between flower initiation to mid bloom stage where yield and nutrients are in optimal stage. The quality of both hay and silage is affected by factors like species, variety within species, maturity at harvest, weed/grass contamination, harvest conditions/procedures, proper moisture, storage method, afternoon vs. morning harvest, etc. Cultivated grasses have sugar contents required for ensiling, while ensiling of tropical range grasses is difficult because of their low sugar contents. Perhaps there is no information on the ensiling of range grasses in the Indian context. At IGFRI under a multidisciplinary project from three years of study, many lines have been identified from more than 100 germplasm each of Cenchrus and Sehima range grasses which contain adequate sugar (>7.0%) required to initiate fermentation. Good quality silage of cultivated grasses-legumes mixtures (2:1 ratio) can be prepared. The silage prepared from Cenchrus genotypes viz. IG96–96, IG96–50, and IG96-401was of good quality as evidenced by their pH (4.07–4.97) and lactic acid values. Identified Cenchrus accessions may be further propagated and multiplied in grazing lands and common areas to produce silage from post-monsoon growth of pasture and grazing lands for subsequent use in lean periods to provide quality fodder to sustain livestock production.

1.9

Matching Nutritive Value with the Animal Requirement

The nutritive value of a forage/feed includes nutrient composition (protein, carbohydrates, minerals, and vitamins), availability (digestibility) of nutrients and energy, and efficiency of nutrient and energy utilization. The digestibility combined with intake is a reasonable determinant of forage/feed quality and is well accepted as an indicator of potential animal production. Animal performance depends on feed availability, nutrient content, intake, the extent of digestion, and metabolism of the digested feed. However, nutrient availability and intake mostly govern animal performance. Energy and protein digestibility is the most common limiting factor and sometimes minerals and vitamins are the nutrients that limit animal performance. The amounts of digestible energy, protein, vitamins, and minerals needed for maintenance are low relative to other animal processes. In general, forages that

26

V. C. Tyagi et al.

contain less than 70% NDF and more than 8% crude protein will contain enough digestible protein and energy, vitamins, and minerals to maintain older animals. Thus, even many low-quality forages and crop residues can meet the maintenance needs of some classes of animals, if protein and minerals are adequate. Grasses mean CP contents (8.0%) are marginally adequate to meet the protein requirement of sheep and goats for maintenance, while mean ME contents (1.60 Mcal/kg DM) are adequate to meet the energy maintenance need of adult sheep and goats and can also meet the energy requirement of growing lambs and kids for moderate growth rate of 50–75 g/day.

1.10

Biomass Yield and Productivity

Globally grasses and legumes have a wide genetic base with 620 and 650 genera and 10,000 and 18,000 species of grasses and legumes, respectively. Out of these only 40 grasses and legumes are used to varying extents in sown pasture establishments. It has been experienced that livestock prefers indigenous species of grasses and legumes that selected varieties of these species in spite of the lower nutritive value and yield of indigenous species. Grasses and legumes are characters to adapt to the different types of climatic and soil conditions making them suitable as forage plants for grazing/mowing and soil water conservation. Characters of grasses’ root systems make them suitable forage species for water conservation and preventing the leaching of nutrients. Legumes, however, through their nitrogen fixation ability improve soil fertility. Biomass yield and productivity of grasses and legumes varied widely depending on the soil type and rainfall including species, accessions, and growing conditions, particularly when N fertilizers are applied.

1.11

Future Research

The role of tropical grass pastures in future environments of variable climates with rainfall variability and changed rainfall patterns, higher temperatures, and higher levels of atmospheric carbon dioxide are likely to be features of future environments. These changes in rainfall patterns, combined with the predicted temperature increases, would favor the establishment and growth of tropical species (Boschma et al. 2010). The potential for tropical pastures is still largely untapped. Demand for red meat is predicted to increase, with greater quantities required for both the domestic and export markets. To achieve this, livestock numbers will need to increase, as will the quantity and quality of pastures to feed them. In the summerdominant rainfall areas, tropical pastures have an important role in contributing to an improved supply of forage throughout the year. Over the past few years, there has been some discussion on the potential of tropical grass pastures, but their persistence and productivity in adverse environments need to be determined. If the summergrowing perennial legumes that are showing promise in current studies continue to persist, then further research will be required to develop the best methods for their

1

Genetic and Genomic Resources of Range Grasses: Status and Future Prospects

27

establishment and management in future environments. Studies are needed to determine techniques for maximizing vegetative growth in-season, as well as making use of excess herbage out-of-season by using supplements and fodder conservation of hay and silage.

1.12

Conclusion

The livestock sector in India is confronted with different problems such as low productivity, high cost of commercial feed, low green fodder production, insufficient availability of dry fodder, and low level of technology. Indian grassland is dominated by a large number of nutritionally rich range grasses which could be fed to the animals in order to meet out fodder demand of the country. There are 60 species of perennial grasses, which make up the fragile ecosystem that supports our cattle. The quantity of fodder that can be produced in India is constrained because the majority of the country’s arable territory is used for food and cash crops. The major fodder crops being cultivated in arable land under non-leguminous fodder are maize, oats, sorghum, bajra, hybrid Napier, Guinea grass, para grass, etc. and under leguminous fodder are berseem, cowpea, lucerne, velvet bean, and others. Increasing the area producing green fodder is difficult because of severe competition from food crops. Moreover, adaptation of forage species particularly perennial species and trees (agroforestry) has the ability to produce more forage yield under changing climate and reduces the ill-effects of climate change through carbon sequestration (Kaul et al. 2010; Dhyani et al. 2010; Rai and Palsaniya 2015; Palsaniya et al. 2011). Considering the limitations of traditionally cultivated fodder crops, it is necessary to introduce various non-traditional fodder crops for growing on marginally productive farms and denuded community lands. There are many hardy grasses that can be grown on wastelands without irrigation. Such species can be established on field bunds, home gardens, and along farm boundaries. Distribution of identified sexual lines between breeders is involved in tropical grass breeding. Diversification of sexual lines in the good agronomic background. Most tropical forage grasses are polyploids in nature, the development of diploid lines will help in understanding the genetics of forage-contributing traits and speed up the breeding program.

References Abdi S, Dwivedi A, Kumar S, Bhat V (2019) Development of EST-SSR markers in Cenchrus ciliaris and their applicability in studying the genetic diversity and cross-species transferability. J Genet 98(4):101 Adams RP (2000) DNA analysis of the genetic diversity of vetiver and future selections for use in erosion control. In: Second International Conference on Vetiver-ICV2, Thailand Adams RP, Daffornli MR (1999) DNA fingerprints of the pantropical grass. AUJT 2(4):173–180

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Adams RP, Turuspekov MZY, Dafforn M, Veldkamp J (1998) DNA fingerprinting reveals clonal nature of Vetiveria zizanioides (L.) Nash Gramineae and sources of potential new germplasm. Mol Ecol 7(7):813–818 Ambiel AC, Guaberto LM, Vanderlei TM, Machado Neto NB (2008) RAPD grouping of accesses and cultivars of three Brachiaria species. Acta Sci Agron 30(4):457–464 ASARECA (2007) Rationalisation and Harmonisation of Seed Policies and Regulations in Eastern and Central Africa. ASARECA Technology Uptake and Up-scaling Support Initiative (TUUSI) Bashaw EC (1962) Apomixis and sexuality in buffel grass. Crop Sci 2:412–415 Bishoyi AK, Sharma A, Kavane A, Geetha KA (2016) Varietal discrimination and genetic variability analysis of Cymbopogon using RAPD and ISSR markers analysis. Appl Biochem Biotechnol 179(4):659–670 Bluma-Marques AC, Chiar L, Agnes DC, Jank L, Pagliarini MS (2014) Molecular markers linked to apomixis in Panicum maximum Jacq. Afr J Biotechnol 13(22):2198 Boschma SP, Lodge GM, Harden S (2010) Seedling competition of lucerne in mixtures with temperate and tropical pasture species. Crop Pasture Sci 61:411–419 Burson BL, Actkinson JM, Hussey MA, Jessup RW (2012) Ploidy determination of buffel grass accessions in the USDA National Plant Germplasm System collection by flow cytometry. S Afr J Bot 79:91–95 Burton GW (1989) Registration of ‘Tifton 9’ Pensacola bahiagrass. Crop Sci 29:1326 Burton GW, Gates RN, Hill GM (1993) Registration of ‘Tifton 85’ Bermuda grass. Crop Sci 33:644 Busey P (1989) Progress and benefits to humanity from breeding warm-season grasses for turf. In: Sleper DA, Asay KH, Pedersen JF (eds) Contributions from breeding forage and turf grasses, CSSA Special Publication no. 15. Crop Science Society of America, Wisconsin, pp 49–70 Carino DA, Daehler CC (1999) Genetic variation in an apomictic grass, Heteropogon contortus, in the Hawaiian Islands. Mol Ecol 8(12):2127–2132 Carlier L, Rotar I, Vlahova M, Vidican R (2009) Importance and functions of grasslands. Notulae Botanicae Horti Agrobotanici Cluj-Napoca 37:25–30 Celestino RDS, Zucchi MI, Pinheiro JB, Campos JBD, Pereira AA, Bianchini FG, Blank AF (2015) Molecular and chemical characterization of vetiver, Chrysopogon zizanioides (L.) Roberty, germplasm. Genet Mol Res 14:3 Chandra A, Dubey A (2007) Transferability of STS markers for studying the genetic diversity within the genus Cenchrus (Poaceae). Curr Sci 92:961–967 Chandra A, Dubey A (2008) Evaluation of genus Cenchrus based on malondialdehyde, proline content, specific leaf area and carbon isotope discrimination for drought tolerance and divergence of species. Acta Physiol Plant 30:53–61 Chandra A, Tiwari KK (2010) Isolation and characterization of microsatellite markers from guinea grass (Panicum maximum) for genetic diversity estimate and cross-species amplification. Plant Breed 129(1):120–124 Chandra A, Saxena R, Roy AK, Pathak PS (2004) Estimation of genetic variation in Dichanthium annulatum genotypes by RAPD technique. Trop Grassl 38:245–252 Chandra A, Saxena R, Roy AK (2006) Polymorphism and genotype-specific markers for Dichanthium identified by random amplified polymorphic DNA. Genet Resour Crop Evol 53(7):1521–1529 Chandra A, Jain R, Solomon S, Shrivastava S, Roy AK (2013) Exploiting EST databases for the development and characterisation of 3425 gene-tagged CISP markers in biofuel crop sugarcane and their transferability in cereals and orphan tropical grasses. BMC Res Notes 6(1):47 Cidade FW, Dall Agnol M, Bered F, de Souza-Chies TT (2008) Genetic diversity of the complex Paspalum notatum Flügge (Paniceae: Panicoideae). Genet Resour Crop Evol 55(2):235–246 Cidade FW, Vigna BB, De Souza FH, Valls JFM, Dall Agnol M, Zucchi MI, Souza AP (2013) Genetic variation in polyploid forage grass: assessing the molecular genetic variability in the Paspalum genus. BMC Genet 14(1):50 Combes D, Pernes J (1970) Variations in chromosome numbers in Panicum maximum Jacq. in relation to method of reproduction. Acad Sci Paris C R Ser D 270:782–785

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Forage Genetic Resources and Scope for Allele Mining of Abiotic Stress Tolerance Brijesh K. Mehta, Surendra Kumar Meena, Nilamani Dikshit, P. Shashikumara, Anup Kumar, Praveen Kumar, Mahendra Singh, Gaurendra Gupta, and Shahid Ahmed

2.1

Introduction

Forage genetic resources play an important role in mitigating the climate change effects. The livestock productivity of India is one of the lowest in the world mainly due to various problems faced by livestock and the forage sector. In India, fodder requirement is mainly fulfilled by three sources: crop residues, fodder crops and pasture or grazing lands. One of the main problems faced in meeting fodder requirement is the uneven distribution of fodder sources throughout the country. Currently, India faces a net deficiency of 35.6% green fodder, 10.95% dry crop residues and 44% concentrate feed ingredients (IGFRI Vision 2050). The main reasons behind this are climate change that has a significant impact on forage production and livestock management by intensifying temperature fluctuation, salinity and rainfall excess or deficit (Perring et al. 2010). Milk and other livestock products play a significant role in providing nutrition to humans. The demand of livestock-based products is increasing and estimated to double by 2050 (Rojas-Downing et al. 2017). However, climate change, mainly triggered by greenhouse gases such as CO2, N2O and O3, adversely impacts the quantity and quality of feed and fodder, thereby threatening livestock sustainability and subsequently human health (Henry et al. 2012). Therefore, the adverse effect of climate change needs to be managed for continuous supply of milk and meat, while maintaining animal health. Forages are a large component of agro-ecosystems worldwide, which provides the major chunk of diet to the ever-increasing livestock population. A wide diversity of plants is cultivated as forages, from succulent legumes to cool- and warm-season perennial grasses and annual crops. Range legumes, grasses and wild relatives of cultivated forages are adapted to harsh climate B. K. Mehta (✉) · S. K. Meena · N. Dikshit · P. Shashikumara · A. Kumar · P. Kumar · M. Singh · G. Gupta · S. Ahmed ICAR-Indian Grassland and Fodder Research Institute, Jhansi, India # The Author(s), under exclusive license to Springer Nature Singapore Pte Ltd. 2023 R. K. Singhal et al. (eds.), Molecular Interventions for Developing Climate-Smart Crops: A Forage Perspective, https://doi.org/10.1007/978-981-99-1858-4_2

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conditions. These forage genetic resources possess a large diversity of functional traits for climate change adaptation (Indu et al. 2022). Screening of forage genetic resources for allelic variation in identified genes of known sequence for abiotic stress tolerance traits by the ‘tilling strategy’ would help in mitigating climate change effects and overcoming forage deficit (Kumar et al. 2010). In this chapter, we discuss different forage genetic resources for abiotic stress tolerance, success of allele mining in model crops and some forages, challenges of allele mining in forages and further scope of allele mining for abiotic stress tolerance traits in forages.

2.2

Impact of Climate Change on Forage Production

Climate change results in erratic and irregular rainfall, frequent drought, terminal heat stress and salinity of soil, which impact the quantity and quality of forage and feed resources, resulting in threats to livestock sustainability. Several workers have reported that climate change affects the production and productivity of forage (Perring et al. 2010), quality of forage (Thivierge et al. 2016), livestock health, water availability for both forage and livestock (Lacetera 2019) and forage species diversity and composition (Yang et al. 2017). Climate change also affects the digestive process of livestock by changing the physic-biochemical properties of forages such as less protein and more fibres (Barbehenn et al. 2004). The biomass and quality of forages are reduced significantly with the rise in temperature and CO2 concentration (Chapman et al. 2012). Decline in crude protein and digestible organic matter has been observed due to the rise in CO2 levels (Melo et al. 2022). Thornton and Gerber (2010) observed vigorous growth of legumes in grasslands with elevated CO2 concentration. The rise in CO2 level has more positive impact on C4 forages than C3 forages as it slows down evapotranspiration and thereby increases the primary production of pastures (Howden et al. 2008). Drought is the major cause of reduction in grassland productivity by triggering withering of leaves, tillers and rhizomes (Lei et al. 2020). Drought stress decreases forage yield and crude protein, increases water soluble carbohydrates and crude fibres, and decreases nitrogen assimilation rate (Liu et al. 2018). The high temperature shortens the life cycle of perennial grasses, leading to about 25% reduction in production, and increases crude fibre content, including lignin (Dumont et al. 2015). Forage yield reduction has been reported in maize and soybean at high temperature (Schlenker and Roberts 2009). High temperature increases lignification of plant tissues that results in slow degradation of plant tissues, and therefore, causes poor digestibility (Dumont et al. 2015). Further, high temperature inhibits photosynthesis by altering chloroplast structure and enzymes inactivation (Cui et al. 2006). Erratic rainfall increases soil water stress, which causes yield penalty and quality reduction in forages (Knapp et al. 2008). Change in rainfall pattern also declines grass biodiversity and creates soil salinity. Fay et al. (2003) reported that a 50% increase in dry spell duration reduces about 10% of the primary productivity. The higher ozone concentration is reported to be harmful for crops like clover and alfalfa

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(Booker et al. 2009). Further, feed quality is reported to be more prone to ozone levels than dry matter content.

2.3

Strategies for Mitigating Climate Change Effects

Diversification of forage crops reduces the chances of outbreak of various pathogens and insect pests (Ghahramani et al. 2019). The effect of heat stress and rainfall variability can be minimized by adjusting cropping pattern and crop cultural practices like sowing, irrigation, cutting and grazing (Fawzy et al. 2020). Adoption of various conservation practices enhances the soil organic content. The emission of greenhouse gases from the livestock sector can be minimized by managing diet composition, grazing management and inclusion of perennial forages in forage production systems (Kaul et al. 2010). Methane emission from livestock can be reduced by improving rumen fermentation efficiency of animals. Efficient nutrients management strategies such as high nutrient use efficiency, use of organic source of fertilizers, use of slow-release fertilizers and inclusion of legumes help in improving forages productivity (Dixit et al. 2014). Apart from management strategies, plant breeding has the potential to improve the characteristics of forages to withstand adverse environmental conditions and to increase their yield potential. The breeding strategies for developing climate-resilient forages include collection and introduction of tolerant forage species, intra- and inter-specific hybridization, polyploidization, mutagenesis, marker-assisted breeding and genome editing (Razzaq et al. 2021). The biggest bottleneck in recombination breeding in tropical grasses is apomictic reproduction behaviour. Thus, identification and generation of sexual lines in apomictic grasses may prove to be new era of research direction. Understanding physiological, genetic and molecular mechanisms that allows range grasses and legumes to adapt a wide range of climate is essential for designing efficient breeding programmes (Taranto et al. 2018). Mapping of climate-tolerant QTLs/genes and their introgression supports the breeding of varieties better adapted to harsh climate (Mehta et al. 2020).

2.4

Forage Genetic Resources

Forage genetic resources provide a valuable source of livestock feed and play a very important role in food security and poverty alleviation. Forages also provide significant ecosystem services by preventing soil erosion, purifying air and water, mitigating greenhouse gas impacts and providing wildlife habitat. Forages may be harvested as hay, grazed pasture, range, green chop or silage. Economic and human food value of forages accrues primarily through dairy, beef and other livestock products, producing staple foods throughout the world. Forage genetic resources include primitive forms of cultivated plant species and landraces, modern cultivars, breeding lines, genetic stocks, weedy types and related wild species. India is rich in genetic diversity of various forage grasses (263 genera and 1506 species) and

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Table 2.1 Forage genetic resources conserved in various institutes in the world Institutes CIAT

Accessions 22,694 and 546 live plant species 18,639 and 1531 live plant species >70,000 >22,299

ILRI APG IITA ICARDA ICRISAT ICAR-NBPGR, New Delhi

>40,000 >62,000 8107

No. of genera and species 127 genera, 700 species >1000 species >2000 species Cowpea: 15,115, maize: 1561, legumes: 6623 Forages: 25,556, grass pea: 4193 Pearl millet: 23,057, sorghum: 39,264 > 145 genera

Table 2.2 Status of germplasm holding at ICAR- IGFRI, Jhansi, India Crop groups Cereal fodder Cultivated legumes Range legumes

Range grasses

Total

Crops Sorghum, maize, pearl millet, oat, barley, finger millet

Accessions 3437

Berseem, cowpea, guar, lablab, lucerne

3233

Stylosanthes, Desmanthes virgatus, Clitoria ternatea, rice bean, grass pea, horsegram, joint vetch, Centrosema, Siratro, Neotonia, Rhynchosia, Subabul, sword bean, jack bean, wild ground nut, zornia, red clover, white clover, sainfoin, shaftal Tropical: Buffel grass, Marvel grass, Sword grass, Rhodes grass, Guinea grass, Setaria, Panicum, Themeda, Urochloa spp., Napier grass, Iseilema, Vetivera spp. Temperate: Tall fescue, Orchard grass, Prairie grass

746

3480

10,896

legumes (1550 species, of which 60 species used as forage) (Kellogg et al. 2020). Diverse forage grasses and legumes such as Panicum, Cenchrus, Pennisetum, Dichanthium, Sehima, Chrysopogon, Bothriochloa, and Lasiurus and legumes such as cow pea, lablab bean, Clitoria and rice bean show a wide diversity in different zones of the country. These wealth of genetic diversity holds vast potential and are among the world’s most essential natural resources; hence, there is a need to collect, evaluate and conserve this diversity for present and future use. The direct connection between genetic resources and humanity’s food supply relies on improving the management and utility of genetic resources important to agriculture. Many institutes viz. CIAT, ILRI, Australian Gene Bank (APG), IITA, ICARDA, ICRISAT and ICAR-NBPGR are engaged in various activities related to enrichment and management of forage genetic resources (Table 2.1). ICAR-IGFRI, Jhansi, conserved about 10,896 accessions of forages and also provides active collection site for forage crops in India (Table 2.2).

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2.5

39

Abiotic Stress-Tolerant Forage Genetic Resources

Forage crops are the major source of animal diet, and nearly 800 million people depend on livestock for their livelihood. Grasses play a significant role in carbon sequestration, water conservation and sustaining wildlife diversity (Rao et al. 2015). Grasses and range legumes possess some inherent traits that enable them to propagate under harsh conditions. The forage grasses, range legumes and wild relatives of cultivated forage crops survived in their natural habitat for millions of years without human interference. During evolution, these species exposed to natural selection pressure against various abiotic stresses and accumulated genes that enable them to resist, tolerate or avoid drought, salinity, flood, extreme temperature and even trampling by animals (Jensen et al. 2012). Perennial grasses and legumes dominate the rangelands and exhibit superiority over cultivated forage crops due to their vegetative propagation, fast regeneration, stress tolerance and higher resource use efficiency for light, water and nutrients (Hall et al. 2020). Besides, perennial forage crops showed deep root system which helps in the extraction of water from deeper layers of soil (Lasisi et al. 2018). Medicago sativa is a model forage crop with salt tolerance ability, while Medicago arborea possesses drought tolerance (Lei et al. 2018). Signal grass is a model grass with tolerance to low soil fertility, drought, flood and other adverse conditions (Rao et al. 2011). Wild legumes possess high phosphorous use efficiency to persist in rangelands and under low fertile soil. Thus, grasses and wild relatives of forage crops offer primary and more often secondary and tertiary gene pool of useful traits to mitigate the adverse effect of climate change in cultivated forage crops (Brozynska et al. 2016). Some climate-resilient species of forage legumes and grasses are provided in Table 2.3. The direct exploitation of wild forages and grasses is very difficult through traditional breeding; however, advanced breeding tools can be used to incorporate abiotic stress tolerance and other useful traits into cultivated forage crops through molecular or genome editing approaches (Chand et al. 2022).

Table 2.3 The list of abiotic stress-tolerant forage grasses, legumes and wild relatives of cultivated forage crops Abiotic stress Drought

Water logging Salinity Heat

Tolerant forage legume/grass Vigna subterranean, Medicago laciniata, Medicago arborea, Medicago truncatula, Medicago ciliaris, Medicago polymorpha, Medicago ruthenica, Austrodanthonia racemose, Austrodanthonia caespitose, Dactylis glomerata, Rytidosperma species, Pennisetum clandestinum, Dactylis glomerata, Lolium perenne Trifolium subterraneum, Trifolium michelianum, Melilotus siculus Melilotus siculus, Hedysarum carnosum, Vigna marina, Vigna luteola, Vigna vexillate, Medicago intertexta, Medicago truncatula Cenchrus ciliaris, Festuca species, Poa pratensis

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Adaptation Strategies of Forages for Abiotic Stress Tolerance

The wild relatives of crops and range grasses and legumes are adapted to harsh climate conditions. The adaption of these species is due to various mechanisms that operate during stress conditions. The range perennial grasses are vegetative propagated, and exhibited high regrowth potential, tolerance to abiotic stresses and high biomass conversion efficiency (Hall et al. 2020). Further, perennial forages can extract water from deeper layers of soil due to their deep root system (Lasisi et al. 2018). The C4 forage crops and grasses showed lower photo-respiration, methane and N2O emission, thus contributing less to global warming compared to C3 crops (Ning et al. 2020). The summer dormancy and root plasticity traits of some forages help them to mitigate climate change effect on their growth and biomass production (Münzbergová et al. 2017). Salt tolerance in Medicago species involves mechanisms like high Na+/ K+ transport, Ca signalling, phytohormone biosynthesis and antioxidant accumulation (Lei et al. 2018). Grasses have evolved many adaptive strategies to survive under stress conditions, which include osmotic adjustment, stomatal regulation, phenotypic plasticity, metabolite production and stress-related proteins (Ma et al. 2017). Signal grass showed flood tolerance due to the formation of aerenchyma for oxygen movement from root to shoot. Some species of signal grass also showed drought tolerance due to their improved leaf area, root length, stomatal movement and water use efficiency (Rao et al. 2011). Drought avoidance mechanism in grasses includes leaf folding, leaf rolling, leaf shedding, accumulation of cuticular wax and summer dormancy (Bolger et al. 2005). In some droughttolerant grasses fructans concentration increased under drought. The wild legume species possessed high phosphorous use efficiency due to their extended root length, diameter, density and hairs (Becquer et al. 2021). The potential forage legume and grasses showing tolerance to abiotic stress along with adaptation strategies/ mechanisms are presented in Table 2.4. These species can serve the valuable genetic material for mining of stress-tolerant alleles and subsequent introgression or modification in cultivated forage crops.

2.7

Allele Mining for Abiotic Stress Tolerance in Forage Crops and Grasses

Allele mining is the process of identifying the allelic variants of a candidate gene in other genotypes or identifying novel alleles for a given trait in the germplasm collections and natural populations (Ashkani et al. 2015). The genome sequencing projects have generated a huge amount of sequence information, which is available in various genomic database. The available genome and gene(s) sequence information facilitate rapid discovery of useful alleles conferring resistance/tolerance to various biotic and abiotic stresses (Chaudhary et al. 2019). Allele mining in available germplasm collections offers a novel genotype to deliver new/superior allele(s) for use in crop improvement programmes. Allele mining has been shown to have wide

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Table 2.4 The potential forage genetic resources and their adaptation strategies for abiotic stress tolerance

Salinity

Forage legume/ grass species Trifolium subterraneum Trifolium fragiferum Medicago arborea Medicago ruthenica Vigna marina

Salinity

Melilotus siculus

Waterlogging

Melilotus siculus

Salinity

Hedysarum carnosum Vigna luteola

Trait Waterlogging Salinity and alkalinity Drought Drought

Salinity Salinity

Salinity

Drought Drought Drought Drought Drought High phosphorususe efficiency Improved soil health Drought tolerance

Medicago truncatula Medicago lupulina Medicago intertexta

Medicago polymorpha Medicago ciliaris Medicago truncatula Medicago arborea Medicago laciniata Trifolium uniflorum N2-fixing forage legumes Pennisetum clandestinum

Adaptation strategies Aerenchyma formation Osmolytes and Na+ accumulation in leaves High RWC and gas exchange; less lipid peroxidation; higher antioxidant capacity Efficient photosynthesis; less H2O2 and more proline accumulation Na+ exclusion; increased stomatal conductance, transpiration rate and photosynthetic rate Greater K+/Na+ ratio in shoots; hard seed coat Aerenchyma and aerenchymatous phellem (secondary aerenchyma) in roots Increased Na+ accumulation in roots Na+ inclusion; increased photosynthetic rate High germination; osmotic and ionic homeostasis in roots and shoots

High germination speed; osmotic and ionic homeostasis in roots and shoots

Greater root development and stem length High RWC; greater root development and water use efficiency High RWC; antioxidative protection; decline in lipid peroxidation High RWC and gas exchange; less lipid peroxidation; higher antioxidant capacity Stomatal regulation; high RWC; proline and K+ accumulation Extended root length, root length density, root hair length and average root diameter Deeper and larger root system; greater SNF capacity; root exudates increase soil aggregation and reduce soil erosion Long roots

References Aschi-Smiti et al. (2003) Jēkabsone et al. (2022) Tani et al. (2019) Wang et al. (2021) Yoshida et al. (2020) Striker et al. (2015) Teakle et al. (2011) Kouas et al. (2010) Yoshida et al. (2020) El-Shafey and Al-Sherif (2020) El-Shafey and Al-Sherif (2020) Badri et al. (2016) Badri et al. (2016) Luo et al. (2016) Tani et al. (2019) Yousfi et al. (2010) Becquer et al. (2021) Abiala et al. (2018) Nie et al. (2008)

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applications such as discovery of superior alleles and new haplotypes, gene diversity analysis, gene prediction and expression analysis, functional marker development and evolutionary studies (Kumar et al. 2010). A generalized allele mining procedure involves (1) identification of germplasm accession with target trait, (2) selection of gene(s) underlying target trait, (3) primer designing for target gene(s), (4) primer amplification of target gene(s) in identified germplasm accessions, and (5) finding allelic variants of target gene(s).

2.7.1

Genesis of Allelic Variation: Natural or Induced

For any crop improvement programme, existence of genetic variation for target trait is the basic requirement. Variations can arise naturally or can be induced using mutagens (Hill et al. 2021). Mutation is the prime source of allelic diversity in any crop species. Mutations in genic region creates new allele by either single nucleotide polymorphism (SNP) or InDel variation. Mutations in coding sequence of a gene may lead to distinct phenotype, while mutations in non-coding sequence do not affect the phenotype. The phenotypic variations may be due to either altered regulatory function or altered protein structure and/or function (Albert and Kruglyak 2015). The reverse genetic approach involving alteration in target gene sequence and analysing its phenotypic effect is a powerful strategy for elucidating gene function and creating new alleles (Irshad et al. 2020). Many studies suggest that the wild relatives of crop species and grasses possessed many superior traits/allele that allow them to survive in extreme stress conditions (Indu et al. 2022). These superior and beneficial alleles were lost in improved varieties of food and forage crops during crop domestication and evolution due to inbreeding and selection (Kyriazis et al. 2021). These superior alleles tend to be retained in wild species, land races and grasses due to their high level of genetic diversity. The transfer of these superior alleles to elite varieties of forage crops may result in increased abiotic stress tolerance. The prime role of these superior alleles in plant adaptation suggests that such alleles could be preserved in land races and related wild and cultivated crop species (Kumar et al. 2010). Hence, exploring the natural superior alleles holds enormous potential to improve forage crops and sexually reproducing grasses. Further, allele mining is expected to boost the pace of identification and characterization of new allelic variants.

2.7.2

Detection of Allelic Variation: Allele Mining Techniques

Eco-TILLING (Targeting Induced Local Lesions in Genomes) and sequencingbased allele mining approaches are generally followed for identification of allelic variations (SNP polymorphism) for the target candidate gene in the natural populations.

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2.7.2.1 Eco-TILLING The TILLING technique was developed by Claire McCallum and their colleague in the late twentieth century (Borevitz et al. 2003). In this technique, induced mutation is used to generate sequence variation (mostly point mutation) in the target gene and these sequence variations are identified by heteroduplex analysis. Eco-TILLING is a variant of TILLING approach in which sequence variations in the target gene(s) are identified in the natural populations or crop gene pools, without the use of mutagens (Comai et al. 2004). In Eco-TILLING, single nucleotide mismatch is introduced in the heteroduplexed DNA using DNA nuclease enzyme (cel-1, S1 nuclease, mung bean nuclease, etc.) and sequence variations between test and reference genotypes are identified through Li-Cor gel assay (Li-Cor, USA). Eco-TILLING is generally used to discover SNPs and small InDels; however haplotyping at target gene and variation in SSRs (simple sequence repeats) can also be detected (Till et al. 2006). Eco-TILLING is an effective approach for discovery of haplotype analysis and SNP discovery. It is applicable to any organism, whether heterozygous or polyploid, and can be applied for searching of superior alleles for abiotic tolerance in forage crops and grasses. Natural alleles with limited genetic diversity in wild relatives can also be detected using Eco-TILLING. However, this technique requires a more sophisticated laboratory, pooling of DNAs of test and reference genotypes, specific conditions for enzymatic cleavage, Li-Cor genotyper and high-throughput sequencer (Kumar et al. 2010). Raghavan et al. (2007) used agarose gel for rapid detection of novel allelic variants of known candidate gene. Wang et al. (2008) developed a modified Eco-TILLING approach, called self-EcoTILLING, for the discovery of SNP polymorphism in a multi-gene family. Eco-TILLING is considered to be cost-effective as it requires sequencing of one individual for each haplotype. The cost of Eco-TILLING can be further reduced in future through the development of low-cost innovative strategies. 2.7.2.2 Sequencing-Based Allele Mining Under current scenario of climate change, advanced breeding techniques in forage improvement programmes will be crucial for increasing forage productivity to feed the livestock. Recent advanced technologies like TILLING through sequencing, sequencing-based allele mining and mapping along with trait-based breeding provide opportunities to develop climate-resilient crops within less time (Kumar et al. 2010). Eco-TILLING has been proposed to be economical approaches for haplotyping and SNP discovery. Under sequencing-based allele mining, alleles of a gene in diversified germplasm are amplified through polymerase chain reaction in a thermocycler. Furthermore, nucleotide variation in the alleles is identified by DNA sequencing. Different useful alleles of genes in a wide range of species can be identified and isolated by this novel genomic tool. This technique can also be used to analyse individuals for haplotype structure and study of diversity to determine genetic association in forage crops (Min et al. 2021). Unlike Eco-TILLING, this approach does not require much sophisticated equipment or tedious steps. For constructing haplotypes, point mutations or SNPs and InDels can be effectively analysed in sequencing-based allele mining. Furthermore, screened genotypes for

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tolerance to abiotic stresses are used for the extraction of genomic DNA by using the CTAB method. In addition, sequencing-based allele mining of specific genes in identified accessions and their association with phenotypic variations provides a tremendous impetus to precision breeding programmes in crop plants (Thudi et al. 2014). The advances in DNA sequencing technologies has resulted in low cost, more read length and high throughput NGS platforms, such as single molecule sequencing (SMS) technology, single-molecule real-time (SMRT) technology, nanopore sequencing, 10X genomics etc (Ashkani et al. 2015). Due to the availability of these NGS techniques, sequencing-based allele mining would result in faster generation of allelic data at a cheaper cost. Further, technological developments also continue to increase sample throughput, which will facilitate large-scale genotyping of genetic resources. Therefore, this approach will deliver a platform for the largescale development of SNPs and haplotypes at loci of interest and it will soon be a cost-effective measure for allele mining of rare alleles in populations in the near future.

2.7.3

Success of Allele Mining for Abiotic Stress Tolerance in Model Crops

A large number of allele mining studies have been performed in recent years for dissection of useful alleles in imparting abiotic stress tolerance in the model crops. Both Eco-TILLING and NGS can be implemented to expand and detect allelic series of functional genes. A number of genes and loci have been implicated in rice, imparting tolerance to salinity (Negrão et al. 2013; Raghuvanshi et al. 2021), drought (Singh et al. 2021), high night temperature (Dhatt et al. 2021) and submergence (Singh et al. 2020). These loci regulate diverse plant functions including ionic balance in shoots, root growth, grain width and osmotic adjustment. Likewise, allele mining has also been applied in other cereals, legumes, oilseeds and fibre crops for different abiotic stress tolerance traits (Table 2.5). Based on the sequence variation in useful alleles, allele-specific markers can be developed for introgression of these useful alleles into cultivated varieties using marker-assisted breeding.

2.7.4

Examples of Allele Mining in Forage Crops and Grasses

Allele mining has been shown to have wide applications such as discovery of superior alleles and new haplotypes, gene diversity analysis, gene prediction and expression analysis, functional marker development and evolutionary studies (Kumar et al. 2010). So far, using allele mining approaches, few genes have been mined in forage crops, particularly in barley, sorghum, maize, alfalfa and red clover (Table 2.6). NGS-based allelic variations has been identified in candidate genes for drought tolerance (Zhao et al. 2010), nitrogen remobilization and dehydration tolerance (Estermann et al. 2017) in maize, lignin content and composition in

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Table 2.5 Successful examples of allele mining in different crops for abiotic stress tolerance Mining approach NGS

S. no. 1.

Crop Oryza sativa

2.

EcoTILLING

3.

Oryza sativa and O. glaberrima Oryza sativa

NGS

Gene/locus Saltol, qGR6.2, qSE3, RNC4, OsHKT1;1, SKC1 and OsSTL1 OsHKTI;5, OsNHX1, SalT, OsRMC Fie1

4.

Oryza sativa

NGS

DRO1

5.

Oryza sativa

NGS

6.

NGS NGS

TaPYL4

EcoTILLING NGS

sHsp26

NGS

Dehydrin (DHN)

NGS

CabHLH10

12.

Triticum aestivum Triticum aestivum Triticum durum Triticum aestivum Cicer arietinum Cicer arietinum Glycine max

Sub1A, Sub1B and Sub1C TaHKT1;5

NGS

qWT_Gm03

13.

Glycine max

NGS

14.

Zea mays

NGS

GmCHX1 (Glyma03g32900 locus) ZmARF31

15.

Zea mays

NGS

ZmTIP1

16.

Brassica oleracea Gossypium herbaceum

NGS

BoCCA1

NGS

GhRD2, GhNAC4, GhHAT22 and GhDREB2

7. 8. 9. 10. 11.

17.

TaHAK25

Trait Salinity tolerance

Reference Raghuvanshi et al. (2021)

Salinity tolerance

Negrão et al. (2013)

High night temperature tolerance Drought tolerance Submergence tolerance Salinity tolerance Drought tolerance Heat tolerance Salinity tolerance Drought tolerance Drought tolerance Water logging tolerance Salinity tolerance

Dhatt et al. (2021)

Low phosphorus tolerance Drought tolerance Freezing tolerance Drought tolerance

Wu et al. (2016)

Singh et al. (2021) Singh et al. (2020) Thiyagarajan et al. (2022) Wu et al. (2022) Comastri et al. (2018) Ma et al. (2022) Kumar et al. (2020) Thakro et al. (2022) Ye et al. (2018) Patil et al. (2016)

Zhang et al. (2020) Song et al. (2018) Li et al. (2020)

maize (Xiong et al. 2020) and sorghum (Emendack et al. 2022), drought and frost tolerance in barley (Shrestha et al. 2022; Guerra et al. 2022). Hufnagel et al. (2018) identified six allelic variants of AltSB gene controlling aluminium tolerance in

Crop Sorghum bicolor

Urochloa species and Megathyrsus maximum

Medicago sativa and M. truncatula

Sorghum bicolor

Lolium perenne

Zea mays

Lolium perenne

Zea mays

Trifolium pratense

Hordeum vulgare

Hordeum vulgare

S. no. 1.

2.

3.

4.

5.

6.

7.

8.

9.

10.

11.

NGS

NGS

NGS

NGS

De novo assembly strategy

NGS

NGS

Eco-TILLING

NGS

Mining approach Amplification refractory mutation system (ARMS) markers RNAseq and bait capture genomic DNA sequencing

Table 2.6 Status of allele mining in forage crops and grasses

Frost tolerance HvCBF14 (Fr-H2 locus)

P5CS1

Nitrogen remobilization and dehydration tolerance Biological nitrogen fixation Drought tolerance

Self-incompatibility

Drought tolerance

Forage digestibility Forage yield Salt tolerance Drought tolerance Cyanogenic glucoside dhurrin Flowering time

Lipid content

Cell wall digestibility

Trait Aluminium toxicity

GRMZM2G179885 and GRMZM2G347043 EFD and MOT1

LpSDUF247

dhn2

LpFT3

4CL, CCoAOMT, COMT, CCR, CAD, GT43, BAHD01 and BAHD05 SDP1, SDP1-like, CGI58, PXA1, PXA1-like, TGD1, TGD2 and TGD3 CAD1 and CCoaOMT CONSTANS-like NHX1 WXP1 CYP79A1

Gene/locus AltSB

Blomstedt et al. (2012) Skøt et al. (2011) Zhao et al. (2010) Veeckman et al. (2019) Estermann et al. (2017) Trněný et al. (2019) Shrestha et al. (2022) Guerra et al. (2022)

Gréard et al. (2018)

Hanley et al. (2021)

Reference Hufnagel et al. (2018)

46 B. K. Mehta et al.

Zea mays

Sorghum bicolor

Hordeum vulgare

Hordeum vulgare

Hordeum vulgare

12.

13.

14.

15.

16.

NGS

Eco-TILLING coupled NGS

Eco-TILLING

NGS

NGS

GAPDH

P5CS1

mlo and mla

bmr12

bm1-E1 and bm1-E2

Salinity tolerance

Lignin content and composition Lignin content and composition Powdery mildew resistance Drought tolerance

Xiong et al. (2020) Emendack et al. (2022) Mejlhede et al. (2006) Xia et al. (2017) Thabet et al. (2021)

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sorghum using Amplification Refractory Mutation System (ARMS) markers. In alfalfa, Gréard et al. (2018) reported 8, 5, 8, 45 and 91 non-synonymous variants of CAD1 (digestibility), CCoaOMT (digestibility), NHX1 (salt tolerance), CONSTANS-like (forage yield) and WXP1 (drought tolerance) genes, respectively. The allelic variants for two candidate nitrogen fixation genes (EFD and MOT1) in red clover have been revealed by Trněný et al. (2019). Eco-TILLING approach has been applied to screen allelic variation at CYP79A1 locus for dhurrin production in sorghum (Blomstedt et al. 2012) and mlo and mla loci for powdery mildew resistance in barley (Mejlhede et al. 2006). In forage grasses, allele mining has been applied to a limited extent in Urochloa, guinea grass and ryegrass (Table 2.5). Hanley et al. (2021) studied allelic diversity of 11 digestibility and 9 lipid content genes in four species of Urochloa (syn Brachiaria) and guinea grass using RNAseq and bait capture genomic DNA sequencing. The study reported 953 non-synonymous polymorphisms across 20 genes and 104 accessions. In ryegrass, candidate gene association genetics approach has been successfully employed for mining of LpFT3 gene for flowering time (Skøt et al. 2011). Veeckman et al. (2019) using de novo strategy discovered 28 novel alleles of LpSDUF247 gene associated with S-locus of self-incompatibility in ryegrass.

2.7.5

Challenges

Genetic resources of forage crops and grasses held in various gene banks harbour numerous climate-resilient traits (Indu et al. 2022). The target genes and their allelic variants for such traits are still undisclosed in many forage crops and grasses. The identification and efficient exploitation of the useful allelic variants posed several unknown and specific challenges in forage crops and grasses. The first major challenge is the selection of germplasm to be used for allele mining. Screening of the entire germplasm of a crop provides more opportunity to detect rare alleles, but this task is considered to be inefficient and cumbersome. Development of core and mini core collection, precise phenotyping techniques and efficient computational tools can maximize the chances of mining the prospective allelic variants (Kumar et al. 2010). The candidate genes for abiotic stress tolerance have not been characterized in majority of grasses. Further, genomic resources (whole genome sequence data) are lacking in most grasses. Handling of genomic resources for exploiting allele mining is another challenge in forage crops and grasses. The demarcation of core promoter region from regulatory elements also poses challenge in mining of promoters (Veerla and Hoglund 2006). The characterization of cis-acting regulatory elements is very difficult, even in the fully sequenced genome, due to transcriptional differences arising from various trans-acting factors and environmental conditions (Dai et al. 2007). Besides the enormous efforts and time required for allele mining, high cost of sequencing platforms is also an important consideration in allele mining. Particularly in forage grasses, achieving an induced desired phenotypic variation is cumbersome due to limited knowledge of mutation (Chen et al. 2016). Long

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generation time, perennial growth habit and small seed size interfere with population development in forage grasses (Manzanares et al. 2016). Allele mining in forage grasses is also complicated by apomixis and self-incompatibility mechanisms, as it hinders self-pollination to develop homozygous recessive phenotype (Ortiz et al. 2013). For example, in Brachiaria grass, obligate outcrossing occurs in sexual lines and apomixis takes place in half of the progenies (Worthington and Miles 2014). In forage grasses, biomass yield is the primary trait, which is governed by multiple genes, generally with epistatic effects. For example, heading date in ryegrass is influenced by allelic variation in LpHD1 gene (homologue of CONSTANS-like gene in Arabidopsis), which also affects forage yield, quality and persistency (Skøt et al. 2007). The polyploidy and highly polymorphic nature of forage crops and grasses are the major challenge for the use of Eco-TILLING (Gréard et al. 2018). Considering the nature of grasses and other factors, allele mining is more challenging in grasses compared to food crops.

2.7.6

Future Scope

Allele mining possesses tremendous potential in effective utilization of genetic and genomic resources for genomics-assisted plant breeding. Efficient allele mining tools and techniques need to be developed to screen the large collections of forage crops and grasses along with analysing gene sequence variation and handling genome sequence data. The success of allele mining largely depends on crop species and diversity of germplasm of a crop. The grasses and wild relatives of forage crops are the great reservoir of valuable alleles for abiotic stress tolerance traits (Tanksley et al. 1996). These traits can be utilized through identification of target gene, analysing sequence variation in gene and subsequent marker-assisted breeding. The refined sampling techniques need to be developed for establishing core and mini core set in forages. Diversity at molecular level can be best exploited in shortlisting of germplasm accessions. High throughput phenotyping of forages helps in development of trait-specific subsets, which could assist in selection of key genotypes for mining of the novel alleles governing the target trait. The genome and gene sequence information of a given species are the basic needs of any efficient allele mining programme. The genomic sequence information of many grasses is not available. Further, the genes underlying the abiotic stress tolerance in most of the grasses has not been fully characterized. This provides opportunity for the genetics, molecular and bioinformatics scientists to work in collaborative manner for generating genome sequencing information and identifying candidate genes and useful allelic variants. The genome/ candidate gene sequence information of one forage species will be helpful in mining of underlying traitspecific genes in other forage species that falls within the limits of genome synteny and conserved sequences (Kumar et al. 2010). The power of this approach can be further enhanced by the use of new technologies that allows faster allele discovery. With the availability of low-cost and high-throughput sequencing platforms and efficient bioinformatics tools, sequencing-based allele mining would be a prime

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choice for scoring allelic variations in candidate genes and developing novel genetic resources for abiotic stress tolerance in forages, particularly in grasses. The outcome of genome sequencing projects in alfalfa, white clover, red clover, Brachypodium and Panicum will be helpful in fine mapping of stress-tolerant loci and successive allele mining and synteny analysis. Gréard et al. (2018) describe the procedure and prospects of allele mining for breeding of auto-tetraploid species using Medicago as model species. In a crop with diploid and polyploid collections, the draft genome sequence of one diploid species can be used for discovering allelic variants of candidate genes. This approach of combining RNAseq and bait capture genomic DNA sequencing was found successful in mining of forage digestibility traits in Brachiaria (Hanley et al. 2021). Same approach can be applied to other genes such as apomixis and spittlebug resistance in Brachiaria and other grasses and forage crops having diploid and polyploid accessions. Candidate gene association genetics approach used for mining of flowering time allelic variants in ryegrass can be used for mining of genes with dominant phenotypic effect (Skøt et al. 2011). The use of whole genome SNP array chips will speed up the identification and characterization of new allelic variants. Moreover, advanced allele mining schemes where the artificial allelic variants with more relevant function are designed by domain or gene shuffling among related genes will offer new opportunities for improving stress tolerance of forage crops and grasses.

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provides insights into the tolerant mechanisms of legume forage to environmental stress. BMC Biol 19:96. https://doi.org/10.1186/s12915-021-01033-0 Worthington ML, Miles JW (2014) Reciprocal full-sib recurrent selection and tools for accelerating genetic gain in apomictic Brachiaria. In: Budak H et al (eds) Molecular breeding of forage and turf. Springer, Cham, pp 19–30 Wu F, Liu Z, Xu J, Gao S, Lin H, Liu L, Liu Y, Lu Y (2016) Molecular evolution and association of natural variation in ZmARF31 with low phosphorus tolerance in maize. Front Plant Sci 7:1076. https://doi.org/10.3389/fpls.2016.01076 Wu BB, Shi MM, Pourkheirandish M, Qi ZH, Ying WA, Yang CK, Ling QI, Zhao JJ, Yan SX, Zheng XW, Zheng J (2022) Allele mining of wheat ABA receptor at TaPYL4 suggests neo-functionalization among the wheat homoeologs. J Integr Agric 21(8):2183–2196 Xia Y, Li R, Bai G, Siddique KH, Varshney RK, Baum M, Yan G, Guo P (2017) Genetic variations of HvP5CS1 and their association with drought tolerance related traits in barley (Hordeum vulgare L.). Sci Rep 7(1):7870. https://doi.org/10.1038/s41598-017-08393-0 Xiong W, Li Y, Wu Z, Ma L, Liu Y, Qin L, Liu J, Hu Z, Guo S, Sun J, Yang G (2020) Characterization of two new brown midrib1 mutations from an EMS-mutagenic maize population for lignocellulosic biomass utilization. Front Plant Sci 11:594798. https://doi.org/10.3389/ fpls.2020.594798 Yang H, Wu M, Liu W, Zhang ZH, Zhang N, Wan S (2017) Community structure and composition in response to climate change in a temperate steppe. Glob Chang Biol 17:452–465 Ye H, Song L, Chen H, Valliyodan B, Cheng P, Ali L, Vuong T, Wu C, Orlowski J, Buckley B, Chen P (2018) A major natural genetic variation associated with root system architecture and plasticity improves waterlogging tolerance and yield in soybean. Plant Cell Environ 41(9): 2169–2182 Yoshida J, Tomooka N, Yee Khaing T, Sunil Shantha PG, Naito H, Matsuda Y, Ehara H (2020) Unique responses of three highly salt-tolerant wild Vigna species against salt stress. Plant Prod Sci 23:114–128 Yousfi N, Slama I, Ghnaya T, Savouré A, Abdelly C (2010) Effects of water deficit stress on growth, water relation and osmolyte accumulation in Medicago truncatula and M. laciniata population. C R Biol 333:205–213 Zhang X, Mi Y, Mao H, Liu S, Chen L, Qin F (2020) Genetic variation in ZmTIP1 contributes to root hair elongation and drought tolerance in maize. Plant Biotechnol J 18(5):1271–1283 Zhao X, Liu C, Yu Y, Shi Y, Song Y, Wang T, Li Y (2010) Allele mining of a drought tolerance candidate gene dhn 2 and association analysis between dhn 2 and some phenotypic traits on maize. J Maize Sci 18(1):20–28

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Breeding for Developing Higher Productive Tree-Based Forage Under Stress Environments Hirdayesh Anuragi, Srijan Ambati, Rajesh Kumar Singhal, Sukumar Taria, Alka Bharati, Kunasekaran Rajarajan, Arun Kumar Handa, and Ayyanadar Arunachalam

3.1

Introduction

The burgeoning population necessitated the demand for resources and dependency on alternative sources of food habits. Livestock is one such alternate source of food mankind is dependent on. India is the leading milk producing country with more than 15% of world’s livestock population from 2.4% of global area. Quality fodder is an essential commodity for good cattle health and enhanced milk productivity and eventually decides the success of the dairy sector. Increase in agricultural production in recent decades has led to the enhanced production and supply of green fodder; however, the supply has always fallen short of the demand for the livestock. During 2020, a deficit of 12% dry fodder and 30% green fodder has been reported in India (ICAR-IGFRI 2021). The demand for dry and green fodder is expected to rise significantly to 631 and 1012 million metric tons, respectively, in the next two decades. Further, the changing climate and its consequences like extreme climates and various abiotic and biotic stress also put a huge challenge on fodder production and availability for livestock (Indu et al. 2022; Chand et al. 2022). Agroforestry is a climate smart technique of effective land-use management where woody perennials and annual crops are deliberately grown together with livestock. This system also includes tree species having greater fodder values for livestock growth and development. Since ancient times, farmers have been traditionally using fodder trees as important sources for livestock. In recent years, the cultivation of fodder shrubs has H. Anuragi (✉) · S. Taria · A. Bharati · K. Rajarajan · A. K. Handa · A. Arunachalam ICAR-Central Agroforestry Research Institute, Jhansi, India S. Ambati Agriculture College Warangal, Professor Jayashankar Telangana State Agricultural University, Warangal, Telangana, India R. K. Singhal Division of Crop Improvement, ICAR-Indian Grassland and Forage Research, Jhansi, India # The Author(s), under exclusive license to Springer Nature Singapore Pte Ltd. 2023 R. K. Singhal et al. (eds.), Molecular Interventions for Developing Climate-Smart Crops: A Forage Perspective, https://doi.org/10.1007/978-981-99-1858-4_3

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enhanced in many areas; despite that, the tree fodder still has a high potential for fodder security in India. Tree-based agroforestry meets additional 9–11% green fodder demand today. Additionally, the agroforestry as a whole provides other major advantages like enhancing the additional 28.724 million hectares of green cover/tree cover which is otherwise not possible through forestry, and thus directly contributing to achieve the minimum requirement of 33% of forest/tree cover and plays a very significant role in mitigating climate change and securing the food, fuel, and livelihood security simultaneously in the country (Dhyani and Handa 2013). Fodder trees have many advantages like they are easier to grow, require lesser labor, land, capital, and provide several by-products in comparison to fodder shrubs. However, tree fodder species face several constraints like nonavailability of quality seed or planting material and lack of cultivation knowledge and skills (Franzel et al. 2014). Further, India is the hub of livestock consisting of over 15% of world’s total livestock population; however, the supply is lagging much behind. Therefore, this huge fodder demand has to be met through encouraging the genetic improvement of genotypes of tree-fodder in addition to the ongoing breeding research programs on annual fodder crops. In most intensely populated countries like India, the pressure side on cultivable area for food grains, oilseeds, pulses etc. is increasing every day making no vacancy for the fodder crops. Hence, the agroforestry systems are supporting to fill this gap in the form of fodder trees. However over a period of time, these existing fodder trees cannot nourish to meet the growing livestock demands. Hence, there is a great need to genetically improve these fodder tree germplasm lines. The development of diverse genetic resources to develop futuristic trait-specific lines is the need of the hour. However, there hasn’t been as much research done on the genetic improvement of fodder trees as there has been on forage grasses and legumes. This chapter attempts to highlight the current status, major limitations, and futuristic tactics of fodder trees genetic improvement in India and abroad (Dhyani et al. 2009).

3.2

Major Tree Fodder Species and Breeding Interventions

Fodders trees are the cheapest source of feed for livestock and ruminants in addition to several other uses. Not all trees can be used as fodder as they need to have palatable leaves and pods with high proteins and other nutrient contents and must be devoid of any toxic or antinutritional factors. There are more than 500 species of fodder trees available in different regions based on their suitability to the respective climatic conditions, though only above 150 species are economically important and only 44 species are more often preferred by livestock. Since ages, these trees have been naturally fed by livestock in the farms or forests to meet their nutritional requirements. However, the quantity and quality of these trees are directly influenced by species, age, season, geographic location, and the management practice (Kshatri 2007). Interestingly, no appropriate and area-based cultivation and conservation practices have ever been followed by the farmers. Some of the major trees having high fodder values are listed in Table 3.1 and depicted in Fig. 3.1.

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Table 3.1 Major tree fodder species preferred as livestock feed S. no. 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12 13. 14. 15.

Species Leucaena leucocephala Acacia Senegal Acacia nilotica Prosopis cineraria Ailanthus excelsa Moringa oleifera Gliricidia sepium Ziziphus mauritiana Bauhinia variegate Albizia lebbeck Morus alba Dalbergia sissoo, Gmelina arborea, Calliandra calothyrsus Sesbania sesban

Vernacular name Subabul Gum Arabic Babul Khejri Ardu Drumstick Gliricidia Ber Kachnar Siris Mulbery Shisham Gamhar Calliandra, spiked powder puff Dhaincha

Preferred regions Semiarid Tropics and subtropics Tropics and subtropics Arid and semiarid Arid and semiarid Tropics and subhumid Subtropics and semiarid Tropics and subtropics Tropics and subtropics Tropics and moist tropics Tropics and moist tropics Tropics and moist tropics Tropics and subtropics Tropics and subtropics Tropics and subtropics

Fig. 3.1 Major potential tree fodders, (a) Leucaena leucocephala, (b) Acacia senegal, (c) Acacia nilotica, (d) Prosopis cineraria, (e) Ailanthus excelsa, (f) Moringa oleifera, (g) Gliricidia sepium, (h) Ziziphus mauritiana, (i) Bauhinia variegate, (j) Albizia lebbeck, (k) Morus alba, (l) Dalbergia sissoo, (m) Gmelina arborea, (n) Calliandra calothyrsus, and (o) Sesbania sesban

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Status of Genetic Improvement Research in Major Tree Fodder Species

Breeding interventions in tree species for improving fodder quality and tolerance to biotic and abiotic stresses is at a nascent stage. In recent years, advanced biotechnological and genomics tools and techniques have opened a new niche for quicker improvement of fodder trees which was otherwise not possible through traditional breeding methods. The progress of breeding research interventions in major tree fodder species is briefly described below.

3.3.1

Leucaena leucocephala

Leucaena genus is an excellent tree leguminous fodder source containing more than 14 species, viz. L. leucocephala, L. collinsii, L. diversifolia, L. lanceolata, L. macrophylla, L. esculenta, L. cuspidata, L. greggii, L. multicapitula, L. retusa, L. pallida, L. pulverulenta, L. salvadorensis, L. shannoni, and L. trichodes. Among these, few have diploid level with 52 or 56 chromosomes and polyploidy level with 104 chromosomes and mostly exhibit natural outcrossing mechanisms of pollination. Among these, L. leucocephala, vernacularly referred to as subabul in India, is most popular and is highly preferred by goats, sheep, and other livestock for their feed. In recent years, efforts on genetic improvement of Leucaena have been initiated for wood energy and fodder however, majorly limited to the divergence studies. The interspecific hybridization among these Leucaena species might be capable of producing hybrids of improved vigor and tolerance to various biotic and abiotic stresses. The collection of useful Leucaena germplasm, their evaluation, and multiplication is highly required for sustainable conservation and production of quality fodder. Recent understanding of its taxonomy (Abair et al. 2019) and advance in molecular markers and genomic resources have enabled effective genetic improvement programs. However, a long juvenile phase, highly heterozygous nature, and high G × E interaction still remain key challenges in developing new cultivars. L. leucocephala exhibits a high level of tolerance to biotic and abiotic stresses. Negi et al. (2011) identified a few Leucaena-specific genes for stress tolerance which might be the useful source of stress tolerance in breeding programs. Another study by Honda et al. (2018) has identified 73 (root) and 39 (shoot) drought responsive genes sequences in Leucaena leucocephala subsp. glabrata. With regard to the fodder quality, the genes were identified in Leucaena under different stress environments by Ageel et al. (2022). Through microarray analysis, a total of 138 gene sequences were found upregulated in the foliage than in roots out of which 13 genes were confirmed by quantitative real-time PCR. These genes coded several proteins involved in the starch, lipids metabolisms, cell wall formation, and biosynthesis of secondary metabolites in L. leucocephala (Honda et al. 2020).

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Acacia senegal

Acacia senegal (L.) Willd. is a Fabaceaeous multipurpose tree species widely popular in African and Asian countries like India and Pakistan. This species provides fuel wood, pale to orange-brown solid Gum Arabic and fodder during dry periods and has the ability to restore the soil fertility through nitrogen fixation and thus became a promising component of agroforestry systems particularly in drylands. Its fodder is highly preferred by goats, camel, sheep, and other cattle. Leaves and pods contain highly digestible proteins and minerals and thus are the most favored plant parts for cattle fed through lopping, cat and carry and range browsing techniques, particularly in drought-prone areas. Besides this, timber, fuel wood, shade, honey, gum, and natural dye are also produced by this species. It also serves as a very good biological fence for protecting the farm area against stray cattle. It is a slow growing and very drought-tolerant species containing a diploid chromosome number of twenty six (2n = 26). The breeding in this species is at a very nascent stage and currently focuses around the conservation, evaluation of provenances, enhancing the seed viability, and high yielding of foliage and Gum Arabic (Cossalter 1991; Shiran et al. 2020).

3.3.3

Acacia nilotica

Acacia nilotica (L.) Willd. is an important multipurpose agroforestry tree species in arid and semiarid regions. It belongs to the family Fabaceae and is popularly known as Babul in India. This tree species is basically medium in size, evergreen in nature, and contains thorns. Being a drought-tolerant leguminous species, it also fixes atmospheric nitrogen and therefore signifies its importance in agroforestry systems in nitrogen-deficit locations for fuel wood and fodder production as well as other environmental benefits. It is the source of timber, fuel wood, shade, nutritious fodder, honey, gum, and natural dye and also serves as a biological fence for protection against stray cattle (Bargali and Bargali 2009; Amadou et al. 2020). The leaves and pods are a good source of fodder for goats and sheep due to their high protein and mineral content particularly in the hot and drought-prone regions like Rajasthan state and Bundelkhand region of Madhya Pradesh and Uttar Pradesh. The genetic improvement research in this species is at a very initial stage and currently focuses on the conservation and evaluation of different provenances for quality planting material development. Genetic variability studies in Acacia nilotica provenances in Central India has been initiated in recent years (Devi et al. 2017; Singhdoha et al. 2017). Recent development of biotechnological tools like molecular markers and genomic resources has enabled to carry out faster and accurate traitspecific genetic improvement programs in slow growing tree species like A. nilotica. However, only a few basic studies have been reported on this aspect. A study by Yadav et al. (2016) has validated the transferability of 30 microsatellite markers in five different Acacia species including A. nilotica for genetic diversity analysis and futuristic breeding programs.

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Prosopis cineraria

Prosopis cineraria is a medium sized, evergreen, multipurpose tree species and thrives well in the dry, hot arid regions due to its xerophytic and highly droughttolerant nature and thus has been referred to as the “kalpavriksha of the desert” or “king of the desert.” It also tolerates the frost and saline condition. This Fabaceous plant is an important source of feed and fodder for camels, goats, sheep, and other livestock. The leaves are gray-green and pods are yellow to reddish brown in color and are produced abundantly under drought conditions. The leaves contained crude protein (12.1%) and crude fiber (20.1%) whereas pods contained crude protein (13.5%) and crude fiber (14.3%). The species is propagated through seeds only. P. cineraria increases soil fertility through fixing atmospheric nitrogen. The decomposition rate of this species is much higher than other arid tree species; therefore it increases the soil organic matter and soluble calcium and phosphorus in the surrounding region. It also provides good quality timber and fuel wood in the desert regions. Honeybees require abundant and prolonged flowering and this species supports good quality honey production in the dry arid regions (Afifi and Al-rub 2018).

3.3.5

Ailanthus excelsa

Ailanthus excelsa Roxb., vernacularly called Mahaneem, is a promising multipurpose tree species that acts as a source of dietary nutrients, alleviates fodder scarcity, and thereby significantly boosts livestock production. It is a deciduous species of Simaroubaceae family and has been referred to as “tree of heaven.” It is originated in China but widely distributed in various types of soils and climate in different tropical countries. It acts as an ideal tree species for fodder in dry areas (Kaushik et al. 2017). Mature leaves are preferred by sheep and other livestock as immature young leaves produce offensive smell. It contains 19.87% of crude proteins and 12.72% crude fibers. Also, leaves contain tannins and some antinutritional factors and thus proper precautions are required. In addition, it acts as a good firewood, soft timber wood, gum and resins and contains several pharmaceutical compounds for improved health. It also provides shelter and avenue during hot summer in the arid regions and can be utilized for boundary plantation as shelterbelts along the field borders. The genetic improvement in A. excelsa currently focuses on the conservation, provenances evaluation, and genetic diversity analysis for identification of superior provenance. Breeding efforts need to be initiated for developing improved clones or varieties with lower antinutritional factors for enhanced suitability to the cattle. Development of sufficient genetic and genomic resources is also a need of hour for speeding up of the breeding program in this species (Deswal et al. 2022). As its wood is soft and predominantly utilized in the production of match splints, Dasgupta et al. (2021) have recently studied the lignin biosynthetic mechanism through transcriptome analysis to understand the generate some useful genomic resources for futuristic wood trait-specific genetic improvement in A. excela.

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Moringa oleifera

M. oleifera Lam. is a multipurpose perennial species belonging to the family Moringaceae and is widely cultivated for food, medicinal, industrial, and fodder purpose. It is popularly known as drumstick, sahjan, horse reddish and has been referred to as wonder tree owing to its extraordinarily high nutritional and pharmaceutical importance (Nouman et al. 2014). All parts of this tree species is consumed or utilized for a variety of purposes. Though it is a well-known species for health utilities since ancient times, it received very good attention worldwide in recent years for its potential to deal with nutrition and health issues (Kumar et al. 2017). It is basically a fast growing, drought- and salinity-tolerant species widely distributed in Asian countries. It can cure up to 300 human diseases and supply high amount of proteins, fibers, minerals, and vitamins. Also it provides fuelwood, food, good quality seed oil, and high-quality fodder for enhancing the nutrition and the health of farm animals. Besides, quality honey can also be produced throughout the year. It can be easily propagated through seeds and cuttings. Moringa is nowadays becoming a very popular fodder species which can produce round the year high-quality green fodder in the first year itself in the dry and arid regions. Both leaves and pods are preferred by livestock and ruminant. First harvest is ready within the first 3 months of sowing and can produce more than 100–120 tonnes green fodder per hectare area in just 4 to 5 cuttings. It adds to the weight, improves the overall health of the animals, and enhances the milking capacity and quality of dairy animals. In the last few decades, the breeding for round the year leaves and fodder production has generated several improved varieties of Moringa like PKM-1, PKM-2, ODC, ODC-3, Bhagya, etc. which not only enhanced the area under cultivation, but also occupied significant place in International Moringa trade (Pandey et al. 2012). These cultivars have a very high potential of supplying huge fodder resource to meet the ongoing demand of fodder in the country and abroad. The government policies and research priorities have generated a very good amount of genetic and genomic resources in this species which are being utilized effectively in trait-specific genetic improvement of Moringa.

3.3.7

Gliricidia sepium

One abiotic factor that affects agriculture in more than 100 nations is soil salinity. Although gliricidia [Gliricidia sepium (Jacq.) Kunth] is a versatile tree with a capacity for adapting to a variety of soils, its tolerance thresholds and reactions to salt stress are still poorly understood. Carvalho da Silva et al. (2022) assessed the morph-physiological responses of young gliricidia plants under salinity stress and generated leaf metabolic and transcription profiles for elucidating the molecular mechanisms and candidate genes governing salt stress tolerance. Using the pairedend method and the Illumina HiSeq platform, RNAs from leaf samples were subjected to RNA sequencing. Leaf samples were extracted for polar and lipidic fractions, which were then examined using an electrospray ionization quadrupole

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time-of-flight high-resolution mass spectrometry (MS) equipment using ultra-high performance liquid chromatography (UHPLC). The platforms OmicsBox, XCMS Online, MetaboAnalyst, and Omics Fusion were used to analyze the acquired data. The substrate salinization technique allowed for the discovery of tolerance and adaptation to salt stress. A group of 5672 transcripts and 107 metabolites that were differently expressed in gliricidia leaves under salt stress were discovered through a single analysis of transcriptome and metabolome data sets. The most impacted pathway was the phenylpropanoid biosynthesis, which had 15 metabolites and three genes that were differently expressed. The outcomes demonstrated that this pathway’s differentially expressed metabolites and genes primarily influence shortterm salt stress (STS). Twelve genes encoding proteins that may be involved in the response of gliricidia to both short-term salt stress (STS) and long-term salt stress (LTS) were discovered by a single investigation of the transcriptome. To understand the mechanisms underlying the adaptive response, more research is required (de Oliveira et al. 2022).

3.3.8

Ziziphus mauritiana

Indian jujube botanically Ziziphus mauritiana is an important multipurpose species of the Rhamnaceae family. It is well suited to the dry tropical and subtropical regions which are hot and drought prone owing to its high stress tolerance potential. The tree is deciduous or almost evergreen bearing dark glossy green leaves and ovoid drupe fruits. The tree is basically grown for fruits though it has multiple benefits. It is a good source of carotene, vitamins A and C, and fatty oils. Indian jujube wood is reddish, fine-textured, hard, and durable. It can be used in rural house construction, posts, and tool manufacturing. It makes excellent firewood. A potential agroforestry species, this thorny tree can be grown to provide windbreaks and living fences. It is browsed by livestock and its leaves are nutritious fodder for sheep and goats (Maruza et al. 2017). Indian jujube can be propagated from seeds, or vegetatively, through in situ grafting or budding on to rootstocks. Trees must be pruned so that new fruits are produced on the next year shoots. In India, the best cultivars yielded 50–80 kg fruit/ year. Leaves are used as nutritious fodder for sheep and goats. Analysis of the chemical constituents on a dry weight basis indicates the leaves contain 15.4% crude protein, 15.8% crude fiber, 6.7% total minerals, and 16.8% starch. In India, the leaves are also gathered as food for silkworms (Gupta et al. 2012). The breeding in this species still focuses on the collection and conservation of the wild types, varietal evaluation, genetic diversity analysis, marker-assisted molecular breeding, etc. Several high-yielding varieties with improved quality have been developed by several institutions in India and abroad. However, the genetic and genomic resources are yet to be developed for effective understanding of the molecular mechanisms and identification of candidate genes controlling the key traits in this species.

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Bauhinia variegata

Bauhinia variegata locally known as Kanchan in Hindi is a member of the Fabaceae family. It is a well-known ornamental tree of tropical and subtropical regions experiencing the frequent dry and hot climate. It is moderately tolerant to drought. Various plant parts of the species like stem, stem bark, leaves, seeds, roots, flower buds, and flowers are used in various indigenous systems of medicine and popular among various groups in India for cure of a variety of ailments. It is also a source of fuel, fiber, timber, gum, tannin, lipids, etc. (Gautam 2012). Leaves make good fodder and are greedily eaten by sheep, goats, and cattle. The crude protein is 14.18% while crude fiber content is around 23.79%. Kanchan is the better source of nutrients, whereas it should be utilized as a feed with or without combination for the livestock (Naeem and Ugur 2019). The average annual fodder yield per tree is 15–20 kg of dry matter. The genetic improvement research has initiated provenance trials, genetic divergence studies, and molecular diversity analysis using RAPD and SSR markers. However, advancements toward molecular and genomics work are yet to be initiated for speeding up genetic improvement programs in the future.

3.3.10 Albizia lebbeck Albizia lebbeck(L.) Benth., also referred to as Koko or lebbeck tree, is an important leguminous fodder tree belonging to the Fabaceae family. Lebbek is native to tropical Africa, Asia, and Northern Australia. In subhumid, semiarid tropics and subtropical regions with a distinct dry season and a dependable rainy season, it has become widely naturalized. It is a multipurpose tree species that plays significant roles in improving the quality of the soil and pastures and controlling erosion. It is also grown as a shade provider and shelter belts (Lowry et al. 1994). Lebbek is a fastgrowing tree that can reach a height of 5 m per year under ideal circumstances, while dry conditions significantly slow growth. Since Lebbek does not readily regenerate, it should not be frequently browsed. As lopping improves coppicing and produces roughly 2500 kg/ha/year of edible material in low-rainfall locations where Leucaena only produces 1500 kg/ha/year, it should be lopped or grazed by cattle twice a year. Since triennial pollarding produces 1700 kg/ha/year, it is also worthwhile to practice. The yields of leaf dry matter could reach 5 t/ha per year. Though it is a promising fodder tree species, no much breeding work has been reported in this species. Recently, genetic diversity and markers-aided molecular characterization have been performed in this species. The genetic and genomic resources have to be developed for identifying fodder-related candidate genes and elucidating the related molecular mechanisms for effective genetic improvement in this species.

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3.3.11 Morus alba White mulberry (Morus alba L.) is a high-yielding, medium-sized tree species of tropical and subtropical regions. The leaves of this economically significant tree (Bombyx mori) are used to feed the monophagous mulberry silkworm in China, India, Thailand, Brazil, Uzbekistan, and other nations of the world. It has historically been utilized as both highly palatable food for the majority of farm animals and fodder for growing silkworms. Mulberry can be grown opportunistically near residential areas, on vacant lots, and along the edges of fields to be used as fodder. Depending on the variety, age of the leaves, and growing conditions, the crude protein concentration in leaves can be compared to that of the majority of legume forages; fiber fractions are lower. Mulberry leaves have a notable mineral concentration, with ash values up to 25%. P and Ca concentrations are typically 0.14–0.24% and 1.8–2.4%, respectively, whereas K values in leaves and young stems are 1.90–2.87% and 1.33–1.53%, respectively. Mg concentrations are typically 0.47–0.63% (in leaves) and 0.26–0.35% (in young stems). Mulberry leaf has a very high degree of digestibility: 78–81% in vivo (in goats) and 90% in vitro. The great palatability of mulberries makes them an ideal fodder crop. The fresh leaves and young stems are devoured first by small ruminants, even if they have never encountered them previously. If the branches are presented uncut, they may then rip off and consume the bark (Kandylis et al. 2009). If the biomass is finely diced, the cattle eat the entire amount. When presented concurrently, animals initially prefer mulberry to the other forages and would even sift through a collection of other forages in search of mulberry. Mulberry trees can be either dioecious or monoecious, and they occasionally transition between the two sexes. Wind pollinates flowers, and certain cultivars will produce fruit even in the absence of pollination; this cross pollination is essentially not required. The main abiotic factors affecting its potential yield and quality of leaves are drought, cold temperatures, excessive salt, and alkalinity, which are all widespread conditions. Although traditional breeding techniques made a significant contribution to the creation of mulberry types that were resistant to abiotic stress, there is still much room for the application of cutting-edge molecular and genomic tools to speed up mulberry genetic improvement programs.

3.3.12 Dalbergia sissoo Dalbergia sissoo Roxb. is an economically important, multipurpose tree species capable of growing under wide agro-climatic conditions. It is commonly known as Indian rosewood, shisham etc. and belongs to the family Fabaceae. It is a very good source of firewood, timber wood, and fodder. Also, its presence checks the soil erosion and can be used as windbreak. It also produces seed oil and tannins which have high industrial value. Its leaves, tender shoots, and pods are an excellent source of fodder for livestock and grazing animals during dry and off season where fresh grasses are not available. Leaves are highly nutritive, palatable, and rich in crude

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protein and essential minerals. It is tolerant to drought and can be easily grown in almost all the agro-climatic zones of India. It suits well under agroforestry either as block plantations or boundary plantations in cultivable and wastelands. Its largescale cultivation by the farmers will fetch them very good economic returns and will ensure fodder security and wasteland restoration (Bhattacharya et al. 2014). Dalbergia sissoo breeding research has been focused so far on the evaluation of plus trees for improved timber yield and quality. Development of tolerance to biotic and abiotic stresses and fodder quality is at a very initial stage. Sharma and Bakshi (2011) reported various growth and heritability estimates among D. sissoo clones from a clonal seed orchard. Recent molecular and genomic research advances have triggered in-depth research for identifying molecular mechanisms and candidate genes for specific traits. Molecular divergence studies using RAPD and SSR markers were conducted with 30 shisham genotypes obtained from different regions of India (Tewari et al. 2022). Singh et al. (2020) described the pattern of variation in pod features, germination behavior, and growth traits among seeds of D. sissoo gathered from various sources. Recently, phylogenic research employing molecular markers has also been carried out in this species (Bal and Panda 2018).

3.3.13 Gmelina arborea Gmelina arborea Roxb., a member of the family Lamiaceae, is a multipurpose tree species with numerous uses world over. It is a fast-growing species that demands good site conditions. Its wood is a source of pulp, particle board, plywood, matches, carpentry, and packing industries. Also, boards, carvings, and musical instruments are prepared using its wood. In addition to this, the leaves and fruits are the source of fodder and also used for rearing silkworms. The chromosome number in this species varies from 2n = 36, 2n = 38 to 2n = 40. The provenances of this species were outperforming than the plants originating from forests or natural origins. Leaf morphological variations are high in this species and leaf length, leaf breadth, leaf area, and petiole length showed significant positive correlation toward each other. Significant levels of morphological and phytochemical variation were found among five accessions that were gathered from various geographical and agroclimatic zones of South India, according to Raghu et al. (2011). Additionally, the leaves of 12 provenances have had their levels of crude protein, soluble sugar, phenol, crude fiber, phosphorus, potassium, calcium, and magnesium analyzed (Dey and Todaria 2015). Irrespective of provenances, the harvested leaves contained an average of 14.40% crude protein, 16.88% soluble sugar, 0.076% phenol, 13.25% crude fiber, 0.206% phosphorus, 1.44% potassium, 1.014% calcium, and 0.561% magnesium. Variability estimates for proximate and mineral nutrient contents in the leaf of different provenances were also computed. The differences between phenotypic and genotypic coefficient of variability indicated that these parameters are sensitive to environmental changes. Using Random Amplified Polymorphic DNA (RAPD) markers, the genetic diversity and relationships among the fourteen accessions of Gmelina arborea

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were evaluated. After comparing the RAPD profiles of all the accessions, a total of 262 scorable bands with fifteen primers were generated, and 243 of these bands were polymorphic. The primers OPJ-19, OPM-02, OPN-02, OPN-05, OPN-16, OPN-18, and OPC-10 revealed the lowest level of the polymorphic band and the maximum number of amplified products, respectively. The range of the similarity coefficient values was 0.58 to 0.74. The molecular relationships between Gmelina accessions that are geographically related were also discovered. The Vazhachal and Rosemala accessions have significant inter-population variation that can be employed as a parent in the Gmelina breeding program to increase productivity and wood quality (Mayavel et al. 2020).

3.3.14 Calliandra calothyrsus The multipurpose Calliandra calothyrsus tree, which is good for energy wood, has a lot of potential as a short-rotation crop. This nitrogen-fixing species can be utilized not only for energy wood by planting once but also is able to secure annual harvest up to 15–20 years from its coppices. Besides that, this species supports earth vegetation cover for mitigating climate change. In optimizing this role, its genetic improvement was undertaken during 2011–2014 to obtain the best energy wood individuals in volume and quality. Hendrati and Nurrohmah’s (2019) study compared five families of genetically modified trees—selected in 2014 from progeny tests in Wonogiri, Central Java—against five families of unimproved trees to determine the growth gain. At the age of 4-month-old seedlings prepared for field planting, significant variances on key growth characteristics have been found. Calliandra’s genetic improvement has been proven to increase the number of leaves, height, diameter, and seedling quality up to 23.4%, 24.3%, 6.7%, and 20% consecutively. Due to the low success rate of its reproduction, study on its reproductive biology was undertaken. The study by Baskorowati et al. (2021) was carried out during the flowering seasons, by examining flower morphology, pollen, and pistil viabilities as well as insect visitors to this species’ flowers. Inflorescences (spikes) bearing an average of 354 individual flowers, which develop acropetally, are how this species’ blooms are borne, according to the study. By the second or third day after they first appear, pistils (the female organ) stop developing, and at that point, flowering has also stopped. Between 9:00 and 10:00 WIB, pistils matured, and pollen is still viable one to three days following pollen opening. Apidae, Vespidae, and Formicidae, three of the flowers’ most frequent visitors, as well as one order of lepidoptera, were discovered. However, it is discovered that C. calotyrsus has a very poor reproductive success rate of only, i.e., 0.024. These preliminary findings supported the theory that this species’ poor reproductive performance is caused by the structure and growth of its blooms, which enable instances of both outcrossing and self-pollination. The short pistil maturation period may potentially be a factor in this species’ poor reproductive performance. In this species, no molecular work has been started.

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3.3.15 Sesbania sesban Sesbania sesban (L.) Merr., also known as “Dhaincha” in India, is a perennial plant that belongs to the Fabaceae family and grows rapidly. The excellent nitrogenous source of feed for goats and other livestock is its leaves, pods, and seeds. Since it is a N2 fixing species as well as more drought- and salt-tolerant in nature, therefore it can be produced in fairly severe climatic and soil conditions to meet the feed requirements besides enriching the soil nitrogen. S. sesban, being a potential tree species for agroforestry systems, its domestication was carried out in eastern and southern African countries (Owino et al. 1994). Sajjad et al. (2009) conducted an experiment to understand its pollination biology and ecology. Three types of pollination treatments, namely wind pollination, self-pollination, and insect crosspollination, each had various yield-attributing components that were measured. Abd Allah et al. (2015) reported the improved growth and systemic acquired resistance (SAR) using arbuscular mycorrhizal fungi under salinity stress in S. sesban. S. sesban has a strong chromium alleviation capacity, tolerance index, and transportation index (Patra et al. 2020), indicating its appropriateness for bioaccumulation programs, especially in mining areas. For forage yield improvement, Zulfiqar et al. (2015) evaluated the eight S. sesban accessions, viz. Shahpur, Sialkot, Sahianwala, Khanewal 1, Khanewal 2, Chiniot Bhowana 1, and Bhowana 2 for drought tolerance using morpho-physiological parameters and identified Shahpur, Khanewal 2, Sahianwala, and Bhowana 2 as superior for livestock in dry and hot climate. At its gene bank in Addis Abeba, Ethiopia, the International Livestock Research Institute (ILRI) keeps a collection of 18,662 feed germplasm accessions of grasses, herbaceous legumes, and browse plants. Determination of genetic diversity and underpinning the development of trait-based subsets of accessions is highly required for the identification of genotypes that can be used as parents to develop superior germplasm. Marker-trait association and genomic prediction enhance the prediction accuracy of superior genotypes and the efficiency of selection of new varieties. Recently developed advanced genomic tools would be highly useful in the trait-specific genetic improvement of this species.

3.4

Major Breeding Challenges and Opportunities

Tree breeding faces major bottlenecks like prolonged breeding cycles, long juvenile phase, high flower drops, high heterogeneity, self-incompatibility, complicated reproductive biology, varied pattern of genetic variability over time, etc. Although vegetative reproduction methods like cutting, grafting, air layering etc. have enabled the faster multiplication of improved clones however, other challenges still remain unsolved. Modern day biotechnological approaches have been emerged as an opportunity for reducing the life cycle, mass multiplication of quality planting material, understanding the genetic diversity and population structures of the tree populations, identifying the ancestral relationships, dissecting the molecular pathways for major biosynthetic pathways through genomic approaches, breeding for biotic and abiotic

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stresses, and higher yield of improved quality of trees which was otherwise not possible through traditional tools and techniques. Because most, if not all, variables of interest have complex multifactorial inheritance, genetic dissection techniques like quantitative trait mapping and association genetics have proved ineffective in guiding operational marker-assisted selection (MAS) in forest trees. High-throughput genomics and quantitative genetics have combined to create two new paradigms that are challenging traditional ideas about tree breeding. A vast number of genome-wide markers are used in genomic selection (GS) to forecast complicated phenotypes. It has the ability to quicken breeding cycles, intensify selection, and enhance breeding value precision. Future GS research should focus on improving methods for updating prediction models, adding functional genomics data that has been validated to increase prediction accuracy, and combining genomic and multi-environment data to predict how genetic material will behave in untested locations or under changing climate scenarios. There should be further chances to improve the application of genomics to tree breeding as phenotypic and genome-wide data accumulate across large breeding populations and breakthroughs in computational prediction of discrete genomic traits.

3.5

Conclusion and Way Ahead

Healthy feeding of livestock has been a major constraint in developing countries like India. Tree fodder is a traditional source of feed and the modern days’ need to support livestock health and productivity. Additionally, tree-based forage also provides better mitigation to the ill effects of environments in comparison to the annual forage crops. Breeding such species would ensure yield and greater feed quality while also creating a scope for restoring degraded lands and harsh biotic and abiotic pressures in the scenario of climate change era. Several tree-based forage sources are available today and the selection for better traits has led to the development of improved varieties and clones. Further, the advanced molecular and next-generation sequencing (NGS) tools have a great scope for accelerating the genetic improvement of these perennial species.

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Impact of Climate Change on Forage Crop Production with Special Emphasis on Diseases and Mitigation Strategies Through Breeding and Molecular Approaches Namburi Karunakar Reddy, Gaurav Rakhonde, Pooja Purushotham, Pooja S. Patel, and Shalaka Ahale 4.1

Introduction

Fodder crops are the plant species that are cultivated and harvested for feeding the animals in the form of forage (cut green and fed fresh), hay (dried green fodder), and silage (stored under anaerobic condition) (Kapoor et al. 2018). These fodder crops are essential for livestock for ruminant production, with grazing land accounting for approximately 60% of global agriculture land (FAO 1997). In India, the total area under cultivated fodders is 8.4 million ha on individual crop basis. The average cultivated area in India under fodder production is only 4.4% of the total cultivated area but the country inhabits 15% of world livestock population on 2% geographical area, which itself is an indication of an extent of livestock pressure on our resources in comparison to other countries (Kapoor et al. 2018). Continued human population growth predicts an increase to 9.7 billion by 2050, which will lead to an increase in demand for animal products together with pressure to decrease pollutant output (Kingston-Smith et al. 2013). The total livestock population is 535.78 million in the country showing an increase of 4.6% over Livestock Census-2012 (Department of Animal Husbandry and Dairying 2019). Composition of livestock is also changing with a shift towards small ruminants due to high growth in the meat sector. By the end of 12th Plan, demand for milk is expected to increase to 141 million tons and for meat, eggs, and fish together to 15.8 million tons (Kumar et al. 2012).

N. K. Reddy (✉) · G. Rakhonde · P. Purushotham · P. S. Patel Department of Plant Pathology, University of Agricultural Sciences, GKVK, Bangalore, Karnataka, India S. Ahale Department of Plant Pathology, Punjab Agricultural University, Ludhiana, Punjab, India # The Author(s), under exclusive license to Springer Nature Singapore Pte Ltd. 2023 R. K. Singhal et al. (eds.), Molecular Interventions for Developing Climate-Smart Crops: A Forage Perspective, https://doi.org/10.1007/978-981-99-1858-4_4

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Population Growth of Livestock Vs. Fodder Production

Due to conflicting land uses, the cultivated area for fodder has remained constant over the last 20 years at about 8.4 Mha. The country’s production of fodder is insufficient to fulfil the demands of the expanding livestock population, and the majority of the forages provided to animals are of poor quality. Currently, the nation has a net shortage of 44% feeds, 10.95% dry crop leftovers, and 35.6% green fodder (IGFRI Vision 2050). Due to the cattle population’s continued growth at a pace of 1.23% in the upcoming years, the gap between supply and demand may widen even more (Kumar et al. 2012). India ranks first in the production and consumption of milk. Due to urbanization, there is a significant change in the feeding habits of various people regarding consumption of milk products, meat, and eggs which resulted into an increase in demand of various livestock products (Ghosh et al. 2016). But livestock productivity of India is one of the lowest in the world mainly due to various problems faced by the livestock and forage sector. In India, fodder requirement is mainly fulfilled by three sources: crop residues, fodder crops, and pasture or grazing lands. One of the main problems incurred in meeting fodder requirement is the uneven distribution of fodder sources throughout the country. In addition to increasing feed availability, forage production is crucial for sustaining the basis of natural resources by stabilizing the soil, avoiding soil erosion, and enhancing soil fertility. However, the current fodder supply of 27 kg per animal per day is still far from enough as each horse needs roughly 40 kg of green fodder daily for appropriate feeding. Increased pressure to produce more food grains, oilseeds, pulses, and commercial crops is anticipated to cause this gap between supply and demand to expand even further, maybe with no growth in the area planted in forage crops. As a result, it is crucial to raise the productivity of pasture and fodder lands by creating new species of fodder and using more waste land for the production of fodder (Kapoor et al. 2018). At present the productivity of cultivated fodder crops is low, due to least attention and allocation of minimal production resources and because of non-availability of the production techniques to stakeholders involved in the forage resource development on the other. This needs to be tackled by educating the farmers about the production packages of fodder crops like selection of appropriate forage species, varieties, and management techniques to sustain forage yields and soil fertility (Kumar et al. 2012). Green fodders are crucial to the success of livestock production as well as the beginning of the white revolution. The success of the dairy industry depends greatly on the availability of green fodder throughout the year because green fodders are the least expensive source of carbohydrates, proteins, vitamins, and minerals for dairy animals. Farmers cultivate fodder crops in accordance with traditional crop rotations, which results in an irregular supply of fodder throughout the year. Therefore, the price of producing milk can be significantly decreased by giving the milk animals more green fodder rather than expensive concentrates and feed. Therefore, maintaining the availability of high-quality fodder in a balanced ratio is the only way to achieve the goal of enhanced milk production. The green fodder availability from cultivated areas includes supplementation from sugarcane tops and seasonal weeds. The availability

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of dry fodder is constituted of straw from crops such as rice, wheat, barley, maize, sorghum, pearl millet, and other crops such as groundnut and chickpea, as well as dry grass from grazing lands and forests (Kumar et al. 2012). Among the various limiting factors, diseases have also been the major constraints for fodder cultivation. Several diseases are responsible for causing serious damage to fodder crops. In India, diseases alone can cause losses up to 74% in cowpea (Chester 1950), 50% in sorghum (Sunderam 1970), 72% in Lucerne (Ahmad 1977), 75% in cluster bean (Chester 1950), 30% in bajra (Ahmad 1969; Sunderam 1970), and 55% in oats (Ahmad 1969). Besides, these diseases also affect the quality parameters of forages.

4.3

Impact of Different Biotic and Abiotic Factors on Forage Crop Production

Forage crops often face an increasing number of biotic and abiotic challenges as a result of global warming and probable climatic irregularities, which significantly impair their development and productivity. Abiotic stresses such as drought, temperature, soil moisture, and other factors have been proven to be more damaging to crop yield when they occur concurrently with biotic stresses at various crop growth stages. Biotic factors that affect agricultural productivity are simply living or biological forces that influence agricultural output or practice. Drought, high and low temperatures, and salinity are all known to impact the prevalence and spread of diseases, insects, and weeds (biotic factors) by affecting plant physiology and defensive responses (Ramegowda and Senthil-Kumar 2015).

4.3.1

Impact of Different Diseases on Forage Crop Production

Diseases continue to impair herbage and seed productivity, as well as the nutritional content and palatability of forage crops. It has an impact on animal health and output, although the precise loss caused by it is difficult to predict (Chakraborty 2021). Soilborne fungi and rusts cause a variety of foliar diseases, and many viruses (Jones 1996) and nematodes (Stanton 1994) have a host range that includes both field crops and forage crops. Pathogen survival and dissemination methods are significant. Many soil-borne pathogens survive for extended periods of time without host plants as latent propagules or inside decaying host tissue and rust spores can travel considerable distances (Nagarajan and Singh 1990). Diseases produced by endemic pathogens are typically not a serious problem for forage crops that have co-evolved with existing flora and fauna. When native forage crops are supplemented with imported grass and legume species to improve animal performance, diseases have influenced the establishment, productivity, quality, and persistence of these improved grasslands (Lenne and Trutmann 1994). The diseases that have devastated introduced grasses are anthracnose (Colletotrichum gloeosporioides) of Stylosanthes in Australia (Irwin and Cameron 1978) and blight

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of Cenchrus in the USA and Australia (Perrott and Chakraborty 1999; Rodriguez et al. 1999). Root and crown rot of alfalfa in Australia is caused by Phytophthora megasperma, Colletotrichum trifolii, Acrocalymma medicaginis, Staganospora meliloti, and Phomopsis sps (Irwin 1989; Nikandrow 1990). Symptoms can differ depending on plant age, and seasonal variation can affect the pathogen suite involved. Similarly, pathogens such as Pythium middletonii, Cordinea fertilis, Fusarium solani, Macrophomina phaseolina, and Aphanomyces euteiches can interact in white clover root causing stolon rot disease (Greenhalgh 1995; Flett and Clarke 1996). The negative effects of forage crop pathogens, such as oestrogenic clover on animal health and reproduction, are of special concern (Latch and Skipp 1987). Production losses in Australia from lupinosis-induced animal toxicity (Gardiner 1975) and annual ryegrass toxicity (McKay 1993) are estimated to be between $10 and $16 million per year (Madin 1993). Annual ryegrass poisoning, caused by the bacteria Clavibacter toxicus and the nematode Anguina funestra, kills animals that graze infected ryegrass and other grasses like Polypogon monospeliensis (McKay and Ophel 1993). Besides, the economic losses caused by pathogens also include damage to soil reactions, nutrients, and nitrogen-fixing organisms (Chakraborty 2021).

4.3.2

Impact of Insects on Forage Crop Production

Insect populations will be altered in a variety of ways as temperatures rise. Higher yearly temperatures will hasten insect life cycles and allow them to expand their geographic range (Walther 2010). Species that may produce several generations per year will not only be able to produce more individuals each growing season, but the existence of multiple generations each growing season raises the risk of resistance to control techniques (e.g. pesticide resistance) (May and Dobson 1986). This is because as most of the forage crops are perennials with multiple growing seasons, they are available to insects for feeding for a longer period of time, affecting future fodder output. As a result, most assessments of insect-induced damage in forage crops have concentrated on yield loss over a short period of time, which serves as the only foundation for most EIL and intervention thresholds (Sulc et al. 2020). Managing pests and diseases in a changing climate will be made more difficult by biological, social, and economic variables that impact the available methods (CAST 2017). Pesticide resistance will restrict the number of products available for pest management. The public’s opinion of pesticides, as well as the expense of creating and bringing new products to market, has hindered the introduction of new pesticides. The stem-boring fruit fly (Oscinella sp.) may reduce the number of tillers in Italian ryegrass during establishment, though there is little evidence of insect damage to forage grasses (Bentley and Clements 1989). Insect infestations have greater influence on the yield of fodder crop and have received significantly less attention on quality in terms of nutritional content. Though insects have a little

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impact on nutrient concentrations in forage crops, they can cause significant decreases in nutrient output per unit of land area due to lower biomass production (Hutchins et al. 1989). But potato leafhopper feeding lowers alfalfa carotene, crude protein, ash, calcium, and phosphate contents (Smith and Medler 1959; Kindler et al. 1973). Stem boring damage caused by the Bermuda grass stem maggot reduces the relative feed quality of late-season Bermuda grass hay by 7% owing to a decrease in total digestible nutrients (TDN) and a slightly decreased dry matter intake (DMI) (Baxter et al. 2017).

4.3.3

Impact of Weeds on Forage Crop Production

Weed species responded relatively strongly to elevated CO2, which could boost growth and make them herbicide-resistant (Ziska 2003). This will increase competition between weeds and forage crops for space, water, and nutrients resulting in increased crop loss. Rising temperatures will aggravate competition by stimulating the growth of most weeds (primarily those in the C4 metabolic pathway) (Ramesh et al. 2017). Weed infestation and growth of forage crops are dependent on the combination of weed and forage crop species (e.g. C3 weed and forage crop, C4 weed and forage crop, C4 weed and C3 forage crop, and any other combinations) (Ramesh et al. 2017). Many agronomic weeds originate in warmer locations (tropical or warm temperate) and are likely to spread when air temperatures rise (Patterson et al. 1999; Rahman and Wardle 1990). It is also demonstrated that a 3 °C increase in temperature could potentially increase itch grass biomass yield by 80%. In comparison to sorghum crop, increased CO2 concentration causes more growth and biomass production of cocklebur (a common weed of sorghum crop) (Ziska 2001). Some weed species, like yellow star thistle and cheatgrass, can grow more quickly with even less precipitation due to their adaptable phenology to drought tolerance, and they have a significant negative impact on the growth of forage grass (Hatfield 2006). Invasive weeds are expected to be a bigger problem in grasslands and forage crops than in crop-producing areas. Weeds such as Sida spp. have taken over certain pastures in Queensland after anthracnose destroyed the vulnerable Stylosanthes cultivars (Chakraborty 2021). Rumex obtusiflius is a noxious weed of grasslands, grows at a very fast rate, and significantly reduces crop yield (Gilgen et al. 2010).

4.3.4

Impact of Temperature on Forage Crop Production

Warmer temperatures influence forage crop growth and development, as well as the survival of plants and grain under high temperatures. The time it takes from seeding to harvesting is controlled by the average temperature and day length (Craufurd and Wheeler 2009). For optimal vegetative and reproductive growth, forage crops have a very restricted temperature range. As the temperatures increase, the time it takes to harvest decreases, at least 25% of deficit can be seen in their production. Extreme heat will become more common as a result of climate change. When hot days

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coincide with a sensitive stage of forage crop development, such as flowering, large reductions in seed or grain yields can be seen due to pollination disturbance (Wheeler et al. 2000). Each 1 °C rise in average growing season temperature is expected to reduce maize production by 8.3% and soybean yield by 13% (Lobell and Field 2007). Wheat yields are lowered when air temperatures are above 31 °C, which lowers pollen and ovule development, resulting in fewer and smaller kernels (Ferris et al. 1998). Higher temperatures cause increased lignification of plant tissues, resulting in decreased digestibility and rate of degradation of plant tissues. It reduces the availability of nutrients to animals, resulting in a decrease in livestock production (Minson 1990).

4.3.5

Impact of Precipitation on Forage Crop Production

Precipitation estimates are less definite than temperature projections, and crop response will be determined by year-to-year fluctuation, precipitation distribution over the growing season, and the quantity of precipitation received in extreme events (Pryor et al. 2014). Around 80% of our agricultural crops and 100% of pasture crops are rainfed (Reilly et al. 2003). The intensity of precipitation is also predicted to fluctuate as the number of heavy rainfall incidents increases. Extreme rainfall regimes have a tendency to lengthen and exacerbate soil water stress, particularly in mesic ecosystems (Knapp et al. 2008). Precipitation obtained during heavy rainfall frequently exceeds the soil’s infiltration rate, and this precipitation may be lost as runoff. Increased runoff increases the chances of soil erosion (Zhang et al. 2012). Another consequence of severe precipitation is that rainwater washes off the surface nutrients making them unavailable for crop cultivation (SWCS 2003). Any change in growing season rainfall results in a decrease in grass species richness (Wilkes 2008) and an increase in soil salinity and degradation (Howden et al. 2008). Increased precipitation may increase forage crop productivity, but drought may reduce both the quantity and quality of pasture (Humphreys 1991). A 50% increase in dry spell duration results in a 10% decrease in primary productivity (Fay et al. 2003). If there is a high temperature along with a precipitation deficit of up to 300 mm, the yield will be reduced by 20–36% (IPCC 2007).

4.3.6

Impact of Water Availability on Forage Crop Production

Forage crop production will be affected by reduced water availability and forage crop production will need careful use of remaining water resources for irrigation (Bathke et al. 2014). Increasing the region’s precipitation may result in enough soil water availability, which benefits forage crop output. With its extensive tap root, alfalfa (lucerne) is more drought resistant than many grasses, but it cannot be grown successfully in all soil types and climates. The major effects of water supply on forage crops are generally related to the relative maturity of the plant at harvest, as

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well as the timing of the stress relative to the plant’s growth stage. Less water supply makes forage crops mature early. The fibre content of plants tends to increase as the protein content of leaves and stems decreases, while the starch, oil, and protein content of seeds increase at the maturity stage. The degree of lignification of the fibrous material present is extremely crucial for herbivores, as lignified plant cell walls are generally less digestible in turn more filling, which limits intake and thus reduces the production rate even further when compared to more digestible feeds (Van Soest 1994).

4.3.7

Impact of Soil on Forage Crop Production

When temperatures are forecast to rise with no change in precipitation, soil moisture is likely to fall by 5 to 10%, resulting in increased crop water stress. These modifications will put greater pressure on water resources where irrigation is carried out. But when both temperature and precipitation are expected to rise, soil moisture will increase leading to an increase in N2O emissions (Doran et al. 1990). As a result, warmer soil temperatures and a longer frost-free period will promote biological activity, altering the nutrient cycle and carbon sequestration. Increased precipitation increases the possibility of nutrient leaching and erosion. Solute leaching, lateral flow, and seepage can all lead to the formation of salty seeps that is detrimental to the growth of forage crops (Halvorson and Black 1974).

4.3.8

Impact of Increased CO2 Concentration on Forage Crop Production

An increase in the concentration of CO2, one of the principal greenhouse gases, boosts forage crop output through increasing photosynthetic rates in all forage crops that employ the C3 photosynthetic pathway but not so in the crops that are undergoing the C4 photosynthetic pathway. The leaf photosynthetic rates of C4 plants are not significantly increased by increasing CO2 concentrations (Drake et al. 1997). Hence, yield improvements in C4 plants cultivated under elevated CO2 are considerably meagre than for C3 plants (Kimball et al. 2002). Higher CO2 concentrations cause partial stomatal closure, which reduces transpiration and improves water use efficiency (Rotter and van de Geijn 1999). There is also a little improvement in the efficiency with which both C3 and C4 crops utilize water under elevated CO2 circumstances (Drake et al. 1997). According to the IPCC, the decrease in the nutritional quality of fodder caused by increased CO2 concentrations is primarily due to increased carbon-to-nitrogen ratio in plants and increased dominance of palatable plant species. Grain cultivation for C3 crops may result in higher yields but inferior quality (protein content) for human and animal nutrition under increased atmospheric CO2 levels (Erbs et al. 2010; Asif et al. 2017). A meta-analysis conducted by Dumont et al. (2015) discovered that higher

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CO2 levels increased non-structural carbohydrates by 25% while lowering nitrogen by 8% in content.

4.3.9

Impact of Increased Concentration of Ozone on Forage Crop Production

A visible injury in the form of burning caused by oxidizing of ozone in the leaf tissue during respiration caused by the severity of ozone gas under environmental temperature (Krupa et al. 2001). However, foliar injury may not always be an accurate indicator of ozone effects on crop dry matter production and quality (Booker et al. 2009). Current ozone gas concentrations in a number of countries around the world have the potential to suppress the growth and productivity of various agricultural plants (Mills et al. 2007). When ozone gas enters a plant’s leaf, it interacts with various cellular processes and inhibits photosynthesis, ultimately reducing crop plant growth and yield. Increased ozone levels reduce maize yield by 2.5–5% (Avnery et al. 2011). According to Cho et al. (2011), the harmful effects of ozone are primarily caused by a combination of chemical toxicity and plant-mediated responses that may amplify or inhibit the injury. The current level of ozone concentration is extremely harmful to forage crops such as lucerne and clover, significantly reducing yield in various parts of the world (Booker et al. 2009). The effects of ozone on feed quality are greater than the effects of feed dry matter content (Muntifering et al. 2000). According to reports, increased CO2 levels may mitigate the harmful effects of ozone on vegetation (Booker et al. 2009).

4.3.10 Impact of Change in Air Composition on Forage Crop Production Changes in air composition may have direct effects on crop diseases, such as effects on host-pathogen interactions either through effects on the host, the pathogen, or the interaction (Eastburn et al. 2011). However, there may be indirect effects mediated by changes in climate caused by rising greenhouse gas concentrations Pachauri and Reisinger 2008) and these changes have been accelerating in the last 20 years (Semenov 2009). Increased concentrations of both wet and dry sulphur led to deposition that may cause plant tissue damage, acidification of the water in which spore germination occurs, and improved crop sulphur nutrition (Fitt et al. 2011).

4.4

Impact of Climate Change on Forage Crop Production with Emphasis on Diseases

Over the last millions of years, the earth’s natural history has seen dramatic changes in the atmosphere and biosphere. These changes were glitchy in the past, but currently, they are pretty fast and significantly impacted by human activity. Climatic

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observations provide growing evidence that the climate is changing. Since the 1850s, the global mean temperature has risen by 0.8 °C (IPCC 2007). Carbon dioxide (CO2) levels in the atmosphere have increased from around 284 mg/kg in 1832 to 391 mg/kg in 2012 (Tans and Keeling 2012), which is primarily attributed by the combustion of fossil fuels, with minor contributions from land-use changes. Through fundamental physics, there is a strong theoretical relationship between higher greenhouse gas levels in the atmosphere and increased global warming. As a result, climate change caused by human activity is projected to result in greater temperatures, changes in rainfall patterns, and an increase in the frequency of extreme weather. The IPCC (2019) special report discussed that rising global greenhouse gas emissions increased land surface and ocean temperatures. The temperature varied from 1.38 to 1.68 °C. Increased GHG emissions associated with extreme climate events will harm the livestock industry by damaging forage crop output directly or indirectly (IPCC 2019). India is currently experiencing a net shortfall of 35.6% green fodder, 10.95% dry agricultural leftovers, and 44% concentrate feed supplements (IGFRI Vision 2050). The root cause of this insufficiency is climate change, which has a negative influence on fodder production. Weather and climate are major factors impacting the lives of flora and fauna on Earth. Agriculture and animal husbandry are innately vulnerable to climatic fluctuation and change that might have been caused by natural or anthropogenic factors. Climate change caused by GHG emissions is projected to have a direct influence on agricultural production systems for food, feed, or fodder and affect livestock health (Wheeler and Reynolds 2013). Human activity has a significant impact on the climate, seas, cryosphere, and terrestrial and marine biospheres. Although change has always existed on our planet, the industrial revolution has hastened its pace (Chakraborty 2021). Temperature and CO2 concentration changes in the atmosphere already has an effect on grassland production and carbon sequestration in soils (McGinn and Wedin 1997). Plant pests (insects and diseases) cause significant forage crop productivity losses, which may be worsened by climate change. Crop productivity losses will rise by 10 to 25% for every degree of global surface temperature increase. Rising temperatures may impact the distribution, intensity of infestation, and life cycles of pest species, as well as crop or forage species susceptibility to disease. Food insecurity might become exceedingly severe as a result of climate change (Uddin and Kebreab 2020). Changes in the gaseous composition of the air can have a direct impact on the severity of crop disease epidemics by influencing the host, the pathogen, or the hostpathogen interaction (Eastburn et al. 2011). The severity of epidemics may be reduced if increased CO2 concentrations while increased O3 concentrations may damage tissue and favour the development of necrotrophic pathogens such as Botrytis cinerea causing grey mould on many hosts (Semenov 2009; Eastburn et al. 2011). While there are interactions between N2O and O3 concentrations, there has been little research on the direct effects of N2O on the severity of disease epidemics (Chipperfield 2009). Climate change could have direct effects on individual plants and plant communities in the absence of diseases, but it may also elicit changes in plants

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that alter their interactions with pathogens (Garrett et al. 2006). Climate change leads to the changes in the genetic makeup of plant population, abundance of certain plant species, and certain species may succumb to pathogens and exhibit symptoms such as wilting, leaf burn, leaf folding, abscission, and pathogen susceptibility. Challenges with preserving traditional land races or varieties, loss of biodiversity, deterioration of source of resistance genes, and cultivars with broader adaptability are required. Climate change will impact host physiology and resistance, as well as the stages and rates of pathogen development (Kumar and Tuti 2016). In addition to influencing forage crop growth and distribution, climate change will also have an impact on fodder quality. Until recently, the potential impact of climate change on plant disease was mostly unexplored (Manning and Tiedemann 1995; Chakraborty et al. 1988, 2000; Coakley et al. 1999). Climate change from experiments and modelling studies shows an influence on crop loss, the efficacy of management techniques, and the geographical spread of diseases. Temperature and precipitation changes will be significantly responsible for changing geographical distribution. Other consequences will be mediated by changes in host-pathogen physiology. In controlled conditions, for example, twice the ambient CO2 lowers germination, germ tube development, and aspersorium formation while increasing fecundity of Colletotrichum gloeosporioides infecting Stylosanthes (Chakraborty et al. 2000). When canopy CO2 concentration is elevated leading to favourable weather for anthracnose growth, the expanded canopy captures more disease spores, which multiply fast due to higher fecundity. As a result, more virulent and aggressive races may emerge faster, threatening the endurance of resistance. Climate change will also have an influence on diseases caused by pathogens found in soil. Perennial ryegrass roots grown at 700 ppm CO2 disintegrate at a slower pace than those produced at 350 ppm CO2, increasing the time of roots exposed to soil-borne diseases (Pennypacker 1997). SO2 in the atmosphere causes rain to fall with a pH as low as 4 and aids in the germination or growth of M. graminicola or P. nodorum on wheat leaves (Chandramohan 2010). Detailed research on the effects of climate change on grassland diseases is severely lacking. Because microorganisms have a short generation time and a huge population size, potential ramifications will be noticed first in these species. They have the potential to function as an early warning system for climate change. Because it typically takes 10–30 years to create a disease-resistant cultivar, mitigation studies must begin immediately. Growing perennial forage crops is a solution for mitigating the effects of climate change.

4.5

Impact of Forage Diseases on Livestock Health

Access to livestock products is a vital issue in global food supply as human population explosion is a continuous process. The ruminants contribute to climate change, so it is possible that climate change will have an impact on ruminant output. Climate change models for the near future forecast increase in temperature, CO2,

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precipitation, and fluctuating weather systems causing stress reactions in field crops. Pre-exposure to changed climatic conditions alters plant cell composition while also priming plant cells to modify their post-ingestion metabolic response to the rumen. In terms of gas generation, varietal variations in fermentation were noticed; however, higher temperature and CO2 had minimal influence (Hart et al. 2022). The diseases have also impact on the quality of the forage and affect the nutritional value of the forage and it ultimately affects the livestock. Various diseases occurred in the forage crops and they directly and indirectly affect the animal health. Compounds like alkaloids, toxins, secondary metabolites, etc. were produced by fungal, bacterial pathogen associated with disease. Sometimes plants also possess such compounds and impact the health and other aspects with respect to the livestock.

4.5.1

Toxicants in Forages

Each plant defends itself against herbivores using physical defences such as leaf hairs, spines, thorns, high lignification and growing habitat, as well as chemical defences such as a diverse variety of complex compounds that are toxic or poisonous in nature. These compounds can be generated by the plant itself or by symbiotic or mutualistic fungi or bacteria that grow alongside it. These are often secondary chemicals like alkaloids that indirectly participate in cellular metabolism but are evidently generated to act as the plant’s defence arsenal. Mycotoxins are chemicals created by fungus that can be produced by fungi growing on or in forage plants. Many grazing animal diseases are caused by mycotoxins. Neotyphodium coenophialum, a tall fescue endophytic fungus, generates ergot alkaloid causing fescue foot, summer fescue toxicosis, and reproductive problems, whereas an endophyte in perennial ryegrass produces lolitrems that induce ryegrass staggers (Mayland et al. 2007). Total livestock losses linked to the tall fescue endophyte in the United States are estimated to be between $500 million and $1 billion per year (Ball et al. 1993). Annual economic impact of a diverse range of toxic plants on cattle output in the western United States is estimated to be in the hundreds of millions of dollars (James et al. 2019). Toxins generated from plants are classified into many major classes, including alkaloids, glycosides, proteins, and amino acids and phenolics. Alkaloids are bitter chemicals that have a heterocyclic ring structure and include N. There are hundreds of different alkaloids and are classified based on the chemical structure of the N-containing ring. Condensed and hydrolysable tannins are phenolic compounds having aromatic rings with one or more hydroxyl groups. Due to their chemical reactivity, hydroxyl groups can react with protein functional groups to form indigestible complexes. Feed intake is reduced by astringent tannin-protein complexes (Min et al. 2003). All plants contain phenolic compounds. In rare cases, their kind or concentration may cause unwanted animal responses. Feed intake and protein digestibility are decreased in bird’s foot trefoil and Sericea lespedeza. Tannins found in oak browse cause oak poisoning. Many tropical agroforestry tree legumes contain enough tannin to impair animal performance (Mayland et al. 2007).

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Table 4.1 Impact of toxins associated with fungus and bacteria in forage crops on animal disorders Sl. no 1.

Forage Red clover

Fungus Rhizoctonia leguminicola

Alkaloid Slaframine

2.

Sweet clover

Mould

Dicoumarol

3.

Wild lupines (silky lupine, tail cup lupine, spurred lupine)

Phomopsis leptostromiformis

Anagyrine, an alkaloid

4.

Perennial ryegrass pastures

Pithomyces chartarum

Hepatotoxin, sporidesmin

5.

Dallisgrass

Claviceps purpurea

Ergot alkaloids

6.

Tall fescue cultivars

Neotyphodium coenophialum

7.

Perennial ryegrass Ryegrass

Neotyphodium lolii Clavibacter spp. and Anguina agrostis

Ergovaline, an ergot alkaloid Lolitrems

8.

Corynetoxins

Impact Excessive salivation, eye discharge, bloat, frequent urination, and watery diarrhoea Vitamin K deficiency, bleeding, and haemorrhaging Teratogenic in cattle. Crooked calf disease to pregnant cows during days 40–70 of gestation. Severe skeletal deformations in the foetuses may occur Facial eczema, sporidesmininduced liver damage and unable to metabolize phylloerythrin and accumulates in blood. Also causes severe dermatitis of face, udder, and other exposed areas Vasoconstriction and reduced blood supply to the extremities, resulting in sloughing of ear tips, tail, and hooves. Neurological effects, including hyperexcitability, incoordination, and convulsions Fescue foot, summer fescue toxicosis, and fat necrosis Inhibitor of neurotransmitters Causes annual ryegrass toxicity, permanent brain damage, convulsions of increasing severity and death

Table 4.1 highlights the impact of toxins associated with fungus and bacteria in forage crops.

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4.6

Mitigation Practices at Farmer or Field Level to Curtail Hazards of Mycotoxin in Forage Crops

4.6.1

Deterrence of Mycotoxin Contaminations of Forage Crops in Field and During Storage

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In general, pre-harvest and post-harvest approaches should be adopted to avoid or mitigate mycotoxin presence in agricultural products. The most significant field measures to combat fungal invasions are opportune crop rotation, tillage, planting date, soil nutrients, crop hybrid or variety selection, chemical and biological infestation control, crop removal, bug and weed controls (Jouany 2007; Jard et al. 2011; Battilani et al. 2013). Farmers are required to eliminate any material suspect of being associated with mycotoxins, including grains appearing clean but may have mouldy areas and grains with moisture less than 13% with low temperatures (Jard et al. 2011). In the case of silages, a little oxygen content and CO2 supplementation are effective in avoiding mould formation (Elferink et al. 2000). Aerobic degradation might result in nutritional and dry matter loss, heating harm to nutrients, unnecessary proteolysis, development of unwanted microbes such as mycotoxigenic fungal infestation and their toxins (Driehuis et al. 2008). The detrimental consequences of aerobic exercise may be more severe in some sections of silage, particularly towards the outskirts of the ensiled crop (Borreani and Tabacco 2010). When the silo is opened, in the front mass, the activity of yeasts and mould gets activated and promotes the growth of potentially toxic fungi. Reducing the area vulnerable to the risk of air penetration is made possible by using precise ensiling procedures. As feasible and affordable alternatives for acid-based additives, lactic acid bacteria (LAB) are being used (Tabacco et al. 2011). The use of heterolactic LAB inoculants, L. buchneri, has demonstrated the potential to advance silage production from easy, moderately difficult, and difficult to ensile materials by reducing pH, ammonia nitrogen, and DM losses due to the greater generation of acetic acid, which has a stronger antimycotic action than lactic acid. This is despite the fact that heterolactic fermentation is less effective than homolactic fermentation in terms of nutrient conservation (Ranjit and Kung Jr 2000; EFSA 2013).

4.6.2

Mycotoxin Detoxification and Biodegradation

Mycotoxins can cause a variety of adverse health effects and pose a serious health threat to both humans and livestock. The adverse health effects of mycotoxins range from acute poisoning to long-term effects such as immune deficiency and cancer and hence It is an extremely challenging task to avoid mycotoxin contamination of feeds before harvest or during storage (CAST 2017). Mycotoxicosis has been successfully reduced in both humans and animals by adding sorbent materials that are supposed to bind the mycotoxins effectively in the gastro-intestinal tract as well as addition of enzymes or microorganisms can detoxify mycotoxins in animal diets (Gallo et al. 2010). Mycotoxin sequestering agents are compounds proficient of binding

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mycotoxins in feeds without separating the toxin-sequestering agent combination, enabling the toxin to transit through the gastrointestinal tract of animals and be eliminated via faeces. Clays are generally used as binding entity to lessen cattle AFB1 (Aflatoxin B1) poisoning and AFM1 (Aflatoxin M1) transfer to breast milk (Masoero et al. 2009). Organic sequestering entities like activated carbon and yeast cell wall products have been shown to effectively lower AFM1 in milk from cows fed AFB1-contaminated diet. Besides, a mycotoxin neutralizing product was tested on lactating dairy cows also caused those animals to become cured when they were naturally exposed to AFs (Pietri et al. 2009) and some Fusarium origin toxins, viz. DON (Deoxynivalenol), ZEA (zearalenone), FB1 (Fumonisins B1), OTA (Ochratoxin A), and T-2 toxin, and it is now approved by the European Union for usage in pig diet (Kiyothong et al. 2012; Murugesan et al. 2015). Ergotism can be avoided by inhibiting grass seed establishment. Jones and Megarrity (1986) discovered that Hawaiian ruminants accustomed to a leucaena diet have mimosine-degrading rumen bacteria that removed the toxicity. In Australia, these bacteria have been introduced into cattle, allowing leucaena to be utilized as a productive source of high protein fodder (Quirk et al. 1988). In Florida, Hammond et al. (1989) documented detoxification of mimosine using native or imported rumen bacteria. Thus, such bacterial inoculum can be utilized and will be a good eco-friendly approach. Three strategies underpin the method of action: (1) Polar mycotoxins (e.g. AFs) remain adsorbed by inorganic entities; (2) other mycotoxins not or poorly engrossed by inorganic entities (e.g. trichothecenes, ZEA) are biotransformed using biological constituents, Eubacterium strain (BBSH 797), and a yeast strain (Trichosporon mycotoxinivorans MTV) capable of transforming mycotoxin structures into non-toxic form; (3) Mycotoxins phycophytic compounds produced from marine alga, Ascophyllum nodosum and plant Silybum marianum extracts protect against mycotoxins (Pietri et al. 2009). Vaccination techniques have recently been investigated as a substitute to the usage of sorbent ingredients in animal diet to avoid deleterious impact of mycotoxin intake in nursing dairy cows and heifers or to limit AFM1 carryover into milk and cheese (Polonelli et al. 2011). On commercial farms, Santos and Fink-Gremmels (2014) discovered that nutritional supplementation with a glucomannan mycotoxin absorbent entity was effective in preventing mycotoxincosis in dairy cattle having mouldy silages. In vitro and in vivo research is mandatory to validate the effectiveness of numerous commercially existing sequestering entities on several of mycotoxins. Moreover, for several contaminated diets, challenge is the possible co-occurrence of numerous mycotoxins; subsequently it is essential to standardize sampling techniques, use specific analytical methods capable of detecting several mycotoxins concurrently and their modified forms, and implement appropriate tactics for fruitfully mitigating the various undesirable things of the extensive variety of mycotoxins contaminating animal diets.

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Breeding for Disease Resistance of Forage Crops

The key biological strategy for raising yields is diversity. Due to an appropriate diversity of genetic resources and a recognized gene pool, it is possible to successfully solve the fundamental problems of breeding to produce fundamentally new, resilient to environmental stress, high-yielding varieties of forage crops that meet the objectives of sustainable development of contemporary animal husbandry and ecological agriculture (Kosolapov and Shamsutdinov 2015). One of the most difficult aspects of breeding is the development of resistant variants. They have substantial adaptation capacities due to their fast reproduction rate and, as a result, the large potential of pathogen variety. The genetics of the host-pathogen connection is a complicated problem in the research of resistance genetics. Plants with the best combination of resistance genes are rare and difficult to find. Furthermore, resistance frequently has a negative correlation with commercially useful plant features (Razgulyaeva et al. 2019). The quest for the development of disease-resistant breeding material is critical. According to N.I. Vavilov (1965), the production of the source material is as important as the selection procedures themselves.

4.7.1

Use/Development of Host Resistance Varieties

Genetic tactics to disease management are the most feasible and successful. Between 1967 and 1985, incorporating Anthracnose resistance into alfalfa cultivars added more than US$240 million per year, while increasing the percentage of Kabatiella caulivora resistant plants in red clover boosted output by more than 30% and better persistence (Casler and Pederson 1996). Selection is persisted as most important source of novel cultivars exploited to surge the quality and availability of grassland feed. Breeding is more expensive and justified for temperate species with larger economic returns, such as alfalfa, clover, and several grasses like ryegrass (Cameron 1983). Breeding for disease resistance in tropical plants has not been widely used, with the exception of legumes like Stylosanthes, which have at least two breeding programmes aimed at enhancing Anthracnose resistance (Cameron et al. 1997; Miles and Lascano Aguilar 1997). An indirect yet precise selection technique based on pathogen life cycle components and race non-specificity has been employed as a potent selection means for more than one species of Stylosanthes against Anthracnose (Chakraborty et al. 1988; Iamsupasit et al. 1993). The capacity of this resistance to slow down the spread of an epidemic has been proved in fields and a unique stochastic model was exploited to elucidate disease progression and the role of weather and host resistance (Chakraborty and Smyth 1995). Nineteen lines possessing race-nonspecific quantitative resistance were found and a recurrent selection effort utilizing these lines resulted in considerable improvements in biomass, seed output, and anthracnose resistance (Cameron et al. 1997). Some elite choices from breeding effort have maintained a high degree of Anthracnose resistance at the heart of host-pathogen

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diversity in Brazil against a more diversified pathogen population than the suite of races for which the material was first chosen and exploited.

4.7.2

Molecular Approaches for Resistance Breeding in Forage Crops

There are potentially a lot of genes accessible for resistance engineering and breeding. Natural R genes, genes implicated in disease resistance response, PR proteins and other antimicrobial proteins, and pathogen-inducible promoters influence the specificity in host-pathogen recognition. Resistance to fungal and a viral disease has been acquired by transforming plants carrying the matching R gene with the pathogen Avr gene and manipulating its expression through a promoter that is inducible by a wide range of fungal infections (Melchers and Stuiver 2000). The use of transgenic plants produces genes that can inactivate pathogenicity factors like toxins (Manners and Dickman 1997). The strategy and processes will differ depending on the host species and pathogen involved and their interactions. Regardless of availability, adoption of genetic methods to promote disease resistance will be influenced by economic, social, and ethical considerations. The high research costs and intellectual property issues of enabling technologies and the use of molecular methods to commercially significant fodder crops are likely to be limited. An increasing number of transgenic agricultural products have fuelled controversy about environmental safety, as well as moral, ethical, and regulatory challenges surrounding GMOs (Flavell 2000). Rapid advances in molecular biological science, genetics, and information technology provide new opportunities for creating fresh methods to combat against diseases. However, since infections continue to outcompete previously effective resistant cultivars or new chemicals, disease control becomes a continual process that necessitates constant research to protect fodder crops against evasive diseases.

4.8

Conclusion

A well-balanced diet that includes green fodder, feeds, mineral mixture concentrate, and other supplements is critical for livestock nutrition. Climate change has been identified as a major threat to the forage production system. The response of different forage crops, grasses, and trees to climate change is complicated because of the interaction of climatic drivers like CO2 concentration, precipitation, and temperature with plant and management factors. The reduction in livestock production caused by diseases, dearth of forage, heat stress, and breeding strategies resulted in massive economic losses. Diseases in forage crops played an important role among these factors because they reduce the quality and quantity of forage crops and have a direct effect on the nutritious quality of the produce. Stand management practises that limit disease development and impact are essentially the same as those recommended in the absence of serious disease problems. Any practise that reduces crop stress (biotic

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or abiotic) and promotes vigour aids in extending the productive life of the stand. This is especially important in the presence of pathogenic organisms. Many of the early gains were made by relatively simple elimination of un-adapted germplasm and plants with low vigour or high disease susceptibility. Germplasm has become more elite and homogeneous as it has improved, challenging breeders to use more relevant plot methods, such as sward plots instead of spaced plants, and family-based selection methods that allow for increased replication or selection. As germplasm has improved, it has become more elite and more homogeneous, challenging breeders to use more relevant plot methods, such as sward plots instead of spaced plants, and family-based selection methods that allow for increased replication or selection intensity, as well as possible incorporation of genomic DNA marker technologies. The Department of Agriculture and Allied Sectors has been providing assistance to the State Governments for the control of animal diseases, scientific management and upgradation of genetic resources, increasing availability of nutritious feed and fodder, sustainable development of processing and marketing facilities, and enhancement of production and profitability of livestock.

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Effect of Nano-Priming on Maize Under Normal and Stressful Environment Sananda Mondal, Bandana Bose, and Debasish Panda

5.1

Introduction

Germination of seeds is the beginning phase of plants life. Specifically, in agricultural and rangeland ecosystems, successful seed germination is an important factor for the survival and conservation of plant species (Manjaiah et al. 2019). Efficient germination of seeds, rapid and homogeneous seedling emergence side by side successful establishment of seedling eventually leads to deep root system which is vital to increase the production of forage and medicinal plants in rangeland and agricultural fields (Azimi et al. 2014). In various parts of the world forage production is decreasing due to problems of germination, overgrazing, drought, and environmental stresses. In India, in the context of green fodder production, a huge deficit was observed between the demand and supply. In the present scenario, 63% deficiency of green fodder and 23.5% deficiency of dry fodder were observed in India. On the contrary, to feed the ever-increasing livestock population, the production and productivity of fodder need to be increased. The cereals and cash crop cultivation are increasing day by day and results the area under fodder cultivation is decreasing. Therefore, a tremendous pressure was faced to feed the livestock on the limited resources. In addition, green forages are rich and the cheapest source of carbohydrates, protein, minerals, and vitamins for dairy animals (Chaudhary et al. 2014). One of the most nutritious non-legume green fodders is maize. As a fodder maize is highly acceptable and can be judged based on the fact that it is free from any anti-nutritional components. Maize is easy and quick growing, high biomass yielding crop, and is highly palatable. It contains enough protein, minerals, and high S. Mondal (✉) · D. Panda Department of Crop Physiology, Institute of Agriculture, Visva-Bharati, Sriniketan, West Bengal, India B. Bose Department of Plant Physiology, Institute of Agricultural Sciences, Varanasi, Uttar Pradesh, India # The Author(s), under exclusive license to Springer Nature Singapore Pte Ltd. 2023 R. K. Singhal et al. (eds.), Molecular Interventions for Developing Climate-Smart Crops: A Forage Perspective, https://doi.org/10.1007/978-981-99-1858-4_5

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concentrations of soluble sugars in its green stage which possesses high digestibility as compared to other non-legume fodders, and it is also most suitable for preservation as silage (Chaudhary et al. 2014). Climate is changing day by day as drought and heat stress are increasing seriously which comes under abiotic stress factors and negatively affects the crop production. However, the environmental stresses like waterlogging, salinity, and heavy metal contamination have detrimental effect and it limits the crop production almost all over the world (Ye et al. 2019). Maize is considered as salt-sensitive crop plants having significant contribution with respect to food and forage worldwide (Farooq et al. 2015), whereas the productivity of maize is restricted by different environmental stresses like drought, floods, heavy metals, etc. but among all the stresses salinity stress is directly or indirectly linked with maize productivity. Increased soil salinity/ salt concentration leads to the significant decrease in the rate of seed germination as well as it delays the development of maize seedling (Zhang et al. 2007). Seed priming is one of the important seed invigoration technologies which results in quick and uniform germination, seedling emergence, and establishment that leads to improved growth of the plants (Ibrahim 2016; Abid et al. 2018; Thejeshwini et al. 2019). Seed priming has several beneficial effects, it ameliorates the germination, decreases the time of seedling emergence, an increment was observed in seedling stand establishment and overall growth of plants, survival, decreases the time to flower, produces strong and healthy plants, ameliorates the tolerance level toward various biotic and abiotic stresses (Mondal and Bose 2019). From the last two decades using nanoparticles as seed priming agent has been found to be very promising way out in comparison to usual seed priming methods for the purpose of germination and overall growth and development of the plants. Based on the problem that the production of forage crops is decreasing day by day due to low germination percentage, overgrazing, and environmental stresses, along with that maize is a major forage crop in terms of production and nutritional aspects. With these views the chapter accumulated the knowledge regarding the performance of nanomaterial primed maize under normal and stressful condition.

5.2

Nanoparticles

Nanoparticles (NPs) are generally microscopic, with a size range of 1–100 nm as recommended by Khan and Upadhyaya (2019). Mostly, NPs enter into the cell via aboveground organs like cuticle, epidermis, stoma, hydathodes, and other openings or by underground organs such as root tips, cortex, lateral roots, wounds, and other openings having various morphological and physiological effects on plants. However, these compound effects are based upon the plant species, growth period and condition of growing, method of application, and the required doses as well as the exposure time (Dietz and Herth 2011; Rizwan et al. 2017). The entry of nanoparticles within the plant roots takes place through osmotic pressure, capillary force, and cell wall pores via plasmodesmatal connections or in a symplastic way. Nanoparticles bind with various transporter proteins like ion

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channels, aquaporin, endocytosis, and sometimes it forms new pores and gets the entry into the plant cell. Once it enters the plant cell, it may be transported from one cell to another by following the route of plasmodesmata or via apoplastic or symplastic pathway (Usman et al. 2020). Based on the pore size of the cell wall the nanoparticles entry varies, the small size particle enters easily (Fleischer et al. 1999) as well as the larger nanoparticles pass via stomata, hydathode, and stigmas (Hossain et al. 2016). In case of seeds, nanoparticles enter via parenchymatous intercellular spaces of seed coat (Lee et al. 2010), whereas aquaporins play an important role in regulating the entry of nanoparticles in the seed coat (Abu-Hamdah et al. 2004).

5.3

Seed Priming with Nanoparticle

Seed priming has the capacity to alter the metabolism of seeds and signaling pathways, which influence their germination and growth; similar kind of effects was also noted in nano-priming. In agricultural sector, nano-priming has a significant impact toward agricultural sustainability (do Espirito Santo Pereira et al. 2021). Scientific studies demonstrated that application of nanoparticle can be able to stimulate the germination and growth of plants in several ways. The effectivity of nanoparticles is due to their small size and unique physio-chemical properties, which make them ideal seed priming agent (Mittal et al. 2020). It is denoted that nanomaterials have a versatile range of physio-chemical properties based on their size, shape, surface area, surface/volume ratio, chemical nature, particle charge, method of production, coating, and so on. A special feature of nanomaterials is that its high surface to mass ratio enables them to improve catalysis and supply materials and adsorb substances of interest. Similarly, it modulates the seed metabolism and helps by enhancing the water uptake, increase the starch hydrolysis rate, loosening the cell wall and weakening the endosperm, faster growth of embryo and development of root-shoot was observed. In addition, it also improves the seedling emergence, growth, production, and seed quality of crops (García-Gómez and Fernández 2019). Seed nano-priming can regulate the metabolism at cellular and molecular level, while plant interacts with the environment and maintains the sustainability in agriculture and natural ecosystems (Prasad et al. 2017; Chen 2018).

5.4

Nano-Priming in Relation with ROS

Studies depicted that, nano-priming promoted seed germination by developing nanopores in the seed coat, generating reactive oxygen species (ROS) in the seed, and using the nanocatalyst at the starch-degrading site activate the hydrolyzing enzyme activity (Mahakham et al. 2017). Although the exact mechanism of seed nano-priming is still not clear, it is assumed that it can induce physiological effects during the phase of seed germination. Due to formation of small pores in the seed coat which leads to increased water uptake, upregulate the gene expression of

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aquaporin and accumulate more ROS during germination in comparison to non-primed and other primed seeds (Kibinza et al. 2011). The antioxidant scavenging enzyme superoxide dismutase scavenges the O2_. and converts it into H2O2, another enzyme catalase transforms it into water. However, the nanoparticlemediated reduction of ROS and a significant increment in the amount of antioxidant enzyme were noted. By studying the interrelationship between ROS and antioxidant enzymes, nano-primed seeds contain high amount of antioxidant enzymes which have nano-priming mediated ROS mitigation mechanism. Furthermore, seed development has been also influenced by aquaporin and ROS-mediated interactions as reported by Maurel et al. (2015). Side by side the aquaporins and ROS can be also responsible for the activation of seed germination. Various studies depicted that aquaporins facilitate in the transfer process of H2O2 and ROS via biological membranes along with water uptake (Bienert and Chaumont 2014), whereas, in nano-priming process, nanopores are formed in the biological membrane which allowed the rapid influx of water into the seeds and activate the aquaporin genes. For seed germination and seedling emergence, seed’s antioxidant systems must activate first and regulate the ROS in order to trigger oxidative signal transduction mechanism. The process of nano-priming leads to increment in soluble sugar levels. Increased level of soluble sugar in the cell reduced the osmotic potential, which also decreases the water potential level. Due to the water potential gradient differences (gradient) between the outside and inside the cells increase which facilitates the movement of water into the seeds. Likewise, in nano-primed seeds increased level of α-amylase activity was observed and as a result soluble sugars content was also increased. In seeds, water uptake was increased due to changes in internal osmotic potential, and the reason behind this is increased level of soluble sugar content (solutes) in it. Whereas Müller et al. (2009) demonstrated that ROS, including OH, contribute to radiation growth, cellular reorganization, endosperm, and testicular wasting. In addition, Cinisli et al. (2019) depicted that the way nanoparticle are intake within the cells, the physiological and metabolic parameters like germination, antioxidant activity, macro and micro-nutrients, chlorophyll content, chloroplast number and photosynthesis in plants are varied Cinisli et al. (2019). According to Itroutwar et al. (2020) nano-priming with ZnONPs is considered relatively environment friendly and economically prudent among the other application mode. Various scientists throughout the world revealed that seed invigoration technologies have a novel impact toward precision farming which improves potentiality of the seeds to germinate in lesser time, high vigor and overall development, photosynthesis, and reproductive growth to ensure enhanced yield (Fig. 5.1) (Farooq et al. 2012, 2019; Chatterjee et al. 2018).

5.5

Maize Nano-Priming in Respect to Environmental Stresses

In the present scenario, to improve the seed quality and agricultural productivity, seed nano-priming is a new frontier (Mahakham et al. 2017; Acharya et al. 2020; Song and He 2021). In silver-nanopriming using silver nanoparticles (AgNPs) not

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Fig. 5.1 Nanopriming in plant growth development and stresses amelioration

only improves the performance of seed germination but also affects the continuous growth of the seedlings and plant defense mechanisms which showed resistance against biotic and abiotic stresses (Mahakham et al. 2017; Noshad et al. 2019; Prażak et al. 2020). For instance, previously, several reducing agents are used for the synthesis of NPs which are highly toxic, but it has been noted that some of the toxic reductants can be replaced by plant-based alternatives like extracts of kaffir lime leaf, turmeric oil, and onion peels. In addition, synthesis of plant-based NP creates lesser toxicity in comparison to chemically synthesized NPs and it is considered as a promising method because of its environmental friendliness and good biocompatibility (Mahakham et al. 2017; Acharya et al. 2020). Similarly, if maize (Zea mays L.) seeds were primed with 1000 mg/L mango peel nanoparticles (nMPs), it was noted that nMPs could mitigate the negative effect of salinity toward the performance of seed germination as well as significantly improved the germination percentage (Elkhatib et al. 2019). Whereas in maize seeds, application of titanium oxide nanoparticles (TiO2NPs) induced improvement of root-shoot lengths and fresh and dry weights of the seedlings was observed in both the cases like in normal as well as salinity stress conditions (Shah et al. 2021). In the same experiment, they primed the maize seeds with TiO2 nanoparticle and it was found more effective in promoting seedling growth than hydro-priming and other treatments. Due to this TiO2NPs treatments, it supported to increase the relative water content, cellular K+ concentration, total proline and phenolic contents and as a result the growth of the seedlings was improved. While using nano-ZnO material as seed polymer coating/priming treatment of fodder maize (var. J-1006) it was noted that it enhanced the vegetative growth, yield of the fodder, and the quality of fiber. Similarly, ZnONPs coating or priming treatments also proved to be better in terms of the improved availability of zinc

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micronutrient in fodder which cultivated under the field conditions as depicted by Tondey et al. (2021). Moreover, Naguib and Abdalla (2019) demonstrated that silica-nano-primed maize (Zea mays) seeds have a higher germination rate and seedling vigor index in comparison to non-primed one. The process of seed nano-priming helps to increase the antioxidant enzymes activity, which can have the capability to suppress the lipid peroxidation mechanism by suppressing the production of ROS under salinity stress condition. In addition, this process promotes the synthesis/activated the stored form of gibberellin in seeds as well as it reduced the abscisic acid content. As a result, such kind of hormonal balances activated the hydrolytic enzymes concentration like amylase and lipase. It was also noted that silica nano-priming increased the metabolic activity while exposed to saline condition of maize seeds. Choudhary et al. (2019) examined the zinc chitosan nanoparticle’s effect on maize plant as a priming agent side by side as foliar application. The outcome of an in vitro study depicted that seed nano-priming with Zn-chitosan nanoparticles improved the germination of seeds and inhibited the growth of fungi. These results depicted that Zn-chitosan nanoparticles have a strong fungicidal activity, which are effective micronutrient fortifier and can stimulate the growth of maize crop (Choudhary et al. 2019). Whereas Saharan et al. (2016) demonstrated that chitosan nanoparticles contain copper while applied to maize seeds (Zea mays L.) it enhanced the seed germination and the vigor index of seedlings. In corn (Zea mays), nano-priming with chitosan mitigates the deleterious effects of salinity (Oliveira et al., 2016). Moreover, Thongmak et al. (2022) examined the potentiality of plasma activated water priming, silver-nanopriming, and green silvernanopriming on the performance of aged and non-aged maize seeds. By applying these priming technologies, it improved the germination performance and seed vigor of new maize seeds.

5.6

Conclusion

This chapter incorporates the background and characteristics of nanoparticles, the beneficial role of seed priming including nano-priming, relationship of nano-priming with ROS, and the effect of nano-priming in forage maize under normal and stressful condition. But very less data is available regarding forage maize with respect to the role of nano-priming. For the enrichment of database further advanced morphophysiological, biochemical, and molecular studies are required in this aspect. Then the required amount of forage production will be increased side by side, nanopriming also mitigates the problem of poor germination under normal and environmental hazardous condition.

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Oxidative Stress and Antioxidant Defense in Mitigating Abiotic Stresses in Forage Crops: A Physiological and Biochemical Perspective Meenakshi Goyal, Archana Kumari, Ankita Kumari, Himanshu Sharma, Pashupat Vasmatkar, and Namrata Gupta

6.1

Introduction

Abiotic variables such as extreme temperatures, drought, flooding, salt, and heavy metal stress are the main determinants which have an impact on the development of crop plants’ yields (Canter 2018; Zörb et al. 2019). Above mentioned stresses affect approximately 90% of arable lands and can result in yield losses of up to 70% for main food crops (Dos Reis et al. 2012; Mantri et al. 2012). According to previous estimates based on the integration of crop yield and climate change models that abiotic stresses have a detrimental impact on livestock sustainability, national economies, food security, and the livelihoods of farmers and their families (Tigchelaar et al. 2018). Forages are plants and the components of those plants are used for the grazing purpose of livestock. They play an important part in the nutrition of ruminants. In addition to being the main source of nutrition for both domesticated and wild animals, forages also benefit human civilization in various ways, such as soil protection through crop over and fertility improvement through the addition of organic matter. The availability of nutritious food for domestic animals as well as forage production is anticipated to be impacted by climate change. Forage crop M. Goyal (✉) Department of Plant Breeding and Genetics, Punjab Agricultural University, Ludhiana, Punjab, India e-mail: [email protected] A. Kumari · H. Sharma · P. Vasmatkar Department of Biochemistry, Punjab Agricultural University, Ludhiana, Punjab, India A. Kumari National Dairy Research Institute, Karnal, Haryana, India N. Gupta Department of Botany, Punjab Agricultural University, Ludhiana, Punjab, India # The Author(s), under exclusive license to Springer Nature Singapore Pte Ltd. 2023 R. K. Singhal et al. (eds.), Molecular Interventions for Developing Climate-Smart Crops: A Forage Perspective, https://doi.org/10.1007/978-981-99-1858-4_6

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production is largely affected by abiotic factors related stresses such as drought, salinity, heat, temperature, cold stress, heavy metal stress, etc. Extreme weather events and abiotic stress are expected to rise as a result of climate change. There has been a concerted effort to find ways to improve crop tolerance and lessen the impact of abiotic stresses on agricultural yield. Numerous crop features that promote abiotic stress tolerance are the consequence of the interaction of multiple genes, making them challenging to study and alter. Additionally, various stress events may result in protein denaturation, osmotic stress, or both, which trigger cellular adaptation responses like the build-up of osmolytes, the activation of stress proteins, and the acceleration of ROS scavenging mechanisms. Tolerance to a variety of future abiotic stress events, such as priming, acclimatization, conditioning, hardening, or cross-stress tolerance, can develop as a result of exposure to a stress factor. There is need to adopt various conventional and genetic approaches to improve stress tolerance of forage crops. Some early successes in applying genetic manipulation methods to promote abiotic stress tolerance in grasses may also be supported by physiological understanding (Tester and Bacic 2005).

6.2

Different Types of Stress in Forage Crops

6.2.1

Heat and Temperature Stress

High temperatures and heat limit the growth and development of plants. High temperature is a powerful moderator of plant growth because it changes the plants’ qualitative and quantitative traits, which in turn impacts their primary and secondary metabolic pathways (Wahid et al. 2007). Modulation in metabolic processes eventually result in the synthesis of new metabolites or altering the concentrations of already present metabolites (Wahid et al. 2007). The ability of plants to tolerate stress depends on a few specific metabolites but changes to the metabolic pathways typically result in forage with varying nutritional value (Anuraga et al. 1993; Mitchell et al. 2001; Grassmann et al. 2002; Norman et al. 2004, Vasconsuelo and Boland 2007; Edreva et al. 2008). For example, high levels of glycine betaine accumulation were reported in maize and sugarcane in response to high temperature, while in contrast, plant species such as rice, mustard, Arabidopsis, and tobacco naturally do not produce glycine betaine under stress conditions (Quan et al. 2004; Wahid and Close 2007). Numerous studies have categorically stated that high temperature stress enhances the synthesis of condensed tannins, which may improve the nutritional content of fodder (Lees et al. 1994; Gebrehiwot et al. 2002). The right amounts of proteins, carbohydrates, fatty acids, and other important elements must also be present in the feed or forage. Heat stress also induces early abortion of tapetal cells, which cause the pollen mother cells to rapidly progress toward meiotic prophase and finally undergo programmed cell death (PCD), thus leading to pollen sterility (Oshino et al. 2007; Sakata and Higashitani 2008; Parish et al. 2012). In sorghum, heat stress reduces the accumulation of carbohydrate in pollen grains and

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ATP in the stigmatic tissue (Jain et al. 2007). Heat stress induces changes in respiration and photosynthesis and thus leads to a shortened life cycle and diminished plant productivity (Barnabás et al. 2008). The early effects of thermal stress comprise structural alterations in chloroplast protein complexes and reduced activity of enzymes (Ahmad et al. 2010). In addition, by causing injuries to the cell membrane, organization of microtubules, and ultimately to the cytoskeleton, heat stress changes membrane permeability and alters cell differentiation, elongation, and expansion (Smertenko et al. 1997; Potters et al. 2008; Rasheed 2009).

6.2.2

Drought Stress

A form of atmospheric anomaly brought on by a lack of rain or recurrently smallerthan-average rainfall, drought is an extreme weather event (Paulo and Pereira 2006). It happens in places that receive both a lot and little rain. It is a complicated phenomenon and it happens frequently with varying intensities in different seasons. According to Łabędzki (2016), a drought in agriculture is a persistent scarcity of soil water in a specific location that affects a particular plant species or cultivar over an extended period of time. Crop yield is decreased and the environment for growth and development is deteriorated. The growth, development, and most importantly, the crop yields of plants are all greatly hampered by the lack of water. The initial effect of water stress is a slowdown of development and cell division. A prolonged stress can cause disruptions in plant metabolism, especially in the plant’s capacity for photosynthetic activity which is most likely due to a drop in Rubisco activity, stomatal conductance, and availability of CO2 (Jones 1998; Kalaji and Oboda 2009; Hura et al. 2007, 2010).

6.2.3

Cold Stress

Cold stress (CS) affects the survivability, geographical distribution, and yield stability of crops. Suitable management and agronomic practices can minimize the crop losses associated with cooler environments. However, agronomic practices alone cannot support plants adequately to withstand the harsh cold. By decreasing the export of photosynthates from the chloroplasts, low temperature stress has a great impact on photosynthesis (Kratsch and Wise 2000). It is primarily caused by the loss of membrane polyps which decrease electron transport and phosphorylation (Bertamini et al. 2007; Rapacz 2007). A previous study report that the at -2.7, electric conductivity of pearl millet, marandu grass, and sorghum increased. Therefore, cold stress damaged the cell membranes of the cold-sensitive species and, thereby, caused electrolyte leaching (Jatimliansky et al. 2004). This occurs because low temperatures cause fatty acids to become unsaturated and, thereby, alter the structure and fluidity of the membrane layer of lipids and proteins (Hasanuzzaman et al. 2013).

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Different crops widely differ in their sensitivity toward cold. Cereals germinate better at low temperature than many other crops (Coffman 1923). In a previous study, corn and soybean cultivars varied significantly in their susceptibility to low temperature—some show low germination after exposure to 17 °C, others were not injured even at 7 °C (Miedema 1982). A number of morphological and ultrastructural lesions were observed in meristematic cells of the primary root in particular, involving destruction of Golgi apparatus and endoplasmic reticulum. The early vegetative stage of growth is very vulnerable to low temperature. The frosts influence the aerial parts of plant by reducing the growth and establishment of photosynthetic area. Generally, low temperature reduces shoot growth more than root growth (Miedema 1982).

6.2.4

Salinity Stress

Soil salinity is also a prevalent stress that alters geographical distribution of plants that is a global land degradation issue (Sadiq et al. 2020). Salt is accumulated in agricultural soils as a result of various environmental factors such as climate change, excessive use of groundwater, poor drainage associated with massive irrigation and intensive farming, and the use of low quality water in irrigation (Machado and Serralheiro 2017). Several countries such as the USA, Spain, Jordan, China, Greece, and Senegal have been using various techniques to tackle soil salinity on their farm lands (Kumar and Trivedi 2018). Excess salt in the soil may adversely affect plant growth either through osmotic inhibition of water uptake by roots or specific ion effects (Khan et al. 2015). Major adverse effects of salinity stress include increased ion-toxicity, osmotic stress, nutritional acquisition, homeostasis, impaired stomatal conductance, increased cell-turgor loss, reduction in leaf water potential, and altered physiological/biochemical processes.

6.3

Physiological Aspect of Abiotic Stress in Grasses

The major shift in carbon dioxide concentration and other greenhouse gases due to human activities is the driving force behind the climate change. Due to gradual change in temperature and other environmental conditions the production of fodder crops has significantly affected. These factors cause abiotic stresses in grasses and other fodder crops. The increase in CO2 and accompanying factors shifts the adaptive response of grasses (Abberton et al. 2008). These conditions can increase or decrease the productivity (Lobell and Gourdji 2012). Combination of different factors with abiotic stresses causes profound consequences on fodder crops (Mittler 2006; Rhizsky et al. 2002). Physiologically, major fodder crops are categorized as C3 species as they follow the C3 photosynthetic pathway and these pathways work well with optimum temperature for root and shoot growth (Beard 1973; Turgeon 2008). The heat wave during the cropping season has an adverse effect on plant growth. These factors directly affect the hydrological cycle due to alteration in

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precipitation patterns of plants. In addition to this, injuries in plant tissues are anticipated due to increase in the ozone escalation caused by environmental pollutants that substantially decrease plant productivity (IPCC 2014). The adverse conditions primarily affect the photosynthesis and stomatal conductance rates in grasses (Long et al. 2004) and stomatal conductance is also affected by the alteration in CO2 concentrations in plants (Farfan-Vignolo and Asard, 2012; Burgess and Huang 2014; Song et al. 2014). This change is caused due to the change in stomatal aperture and stomatal density (Lammertsma et al. 2011). This further inhibits the photorespiration which is accompanied by the increased carboxylation and electron transport rates (Reddy et al. 2010). These conditions have affected the photosynthetic rates in ryegrass, Kentucky bluegrass, and tall fescue (Song et al. 2014; Yu et al. 2012). Burgess and Huang (2014) have reported the increase in the photosynthesis in the creeping bentgrass with increasing CO2 concentration. This is due to the higher availability of substrates for the rubisco which improves the activation state of enzyme. Such conditions did not affect the efficiency of photosynthesis II in ryegrass (Farfan-Vignolo and Asard 2012). The increase in temperature due to heat stress causes increase in threshold that led to the damage of plant metabolism and productivity (Porter 2005). Among the grasses, Kentucky bluegrass and creeping bentgrass are susceptible to heat stress. Stomatal conductance rates were increased in perennial ryegrass with increase in temperature (Farfan-Vignolo and Asard 2012). Heat stress causes the reduction in photosynthesis due to deactivation of rubisco in ryegrass. It also negatively affects the chlorophyll and carotenoid content. Chlorophyll degradation was explained by Jespersen et al. (2016) in bentgrass. Heat stress also affects the ratio of Chl a/b in tall fescue which shows differential effects in plants. This further causes the alterations in the light harvesting complex of PSII to reduce the photo-oxidative damage (Spundova et al. 2003). Decrease in photosynthetic rate led to the alteration in carbohydrate accumulation in plants. During initial stress, glucose, fructose, and galactose content increase in leaves of bluegrass (Du et al. 2013). Studies also suggested the upregulation of sucrose synthase to maintain the positive carbon balance under heat stress. Water stress is the major abiotic factor in plant growth and productivity around the world. This causes the reduction in the water potential and turgor pressure which ultimately affects various functions in plants. However, severity of drought stress depends on the severity and duration of stress. Drought escape, avoidance, and drought tolerance are strategies adopted by plants to handle the stressed conditions. Mediterranean fescue moderates its growth and development according to the availability of water (Humphreys et al. 2005). Rhizomes of bluegrass play an important role in plant growth during limited water supply. During drought avoidance plants maintain the high tissue water potential under limited water supply. This further helps in accumulation of variable solutes such as carbohydrates, ions to maintain osmotic balance of plant cells.

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Biochemical Effects of Abiotic Stress

Reductions in carbon assimilation during drought stress result in an imbalance between electron excitation and utilization through photosynthesis, leading to the generation of ROS. Several biochemical parameters have been established for stress tolerance in plants based on osmolytes accumulation, ROS production, and antioxidant enzyme activities (Swathi et al. 2017). ROS that are produced in response to various biotic and abiotic stresses act as messenger to activate defense mechanisms in plant. Enzymatic antioxidants comprise catalase (CAT), superoxide dismutase (SOD), peroxidase (POX), ascorbate peroxidase (APX), dehydroascorbate reductase (DHAR), glutathione reductase (GR), and monodehydroascorbate reductase (MDHAR) and non-enzymatic antioxidants include glutathione, ascorbate, tocopherols, carotenoids, phenolics, and ascorbic acid (Sahitya et al. 2018).

6.4.1

Osmolytes

Osmolytes or compatible solutes are small molecules having low molecular weight that are electrically neutral, highly soluble, and non-toxic at molar concentrations that enhance the cell potential to maintain turgor potential without hampering the normal physiological processes. Thus, damaging effects of drought are minimized by accumulation of solutes in cellular cytoplasm and vacuole (Taiz and Zeiger 2006). Various osmoprotectants identified in plants can be grouped as quaternary ammonium compounds such as polyamines, glycine betaine, β-alanine betaine, dimethyl-sulfoniopropionate, and choline-O-sulfate, sugars and sugar alcohols including trehalose, fructans, mannitol, and D-ononitol and sorbitol and amino acids such as proline and ectoine (Sharma et al. 2019). These osmolytes get accumulated under drought stress and confer tolerance to cell without interfering with the cellular machinery of the plant (Anjum et al. 2017).

6.4.2

Reactive Oxygen Species (ROS) and Other Stress Markers

The ROS mainly damage cell membranes, proteins, and nucleic acids, causing oxidative stress. The intercellular concentration of malondialdehyde (MDA) indicates the extent of oxidative stress Abiotic stress induces imbalances in the redox cell homeostasis due to the accumulation of ROS. Different ROS are produced by the unavoidable leakage of electrons on to O2 from the electron transport systems of chloroplasts, mitochondria, and plasma membranes or as a consequence of various metabolic pathways (Foyer 1997). Drought stress causes oxidative stress by decreasing stomatal conductivity that reduces the leaf internal CO2, leading to increased photorespiration and the formation of ROS which includes superoxide anion (O2•-), hydroxyl radical (•OH), hydrogen peroxide (H2O2), and singlet oxygen (1O2) (Table 6.1).

CM, Chl, Mt.

1 nm

1 μm

30 nm

1 ms

1– 4 μs

CM, Chl, Nu

Sources Apoplast, CM, Chl, Mt., Per CM, Chl, Mt

MD 30 nm

t1/2 1– 4 μs 1 μs

Oxidizes proteins, PUFAs, DNA

Reacts with proteins and forms •OH via O•-

Mode of action Reacts with double bond containing compounds such as (Fe-S) proteins Extremely reactive with biomolecules

G

No

Rapid

W, H, Y, M, C

C

Rapid

Reaction with RNA Protein No Rapid

PUFA

Low

Rapid

DNA Low

Carotenoids, α-Tocopherol

Flavonoids, proline, sugars, AsA APX, CAT, GPX, PER, PRX, AsA, GSH

Scavenging systems Sod, Flavonoids, AsA

CM cell membranes, Chl chloroplast, Mt mitochondria, Nu nuclei, Per peroxisomes, MD migration distance, t1/2-half-life, APX ascorbate peroxidase, CAT catalase, GPX glutathione peroxidase, PER peroxidase, PRX peroxiredoxin, SOD superoxide dismutase

ROS Superoxide (O2•-) Hydroxyl radical (•OH) Hydrogen peroxide (H2O2) Singlet oxygen (1O2)

Table 6.1 Properties of different reactive oxygen species (ROS) produced during drought stress

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Fig. 6.1 Structure of different ROS

Generation of ROS is a fundamental process in higher plants to transmit cellular signaling information in response to the changing environmental conditions. Production and structure of different ROS are shown in Fig. 6.1. Reactive 1O2 can be efficiently quenched by small lipophilic compounds such as β-carotene and tocopherol or by a scavenging action of D1 protein present in photosystem II (KriegerLiszkay 2005). However, minor portion of 1O2 is able to diffuse to a smaller distance, where it can react with diverse biomolecules and potentially mediate it signaling pathways (Dogra et al. 2018). Reactive 1O2 can readily induce oxidative modifications in various biomolecules, thereby altering their function permanently (Table 6.2).

6.4.2.1 Superoxide Anion Radical (O2• -) The superoxide radical (O2•-) is formed mainly in the thylakoid-localized PSI during non-cyclic electron transport. It is mediated by NADPH oxidases, belonging to the respiratory burst oxidase homolog (RBOH) family (Sang and Macho 2017). Usually, H2O2 is generated when cytochrome c oxidase interacts with O2, sometimes it reacts with the different components of ETC to give O2•-. Reactive O2•- often undergoes further reactions to generate 1O2, H2O2, and •OH (Janku et al. 2019). The reactions through which O2•- and iron rapidly generate •OH are shown as follows: O2• - þ Fe3þ → 1 O2 þ Fe2þ O2• - þ 2Hþ → O2 þ H2 O2 Fe3þ

ðHaber–Weiss reactionÞ

Fe2þ þ H2 O2 þ Fe3þ → Fe3þ þ OH - þ • OH

ðFenton reactionÞ:

This is mainly due to spin restriction in O2 which cannot accept one electron at a time and hence during reduction of O2 stable intermediates are formed in the

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Table 6.2 Biomolecules oxidized by 1O2 and their corresponding oxidation products Compounds/ biomolecules Linoleic acid/ linolenic acid β-Carotene α-Tocopherol Ascorbate Cysteine Tryptophan

Glutathione DNA

Type of reaction Ene reaction

Oxidized product Allylic hydroperoxides (10- and 12-HOD and 10- and 15-HOT) Endoperoxides (β-CC, β-I, dhA)

Diels–Alder reaction 1,4cycloaddition Electron transfer reaction Direct oxidation

References Mueller et al. (2006) Ramel et al. (2012) Kim et al. (2006) Kramarenko et al. (2006) Buettner and Hall (1987) Dreaden et al. (2012, 2011)

Di-epoxides (tocoquinone and tocopherol) Dehydroascorbic acid (DHA) and H2O2 Cysteine disulfide

1,2cycloaddition reaction Direct oxidation

Dioxetanes (N-formyl kynurenine, kynurenine)

Diels–Alder reaction

Endoperoxide/deoxyguanosine, 8-oxo-dG and 4-OH-8-oxo-dG

Glutathione disulfide

Devasagayam et al. (1991) Cadet et al. (1994)

stepwise fashion. O2•- is the primary ROS formed in the cell. A way to overcome spin restriction, O2 molecule interacts with another paramagnetic center. Transition metals like Fe or Cu frequently have unpaired electrons. O2

e-

-

-

-

→ O.2- → e2Hþ H2 O2 → eHþ H2 O þ .OH - → eHþ H2 O

6.4.2.2 Hydrogen Peroxide (H2O2) Wide range of stressful conditions such as drought, chilling, UV irradiation, exposure to intense light and wounding induce oxidative stress by increasing the levels of H2O2 (Molassiotis et al. 2016). Electron transport system of chloroplast, mitochondria, and processes such as β-oxidation of fatty acid and photorespiration are major sources of H2O2 production in plant cells. It is also produced in tissues where it is required as a substrate for biosynthesis of lignin and suberin. O.2- þ 2Hþ

SOD

→ H2 O2 þ O2

Several recent studies have demonstrated that H2O2 is involved in stress signaling pathways, which can activate multiple responses that strengthen resistance to various biotic and abiotic stresses (Dang et al. 2019). H2O2 can oxidize the protein kinases, phosphatases, enzymes of Calvin cycle, and transcription factors containing thiol groups. H2O2 also acts as a regulator of numerous physiological processes like phytoalexin production, cell wall

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strengthening, photosynthesis, senescence, stomatal opening, and the cell cycle (Petrov and Breusegem 2012).

6.4.2.3 Hydroxyl Radical (•OH) With a single unpaired electron, •OH is the most reactive ROS, capable of cellular damages, including changes in protein structures, lipid peroxidation, and membrane destruction (Demidchik 2015). However, it can also facilitate seed germination, growth, stomatal closure, reproduction, plant cell death, immune responses, and adaptations to stress conditions (Richards et al. 2015). The formation of •OH is dependent on both H2O2 and Fenton reaction and thus its formation is subject to inhibition by both SOD and CAT. Highly reactive •OH promotes oxidative cleavage of pectins and xyloglucans and thus causing cell elongation by loosening the cell wall. Drought stress increases Fe and Cu availability for Fenton reactions and could cause increased •OH production (Moran et al. 1994).

6.4.3

Nitric Oxide (NO)

NO possesses antioxidant properties and plays an important role in resistance to salt, drought, temperature (high and low), UV-B, and heavy metal stress (Begum et al. 2019). NO also reacts with the superoxide radical to form ONOO-. NO plays an important role in cytoprotection by regulating the level and toxicity of ROS (Palmieri et al. 2008). Nitric oxide (NO) is a ubiquitous gaso-transmitter involved in redox homeostasis that regulates plant life cycle, its growth and development, and its biotic and abiotic stress tolerance mainly by post-translational protein modifications (Sanchez-Vicente et al. 2019). Consisting of S-nitrosation of cysteine residues and metals, and nitration of tyrosine residue (Table 6.3). NO promotes the transformation of O2- to H2O2 and O2 and thus inhibits the plants from oxidative damage (Zheng et al. 2009). NO is also an important player inside the network required for stomatal closure since nitrate reductase and NOS-like activities linked to NO production are mandatory for the ABA signal transduction cascade in guard cells to close the stomata. Many studies have reported about the role of NO during plant growth and development under drought stress in many crops. Furthermore, NO has emerged as a key element in drought tolerance of several plant species mainly by enhancing the antioxidant systems, proline, ROS, and osmolytes metabolism (Fig. 6.2) (Filippou et al. 2014; Fancy et al. 2017).

6.4.4

Malondialdehyde (MDA)

Membrane lipid peroxidation in plants is detected by measuring the content of malondialdehyde (MDA). The content of MDA content was increased significantly in cold, salt, drought stress, and disease (Laxa et al. 2019) resulting in decreased membrane fluidity and destroyed ion homeostasis of plants (Huang et al. 2019). However, the breakage of membranes because of lipid peroxidation affects

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Table 6.3 Target and effects of protein nitration described in plants Target protein Catalase Glyceraldehyde-3phosphate dehydrogenase Complexes of PSI and PSII Ferredoxin-NADP oxidoreductase O-Acetylserine (thiol) lyase 1 Glutamine synthetase NADP-isocitrate dehydrogenase NADHhydroxypyruvate reductase Ascorbate peroxidase

Monodehydroascorbate reductase Superoxide dismutases

Mechanism Inhibition of activity against pathogens Inhibition of activity

Reference Clark et al. (2000) Lozano-Juste and León (2011)

Inactivation and disassembly of complexes dependent on light conditions Inhibition of activity, causing changes in photosynthetic activity Inhibition of activity under stress conditions to regulate cysteine and glutathione metabolism Inhibition of activity to regulate N metabolism in nodules Inhibition of activity for the reprogramming of metabolism and redox homeostasis during senescence Inhibition of activity, changes in peroxisomal metabolism

Galetskiy et al. (2011)

Inhibition of activity

Clark et al. (2000) and Begara-Morales et al. (2014) Begara-Morales et al. (2014) Holzmeister et al. (2015)

Inhibition of activity Inhibition of activity

Chaki et al. (2011) Alvarez et al. (2011)

Melo et al. (2011) Begara-Morales et al. (2013) Corpas et al. (2013)

polyunsaturated fatty acids (PUFAs) and leads to the production of cytotoxic lipid aldehydes such as MDA (Fig. 6.3). These lipid peroxidation products can easily diffuse across membranes and cause further cellular damage by reacting with other lipids, proteins, and nucleic acids far from their site of origin (Ayala et al. 2014). The compounds resulting from lipid peroxidation mostly react with DNA showing both genotoxic and mutagenic action. Among them MDA is the most mutagenic (Zarkovic et al. 2013). The effect of lipid peroxidation is also the loss of membrane integrity by alteration of its fluidity and inactivation of membrane proteins and ion channels, which make it permeable to substances that do not normally cross it. Several studies have investigated MDA of plants under different stress conditions (Jbir-Koubaa et al. 2015).

6.4.5

Antioxidant Enzymes

The upregulation of antioxidant enzymes represents an important marker for drought stress (Laxa et al. 2019). In order to cope with the oxidative stress, plants usually rely on the antioxidant defense enzymes such as SOD, CAT, and POD. These either

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Fig. 6.2 Cellular localization and source of ROS in plant cell

Fig. 6.3 Synthesis of MDA in plants

directly scavenge the ROS or protect plants indirectly by managing non-enzymatic defense (Anjum et al. 2011).

6.4.5.1 Superoxide Dismutase Superoxide dismutase (SOD) is the first detoxification enzyme that acts as a component of first line defense system against reactive oxygen species (Ighodaroab and Akinloye 2018). SODs are a class of metalloenzymes that catalyze the dismutation of two molecules of O2●- into O2 and H2O2. In plants, there are three main groups of SODs: Cu, Zn-SODs, Mn-SODs, and Fe-SODs. Among these, Cu/Zn SOD, the most important in oxidative stress is composed of two subunits combined with Cu and Zn atoms, respectively. The activation of SOD isoforms to counteract O2●accumulation in diverse cell compartments under drought stressed Arabidopsis (Jung 2004) and wheat (Cheng et al. 2016) has been reported. SOD removes O2●-

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Table 6.4 Types of SOD isoforms present in plant, their subcellular location, and sensitivity SOD isozymes Cu/ZnSOD MnSOD Fe-SOD Ni-SOD

Structure Subcellular localization Homodimeric and Cytosol, chloroplast, peroxisome, mitochondria homotetrameric Homodimeric and Mitochondria, peroxisome homotetrameric Homodimeric and Cytosol, chloroplast, tetrameric peroxisome, mitochondria Only reported in prokaryotes

Sensitivity H2O2 and KCN CHCl3:CH3CH2OH but not to H2O2 and KCN H2O2 but not to KCN

and hence decreases the risk of .OH formation via the metal catalyzed Haber Weisstype reaction (Gill and Tuteja 2010). SODs are encoded by nuclear genes and targeted to their respective subcellular localization. They also show different sensitivity toward the inhibitors (Table 6.4). Overexpression of Cu/Zn SOD in tobacco (Zhang et al. 2017), sweet potato (Lu et al. 2010), rice (Prashanth et al. 2008), Arabidopsis (Wu et al. 2016), and of Mn-SOD in alfalfa improved the capacity of drought tolerance and recovery in plants.

6.4.5.2 Catalase Catalase (CAT), also known as a H2O2 oxidoreductase, is a heme-containing enzyme that catalyzes the dismutation of H2O2 into H2O and O2 mainly in peroxisomes (Vellosillo et al. 2010). Numerous studies demonstrated that expression of abundance of CATs increased under various stresses for scavenging of ROS (Gupta et al. 2018). The features that distinguish CAT from other H2O2 metabolizing enzymes such as ascorbate peroxidases (APX), peroxiredoxins (PRX), glutathione/ thioredoxin peroxidases (GPX), and glutathione S-transferases (GST) are that CAT does not require a reductant. It has been reported that there is a concomitant increase in catalase activity in plants in response to the different stresses (Ma et al. 2017). CAT is highly sensitive to light and has a rapid turnover rate which may be the result of light absorption by the heme group. Various researchers investigated the role of CAT in pathogen defense by either overexpressing or suppressing CAT in transgenic plants (Vandenabeele et al. 2004). Increase in CAT activity is supposed to be an adaptive trait possibly helping to overcome the damage to tissue metabolism by reducing toxic levels of H2O2. 6.4.5.3 Peroxidase Peroxidases (POXs) are oxidoreductases that transform a variety of compounds via a free radical mechanism into oxidized or polymerized products. In plants, the level of ROS significantly increases under drought conditions. The enzymatic antioxidant system such as SOD, CAT, and POXs such as APX, GPX, and guaiacol peroxidase protect the cells against damaging ROS (Manavalan and Nguyen 2017). These POXs scavenge the H2O2 produced during oxidative stress. POXs are reported from all plants, animals, and microbes and are essential for living systems. Many

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peroxidases contain an iron-porphyrin derivative (heme) in their active site. POXs are involved in many physiological and biochemical processes, including the crosslinking of molecules in the cell wall, lignin, and suberin formation by oxidation of cinnamyl alcohol (Siegel 2003).

6.4.5.4 Ascorbate Peroxidase (APX) Ascorbic acid-glutathione (AsA-GSH) cycle is essential metabolic pathway for protection against ROS and regulation of the cellular level of H2O2 in plants. This pathway includes ascorbate peroxidase (APX), dehydroascorbate reductase (DHAR), and glutathione reductase (GR), monodehydroascorbate reductase (MDHAR) with antioxidant metabolites ascorbic acid (AsA), and reduced glutathione (GSH). It plays an important role in maintaining redox homeostasis in plants to protect them from oxidation damage (Foyer and Noctor 2016). Ascorbate peroxidase (APX) belongs to the family of heme-containing peroxidases which has high affinity for H2O2 and with AsA as an e- donor, it reduces H2O2 to H2O in chloroplasts, cytosol, mitochondria, and peroxisomes. At least five different isoforms, including cytosolic forms (cAPX), peroxisomal forms (prAPX), chloroplast stromal soluble forms (sAPX), and thylakoid (tAPX) belong to the APX family. All these isoforms originate from alternative splicing, which contributes to the differential regulation of expression of various isoforms (Caverzan et al. 2012) (Table 6.5). Increased activity of APX was reported under drought stress in maize (Jiang and Zhang 2001) and soybean (Heerden and Kruger 2002). APX has a higher affinity for H2O2 than that of CAT and POD and it may have a more crucial role in the management of ROS during stress. 6.4.5.5 Glutathione Reductase (GR) Glutathione reductase (GR), also known as glutathione disulfide reductase (GSR), is a flavoprotein belonging to the family of NADPH-dependent oxidoreductases. It catalyzes the reduction of GSSG to GSH and plays an important role in cell defense

Table 6.5 Localization and role of APXs isoforms in different stresses APX form Cytosolic (cAPX)

Peroxisomal (prAPX) Stromal (sAPX) Thylakoid (tAPX)

Stress Salt/ chilling/ drought/ heat Heat/ drought/ salt Heat/salt

Role Cellular response to oxidative stress, ROS salinity stress tolerance, pathogen attack

Plant Tomato, rice Arabidopsis

References Qian et al. (2014)

Salinity and drought tolerance

Barley, tobacco

Li et al. (2009)

Response to salinity stress

Arabidopsis

Salt/ chilling/ high light

Involve in water–water cycle

Tobacco, tomato, wheat

Hirooka et al. (2005) Liu et al. (2013)

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against ROS (Hernandez et al. 2017). GR has been mainly localized in chloroplasts, mitochondria, and the cytosol but also found in peroxisomes (Kataya and Reumann 2010). An increase in GR activity in plants results in the accumulation of GSH and ultimately confers stress tolerance in plants. Various studies have demonstrated that drought stress increased GR activity in tobacco, barley maize wheat, rice, and pea (Sharma and Dubey 2005a).

6.4.5.6 Monodehydroascorbate Reductase (MDHAR) Monodehydroascorbate reductase (MDHAR) is one of the key enzymes in the conversion of oxidized ascorbate back to reduced ascorbate in plants. MDHAR is present across the entire plant kingdom and is localized in chloroplasts, mitochondria, peroxisomes, and cytosol (Kavitha et al. 2010). In higher plants MDHARs belong to a multigene family constituting several subcellular isoforms. The exposure of plants to environmental stress conditions like high light leads to very quick oxidation of AsA to MDHA in chloroplast. Reddy et al. (2004) found that the enzyme activity of MDHAR was significantly high in the water-stressed leaves of mulberry cultivars, while contrary to this finding. Lu et al. (2007) reported that the activity of MDHAR decreased significantly in the heat and drought treatments in Crofton weed. MDHAR activity was enhanced under various other stresses such as salt, high light, and UV radiations. Sharma and Dubey (2005b) reported that the activities of MDHAR, DHAR, and GR were higher in drought stressed rice seedlings. 6.4.5.7 Dehydroascorbate Reductase (DHAR) Dehydroascorbate reductase (DHAR) is a monomeric enzyme, which is a member of the GSHS-transferase superfamily which maintains AsA in its reduced form by catalyzing the reduction of dehydroascorbate (DHA) to AsA using GSH as a reducing substrate (Hasanuzzaman et al. 2019). It is present in various plant tissues and its modulation activity has been reported in various plant species. AsA recycling through MDHAR or DHAR is critical for maintaining the AsA level and redox state in the adaptation of plants to environmental conditions. Knockout mutants of Arabidopsis DHAR were unable to show any significant differences in total AsA content until challenged with the abiotic stress (Das et al. 2016), which showed the importance of DHAR in reducing the DHA under stress conditions. Overproduction of DHAR during stress conditions not only allows the plant to recycle DHA but also increases the AsA pool in cell. 6.4.5.8 Glutathione S-Transferase (GST) Glutathione S-transferases (GSTs) are multigene superfamily with cytosolic, mitochondrial, and microsomal localization in plants (Kumar and Trivedi 2018). Based on their degree of sequence identity, GSTs have been divided into several classes, i.e., tau, lambda, phi, dehydroascorbate reductase (DHAR), theta, zeta, tetrachlorohydroquinone dehalogenase (TCHQD), and elongation factor1 gamma. Among these, four classes, phi, tau, lambda, and DHAR, are plant specific (Edwards and Dixon 2005). In plants, GSTs have been exhaustively studied in terms of

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herbicide detoxification and specific members of this family have been reported to provide tolerance toward herbicide in many crops (Chronopoulou et al. 2017). GSTs are involved in safeguard of the cells against biotic and abiotic stresses and provide tolerance by catalyzing S-conjugation between the thiol group of GSH and electrophilic moiety in the hydrophobic and toxic substrate (Nianiou-Obeidat et al. 2017). The conjugate formed is either sequestered in the vacuoles or are exported from the cells by transport systems. Enzymatic activity of GSTs is induced by various fungal elicitors, wounding as well as by cold, drought, oxidative stress, heavy metals, and salt (Srivastava et al. 2019). Drought tolerant and sensitive sorghum genotypes showed efficient H2O2 scavenging mechanisms with significantly higher activities of GSTs (Jogeswar et al. 2006).

6.4.6

Antioxidants

Antioxidants play a key role in the detoxification of ROS induced by drought stress. Superoxide produced by the Mehler reaction can also be directly reduced by ascorbate which is present in large amount in the chloroplast. Moreover, glutathione participates in ROS scavenging or avoidance under normal and drought stress conditions. Their accumulation under drought stress relates to the drought tolerance of the plant species. α-tocopherol not only prevents the formation of singlet oxygen and the hydroxyl radicals but also scavenges lipid peroxyl radicals.

6.4.6.1 Ascorbic Acid (AsA) Ascorbic acid (AsA) is a water-soluble vitamin and antioxidant molecule for the detoxification of reactive oxygen entities (Smimoff 2000). Physiologically active form of AsA is the resonance stabilized anionic form, which is formed due to deprotonation of the C3-OH group. AsA was reported to protect lipids and proteins, photosynthesis, transpiration, oxidative defense potential, improve tolerance against various biotic and abiotic stresses, and induce plant growth (Alamri et al. 2018). AsA has the capacity to directly eliminate several different ROS including 1O2, O2•-, and • OH (Hasanuzzaman et al. 2019) and maintains the membrane-bound antioxidant α-tocopherol in the reduced state and indirectly eliminates H2O2 through the activity of AsA peroxidase. 6.4.6.2 Glutathione (GSH) Glutathione is a low molecular weight thiol (γ-glutamylcysteinyl glycine) and one of the most important metabolites of the living systems because of the stability of the disulfide bridge in GSSG (Noctor et al. 2012). In most plant tissues it is maintained in GSH form except under stress conditions when its redox status changes and the relative amount of GSSG improved significantly (Wingsle et al. 1999). The efficiency with which GSSG can be re-converted to GSH during the reductive inactivation of peroxides highlights the importance of GSH in antioxidant defense.

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Secondary Metabolites and Enzymes

Phenolic compounds are derived from the phenylpropanoid pathway, mainly from aromatic amino acid phenylalanine in most plants or tyrosine in few cases. Enzymes involved in the phenylpropanoid biosynthetic pathway include lyases, transferases, ligases, oxygenases, and reductases, many of which are encoded by gene super families.

6.4.7.1 Phenylalanine Ammonia-Lyases (PAL) In plants, phenylalanine ammonia-lyases (PAL) catalyze the first step of the phenylpropanoid pathway, i.e., conversion of L-Phe to trans-cinnamic acid, which supplies the precursors for many phenolic compounds (Deng and Lu 2017). PALs are ubiquitous in plants and also found in fungi but have not yet been detected in animals (Hou et al. 2010). They are encoded by a small multigene family. Studies have shown that PAL activity responds to various stresses, such as wounding, drought, salinity, heavy metals, and infection by viruses, bacteria, or fungi (MacDonald and D’Cunha 2007). 6.4.7.2 Tyrosine Ammonia-Lyases (TAL) Tyrosine ammonia-lyases (TAL) convert L-tyrosine to ammonia and p-coumaric acid. TAL has been studied less than PAL, and it remains unclear whether TAL activity is due to a capability of PAL to accept tyrosine as a substrate or due to the activity of a specific enzyme (Jendresen et al. 2015). The rarity of TAL is likely a reflection of its specialized role in 4-coumaric acid biosynthesis, which is used as the cofactor for photoactive yellow protein (PYP) in Rhodobacter and initiates the conversion of tyrosine to N-(m, p-dihydroxy cinnamoyl) taurine moiety of saccharomicin antibiotics. 6.4.7.3 Polyphenol Oxidase (PPO) Polyphenol oxidase (PPO) genes play an important role in plant defense mechanisms against biotic and abiotic stresses. PPOs are di-copper enzymes that use molecular oxygen to oxidize monophenols and/or ortho-diphenols to ortho-quinones which cause browning reactions following tissue damage and is important in plant defense (Constabel and Barbehenn 2008). PPOs are widely distributed in bacteria, animals, plants, and fungi (Boeckx et al. 2015). PPO has been involved in the formation of pigments, oxygen scavenging, defense against plant pathogens, and herbivores (Constabel et al. 2008). When challenged by the bacterial pathogen Pseudomonas syringae pv tomato, PPO overexpressing plants showed reduced bacterial growth, whereas PPO anti-sense-suppressed lines supported greater bacterial numbers (Thipyapong et al. 2004). During various abiotic stresses (cold, heat, and drought), there was significant increase in phenolic compounds and oxidation of these accumulated phenolics was proposed to be inhibited by significant decreases in PPO. This decrease in PPO activity, following abiotic stress was associated with improved antioxidant capacity of plant under stress (Sofo et al. 2005).

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6.4.7.4 Phenolic Compounds Phenols are plant secondary metabolites that hold an aromatic ring bearing at least one hydroxyl group which is synthesized by the shikimate-phenylpropanoid biosynthetic pathway (Tungmunnithum et al. 2018). Secondary metabolites are generally classified into two groups: nitrogen compounds (alkaloids, nonprotein amino acids, amines, alkamides, cyanogenic glycosides, and glucosinolates) and non-nitrogen compounds (monoterpenes, diterpenes, triterpenes, tetraterpenes, sesquiterpenes, saponins, flavonoids, steroids, and coumarins). Structurally, phenolic compounds are composed of the aromatic ring bonded directly to at least one (phenol) or more (polyphenol) hydroxyl groups (-OH) and other substituents, such as methoxyl or carboxyl groups which cause the polar character of the compounds and allow dissolution in water (Michalak 2006). Phenolic compounds are usually divided into two groups: simple phenols and more complex derivatives, often containing several aromatic rings linked together. Phenolic compounds have ability to donate electron, hence their hydroxyl groups can directly contribute to antioxidant action and by stimulating the synthesis of endogenous antioxidant molecules in the cell (Bendary et al. 2013). Phenolic compounds chelate iron and copper ions due to the presence of suitable functional groups: hydroxyl and carboxyl, while some phenolic compounds also inhibit membrane lipid peroxidation by “catching” alkoxyl radicals. These activities of phenolic compounds are dependent on the structure of molecules and the number and position of hydroxyl groups. Under drought stress, increase in the amount of flavonoids and phenolic acids was reported (Akula and Ravishankar 2011). 6.4.7.5 Flavonoids They are synthesized through the phenylpropanoid pathway. Chalcone synthase (CHS), chalcone isomerase (CHI), and flavanone 3-hydroxylase (F3H) are key enzymes in this pathway (Hodaei et al. 2018). There are different classes of flavonoids depending upon the level of oxidation and pattern of substitution of the C ring, while individual compounds within a class differ in the pattern of substitution of the A and B rings. Important flavonoids in plants are flavones, flavanones, isoflavones, flavonols, flavanonols, flavan-3-ols, chalcones, and anthocyanidins. Until now, more than 5000 different flavonoids have been described which are classified into six major subclasses, such as flavones, flavonols, flavanones, flavanols, anthocyanidins, and isoflavones (Ross and Kasum 2002). Their structural variation in each subgroup is partly due to the degree and their hydroxylation, methoxylation, prenylation, and glycosylation pattern. Flavonoids can directly scavenge ROS by donating hydrogen atom, thus inactivate ROS. Flavonoids themselves get converted to phenoxyl radical which can further react with other free radicals to form stable quinone structure (Treml and Smejkal 2016).

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Recently many efforts have been made to improve stress tolerance through molecular biology and biotechnology techniques. Several genes involved in stress tolerance have been recognized and gene constructs have been prepared to achieve the genetic transformation to mitigate stress tolerance in forage plants. Molecular biology mainly utilizes the sequence-based, hybridization-based, and gene-inactivation or gene-editing-based approaches to study and alleviate plant abiotic stress (Kumar et al. 2019). These techniques have nurtured a well understanding of the biochemical and genetic bases of plant abiotic stress as a premise to develop high-level transgenic techniques to make plants stronger against abiotic stresses.

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Forage Cropping Under Climate Smart Farming: A Promising Tool to Ameliorate Salinity Threat in Soils Eetela Sathyanarayana, B. Prem Kumar, Rupesh Tirunagari, G. Keerthana, Vilakar Kayitha, J. Bharghavi, S. Saranya, M. Rajashekhar, B. Rajashekhar, K. Charan Teja, and Saideep Thallapally

E. Sathyanarayana (✉) Department of Soil Science and Agricultural Chemistry, Agricultural College, Palem, PJTSAU, Hyderabad, India B. P. Kumar · R. Tirunagari Division of Soil Science and Agricultural Chemistry, ICAR-IARI, New Delhi, India G. Keerthana Department of Soil Science and Agricultural Chemistry, College of Agricultural Sciences, JKKM, Erode, India V. Kayitha Keladi Shivappa Nayaka University of Agricultural and Horticultural Sciences (KSNUAHS), Shimoga, India J. Bharghavi Department of Crop Physiology, Agricultural College, Warangal, PJTSAU, Warangal, India S. Saranya GD Goenka University, Gurugram, Haryana, India M. Rajashekhar Agricultural Entomology, PJTSAU, Hyderabad, India B. Rajashekhar KVK, Palem, PJTSAU, Hyderabad, India K. C. Teja Department Agronomy, Agricultural College, Palem, PJTSAU, Hyderabad, India S. Thallapally Forest College and Research Institute, Mulugu, Hyderabad, Telangana, India # The Author(s), under exclusive license to Springer Nature Singapore Pte Ltd. 2023 R. K. Singhal et al. (eds.), Molecular Interventions for Developing Climate-Smart Crops: A Forage Perspective, https://doi.org/10.1007/978-981-99-1858-4_7

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Introduction

Drought, salinity, and heavy metals are the most emphasized abiotic stress factors that impede crop growth drastically and thereby leading to the decline in productivity (Desoky et al. 2020a, b; Ma et al. 2020). Soil salinity is one of the most abiotic stresses that hampers both the vegetative and reproductive growth in cultivated crops around the world, most importantly the salt sensitive or intolerant crops (Semida et al. 2021). About 800 Mha of arable land are classified as salt-affected soils worldwide (Shahid et al. 2018) and also it is increasing at a rate of over 2 Mha. In areas where precipitation is far less than evapotranspiration rates (hot and dry climate), the severity of salinity stress is more evident that the agricultural productivity gets drastically reduced (Wang et al. 2011; El-Mageed et al. 2020) and the soil properties also get adversely affected. This yield reduction in plants suffering from salinity stress is due to biochemical and physiological disruption such as ion toxicity by increased salt concentration, osmotic stress in plant cells, nutrient deficiencies, and ion imbalances (Rady et al. 2016; Desoky et al. 2020a, b). This induces a significant decrease in germination of seeds, growth of seedlings, grain yield, and finally the overall productivity of the crops (El-Mageed et al. 2021). Salinity causes three major stresses on the plant growth: (a) Osmotic pressure increase in the soil solution, which reduces water availability; (b) high concentration of toxic ions, especially sodium (Na+) and chloride (Cl-) and ultimately accumulation of these ions in leaves; (c) nutrient disorder and deficiency in plants (Zhang and Shi 2013). The effect of salinity stress on forage crops is highlighted in Fig. 7.1. Salt tolerance of a plant is controlled by several factors including soil, water, plant, and environmental conditions (Ortas and Ra 2021). There are numerous mechanisms and pathways that enable plants to grow well under salinity conditions that include ion vacuolation, accumulation of adaptive osmolytes, osmotic adaptation and adjustment, selective transport and uptake of ions, salt exclusion, ion homeostasis, and salt excretion in plant organs such as leaves (Flowers and Colmer 2008). Physiological limitations of salinity stress in cultivated crops (Kapoor et al. 2020): 1. Under water deficit environment created by salt stress, the decrease in relative water content (RWC %) due to some dehydration in protoplasm is prominent. By this, further reactive oxygen species (ROS) is generated causing oxidation of lipids and membrane injury. Finally, cell membrane stability (MSI %) in crop plants gets decreased. 2. Reduced stomatal conductance, photosynthesis, and different physiological and biochemical processes. 3. Decreased chlorophyll and in turn photosynthetic capacity due to limited water availability hinders growth and yield of crops by imbalanced water and nutrient uptake in plants.

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Fig. 7.1 Highlights the major effects of salinity stress on forage crops

7.2

Forage Crops as Salinity Ameliorants

Salinity stress in soils can be ameliorated by several physical, chemical, and cultural methods (Shrivastava and Kumar 2015). Although they proved sustainable management of soil salinity, practical implications still persist like the huge cost involved and availability of resources. As salinity stress poses a major threat under dry soil conditions, it is important to contain the water level in the rhizosphere region and impede its movement down the root zone. For this, perennial crops, mostly the perennial grasses, can be adopted and included as one of the components in the cropping system. For this, high-quality fodder grass such as Panicum sp. is identified as the most promising species under saline conditions in sandy loam soil (Tomar et al. 2003; Khan et al. 2009). Though the field or greenhouse tolerance of many perennial grasses like Cenchrus and Panicum is well established (Arshad et al. 2007; Alfaidi et al. 2017; Fraser et al. 2017), they are evaluated sparsely under harsh saline environments. Forage plants by physical action of plant roots improve the soil structure and provide channels for infiltrating water in soil. As a result, there is an increase in dissolution of CaCO3 (lime) in the presence of CO2 released from root respiration and decomposition of organic matter and increase in organic matter by contribution from below ground plant material that consists of roots of forage plants (Qadir et al.

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1996). The genetic potential of forage plants for their adaptability to adverse saltaffected soil condition is based on the identification and intensive cultivation of salttolerant plants. The sustainable approach to biological amelioration of salt-affected soils is utilization of improved salt-tolerant forage grass which is one of the new tools that will help farmers to increase production of crops (Kushiev et al. 2005). Besides growing an identified salt-tolerant forage grass in addition for bioremediation is an added advantage as it requires low investment, improves the soil health and quality, and that the produced forage grass can also be used as animal feedlots. Akhter et al. (2003) in their field experiment found that soil salinity, sodicity, and pH decreased exponentially by growing salt-tolerant species kallar grass (Leptochloa fusca L.) as a result of leaching of salts from surface to lower depths. Concentrations of soluble cations (Na+, K+, Ca2+ and Mg2+) and anions (Cl-, SO42and HCO3-) were reduced through to greater soil depths. A significant decline in soil pH was attributed to the release of CO2 by grass roots and solubilization of CaCO3. Both soil salinity and soil pH were significantly correlated with Na+, Ca2+, Mg2+, K+, Cl-, HCO3- and sodium adsorption ratio (SAR). In contrast, there were negative correlations between soil organic matter content and all chemical properties. The restorative effects on the soil chemical environment were noticed after 3 years of growing kallar grass. Kallar grass cultivation enhanced the leaching and soil chemical properties and thus restored soil fertility. The soil environment showed improved characteristics with further growth of the grass up to 5 years suggesting that growing salt-tolerant plants is a sustainable way to biological amelioration of saline wastelands.

7.3

Forage Quality and Yield

Unlike other cereal, pulse, oil seed crops, etc., salinity stress does not affect the crop quality of forage; in turn forage crops improve moderate levels of salinity. Feeding value of forages by cattle mainly depends on total digestible nutrients and crude protein (CP) contents. Salinity had a significant effect on the forage quality as with increase in salinity level that acid-detergent fiber (ADF) of the fodder reduces, while crude protein (CP) increases (Suyama et al. 2007; Al-Dakheel et al. 2015). This indicates improved digestibility and quality of crop with increasing salinities were commonly noticed among all the grasses. Cation absorption under salinity varies among different fodder species with different ions casing salinity. With saline water irrigation, concentration of Na+ and K+ is elevated in shoot tissues emphasizing its direct correlation with salinity (Abu-Alrub et al. 2018). When there is a higher accumulation of such ions in shoots, their sensitivity to salinity is higher. For example, P. maximum accumulated the higher Na+ levels, indicating it as more salt sensitive cultivar than Cenchrus spp. However, sometimes even salttolerant species (like Chloris gayana) shows highest shoot Na+ that is related to the presence of salt glands in such grasses that take in excess salt and transport it to the leaves and the salt crystals get excreted (Oi et al. 2012; Ribotta et al. 2013).

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Toxicity of Na+ with high saline irrigation treatments produced the lower yields as in several grass species is due to the growth inhibition as a result of soil salinity (Saqib et al. 2008; Munns and Tester 2008). Consequently, species with low Na+ content in tissues resulted in higher biomass and better salt tolerance in saline conditions. Among all the ions causing salinity, K+ accumulation in the shoots is the highest under saline condition. Such forage crops innately has the capacity to accumulate high levels of K+ despite competition for Na+ accumulation in its tissues (Hanin et al. 2016; Chakraborty et al. 2016). This can be evaluated using the K:Na ratio accumulated in forage plant shoots which tells the tolerance of species towards salinity, where less K:Na ratio indicates that the species is more tolerant against salinity. Different species possess different accumulation potentials and hence tolerance to salinity response (Nadaf et al. 2008). Also, with increasing salinity, shoot Ca2+ accumulation declined, while shoot Mg2+ increased. Salinity stress improved the growth of forage crops at less and moderate levels of salinity like 100 mM NaCl with Panicum turgidum and 6000 ppm salinity for C. ciliaris, P. maximum, and Chloris gayana (Tomar et al. 2003; Koyro et al. 2013). Such stimulation of growth is due to the physiological processes like cell elongation as a result of osmotic adjustment under salinity (Flowers and Colmer 2008). The salt tolerance as a result of ion accumulation is beneficial in growth stimulation and improving the quality only up to a certain threshold above which severe reduction in dry yields occurs. Beyond 7000 ppm, growth and yield reductions were prominent and declined gradually with increasing salinity (Ruiz and Taleisnik 2013). The performance of different forage crops for salinity tolerance is highlighted in Table 7.1.

7.4

Conclusion

To exploit the maximum use of forage crops under saline soils, more effective methods for reducing salinity should be identified. Different forage species that are climate smart and location specific that can be used efficiently to reclaim the areas affected with salinity stress must be explored to improve crop productivity in those areas and increase the profitability of the farmers. This not only corrects salt stress but also increases production, ensuring food security thereby the socio-economic well-being of the farmers.

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Table 7.1 Performance of different forage crops to ameliorate salinity stress Forage crop Barley and oats C. ciliaris, P. maximum cv. Guinea Chloris gayana Forage corn

Leptochloa fusca (L.) Kunth (kallar grass) Melilotus indicus

Melilotus indicus Porteresia coarctata Reaumuria hirtella

Rhodes grass (Chloris gayana Kunth) Sahelian fodder grass (Burgu) Sorghum bicolor var. Jambo

Sorghum (Sorghum bicolor) Sorghum

Performance under salinity stress Acinetobacter spp. and Pseudomonas sp. produce ACC deaminase and IAA Capacity of plants to maintain high levels of K+ despite Na+ competition Salt gland present on leaves secretes excess salts entering into it Supplementary P application reduces the adverse effects of high salinity on growth and development Salt taken up were excreted by the leaves or extruded to the nutrient solution by roots Salt inclusion mechanism for maintaining growth under saline conditions as it accumulated high levels of Na+ and ClTolerance of M. indicus is associated with ion (Na+ and K+) inclusion rather than exclusion A salt-secreting microhair helps in tolerating salt concentrations of 25% of sea water Diurnal pattern of salt excretion from leaves Salt secretion gradually decreased toward the midday and showed a negative correlation with the daily transpiration pattern The salt glands of Rhodes grass reportedly have a high ability to secrete excess Na+ and a low ability to secrete Mg2+ Natural dealkalinization

References Chang et al. (2014) Hanin et al. (2016), Chakraborty et al. (2016) Oi et al. (2012) Bouras et al. (2021)

Bhatti and Wieneke (1984)

Al Sherif (2009)

Ashraf et al. (1994)

Flowers et al. (1990)

Ramadan (1998)

Liphschitz et al. (1974), Liphschitz and Waisel (1982), Kobayashi and Masaoka (2008) Barbiero et al. (2001)

High peroxidases activity might be involved in maintenance of cell membrane integrity and regulation of seedling growth under salt stress conditions Na+ exclusion from the shoot aids in salt tolerance

Gaspar et al. (1991)

Increase in leaf K/Na ratio

Boursier and Läuchli (1990)

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Part II Sustainable Forage Production

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Forage Cultivation Under Challenging Environment Jasmina Milenković, Mirjana Petrović, Snežana Andjelković, and Debasis Mitra

8.1

Introduction

The problem of climate change was recognized in the twentieth century. According to it, more than 130 countries have already included Nature based solutions actions such as green infrastructure, reforestation, sustainable agriculture, and aquaculture in their national climate plans under the Paris Agreement (WWF 2020). Climate change has a significant impact on agriculture, and meeting market demands for food and fiber necessitates new solutions. Communities with a high proportion of fodder plants, in particular, create oxygen for clean air, assist in preventing silt from entering waterways, reduce soil erosion, provide food and shelter for wildlife, and enhance our surroundings with an ornamental diversity of flowers and foliage. Row crops fields are 10–50% more prone to erosion than grasslands and forage-filled meadows (Hart et al. 2022). Photosynthetic assimilation of CO2 is central to the metabolism of plants and if atmospheric concentrations of CO2 rise, it will affect the plants we depend on (Taub 2010). Increases in temperature, CO2, very low/high precipitation, and changing weather patterns are predicted by models of near-term climate change (2050), which will cause plant stress as a response. Atmospheric concentrations of carbon dioxide have already been rising from approximately 315 ppm in 1959 to atmospheric average of approximately 385 ppm in 2007 (Keeling and Whorf 2002). Increases in precipitation in some areas (up to 33%) and altered weather patterns, such as extreme drought and flooding, may also occur in some regions of the world (Loka et al. 2019; Yeung et al. 2019).

J. Milenković · M. Petrović · S. Andjelković (✉) Institute for Forage Crops Kruševac, Kruševac, Republic of Serbia e-mail: [email protected] D. Mitra Department of Microbiology, Raiganj University, Raiganj, West Bengal, India # The Author(s), under exclusive license to Springer Nature Singapore Pte Ltd. 2023 R. K. Singhal et al. (eds.), Molecular Interventions for Developing Climate-Smart Crops: A Forage Perspective, https://doi.org/10.1007/978-981-99-1858-4_8

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World Forage Production and Climate Change

To meet the demands imposed by climate change, increased forage plant production must be adapted. Ecological, social, and economic needs should all be balanced in sustainable development. Ruminant production requires forage crops for feed, and grazing areas make up around 60% of all agricultural land in the world. The characterization of physiological changes in forage plant varieties as a result of climate change has to be one of the main focuses of future research. Forage is the common name for many different species commonly used for animal nutrition. All of them have different demands for climatic, soil, and water conditions. Forages play an important role in the production of food and fiber, as well as in preserving a healthy natural environment. All of them require specific ways of cropping, tillage, harvest, storage, and keeping. According to that, they are grown in a wide range of areas in the world. Forage crops can be classified in many different ways: • • • • •

Annual or perennial Legumes or grasses Coarse-grained or fine-grained species Production for grain or for fodder The way of use (for fresh or dry biomass)

In spite of this wide range of features and many specific relations to the climate, many agricultural areas can be used for fodder production. In the worldwide forage crop production, we have to define the limiting climate conditions for every species and its production. It is necessary to define the species that thrive in a certain climate, taking into account other factors such as soil, and local needs for this type of food. The production of fodder plants is closely related to animal husbandry. The fact is that currently, in almost all climatic conditions, some of the forage species can be grown. However, given the world’s growing need for food, increasing agricultural production may be threatened by the much-heralded worsening of climate conditions. This depends on the current stage of plant development and the length of the vegetation period of each species. The effect of environmental conditions is important on a certain period of plant growth: • • • • • • • •

Before planting Seed germination in the soil Early vegetative growth Flowering Production of seeds Drought and second/third mowing Wintering Seasonal effects

Growth under unfavorable conditions of high temperature, drought, or flood, however, can cause stress responses in the plant species that could affect not only

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production but also chemical composition, and thus nutritional value. Plant breeding should strike a compromise between the desired traits (highest yield and quality) and tolerance for deteriorating soil and weather conditions.

8.3

Different Types of Environmental Stress Which Could Affect Forage Production

8.3.1

Moisture Stress

Physiological, biochemical, metabolic, and molecular mechanisms are used by plants to respond to biotic and abiotic stress. For instance, it has been demonstrated that high CO2 speeds up photosynthesis in plants and reduces the impacts of drought stress by boosting water conservation, carbon fixation, and fructan accumulation. However, it has been demonstrated that high temperatures prevent photosynthesis by changing the structure of chloroplasts and by causing oxidative stress, which renders chloroplast enzymes inactive. One solution is to introduce species with higher antioxidant activity, such as Trifolium pannonicum Jack., into larger production (Petrović et al. 2016). Water soluble carbohydrate (WSC) levels have been demonstrated to be impacted by drought stress. Plants’ morphological and physiological properties alter in response to a lack of water or frequent floods, making it possible to develop new suited genotypes. Individual features of fodder species vary so widely that initial testing of large samples is required for breeding to be successful. This was demonstrated in the study of Jahufer et al. (1997), who examined 439 samples of white clover.

8.3.2

Temperature Stress

Temperature is a measure of the thermal state, and it is mostly determined by the sun. Human activities, particularly the growth in the amount of greenhouse gases, have had a negative impact on the worldwide release of heat. This thermal energy retention affects the entire plant cover and thus the fodder species, whose production is reduced in warmer parts of the world; on the other hand, the distribution area is expanding, so there are attempts to grow alfalfa in Scandinavian countries for example (https://www.soilcare-project.eu/). If we make a comparison with warmer climate zones, where NDF values are greater and protein content is lower relative to colder regions of the world (Lee et al. 2017), the chemical composition of forage will alter in that direction as the planet’s temperature rises.

8.3.3

Soil Stress

Forage production in changed climatic conditions must be viewed from different aspects because changed climatic conditions will have different effects on soil, and

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Fig. 8.1 Nodules on alfalfa root

other living beings in the soil: microorganisms, weeds, pests, other plants and animals. Microbial communities in soil are characterized that they can respond rapidly to changes in the environment and what directly affects soil fertility (Meisner et al. 2013). The previous application of nitrogen fertilizers has caused various problems of an economic (high price of fossil energy) and ecological nature (degradation in the quality of soil and water), as well as bad effects on the health of living organisms (FAO 2022). Soil in system sustainable production must be protected and developed, using good method management, because consumption and preservation of the natural balance are of great importance (Hasanuzzaman et al. 2020). Rhizospheric microorganisms represent an important link in the soil-plant system, contribute to the enhancement of soil fertility, and may be influenced by important variables such as soil type or chemical status, nutrient quantities and type, pH, humidity, etc. (Stamenov et al. 2016). They ensure the circulation of different substances in nature and, after many processes of transformation in the soil, the nutrients available to plants (Majumder et al. 2018; Andjelković et al. 2020). Also, they could increase tolerance to abiotic stresses of various types (heavy metals, drought, cold, salinity, water logging, etc.) (Inbaraj 2021). The plant that live in symbiosis with nitrogen-fixing bacteria significantly increase the nitrogen balance of the soil, and they provide the possibility for improving nitrogen use efficiency (Lassaletta et al. 2014). Paul and Clark (1989) indicate that the symbiotic community of legumes and rhizobia represents the most significant biocatalytic link between the living world and the flow of nitrogen from the atmosphere (Fig. 8.1). In addition to providing its macrosymbionts with nitrogen, rhizobia also synthesizes various bioactive substances, have properties of biocontrol agents, and may be applied to promote the growth of plants (Baset and Shamsuddin 2010; Stajković et al. 2011; Datta et al. 2015). Also, microorganisms with amazingly very versatile metabolism have a role in mitigating climate change. This influence manifests on the physiological processes of plants, nutrient cycling, and reduction of

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toxic compounds. These microorganisms are present, in higher or lesser abundance, in the soil, but application of selected microorganisms for seed inoculation leads to better results in crop production (Andjelković et al. 2014; Mitra et al. 2021).

8.3.3.1 Seed Dormancy and Seed Storage Seed dormancy is a state in which seeds are prevented from germinating even under environmental conditions normally favorable for germination (Copeland and McDonald 2001). In unfavorable conditions, however, this trait can be much more pronounced. There are various ways to improve the germination of dormant seeds, but such manipulation increases the cost of seed production. Temperatures during seed storage could significantly affect seed quality, and consequently, forage production (Stanisavljević et al. 2020). Also, the ability of the seeds to germinate, as well as the seed vigor, is greatly influenced by the storage conditions (Stanisavljevic et al. 2011). 8.3.3.2 Feed Conservation Changed climatic conditions can significantly affect the storage conditions of fodder and silage. Inadequate temperature and humidity can increase the risk of harmful factors (insects, microorganisms, etc.) and significantly reduce quality (Lee et al. 2017).

8.4

How to Avoid the Negative Impact of Climate Change

In the future production of forage species, we will certainly face a decrease in yield due to unsuitable climatic conditions. Also, it will be difficult to ensure the stable production of fodder plants in bad climate conditions because seed production will also be aggravated. Fodder plants can also thrive on less productive land, and this may be the chance to maintain stable production in the future. A diversity of forage plants can significantly contribute to a more successful fight.

8.4.1

Forage Grasses

Pastures occupy about 60% of global agricultural land and as part of forage crops are necessary for the nutrition of ruminants (FAO 1997). Changes in growing conditions that affect forage grass yield may have great economic consequences. The impact of climate change on grassland agriculture may affect yield, livestock numbers, winter housing, slurry, storage, land spreading, and the production system (Holden and Breton 2002). In the future, the environment will have a great influence on the creation, production, and quality of new varieties of forage plants, on the productivity of fodder, and consequently on the production of livestock. Perennial fodder grasses varied in their rates of survival and recovery throughout the protracted Mediterranean drought (Volaire et al. 1998), suggesting that in the future we might see the cultivation of species that are more tolerant of harsh environmental

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conditions than the species that are currently common. According to Kemp and Culvenor (1994), the improvement of grazing and drought tolerance of temperate perennial grasses is an important issue. Their conclusion is that any future developments must acknowledge that the environment a species has come from will set some limits on how far we can go in selecting more drought- and grazingtolerant plants from within that species. Cool-season grasses native to cold climate zones are economically and ecologically among the most valuable species due to their key role in carbon fixation and as healthy forage for livestock feeding (Falloon and Betts 2010).

8.4.2

Forage Legumes

Alfalfa and clovers have been the foundation of developed animal husbandry. The amount of nitrogen that forage species leave in the soil during one season ranges from 200 kg/ha (Issah et al. 2020) for alfalfa to 133 kg/ha red clover via symbiotic nitrogen fixation, or 32–149 kg/ha as a green manure (Mueller and ThorupKristensen 2001). Benefits are through nitrogen fixation, soil structure change, and the production of high-quality biomass with high protein content. Legumes use symbiotic rhizobial bacteria to fix atmospheric nitrogen, which eliminates the requirement for inorganic nitrogen (N) fertilizers (Oldroyd et al. 2011). The cohesive action of the legumes we use distinguishes them and makes them an inseparable part of modern sustainable agriculture. As a result, the EU funded several projects on this topic over the last decade, including Legume Plus, Legume Futures, MultiSward, and EUCLEG. Sainfoin (Onobrychis viciifolia) is one of the species that appeared from these studies, with excellent persistence in suitable mixtures but a scarcity of varieties on the market (Kolliker et al. 2017). The bird’s-foot trefoil (Lotus corniculatus) is also a minor species, but it is very tolerant of soils with low P content and pH value (Cuitino and Rebuffo 2013). Legumes (Fig. 8.2), and also sainfoin, are highly desirable for beekeeping. There are some species that are grown on a limited scale but possess the potential: Trifolium alpestre L.—a permanent resident of arid habitats, Trifolium pannonicum Jacq.—long-lasting species (up to 16 years), Melilotus albus Medik.—a desirable honey-bearing species that can be utilized as green manure or improving soil quality. Also, Astragalus cicer L. is a species suitable for dry habitat cultivation in combination with other fodder species. Vymyslicky (2014) stated that Trigonella foenumgraecum L. is a fantastic Mediterranean species grown for its high-quality biomass as well as the medicinal properties of its seeds.

8.4.3

Grain Legumes

Grain legumes have been utilized for human nutrition and cattle feeding for millennia because they are high in protein, dietary fiber, vitamins, and minerals (Mudryj et al. 2014). Legumes symbiotic rhizobial bacteria fix atmospheric nitrogen,

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Fig. 8.2 Minor species: (a) Trifolium alpestre, (b) Trifolium pannonicum, (c) Astragalus cicer, (d) Melilotus albus

which eliminates the requirement for inorganic nitrogen (Oldroyd et al. 2011). The dominant crop in the world, soybean, has a significant impact on the production of legumes. Today, numerous species were introduced into the diet, considerably enhancing the gene pool, and are today sold as varieties on the international market. With new scientific study, which is important to provide new knowledge and continued development, it is necessary to promote increased diversity of grain and legume crops (Magrini et al. 2019).

8.4.4

New Approaches

Bearing in mind the variety of plants for fodder, breeders should focus on new plant species as well as improved varieties, which tolerate drought well and have satisfactory quality. Studies on the molecular mechanisms underlying forage and turf grass

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stress tolerance are considerably behind those on other crop species. According to some data, the genes that make rice and other cereals resistant to abiotic stress have a similar role in forage and turf (Jiang and Huang 2002; Humphreys et al. 2004). Recently, demands for more sustainable production systems have required high yielding, high-quality forages that enable efficient animal production with minimal environmental impact (Kingston-Smith et al. 2012). Some recent reports indicated the heat sensing and signaling pathways that activate transcriptional cascades and future directions of promoting crop tolerance to HS using these factors or other strategies for agricultural applications (Zhao et al. 2020; Ohama et al. 2017). Using forage legumes as intercropping species is another strategy to combat environmental change. Cowpea (Vigna unguiculata) is a species that is commonly intercropped with cereals such as corn, sorghum, and millet in Africa, and it has a favorable impact on biomass and grain yield (Hassen et al. 2017). Also because of the existence of environmental problems and climate change, it is necessary to develop and strengthen alternatives that complement or replace the use of mineral fertilizers. Microbiological fertilizers with rhizobial and other bacteria represent a powerful tool in economic profitability and environmental protection. The effect of inoculation depends on host plant activity, species, strain and cell concentration of microorganisms in the inoculum, the structure of indigenous populations, soil health, microbial diversity, and soil disturbances caused by management practices (Walker et al. 2003; Andjelković et al. 2018) of soil fertility. The main goal of that should be in function of producing sufficient quantities of healthy and quality food, directing microbiological processes in the desired direction. Also microorganisms with amazingly versatile metabolism have a role in mitigating climate change. Application of this agrotechnical measure for forage legumes enables improvement of structure and preservation of soil fertility. A good mechanism to solve climate change is connection of the microbial community and biogeochemical cycles. The symbiotic community rhizobia and legumes are a cheaper, ecologically sustainable process using renewable resources (Peoples et al. 1995). Plants have the ability to “remember” past occurrences and to adapt to new environments (Kinoshita and Seki 2014) and also have a kind of memory and signals (Hilker and Schmülling 2019). In the coming period, the use of wild species and other plants for feed should also be considered because weed plants and wild relatives of cultivated plants are more tolerant to stressful environmental conditions. Accordingly, the possibilities of using other species for fodder (e.g., prairie grasses, weeds) should be highlighted (Milenković et al. 2018; Bates et al. 2007). Phalaris is arguably the more droughttolerant of commonly sown species, but its roots still require contact with water for the plant to survive (Kemp and Culvenor 1994). A component of this problemsolving is, also, increasing regional production of leguminous crops that are high in protein (Shepon et al. 2018). Alternative forage crops could become more suitable for winter feed conservation (Holden and Breton 2002). Some of the minor pulses in European temperate regions as a complement to common pulses could adapt to different pedoclimatic conditions, given their physiological adaptation capacity, and these pulses might be of interest for the development of innovative local food chains in an EU policy context targeting protein autonomy (Gotor and Marraccini 2022).

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Acknowledgments The research was financed by The Ministry of Science, Technological Development and Innovation Republic of Serbia (451-03-47/2023-01/200217). We also thank Mgr. Helena Hutyrová for providing pictures of plants.

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Potentials and Opportunities of Agroforestry Under Climate Change Scenario Manjanagouda S. Sannagoudar, G. K. Prajwal Kumar, Vanitha Khandibagur, Avijit Ghosh, Amit K. Singh, G. A. Rajanna, Hanamant M. Halli, V. K. Wasnik, B. R. Praveen, and R. T. Chethan Babu 9.1

Introduction

The term “agroforestry” is defined as sustainable land use systems and technologies of woody perennials (trees, legume shrubs, etc.) are deliberately grown on the same land management system as arable crops and livestock systems in a spatial arrangement or in temporal sequence or both; there are typically ecological and economical interactions between the components with other system (Lundgren 1982). Agroforestry provides both tangible and intangible benefits that are essential to the growth of Indian economy. In reality, agroforestry has tremendous potential for concurrently achieving three key goals: conserving and stabilising the bio systems, providing a greater level of economic production, increasing income and access to basic necessities for the rural people (Anon., 2011). Agroforestry helps in the restoration M. S. Sannagoudar (*) ICAR - Indian Grassland and Fodder Research Institute, Jhansi, India ICAR - Indian Institute of Seed Science, Regional Station, Bengaluru, India G. K. P. Kumar Karnataka State Department of Agriculture, Bangalore, India V. Khandibagur University of Horticultural Sciences, Bagalkot, India A. Ghosh · A. K. Singh · V. K. Wasnik ICAR - Indian Grassland and Fodder Research Institute, Jhansi, India G. A. Rajanna ICAR - Directorate of Groundnut Research, Anantapur, AP, India H. M. Halli ICAR - National Institute of Abiotic Stress Management, Baramati, MH, India B. R. Praveen · R. T. C. Babu Agronomy Section, ICAR - National Dairy Research Institute, Karnal, Haryana, India # The Author(s), under exclusive license to Springer Nature Singapore Pte Ltd. 2023 R. K. Singhal et al. (eds.), Molecular Interventions for Developing Climate-Smart Crops: A Forage Perspective, https://doi.org/10.1007/978-981-99-1858-4_9

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of degraded lands, resulted in increased farm output. Presently, agroforesrty provides nearly 50% of the fuel wood requirement, 66% of the small timber requirement, 70–80% of the wood required for plywood, 60% of the raw material paper industry, and 9–11% livestock feed in the form green fodder, in addition to providing for households’ daily needs of food, fruit, fibre, medicine, etc. Generally, agroforestry plays an important role in preservation of the resource base and the total production in arid and semi-arid regions under rainfed situation (Anon., 2013). Agroforestry is presently being employed to focus on shifting priorities in areas like carbon sequestration, green energy, and agricultural production improvement. It is also acknowledged that the goal of raising forest cover from its current level of less than 25–33% can only be achieved through agroforestry (Anon., 2019). Environmental amenities including phytoremediation, watershed protection, biodiversity preservation, and climate change mitigation will all be significantly aided by agroforestry (Bhagwat et al. 2008). By expanding the forest and tree cover before 2030, it will especially help to achieve the goals of reducing emission intensity and adding new carbon sinks for carbon dioxide. Early in the 1970s, ICAR institutions invited industry to participate in the planting of commercial tree species, which marked the beginning of organised agroforestry research in India. International Centre for Research in Agroforestry participated in the inaugural agroforestry seminar held in Imphal in 1979 (ICRAF). As a follow-up, the ICAR launched the AICRP on Agroforestry in April 1983. The council established the National Research Centre for Agroforestry (NRCAF) in 1988 in Jhansi after taking the initiative to pursue organised research in agroforestry initially through a number of co-ordinated projects. The centre was created to meet the demands of basic, strategic, and applied agroforestry research.

9.2

Major Agroforestry Systems and Practices

Systems and practises are frequently used interchangeable in agroforestry literature. An agroforestry system is a particular local example of a practice, distinguished by the environment, plant species, and how they are arranged, managed, and operated in terms of the socio-economic system. An agroforestry technique, in turn, designates a unique configuration of elements throughout time and location (Handa et al. 2019). Details of the different agroforestry systems and major components are presented in Table 9.1.

9.2.1

Shifting Cultivation

Shifting cultivation is also known as Jhum cultivation, it is one of the oldest traditional agroforestry techniques, is a type of farming practice used in the tropics and subtropics where land covered by native forest vegetation is removed by using the slash-and-burn technique, planted with arable crops for a period of time till loss of native soil fertility afterwards natural vegetation will regenerate. The fallow

Agroforestry practice Shifting cultivation

Taungya cultivation Multipurpose trees on crop lands

Plantation-based cropping systems

Alley cropping (hedgerow intercropping)

Home gardens

Trees on grazing/pasturelands Afforestation for land reclamation and soil conservation (improved) Shelter belts, windbreaks, and live hedges Woodlots Cut and carry systems (protein banks)

Multi-enterprise farming systems (improved)

Others (improved)

S. No. 1.

2. 3.

4.

5.

6.

7. 8.

9. 10. 11.

12.

13.

Table 9.1 Classification of agroforestry systems in India Arrangement of major components Quick growing legume trees are planted after burning natural vegetation and arable crops are grown for period of time till loss of native soil fertility Combined planting of leguminous woody and field crops Trees planted on corners of fields, boundary plantations, and live hedges (forest, fruit trees, arable crops) (a) Integrated multi-storied planting of plantation crops (b) High value plantation crops along with shade trees in field and boundaries (c) Tree planting + arable crops (d) Arable or fodder crops as intercropping in fruit trees (e) Plantations crops on permanent pastures/grazing lands Leguminous trees or shrubs planted in narrow rows as alleys and field crops or grasses between the alleys Around homesteads, there are multi-storied combinations of fruit and plantation trees, shrubs, vines, shade crops, vegetables, spices, etc. Scattered woody trees or shrubs (used as fodder + shade) on grazing lands (involves livestock) Woody trees, shrubs, and grasses on highly problematic lands including eroded, mined, saltaffected, and waterlogged soils Tall woody trees and shrubs sometimes succulent cactus, etc. are used as live fence Mostly on public lands, along roads, railway lines, avenue trees, sacred groves, etc. Usually, legume forage and woody trees or shrubs and fodder cereals and legumes on community lands Forest trees and fruit crops + food crops + vegetables + fodder crops + livestock + poultry + duckery + piggery + fishpond + floriculture + apiculture + natural gas/solar energy The selection of attractive plants from forests lands and wild regions, grazing in forests, aquaforestry, honeybee farming, and medicinal and aromatic plants with fruit trees and forests; fish culture with mangroves; etc.

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period was historically 10–20 years, but it is presently just 3–5 years. In 48 districts across India, around 600,000 households are engaged in shifting agriculture over an area of 2.27 million acres.

9.2.2

Taungya

A precursor of agroforestry, the tropical taungya system is similar to a planned and carefully maintained shifting agriculture. According to Blanford, the word’s origin was in Myanmar (formerly Burma), where tauang means hill and ya means agriculture. Originally used to indicate shifting farming, it was later also used to denote afforestation.

9.2.3

Home Gardens

Home gardens are transitional phase between a tropical forest setting and an arable crop that sustains both sustainable agriculture and forest biomes. It safeguards natural biodiversity as well. Home gardens have been discussed extensively, and experts have employed many different terminologies. These include home gardens, compound farms, kitchen gardens, family gardens, Javanese home gardens, mixedgarden horticulture, and homestead agroforestry.

9.3

Plantation-Based Cropping Systems

A sizable amount of data has recently been produced in India from cropping systems based on coconuts. This information relates to factors that favour intensifying land use pattern with coconut, multi-storied plantation crops, mixed farming, grazing practices under coconut trees and growing other fruit crops, arable crops as an intercrop for effective space utilisation.

9.3.1

Scattered Trees on Farmlands

The practice of cultivating arable crops on farmlands under solitary trees is fairly old and does not appear to have altered much through the years. In tree-crop mixed farming, trees were valued higher than crops in ancient India. Even today, trees can be seen growing haphazardly amid fields of crops for a variety of purposes, including shade, fodder, fruit, fuel wood, vegetables, medicinal purpose, and minor timber. Some of the techniques are extremely sophisticated and broad.

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9.3.2

165

Trees on Field Boundaries

The custom of growing agricultural crops on farmlands under scatted trees is fairly old and does not appear to have altered much through the years. In tree-crop mixed farming, trees were valued higher than crops in ancient India. Even today, trees are growing haphazardly amid crops fields for various purposes, including shade, vegetables, fodder, fruit, fuel wood, minor timber, and medicinal purpose. Some of the procedures are quite sophisticated and broad.

9.3.3

Woodlots

In many areas, farmers also have agricultural fields and separate woodlots where they cultivate trees. Due to the lack of firewood and the demand for poles or pulpwood in industry, the practice is now multiplying. For instance, bamboo poles and Eucalyptus and Populous are both in high demand for WIMCO Industries and orange plantations in the Nagpur area.

9.4

Different Approaches for Soil Conservation

In India, 121 million acres of land area are affected by various problems with deterioration, including severe water and wind erosion, salinity problems. The easiest way to regulate the deep and narrow gullies is to permanently cover them with vegetation after closing them off to grazing. The stabilisation of gullies and ravines is aided by the planting of appropriate tree species like Acacia nilotica, Acacia eburnea, Butea monosperma, Azadirachta indica, Dalbergia sissoo, Prosopis juliflora, Tectona grandis, varieties of Bambusa and Dendrocalamus.

9.4.1

Shelter Belts

The entire year, very high wind speeds are seen in arid places, and sand can start particle movement as early as 12–14 km/h. In Rajasthan, for example, farmers construct kana bandis, or sand-stopping barriers, by utilising either tiny deadwood fragments or indigenous flora to keep wind speeds within safe ranges. Aerva javanica, Leptadenia pyrotechnica, and Crotalaria burhia are arranged in rows with 20–25 m wide against wind direction. On the leeward side of each break, grasses viz., Cenchrus ciliaris, Lasiurus scindicus, and Cenchrus setigerus are planted between the lines of these under plants. This persistent vegetation aids in the accumulation of sand nearby, which is later dispersed over the field. Along these lines, this contributes to higher agricultural yields.

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Fig. 9.1 Silvipasture system in tropical areas: (a) Ficus infectoria based, (b) Subabul based, (c) Hardwickia binata based, (d) Acacia nilotica based

9.4.2

Trees on Rangelands

The most prevalent trees in common communal grazing lands in dry places are Salvadora oleoides, Salvadora persica, Acacia leucophloea, Tecomella undulata, Capra decidua, Prosopis cineraria, Acacia nilotica, and recently, Prosopis juliflora. The coconut tree is the most prevalent one on pasture grounds in coastal locations. Trema tomentosa, Moringa oleifera, Morinda citrifolia, Gliricidia sepium, Albizia lebbeck, Pongamia pinnata, and Ficus spp. trees were lopped for fodder purpose. Generally, grazing on these pastures is a part of livestock production. In arid zones, an organised silvopastoral system ensures 10 tons/ha/year forage biomass production (as opposed to 1 tons/ha/year from natural stands), as well as soil conservation, carbon sequestration, and job creation. The major silvopastoral system of tropical regions is highlighted in Fig. 9.1.

9.4.3

Aquaforestry

Aquaculture and prawn culture in salt-affected water along with growing of coconut trees on bunds of ponds are commonly practised by the coastal region farmers especially by Andhra coast. Coconut trees by-products are used as feeding material

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for the fish and prawns. Presently fish culture under mangroves proved to be better nutrition to the aquatic animals and favourable condition for the breeding of prawn, juvenile fish, turtles, and other aquatic animals. Another successful venture in these locations is poultry. A system of animal husbandry that is well-balanced and includes goat rearing, poultry, donkey, turtles, and fish in tiny ponds creates a system with optimum moisture, energy generation, and nutrient use efficiency per unit area.

9.4.4

Apiculture with Trees

Natural beehives are the source of harvesting of honey in the coastal regions. Apiculture is now recognised as one of the profitable agriculture professions throughout the country. Nowadays many farmers are sowing their interest in apiculture as a business enterprise through different kinds of plantations. Eventually, huge floral diversity in agroforestry system makes bee keeping easy and more productive. Inclusion of apiculture in farming system generates regular income and makes system sustainable. According to estimation, irrigated agroforestry with commercial/industrial plantations occupies 7 Mha, agri-silviculture occupies 2.63 Mha, 2.79 Mha under fruit crop-based cropping systems, and trees on boundary or bunds of cultivated fields occupy 1.58 Mha by social forestry. About 10.6 Mha area is under rainfed agroforestry with agri-silviculture (2.4 Mha), agri-horticulture or fruit trees-based cropping systems (1.86 Mha), and trees on grazing or rangelands and field boundaries (6.32 Mha). In addition to this, rehabilitation of saline soils, mined wastelands make possible 3 Mha area is under tree cover. Considering the significance of agroforestry system in India, the Planning Commission has stressed in its annual report about increasing the additional area under irrigated and rainfed agroforestry 10 Mha and 18 Mha, respectively. Out of this about 7.73 Mha has been brought under agroforestry through various agroforestry schemes and there is potential to add additional 28.0 Mha to the area reported for agroforestry in the future, which might include the remaining 20.27 Mha area that was originally designated by the Planning Commission in 2001 in its report. The majority of the land that will be converted to agroforestry consists of fallows, cultivable fallows, pastures, groves, and problematic soil reclamation. So, in future, agroforestry could potentially cover 53.31 Mha (or about 17.5% of the total reported geographic area, or TRGA), making it India’s third-largest area occupied, behind agriculture and forestry (which together occupy 141 Mha, or 46% of the TRGA and 69.63 Mha, respectively) (Table 9.2). Generally agroforestry system consists of five basic systems based on components which are managed. Basically, the system is categorised as: 1. Agri-silviculture 2. Agri-horticulture 3. Silvi-pastoral

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Table 9.2 Country’s estimated area for agroforestry Category Irrigated land agroforestry Agri-silviculture Agri-horticulture Trees on field boundary/bunds Subtotal (A) Rainfed land agroforestry Agri-silviculture

Area (m ha)

Remarks

2.63 2.79 1.58 7.00

Industrial raw material Fruit gardens or fruit-based cropping systems Live fences and social forestry

2.40

Dispersed trees in fields, bunds, and field boundaries Fruit crops or plantation-based cropping systems Social forestry, live fences, etc. Trees on grazing or rangelands and community lands

Agri-horticulture Trees on field boundary/bunds Silvopastoral

1.86 0.74 5.58

Subtotal (B) Other land uses Homestead gardens Jhum cultivation

10.58

Afforestation on problematic soils Trees planted on community lands Subtotal (C) Sum total of agroforestry area (A + B + C)

2.12

2.42 2.27

0.92

Mostly observed in coastal areas and N-E states Majorly seen in N-E States, Andhra Pradesh, and Orissa Plantations on salt-affected soils, mined lands, bare lands On public lands, along the public roads and railways tracks, etc.

7.73 25.31

4. Agri-silvipastoral 5. Other systems Need and Scope of Agroforestry in India • Especially in areas with little forest cover, agroforestry systems are likely the only way to achieve the desired level of tree cover nationwide. • Agroforestry is a method of cultivating food crops along with perennial woody trees, animals, and other plants to maximise the utilisation of natural resources while simultaneously providing a basis of renewable energy and lowering the ecological degradation and global climatic aberrations. • The livelihood security provided by agroforestry and its potential to provide food, fodder, fuel, and employment are known established advantages. • Agroforestry systems ensure nutritional security to the communities by diversified food grains, vegetables, and fruit crops. • Fodder components in agroforestry system ensure good milk, meat, and animal production. Diversification in cultivation like food crops, pulses, and oil seeds provides nutritional security of the community.

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• Growing of trees, industrial processing, marketing of tree and other by-products is anew-vistas in agroforestry research field and development. • Multi-enterprise agricultural systems must go a long way to help small and marginal stakeholders to achieve sustainability and livelihood security. • The development of techniques to support the rural population for the watershed restoration, conservation of biodiversity, and carbon sequestration that they contribute to society is also playing a significant role in environmental remediation. • Agroforestry is one of the potential climate smart agriculture practices. It withstands various climatic aberrations through established mechanism.

9.5

Recent Advances in Agroforestry

9.5.1

Agroforestry and Natural Resource Conservation

Agroforestry techniques have been developed for the various productive and unproductive lands such as arable, non-arable lands, torrent control, boulder riverbed land, landslide and landslip stabilisation, deserted mined area rehabilitation, and also as a substitution system to jhum cultivation. Agroforestry system has also illustrated their effectiveness in the treatment of saline and alkaline lands as well as the prevention of floods, the recovery of waterlogged and stagnated areas, the restoration of wastelands, the rehabilitation of ravines, sea erosion management, and the prevention of desertification (Narain, 1997). The use of saline soils and water for crop production in order to meet the rising need for food and fodder. This is another instance where agroforestry played a significant role in improving the productivity of agriculture land, addressing the food security and environmental problems. Since it is neither practical nor possible to eliminate accumulated salts on the soil surface, efforts have been made to reduce their negative effects on crops by creating various agrotechniques. CSSRI in Karnal (Haryana) has developed specific planting methods for sodic and saline soil for proper establishment and establishment of multipurpose trees. The agrotechnique guarantees more than 80% tree survival capacity more than one decade in highly salt-affected soils. According to a CAFRI study, expanding integrated watershed management in drought-prone areas with supportive institutional and regulatory frameworks would boost equity and livelihood opportunities, strengthen diverse ecosystem services, and lower poverty in the semi-arid tropics (Chavan et al. 2016).

9.5.2

Soil Organic Carbon Fractions

Various trees such as (Acacia, Ficus, Leucaena, and Morus) and grass species (Cenchrus, Chrysopogon, and Panicum) significantly affect the soil organic carbon content by interacting with rhizosphere soil.

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Enzyme Activities

Different land restoration strategies had different effects on how soil enzymes responded to eco-restoration techniques. DHA, ALP, FDA, and URE activities were greater in subsurface soils for acacia-occupied fields, while they were highest in surface soils for eco-restored lands employing Morus. However, Cenchrus may have more biomass production from its root and a greater supply of substrate, which may be the reasons for all soil enzymes’ higher activity. The hydrolytic enzymes activity was driven by increased total microbial biomass and an equal increase in the substrate supply from litter, and they may have been amplified even more by labile root exudates (Ghosh et al. 2021).

9.5.4

Carbon Management and Biological Activity Index

Strategies for eco-restoration greatly increased CMI. Therefore, even though grasses increased CMI than fallow region, their effects were comparable at the surface soil layers. However, in areas covered with Ficus, Acacia, and Cenchrus, CMI was greater in subsurface soils than surface soil. According to ERE values, Panicum sp. was the most effective grass for restoring degraded lands in the tropical environment of India’s Bundelkhand region, while Morus and Acacia were the most effective trees (Ghosh et al. 2021). The efficiency of paired rows of Leucaena and Eucalyptus trees and Panicum maximum barrier of 0.75 m wide planted at 1.0 m interval of maize crop were compared in N-W Himalayas in Doon Valley watersheds receiving 1740 mm of annual rainfall. It was discovered that all of these systems reduced runoff from 16% to 43% of rainfall and soil loss from 7 to 21 Mg/ha/year. With a concomitant decrease in soil loss, there is noticeable increase in deposition of sediment all along the hedgerows and tree rows. Because of nitrogen fixation, applying Leucaena leaf mulch before or during the harvest of maize boosted moisture retention and wheat production. Agroforestry has a considerable role in the social, economic, and environmental sustainability as well as the livelihood component in a watershed area when used in combination with traditional conservation practices in a watershed area, including ravine erosion lands. In addition to the catchment’s agroforestry constituents (crops, grasses, and trees) improving, IWM has produced outstanding groundwater recharge and water table rise in various regions across the country. Sod grasses like Dichanthium annulatum, Panicum antidotale, and Cenchris ciliaris stabilise the majority of the periphery bunds. Grassed ramps or piped outlets are used to safely dispose of the extra runoff. It has been discovered that graded bunds in combination with grassed waterways and chute drop water control structures for the safe removal of excess water in the Chambal ravines, where the soil is medium fine and the rainfall is greater, are preferable to contour bunds. It has been determined that integrated watershed development has increased in situ water conservation for higher biomass production and has decreased runoff or regulated flooding of downstream areas.

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Table 9.3 Soil improvement through Aonla based agroforestry (Uthappa et al. 2015) Years 1996 (Seeding) 2012 (18 years) % Deviation from initial

pH 7.00

EC (dS/m) 0.16

Available N (kg/ha) 167.70

Available P (kg/ha) 13.20

Available K (kg/ha) 120.60

Organic Carbon (%) 0.32

7.30

0.16

265.00

15.40

162.50

0.60

-7.59

-25.00

+64.32

+16.67

+3.74

+87.50

Table 9.4 Recommended tree species for problematic area (Uthappa et al. 2015) Problematic areas Sodic soils (pH 8.62–10)

Wetland

Waterlogged area

Sand dunes stabilisation Coastal area

Ravine area

Acid soils

Recommended tree species Acacia nilotica, Casuarina equisetifolia, Acacia auriculiformis, Prosopis cineraria, Prosopis juliflora, Tamrix articulate, Salvadora persica, Pithecellobium dulce, Pongamia pinnata, Terminalia arjuna, Poplars, Sesbania sesban, Eucalyptus tereticornis, Butea monosperma, and Azadirachta indica Alnus trabeculosa, Alnus cremastogyne, Salix xuchonensis, Salix babylonica, Paulownia tomentosa, Taxodium distichum, Taxodium scandens, and Morus alba Bambusa arundinacea, Eucalyptus spp., Corymbia tessellaris, Casuarina equisetifolia, Terminalia arjuna, Tamarix, Lucerne, Syzygium cumini, Barringtonia acutangula, Ficus glomareta, and Psidium gujava Prosopis cineraria, Prosopis juliflora, Acacia tortilis, Acacia nilotica, Azadirachta indica, Acacia senegal, Ziziphus spp., Tecomella undulata, Capparis decidua, Ailanthus excelsa, and Parkinsonia Aculeta Anacardium occidentale, Casuarina equisetifolia, Ailanthus malabarica, Pongamia glabra, Terminalia catappa, Calophyllum inophyllum, species of Pandanus, Nypa fruticans, Salicornia bigelovii, Sesbania aculeate, Salvadora persica, S. oleoides, and Borassus flabellifer Acacia catechu, Acacia nilotica, Dalbergia sissoo, Morus alba, Dendrocalamus strictus, Azadirachta indica, Albizia spp., and Holoptelia integrifolia Pinus roxburghii, Pinus wallichiana, Celtis australis, Cassia, Acacia mearnsii, Acacia decurrens, Acacia dealbata, Acacia pycnantha, Manilkara hexandra, Phoenix spp., Acacia auriculiformis, Pterocarpus marsupium, Shorea robusta, Hardwickia binata, Terminalia spp., Tectona grandis, Gmelina arborea, Xylia xylocarpa, Bamboo spp., Alnus spp.

Tree-based farming methods are crucial for preserving the physico-biochemical characteristics of soil and consequently its overall productivity (Table 9.3). The best option for crop diversification is to introduce tree-based farming in order to maintain agriculture’s growth and production. Degraded fields can be improved with the inclusion of tree components, allowing them to be used for regular cropping in the future (Table 9.4). Agroforestry helps in improving and maintaining the soil sustainability for the small and marginal farmers in win–win situation.

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Agroforestry and Conservation Agriculture (CA): Harmonising Agriculture Practices for Sustainable Development

Conservation agriculture (CA) is a sustainable method of agricultural production, is to maximise yields while preventing soil erosion and degradation, enhancing soil quality and biodiversity, and preserving water and air resources. CA consists of three principles which are interlinked to each other, namely: 1. Minimum soil disturbance 2. Permanent soil cover with crop residue or live mulch 3. Diversified cropping systems and crop rotations

9.5.5.1 Conservation Agriculture as a Forest Mimic Principles of conservation agriculture consists of minimum mechanical soil tillage and permanent organic soil cover with crop residues, CA tends to imitate natural systems, especially that of the rainforest. In forest systems, nutrients are recycled through leaf fall and its degradation by the microbial activity in the soil, this process enriches the soil biota. Leaving the soil surface bare results in the destruction of the natural pores for water penetration and gaseous exchange, thus soil damaging tillage practices are to be replaced by the conservation agriculture practices for the crop production (Ramulu et al. 2017). In addition to protecting the soil from the harmful effect of wind and water erosion, permanent soil cover offers additional significant advantages to the soil, including the regulation of soil moisture content and soil temperature. As has already been mentioned, degradation rates under conservation agriculture and agroforestry and forest systems were practically zero (Roy et al. 2011). Details of the potential advantages, synergistic effects, and compatibility of AF and CA are presented in Table 9.5.

9.5.6

Sloping Agricultural Land Technology (SALT)

The majority of the Philippines land area is considered sloppy land, which accounts for 60% of its total land area. The majority of the 17.8 million Filipinos who reside in these regions engage in shifting cultivation. Deforestation of uplands causes environmental hazards such as soil erosion, river siltation, floods, and droughts. The Mindanao Baptist Rural Life Centre created a Sloping Agricultural Land Technology strategy for the uplands to prevent them from being completely destroyed (SALT). Its primary goals include providing food for the farm family and the upland community as a whole in addition to making upland farming sustainable (Tacio 1993). The SALT technological innovation combines a number of soil conservation techniques with food production. The basic idea behind the SALT approach is to plant or sowing of atmospheric nitrogen-fixing trees and bushes in double rows along the contour, followed by narrow strips of field crops and perennial crops that

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Table 9.5 Advantages, synergistic effects, and compatibility of agroforestry and CA (Elevitch 2004) Concept Efficiency of natural resource use

Constituents Soil nutrients

Solar energy

Water

Favourable environment for sustained production

Shade

Wind protection

Soil conservation

Nutrient cycling

Habitat diversity

More profitable systems

Reduced costs

Diversified products Continuous flow of products

Potential of AF and CA Woody perennial trees boost the nutrient movement and atmospheric nitrogen fixation. Crop rotation of main crops with crops with different rooting depths AF systems and CA have an advantage of nutrient cycling and Leguminous cover crops also fix N Multi-storied cropping trap and utilise solar radiation at different levels. Although the same advantage is better explained by agroforestry systems, crop associations in CA illustrate similar efficiency Both AF and CA enhance water infiltration into soil and moisture holding capacity by minimising the surface runoff Systems that use AF (and some CA) can offer diffused shade that conserves water, lowers evapotranspiration, keeps the topsoil cool, and supports healthy soil bioactivity Tree windbreaks safeguard soil from wind erosion and dryness as well as crop harm. Combining windbreakers with CA.provides more comprehensive protection Undisturbed trees, crop, cover crop roots, and Mycorrhizal fungi in rhizosphere bind soil prevent erosion and minimise leaching. Enhancing the physical, chemical, and biological conditions of the soil, CA soil cover and tree leaf litter makes soils more resistant to erosive forces Trees, shrubs, and cover crops encourage more closed nutrient availability and more effective use of nutrients by nutrient uptake from deep soil layers and nitrogen-fixing organisms The provision of habitats for a variety of biota by CA and AF, but especially in conjunction, helps to increase biodiversity and the balance of predators and pests in the system Cover crops and trees reduce the demand for external fertiliser inputs by nutrient cycling. Compared to plough-based agriculture, CA systems require less fuel and labour Usually, mixed cropping systems produce more profitable goods. For instance, tree fruits and wood from AF, whereas legume seeds from CA Multiple cropping in both AF and CA can ensure a more consistent supply of goods all year long (continued)

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Table 9.5 (continued) Concept

Constituents Greater selfreliance

Environmental improvement

Reduced pressure on natural forests Species diversity

Resource conservation Carbon sequestration Decreased pollution

Potential of AF and CA AF and CA are able to lessen the farm family’s reliance on imported goods as well as their sensitivity to shifting market conditions, particularly for mono-cropping systems This is especially beneficial for AF systems because it lowers demand for forest products Enhancing habitat and supporting biodiversity for macro and micro fauna are both goals of AF and CA AF and CA hasten soil conservation, nutrients recycling, and water holding capacity in the soil Trees and especially soils store C and so reduce GHG emissions Nutrient cycling can minimise the external synthetic fertilisers application and control soil erosion and excess runoff means that nutrient loss approaches zero

are 3–5 m wide. These prevent soil erosion and conserve the soil fertility. Legumes, cereals, and vegetables are among the field crops, while cocoa, coffee, bananas, citrus fruits, and fruit trees are the major perennial crops. A stable environment can be established with the help of SALT. Soil erosion can be reduced by quick growing greater foliage leguminous shrubs or trees in double hedgerows. For the purpose of increasing the soil’s fertility, their branches are trimmed every 30–45 days and re-incorporated. The crop offers a constant vegetative cover that helps with soil and water conservation. The soil and air temperatures are kept at levels where different agricultural crops can thrive more effectively by the legumes and perennial crops. The following hedgerow plant species are advised for use in SALT in the Philippines: Flemingia macrophylla, Desmodium rensonii, Gliricidia sepium, Leucaena diversifolia, and Calliandra calothyrsus.

9.6

Agroforestry and Ecosystem Services

The provision of supplemental, secondary habitat for species that tolerate a certain level of disturbance, the creation of a more benign and permeable “matrix” between habitat remnants compared with less tree-dominated land uses, and the reduction of rate of natural habitat conversion in some cases which may support the integrity of these remnants and the conservation of their populations are three roles of AF in biodiversity conservation on a landscape scale. These systems, which frequently resemble tiny forest remnants and integrate multiple species into a single system with a structurally complex canopy (unlike monocrop systems), are able to provide

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ecological services akin to those of forests. However, the scope of these services varies greatly (Roger 1998). It has been demonstrated that the number of flowers in an agroforestry system positively correlates with the abundance of pollinators (honeybees and bumblebees). Additionally, agroforestry has been demonstrated to benefit insects that consume agricultural pests, lowering the need for pesticides. Some of the elements claimed to have good effects on insect diversity include variations in tree-crop combinations and spatial layouts, plant diversity, tree density and canopy height, proximity to forests, and an abundance of food supplies such as hosts, prey, and nectar. These studies show how heterogenic agroforestry systems can preserve arthropods without significantly destroying or changing microhabitats and microclimates (Samra & Narain, 1998).

9.7

Agroforestry and Livelihood Security and Income Generation

For centuries, India has dependent on trees to support a sustainable way of life. Farmers are compelled to employ tree-based systems because of the challenging weather conditions in order to protect their life and income. For the people of drought-prone Bundelkhand region of India, harvesting, collecting, and value addition of non-timber forest products (NTFPs) are generating a number of job opportunities. This article seeks to show how farmers, tribal people, and labours depend on trees for their source of revenue. An overview of the reliance of various rural poor communities on non-timber forest products such as gum, lac, dona pattal, brooms, jaggery, and baskets and sticks from bamboo and Phoenix, floral and seeds from mahua, bidi leaves from tendu for to lead their life was provided by surveys and interviews in the Bundelkhand region. Greater emphasis should be provided for sustainable harvest NTFPs for their healthy livelihood (Hala Naik 2003). Agroforestry is another effective method for managing carbon emissions in damaged natural forests which sequester carbon in vegetation and possibly in soils. The carbon pools of agroforestry component are presented in Table 9.6. Additionally, slash-and-burn method of farming practice is reduced with more intensive land usage for agriculture, and to that extent, agroforestry raises farmers income. The projected average sequestration potential for carbon in India’s agroforestry is 25 Mg C/ha over 96 million ha, but there is a significant regional variance depending on biomass production. Evidence for the effectiveness of agroforestry systems as management strategies to boost soil and above-ground carbon stocks and reduce greenhouse gas emissions is now starting to emerge (Subbulakshmi et al. 2019). Recent studies have reported that tropical agroforestry systems have a carbon sequestration potential of between 12 and 228 Mg/ha with a median value of 95 Mg/ ha (Newaj et al. 2017). Different alternative land use systems recorded higher soil organic carbon (SOC) status than agricultural land and fallow land (Reddy 2002). Among various agricultural land use systems, higher status of SOC (Mg/ha) was recorded under agri-silviculture system (19.93) followed by silvipasture system

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Table 9.6 Carbon pools in agroforestry Pools Living biomass

Aboveground Belowground

Dead organic matter

Decomposed wood Litter

Soils

Soil organic matter

Descriptions All plant material above the soil, including stems, bark, branches, pods, twigs, foliage, and seeds, whether it is woody or herbaceous All living roots make up the biomass, with the exception of very small roots (less than 2 mm in diameter), which are frequently difficult to distinguish from litter or soil organic materials All woody biomass on the surface and subsurface soil Includes on-living biomass with size more than organic matter (2 mm) but less than the minimum diameter considered as for deadwood (e.g. 10 cm), deadwood under various stages of degradation of organic or mineral soil Includes organic carbon applied regularly over the course of the time series at a predetermined depth in mineral soils where they cannot be differentiated from it empirically, live and dead root system in the soil (less than the proposed diameter limit for belowground biomass)

(17.47), agri-silvi-horti system (17.02), Leucaena leucocephala (15.68), Acacia albida (15.23), Eucalyptus camaldulensis (13.22), Tectona grandis (12.54), Dendrocalamus strictus (11.65), Azadirachta indica (11.43), and agricultural land (9.4). Agri-silviculture system has high (4.23) carbon mitigation potential as compared to that in fallow land. The details of the carbon sequestered in different agroforestry based land use systems are presented in Table 9.7.

9.8

Agroforestry and Geospatial Technologies

The CARI (Central Agroforestry Research Institute) in Jhansi, Uttar Pradesh, has designed a procedure for mapping and calculating the area covered by agroforestry system by using medium resolution remote sensing data. First, a supervised approach must be used to classify the land use and cover of the LISS III data. Then, from this categorised image, crops and fallow land are recovered for use in masking an analogous region from the false colour composite. Even when many features or land covers are present, pixel-based approaches only take into account the single significant characteristic that occurs in a pixel. Additionally, using pixel-based approaches could result in certain incorrect classifications. Agroforestry only occurs on agricultural land; hence the sub-pixel classification approach was used there (Rizvi et al. 2019). The benefit of utilising a sub-pixel classifier is that it not only solves the issue of sugarcane and other young plantations crops intermixing, but it also provides results in the form of percent tree canopy cover within pixels. All sorts of agroforestry systems, including boundary plantations trees, scattered trees on agriculture fields, and block plantations on farm are included in this tree cover (20–100%). Using the produced signatures to apply a sub-pixel classifier to the

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Table 9.7 Total carbon sequestrated in various agroforestry systems of India (Anon., 2007) Zone Semi-arid

Agroforestry system Silvipasture (5 years old)

Northwestern India

Silvipasture (6 years old)

Central India

Block plantation (aged 6 years) Agri-silviculture (aged 8 years)

Arid zone

Semi-arid

Agri-silviculture (aged 11 years)

Components Acacia nilotica + natural pasture Acacia nilotica with established pasture Dalbergia sissoo + natural pasture Dalbergia sissoo + established pasture Hardwickia binata with natural pasture Hardwickia binata + established pasture Acacia/Dalbergia/ Prosopis + Desmostachya Acacia/Dalbergia/ Prosopis + Sporobolus Gmelina arborea Emblica officinalis + Vigna radiata Hardwickia binata + Vigna radiata Colophospermum mopane + Vigna radiata Dalbergia sissoo + crop

Total carbon storage (ton C/ha) 9.5–17.0 19.7 12.4 17.2 16.2 17.0 6.8–18.5 1.5–12.3 24.1–31.1 12.7–13.0 8.6–8.8 4.7–5.3 26.0

extracted agricultural region, the resulting image will contain pixels from five categories. Icons that cover trees and cropland, II icons that cover fallow land and trees, III icons that cover trees only, IV icons that cover cropland only, and V icons that cover fallow land only. Agroforestry will be accurately represented by the first three kinds of pixels, which contain dispersed trees, border plantings, and block plantations (Ellis et al. 2004; Rizvi et al. 2020).

9.9

National Agroforestry Policy

Major policy initiatives, such as the National Forest Policy of 1988, the National Agriculture Policy of 2000, the Planning Commission Task Force on Greening India in 2001, the National Bamboo Mission in 2002, the National Policy on Farmers in 2007, and the Green India Mission in 2010, emphasise the importance of agroforestry for effective nutrient cycling, the addition of organic manure for sustainable

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agriculture and for improving canopy cover. However, due to a number of circumstances, agroforestry has not attained the required prominence as a strategy for resource development. In February 2014, the cabinet approved a policy that addresses issue the agroforestry sector faces, such as unfavourable legislation, limited markets, and a lack of institutional financing. India became the first nation in the entire world to establish a thorough agroforestry policy. Basic Objectives • Promote tree planting and extend it in a complementary and integrated way with crops and livestock to boost rural family production, employment, income, and livelihood, especially that of small-scale farmers. • To reduce the danger during catastrophic climatic events, safeguard and stabilise ecosystems and encourage resilient agricultural and farming systems. • Meet the needs of wood-based industrial raw material and lessen the outsource purchase of wood and wood-related by-products to conserve foreign revenue. • Increase the rural and tribal inhabitants’ access to agroforestry products such as food, fodder, firewood, non-timber forest products and timber, easing pressure on existing forests. • Support the effort to increase forest and tree cover, particularly in areas that are vulnerable to it, in order to promote ecological stability. • To accomplish these goals and reduce pressure on already-existing forests, it is important to build up agroforestry research capability, strengthen agroforestry research, and mobilise a large-scale people’s movement.

9.9.1

Tree-Based Farming in the Country

The government is supporting tree-based farming to maximise agricultural yields and give farmers a sustainable means of subsistence, which includes 1. To promote the planting of trees along with crops and cropping systems on farmland known as “Har Medh Par Ped,” a sub-mission on agroforestry was established in 2016–2017. On farms, there is a provision for financial support for nursery development and plantation, which will increase the farmer’s income and increase the farming system’s resilience to climate change. 2. The NMOOP (National Mission on Oilseeds and Oil Palm), a National Food Security Mission division, promotes tree-borne oilseed intercropping, maintenance, and planting. Olive, neem, karanja, Mahua, and others. 3. The National Bamboo Mission (NBM) was restructured and launched in 2018–2019 with the goal of creating a full value chain, including plantations on public and corporate lands that is not under forest area as well as a market for growers.

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179

Tree Insurance (Vriksh Bheema)

Tree insurance is one-way farmers may sustain farm revenue and investment while protecting themselves from the terrible effects of losses caused by natural disasters or poor market pricing. Giving farmers a bare minimum of protection lessens the impact of crop losses. Single-peril coverage and multi-peril coverage are the two main types of agriculture insurance. Protection against a single hazard is provided by single-peril coverage, while multiple perils provide protection against many perils. A multi-peril tree insurance programme is being developed in India due to the damaging impacts of nature on society and the agricultural sector. Farmers typically lose a lot of money due to things like fires, cyclones, floods, etc., which damages years of labour. This insurance policy can serve as a support system for these farmers. The average premium rate for a fundamental plan would be 1.25% of input costs. According to the policy, depending on the input cost of the relevant tree species, the premium for a plantation of 1 acre would cost between Rs. 300 and Rs. 600. It would cover losses caused on by invasions by wild animals as well as losses from forest and bush fires, lightning, riots and strikes, storms and cyclones, floods and inundations. If any of the aforementioned hazards caused damage, the farmer would be compensated with the input cost of the appropriate trees. According to the covered crop, the sum insured will be calculated using the cost of cultivation, input cost, or the cost of raising/developing the insured tree(s), whichever is appropriate. The first step in promoting climate savvy and resilient technology is tree insurance. This plan is anticipated to make it easier to expand the area covered by agroforestry and provide a consistent supply of high-quality raw materials to the sector. As a by-product of the programme, carbon sequestration and the preservation of India’s reserve forests would be achieved. Our farmers are the most at risk for climate-related problems, and tree insurance will serve as a safety net to safeguard and thrive our farming community.

9.10

Conclusion

Increasing production and effectively conserving resources through scientific agroforestry interventions promoting sustainable development is possible. Globally, agroforestry systems provide many benefits and a reciprocal link with livelihood and biodiversity in multifunctional landscapes. The cultivation of tree-borne oil seeds has been found to have the highest potential for creating jobs, followed by silvipasture systems. Agroforestry systems provide the climate change mitigation mechanism through CO2 assimilation, and sequestration provides biological and economic gains. Geospatial technologies can be used for area estimation to assess carbon sequestration potential at state or regional level under different agroforestry systems.

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References Anonymous (2007) Perspective plan: vision-2025. National Research Centre for Agro-Forestry (NRCAF), Jhansi Anonymous (2011) State of forest report. Forest Survey of India, Ministry of Environment and Forests, Govt. of India, Dehradun, New Delhi Anonymous (2013) NRCAF vision 2050. National Research Centre for Agro-Forestry, Jhansi Anonymous (2019) Annual report. All India Coordinated Research Project for Dryland Agriculture, University of Agricultural Science, Bangalore, p 98 Bhagwat SA, Willis KJ, Birks HJB, Whittaker RJ (2008) Agroforestry: a refuge for tropical biodiversity? Trends Ecol Evol 23(5):261–267 Chavan SB, Uthappa AR, Sridhar KB, Keerthika A, Handa AK, Newaj R, Kumar N, Kumar D, Chaturvedi OP (2016) Trees for life: creating sustainable livelihood in Bundelkhand region of central India. Curr Sci 111:994–1002 Elevitch CR (2004) The overstory book: cultivating connections with trees. PAR, Honolulu, HI Ellis EA, Bentrup G, Schoeneberger MM (2004) Computer-based tools for decision support in agroforestry: current state and future needs. Agroforest Syst 61(1):401–421 Ghosh A, Kumar RV, Manna MC, Singh AK, Parihar CM, Kumar S, Roy AK, Koli P (2021) Eco-restoration of degraded lands through trees and grasses improves soil carbon sequestration and biological activity in tropical climates. Ecol Eng 162:106176 Hala Naik AL (2003) Economic analysis of agro-forestry systems in central dry zone and south transition zone of Karnataka. Ph.D. thesis. University of Agricultural Sciences, Bangalore, p 134 Handa AK, Dev I, Rizvi RH, Kumar N, Ram A, Kumar D, Kumar A, Bhaskar S, Dhyani SK, Rizvi J (2019) Successful agroforestry models for different agro-ecological regions in India. CAFRI, ICRAF, Jhansi, New Delhi Lundgren B (1982) Introduction. Agroforest Syst 1:1–12 Narain P, Singh RK, Sindhwal NS, Joshie P (1997) Agroforestry for soil and water conservation in the western Himalayan Valley Region of India 1. Runoff, soil and nutrient losses. Agroforest Syst 39(2):175–189 Newaj R, Rizvi RH, Chaturvedi OP, Alam B, Prasad R, Kumar D, Handa AK (2017) A country level assessment of area under agroforestry and its carbon sequestration potential. Tech Bull 2(2017):1–48 Ramulu I, Ramachandrappa BK, Maruthi Sankar GR, Sathish A, Sandhya Kanthi M, Archana AM (2017) Assessment of changes in soil infiltration, water-holding capacity, bulk density and fertility parameters under different tree-and crop-based systems in semiarid alfisols. Commun Soil Sci Plant Anal 48(5):477–500 Reddy NS (2002) Evaluation of different land use systems in carbon sequestration. M.Sc. (Agri.) thesis. Acharya N. G. Ranga Agricultural University, Hyderabad, p 112 Rizvi RH, Newaj R, Handa AK, Sridhar KB, Kumar A (2019) Agroforestry mapping in India through geospatial technologies: present status and way forward. Tech Bull 1(2019):1–35 Rizvi RH, Alam B, Handa AK, Chavan SB, Prasad R (2020) Mapping of tree species and assessment of area under agroforestry systems in Karnataka, India. Indian J Agroforest 22(2): 16–20 Roger RBL (1998) Agro-forestry for biodiversity in farming systems. In: Collins WW, Qualset CO (eds) Biodiversity in agroecosystems. CRC Publishers, New York, NY, pp 127–145 Roy MM, Tewari JC, Ram M (2011) Agroforestry for climate change adaptations and livelihood improvement in Indian hot arid regions. Int J Agric Crop Sci 3(2):43–54

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Samra JS, Narain P (1998) Soil and water conservation. In: Singh GB, Sharma BR (eds) 50 years of natural resource management research. ICAR, New Delhi, pp 145–176 Subbulakshmi V, Sheetal KR, Renjith PS, Keerthika A, Gupta DK (2019) NTFP based agroforestry to sustain income and employment generation activities of arid regions of Rajasthan. Jaya Publishing House, Delhi Tacio D (1993) Sloping Agricultural Land Technology (SALT): a sustainable agro-forestry scheme for the uplands. Agroforest Syst 22:145–152 Uthappa AR, Chavan SB, Dhyani SK, Handa AK, Newaj R (2015) Trees for soil health and sustainable agriculture. Indian Farm 65(3):2–5

Climate Change Impact on Forage Characteristics: An Appraisal for Livestock Production

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Pooja Tamboli, Amit Kumar Chaurasiya, Deepak Upadhyay, and Anup Kumar

10.1

Introduction

Exponentially increasing human population results in increased demand for agriculture and livestock products. World human population may accelerate up to 9.8 billion by 2050 and 11.2 billion in 2100. This consequently may lead to increase demand by 70% approximately, for agricultural and livestock products (UN 2017). Rural farmers mainly rely on agriculture and livestock farming for their livelihood security. Animal husbandry contributes to regular employment generation and socioeconomic security. Further, both the sectors, i.e., animal husbandry and agriculture are climate sensitive. Climate change possesses tremendous challenge to animal husbandry and agriculture sector at global level. Livestock production suffers from both direct and indirect effect of climate change. Direct effect includes stresses due to meteorological variables like temperature, humidity, wind, rainfall, and solar radiation. Conversely, indirect effects consist of deterioration of forage characteristics, shortage of animal feed, water, grazing land, and vector-borne diseases. Different varieties of forage crops are grown in various agroclimatic regions in our country. Forage pastures are thought to make up around 26% of the total land area and about 70% of the agricultural area, according to the Food and Agriculture Organization (FAO 2010). Although forage grasslands are a costeffective source of nutrients for livestock production, they also contribute to the preservation of the quality of the air, the water supply, and the soil (Chaudhry 2008). Long-term environmental changes impair forage characteristics. It causes alteration in growth rate and forage species components in the pasture (Thornton et al. 2015).

P. Tamboli · D. Upadhyay · A. Kumar Plant Animal Relationship Division, ICAR-IGFRI, Jhansi, UP, India A. K. Chaurasiya (✉) Department of Animal Nutrition, NDVSU, Jabalpur, MP, India # The Author(s), under exclusive license to Springer Nature Singapore Pte Ltd. 2023 R. K. Singhal et al. (eds.), Molecular Interventions for Developing Climate-Smart Crops: A Forage Perspective, https://doi.org/10.1007/978-981-99-1858-4_10

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Climate change drives threats to natural resources available in the earth. Fluctuating ambient temperature, relative humidity, air pressure, etc. causes deterioration of nature and natural resources. Further, increasing temperature causes accumulation of cellulose, hemicellulose, and lignin in the plant species (Polley et al. 2013) and hampers their quality. This low-quality fodder is poor in digestibility projecting to less nutrient absorption in animal body (Thornton et al. 2009). Further, occurrence of natural calamities like floods and drought creates breakdown of plant root, impairs plant structure and function which lowers vegetation quantity (Baruch and Merida 1995). These cumulative effects tend to cause degradation of soil, alter rainfall patterns, and reduce availability of water. India is bestowed with livestock population with 536.76 million, showed an increase of 4.6% over Livestock Census-2012. The huge expanding animal numbers exert pressure on feed and fodder. Further, according to literature, there is deficiency of green fodder by 11.24% and dry fodder by 23.4% (Roy et al. 2019). Also, there is increase in energy expenditure and reduction in feed intake as well as digestibility of fodder during hot summer season. Seasonal fluctuation causes reduction in the forage quality, thereby possessing threat on the performance and survival of livestock. This creates undernutrition and nutritional imbalance in the animal body which results in decrease in growth, production, and reproduction performance of animals. This is major problem in arid or semi-arid region where animals are managed in extensive systems by the rural farmers. Thus, local governments need to take some efficient adaptation strategies, such as setting the grazing intensity according to the grassland forage production and forage quality to avoid rangeland degradation, especially in dry regions. Conservation of biodiversity, environment, and natural resources is need of the hour to improve health and overall performance of animals. Therefore, proper management of the grassland in the direction of creating diversity in the vegetation with good quality nutrient rich fodder and maintaining forage availability is pre-requisite to promote sustainable, economical, and climate-smart livestock farming. Efforts to mitigate impact of climate change will contribute to increasing profitability of the animal farms by accelerating productivity. Keeping these points in view, the chapter summarises the impact of climate change on the forage characteristics and availability in the prospect of livestock production as well as effect on soil and water. In addition, it also encompasses management protocols for grazing land, climate smart feed and fodder production, as well as strategies for animal adaptation to climatic stress through nutritional interventions.

10.2

Environmental Variation: Implications in Natural Resources

Global population expansion is concurrent with a threat to soil and water sources from resource depletion and climate change. In particular, precipitation, evapotranspiration, temperature, stream flow, ground water, and surface runoff are all impacted by climate change, according to a large number of academics. Increased floods and

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drought brought on by these changes will have a profound influence on the availability of soil and water resources. Intricate relationships exist between soils and the climate system because of the nitrogen, carbon, and hydrologic cycles (Daba et al. 2018). Climate change will have an impact on soil processes and qualities. Because carbon and nitrogen are substantial components of soil organic matter, climate change and its hydrological effects may significantly alter soil conditions (Brady and Weil 2008). The amount of organic matter in soils has a significant impact on soil water retention. Additionally, the amount of water in the soil has a significant impact on crop productivity, plant growth, and the health of the soil ecosystem (Horel et al. 2022). Changes in worldwide rainfall volumes and distribution patterns will accompany changes in temperature. The most significant long-lived greenhouse gases are nitrous oxide (N2O), methane (CH4), and carbon dioxide (CO2) (Hansen et al. 2007). The principal greenhouse gases, CO2, methane, and nitrous oxide are on the rise. Additionally, during the coming decades to centuries, there will be brief variations in temperature, precipitation, and other climatic variables. The most trustworthy indicator of global atmospheric change is the rising CO2 content in the atmosphere. Level of atmospheric CO2 can reach between 421 and 936 ppm by the year 2100 based on measurement of global emanations (IPCC 2013). Due to an increase in greenhouse gas emissions and global deforestation, the average global temperature is predicted to rise by at least 1.5 °C by 2050 (Arora 2019; IPCC 2022). An increase in air temperature could result in a 3–15% rise in evaporation and transpiration rates. A warmer environment could also hold more water (Ragab and Prudhomme 2002). Temperature gradients between the soil surface and the atmosphere also rise with warming (Xue et al. 2014). This resulted in increased evaporation and decreased soil moisture in the upper layers, where the majority of plant roots are found. Regarding soil temperature and water content, warming can create an ecosystem below ground that is less favourable for some plant species (Li et al. 2018). An increase in temperature may encourage plant development, but the resulting decrease in soil moisture may counteract the beneficial effects on plant biomass (Li et al. 2018). The climate is substantially impacted by changes in land use. Through photosynthesis, vegetation absorbs carbon dioxide, while soils serve as carbon sinks. The loss of vegetation can impair photosynthesis and the ability of the land to operate as a carbon sink by causing soil erosion, nutrient leaching, and soil deterioration. Nearly 20% of the earth’s surface is publicly accessible for animal grazing or overgrazing, and these regions are sometimes referred to as common property resources. Overgrazing can result in soil erosion, which lowers organic matter levels, soil fertility, and water-holding capacity. More carbon dioxide or greater GHG levels are added to the atmosphere as a result of the soil’s diminished capacity to store carbon dioxide.

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Climate Change Impact on Forage Crops

Adverse climate leads to considerable reduction or complete removal of forage crops. Extreme environment hampers both fodder and concentrate crops by reducing their yield, quality, and ultimately price, thereby influence feed platform of the animal farm. According to a number of studies, climate change may have an impact on the quantity and consistency of forage production, forage quality (Thivierge et al. 2016), thermal stress on livestock, water requirements for both growing forage and meeting animal needs (Lacetera 2019), and the structure and composition of largescale plant communities (Yang et al. 2017; Lee 2018). Additionally, the physical and chemical properties of forages are altered by climate change, which also has an impact on the digestive systems (Dumont et al. 2015). Dry matter (DM), crude protein (CP), neutral detergent fibre (NDF), acid detergent lignin (ADL), acid detergent fibre (ADF), ether extract (EE), mineral ash (Ash), and the digestibility of dry and organic matter (DMD and OMD, respectively) are some of the agronomic nutritional parameters that are likely to be impacted by climate change (Lee 2018). According to research by Dumont et al. (2015), increased temperature increases the NDF, ADF, and ADL contents while decreasing the CP and digestibility contents (Melo et al. 2022). Elevated CO2 lowered forage nitrogen (N) content by 8% while increasing forage tissue’s total non-structural carbohydrates by an average of 25% (Dumont et al. 2015). Various fodder crops are cultivated in various agroclimatic areas of India. Forage quantity and quality are impacted differently by temperature changes, water availability, and an increase in carbon dioxide. The region in question and the length of the growing season have the greatest influence on temperature effects (Polley et al. 2013) For instance, a 2 °C rise in temperature is predicted to limit the availability of feed in dry and semi-arid regions while improving it in humid temperate regions. Plants that use the C3 carbon fixation pathway, such as wheat, rye, oats, rice, cotton, sun sunflower, and chlorella, to convert carbon from CO2 into organic equivalents, experience alterations in their growth as a result of the higher CO2 concentration (Thornton et al. 2015). Positive effects on C3 species result from the induction of partial stomatal closure and reduction in transpiration (Wand et al. 1999). Changes in optimal growth rates caused by changes in temperature and CO2 levels change the dynamics of species competition, which impacts the composition of pastures (Thornton et al. 2015). The supply of water varies throughout the year, which has an impact on this rivalry. Increases in temperature are likely to cause lignin and other cell wall constituents to accumulate in plants (Polley et al. 2013), which affects digestibility and reduces cattle access to nutrients (Thornton et al. 2009). With increased frequency of floods and droughts as well as changing rates of leaf growth, it is anticipated that roots’ structure and function will change, resulting in lower yields (Baruch and Merida 1995). According to Xue et al. (2017), warming can make soil drought stress on plant growth under dry settings worse. Additionally, in locations with little soil moisture, this can result in a decline in grass species but an increase in forb species (Li et al. 2018). A significant ecosystem service of agricultural grasslands is the provision of forage for cattle and dairy animals. Forage provision has two components: quantity (yield or production)

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and quality (the nutritional value for livestock), which together determine carrying capacity and performance of cattle (Schauer et al. 2005; Beeri et al. 2007). Both the intensity of land management and ongoing climatic changes are likely to alter the availability of fodder (Martin et al. 2014). Maintaining a suitable amount of energy and nutrients for animals from plants depends on the quality of the forage. Reduced economic value of grasslands and even a threat to food security might result from failure to mitigate and adapt agriculture to changing environmental conditions (Berauer et al. 2020). We must comprehend how climate and land use affect fodder quality if we are to sustain the ecological and economic benefits of grasslands in the face of future change. Forage quality in grasslands improves as protein and fat content rises but declines as fibre level rises (Xu et al. 2018). Forage quality is influenced by species composition and abundance (Khalsa et al. 2012) and the availability of soil resources (Niu et al. 2016), both of which are influenced by climate factors and the intensity of land use. In the end, land-use intensification weakens ecological stability due to homogenisation of landscapes and biodiversity loss (Blüthgen et al. 2016). Additionally, land-use intensity has a negative impact on the chemical, physical, and biological characteristics of soil, including nutrient mining, compaction, and soil biodiversity (Smith et al. 2016). In addition to increasing gross nitrogen turnover and changing above- and below-ground plant communities (Blankinship et al. 2011), climate change also reduces soil organic carbon (Puissant et al. 2017) and gross nitrogen turnover (Wang et al. 2016). Therefore, there may be both positive and negative interactions between land management and climate change (Karlowsky et al. 2018). Nutrition is a significant predictor of intestinal methane output and is favourably correlated with weight gains, milk production, and reproductive potential. The nutritional value of the forage was decreased at higher temperatures and boosted by the addition of nitrogen fertiliser. Elevated temperatures decrease the nutritional content of grass, and may, as a result, cause an increase in the generation of bovine enteric methane of 0.9% with a 1 °C temperature rise and 4.5% with a 5 °C temperature rise. Seasonal fluctuations have an impact on the nutrients in forage, which have an impact on feed intake, digestibility, and energy released to livestock after intake (Adesogan et al. 2006).

10.4

Livestock Sector Vis-a-Vis Climate Change

10.4.1 Consequences of Climate Change on Livestock Climate change has become a major international issue, as evidenced by the United Nations General Assembly’s creation of the Intergovernmental Panel on Climate Change (IPCC) (Resolution 43/53, 1988). The scientific community has worked to improve its understanding of the underlying mechanisms governing the Earth’s climate system as well as the implications and effects of climate change throughout the last few decades. Agriculture consumes around 70% of freshwater resources globally, making it the major consumer of these resources. By 2025, it is anticipated that 64% of the world’s population will have to work in water-stressful conditions

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due to the competition in global water consumption. Issues with water supply are hitting all industries extremely severely, and this includes the animal industry (Thornton et al. 2009). A rise in temperature is increasing animal water use by a factor of 2–3, accounting for around 8% of all human water usage in the animal husbandry sector (Nardone et al. 2010). As sea levels rise, more salty water is leaking into freshwater aquifers along the shore (Karl et al. 2009). The animals’ metabolism, digestion, and fertility are negatively impacted by the salinity of the water. In addition to reducing production hygiene, chemical pollutants and heavy metals damage essential organs connected to important bodily systems (Nardone et al. 2010). The direct consequences are brought on by a rise in temperature, which also raises the rates of morbidity and mortality. The indirect repercussions include microbial proliferation, an uptick in vector-borne and food-borne illnesses, a decline in host resistance, a shortage of fodder and water (Thornton et al. 2009). Animal disease outbreaks are considerably exacerbated by erratic rainfall and temperature changes. Disease outbreaks, such those caused by foot and mouth disease, haemorrhagic septicemia, and avian influenza, have a negative impact on a large number of animals, the environment, and the health and way of life of the surrounding community. Animal health and welfare are undoubtedly impacted by heat stress. The THI index, also referred to as the discomfort index, or the combined impact of temperature and humidity, is a significant factor in the decreased milk productivity. The dairy industry is impacted by heat stress, which lowers milk output and quality, which has an impact on cheesemaking as well. Additionally, the casein and milk protein contents both tend to decline. The main factor contributing to dairy cow’s reduced conception rates and ineffective reproduction is heat stress. Reduced oocyte competence, which in turn affects the development of the resultant embryo, is the main cause in reproductive inefficiency brought on by heat stress (Naqvi et al. 2012). It has been noted that dairy cow conception rates can decrease by up to 20–27% during the summer. The populations with low genetic diversity are at risk because climate change has the potential to have an adverse effect on biodiversity and can result in the extinction of 15–37% of all species worldwide (Thomas et al. 2004).

10.4.2 Livestock Role on Climate Change Due to the increased market demand for milk and other animal products, the population of livestock is steadily growing. Due to intense production pressure, the number of buffaloes and crossbred cattle is constantly rising, while the number of native cattle is steadily declining. Most of the available feed for ruminants consists of unusual sources or crop leftovers that cannot be readily broken down by enzymes. In the course of rumen fermentation, hydrogen is produced (4H+ during the synthesis of acetate and 8H+ during that of butyrate in the rumen wall), which when combined with carbon dioxide yields methane. Thus, the creation of methane is a crucial component of enteric fermentation. Animals directly responsible for climate change excrete undigested food as manure. Two significant sources of greenhouse gas

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emissions from livestock are enteric and anaerobic fermentation of farmyard manure, which produce CH4 and N2O (Sejian et al. 2012). Globally, livestock accounts for around 15% of the anthropogenic-induced GHG generation, producing about 7.1 Gt CO2 eq GHGs annually. Additionally, 5.7 Gt CO2 eq GHGs are generated by ruminant animals. Methane emissions from enteric fermentation amount to around 90 Tg, while those from excreta fermentation amount to about 25 Tg (Sejian et al. 2016). Our nation produces 15.1% of the world’s methane through livestock. Methane from enteric sources makes for 91.8% of all greenhouse gas emissions, followed by methane from manure (7.04%), and nitrous oxide (1.15%) from manure.

10.5

Strategies for Climate Smart Feed and Fodder Production

Climate change impacts the crop to a greater extent. Higher ambient temperature causes accelerated growth which consequently causes reduction in life cycle of forage crop. Therefore, it is essential to grow the varieties whose growth season is longer in order to maximise yield as the same crop area can be harvested many times in the particular year (Babinszky et al. 2011). Photosynthesis is of three types: C3, C4, and CAM (Crassulacean Acid Metabolism). Majority of the plant follows the C3 photosynthesis pathway. The crop evolved with C4 and CAM photosynthesis are more adapted to arid and semi-arid climatic conditions. In comparison to C3 plant, rate of photosynthesis is faster under high temperature and light in C4 plants, while energy and water retaining capacity is higher in CAM plants during harsh weather. Hence, C3 plants are more susceptible to alteration in temperature or precipitation, accordingly their drought-resistant varieties should be promoted. Nutritionists should incorporate drought-resistant crop varieties such as bajra-Napier hybrid in the feed formulation to overcome environmental pressure on livestock health at the same time ensuring adequate production, quality, and safety of animal products. Fodder production technologies should focus on the utilisation and selective breeding of the hybrid crop or varieties which are more resilient to adverse climate like drought, extreme hot or cold weather conditions. As described earlier, the C4 forages (sorghum, millet, and corn) are more impervious to climate change than those of C3 crop (oat, barley, wheat, sunflower, alfalfa). Variation in the plant species, yield and their characteristics as a result of climate change forces to spread this information to a wider scale up to farmers, researchers, policy makers, entrepreneurs, and other stakeholder for applicable ecosystem management, action plan, and other pertinent resolutions to circumvent impact of changing climate. Different Brachiaria grass cultivars are more tolerant of soils with low fertility and drought. Grass that is incredibly nourishing, tasty, and digestible. Brachiaria grass decreases GHG emissions, improves nitrogen use efficiency, and alleviates feed shortages for animals. It is always thriving, offers a steady source of animal feed, and is simple to dry and store as hay (ILRI 2016). In extensive systems, correct diet formulation can be influenced by both managing grazing and boosting forage quality by switching up the forage species. This can significantly boost feed efficiency and

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productivity. Precision feeding, which integrates the animal’s genetics with feeding and grazing management, calls for cutting-edge technological infrastructure to precisely monitor the animal’s needs and appropriately manage pasture and forage production. It can be implemented in high-value farm systems that use highly technological systems (Garg 2013). Legumes used as feed can be interplanted with grains to increase forage and, in turn, livestock productivity. It has been demonstrated that this intercropping increases the quantity and quality of feed and crop leftovers, improving system effectiveness (Ayarza et al. 2007). Legumes may contribute to the decrease in GHG emissions from ruminant systems. Methane emissions are decreased by using forages that contain tannin and breeding forage species with higher tannin content (Belay 2019). To guarantee a steady supply of feed for animal husbandry, improved fodder management practices are crucial. Droughts may result in a lack of feed and animal losses if pasture productivity is solely dependent on rainfall. A surplus of forage cannot be used as feed in drought conditions without storage methods like creating hay or silage. In addition, stall feeding rather than grazing is made possible by the production of hay and silage, which gives meadows time to regenerate and increases productivity due to the higher nutrient content. But before that, a good pasture management is crucial to ensure good quality forage production. Some species such as Desmodium or Napier grass function as a natural pest control. Adequate fodder species also protect the soil below functioning as a vegetation cover that prevents erosion and drying out. The removal of a vegetative cover and soil erosion reduces not only the pastureland’s productivity but at the same time its capability to sequestrate carbon dioxide. Thus, it further releases soil-carbon into the atmosphere and fosters climate change (FAO 2017). The three foundational tenets of climate smart agriculture (CSA) are promoted by upgraded fodder management methods, which include: 1. Improved soil cover and high-yielding, high-quality pastures can boost animal productivity and, consequently, income, if they are used effectively. 2. Increasing the ecosystems’ and livelihoods’ resistance to climatic extremes: The resistance of livestock keepers to drought is increased by providing sufficient, less drought-affected feed species. A certain stock is still accessible to withstand the hardest drought months since to storage techniques like making hay or silage, even if the feed crops dry out. 3. Reducing and eliminating GHG emissions from the atmosphere: The ability of pastureland to sequester carbon dioxide is protected through fodder management that preserves vegetative cover and prevents soil erosion. High-yielding, superior pastures lower ruminant methane emissions even more. Additionally, FAO anticipates significant reductions in CO2 emissions and very high beneficial effects on nitrogen (FAO 2017).

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Climate Adaptation Protocol Addressing Nutritional Interventions

Reverberation of climate change results in drastic change in weather condition all over the globe which is consequently responsible for frequent occurrence of drought, salinity, storm, and irregular changes in ambient temperature. Animal, mainly small ruminants under extensive or pastoral systems experiences nutritional tension. Therefore, it is very essential to implement remedies not only in agriculture but also in livestock sector to combat stress and loss due to extreme weather. Development of various strategies to maintain and augment production potential of livestock to meet demand of rapidly accelerating human population irrespective of vagaries in environment is imperative in the present era. It is necessary to provide proper nutrition, follow suitable health care and other management practices in the animal farm to protect them from environmental stress because forage availability and their quality decline due to changing climate scenario (Soren 2012). After carbon dioxide, methane is the second-most significant gas, and it accounts for roughly 16% of all GHG emissions. The greenhouse effect is 25 times more powerfully induced by methane than by carbon dioxide. Apart from enteric CH4’s contribution to global warming, cattle eject 250–500 L of methane each day, which resulted in a loss of biological energy (between 2% and 15% of intake) that the host animal would have otherwise used for a variety of productive activities. In developing nations like India where feed and fodder are already in short supply, reducing energy loss in the form of enteric CH4 is essential. Therefore, ruminant methane output must be reduced for both environmental and economic reasons. Modifying the meal will change the amount of ruminal methane generation. Increased feed quality results in a significant decrease in methanogenesis.

10.7

Mitigating the Impact of Livestock on Climate

More methane is produced by ruminants fed low-quality roughages than by those fed high-quality roughages. Forage can be processed into feed by chopping, grinding, or pelleting, which increases digestibility and reduces fermentation and methane output. Plants produce more methane and have a higher fibre content as they get older. Crop residues treated with urea increase their quality and digestibility, which lowers enteric methane emission (FAO 2017). More methane is produced by non-legume than by legumes. Methane generation will be reduced as a result of silage preparation of green feed. Concentrate quality and amount have a detrimental impact on methane output because they slow down rumen fermentation. Rumen-protected, protein- and carbohydrate-enriched concentrate lowers animal methane output. Supplementing with lipids reduces rumen fermentation and, thus, methane output. The best saturated fatty acids for reducing methane generation are medium-chain (C8-C14). However, the use of lipids for methane mitigation is constrained by their high cost and detrimental effects on milk fat. Exogenous enzymes that increase fibre digestibility, such cellulase and hemicellulase, often reduce the acetate:propionate

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ratio in the rumen, which ultimately lowers methane generation. Condensed tannins and saponins, two plant secondary metabolites, are primarily linked to antibacterial properties that destroy bacteria, protozoa, and fungi in the rumen, lowering methane production.

10.8

Mitigating the Impact of Climate on Livestock

Heat stress in livestock is brought on by global warming, which raises the ambient temperature. It is important to employ some nutrition-based interventions for climate-smart livestock husbandry. A balanced diet offers enough energy to lessen the issue of herd health and reproduction brought on by decreased total digestible nutrients (TDN) and dry matter consumption (DMI) intake during heat stress. To boost DMI, fresh, delicious feed should be offered in the meal. High digestible elements for feed should be chosen as they reduce the animal’s production of heat. Intake of dry matter is encouraged by well-balanced total mixed ration diet formulations at a minimum fibre level. Animals under stress require carbohydrates that may easily be fermented. Additionally, giving higher unsaturated fatty acids (up to 5% of total DMI) to animals under heat stress has demonstrated advantages. Because heat-stressed calves frequently have a negative nitrogen balance, it is important to increase the quality and quantity of protein in the diet. To satisfy the animal’s need for dry matter, feeding frequency should be increased. When under heat stress, one drinks 20–50% more water. Therefore, the animal should always have access to fresh, cool water along with enough watering facilities. Increased dietary mineral concentration is necessary in hot weather due to decreased dry matter intake and elevated lactation demand. Antioxidants like vitamin A, vitamin E, selenium, and zinc are fed to animals to boost immunity and reduce heat stress. To maintain a normal rumen environment by reducing the incidence of acidosis in the rumen, which is a regular incident during hot weather, feed ingredients should include the buffering capacity or some buffers such as sodium bicarbonate, magnesium oxide, and sodium sesquicarbonate. On the other side, the body tries to make up for the increased heat loss experienced during cold stress by increasing the pace of heat generation, which includes using more maintenance energy. However, heat is also produced during the process of breaking down and converting food into energy (the thermic effect of diet), which helps to keep body temperature stable in situations below the lower critical temperature. As a result, feeding animals diets with a high thermic effect will help the animals survive in an environment that is too cold. So, for instance, a significant amount of energy is lost as heat when high-fibre foods are fermented by colon bacteria; similarly, oxidising proteins or amino acids to produce energy also generates a significant amount of heat. As a result, animals that consume feeds rich in fermentable fibres or extra protein produce more heat when the weather is cold. However, in actual feeding, overfeeding on protein is not advised from an economic or environmental standpoint (Babinszky et al. 2011).

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Conclusion

In conclusion, it can be said that we should anticipate long-term environmental changes brought on by climate change, which will have an impact on the growth of forage crops and farm animal productivity. It is imperative to develop heat-tolerant plant cultivars. Crops that have developed with C4 and CAM photosynthesis are more suited for dry and semi-arid climates. Because C3 plants are more sensitive to changes in temperature or precipitation, it is important to cultivate drought-resistant cultivars of these plants while still preserving their average yields and nutrient contents. It will be our responsibility as nutritionists to formulate diets using these better feed crop kinds in a highly targeted and expert manner. Globally, the productivity and profitability of the production of grasslands and livestock are projected to be negatively impacted by atmospheric and climate change. Enteric methane emissions per unit of meat or milk are comparatively large when feeds are of poor quality and are difficult to digest. Through better grazing land management, improved pasture species, changing forage mix, and increased use of feed supplements to achieve a balanced diet, including processing of crop residues and crop by-products, it is possible to improve feed digestibility and energy content as well as better match protein supply to animal requirements. These actions can enhance nutrient absorption, raise animal productivity and fertility, and hence reduce methane emissions per unit of production. Animal productivity and output quality, as well as the environment and the animal’s energy metabolism, are all closely related. We can manufacture wholesome meals that meet people’s needs using the knowledge outlined above without increasing the production’s environmental impact, so reducing the stress brought on by climate change.

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Sustainable Use of Paddy Straw as Livestock Feed: A Climate Resilient Approach to Crop Residue Burning

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B. R. Praveen, Manjanagouda S. Sannagoudar, R. T. Chethan Babu, G. A. Rajanna, Magan Singh, Sandeep Kumar, Rakesh Kumar, and V. K. Wasnik

11.1

Introduction

Agriculture growth is significantly influenced by livestock farming and livestock is not only a source of money for farmers but it also helps to produce food for the general people. Apart from livestock derived goods like dung as organic manure and fuel or biogas for household, livestock including cattle and buffaloes made significant role in milk production and sheep and goats are economically significant in production of meat. Livestock are able to completely rely on crops for nutrition in order to maintain normal growth, productivity and reproduction. Their rumen ecosystem dynamics offer a peculiar environment for microbes to flourish and proliferate in order to break down nutrients particularly fibrous materials from the fodder which is subsequently converted into nutrient-rich meals like meat and milk and it depends largely on the quality and quantity of the fodder or feed that is provided to the livestock. Due to a lack of practical solutions, farmers on the Indo-Gangetic plain are faced with the laborious task of managing extra paddy residue sustainably. Farmers eventually choose to carelessly burn it. Because paddy straws can be converted into value-added products and are nutrient-rich, sustainable residue management is

B. R. Praveen · R. T. C. Babu · M. Singh · S. Kumar · R. Kumar Agronomy Section, ICAR - National Dairy Research Institute, Karnal, Haryana, India M. S. Sannagoudar (*) ICAR - Indian Grassland and Fodder Research Institute, Jhansi, India ICAR - Indian Institute of Seed Science, Regional Station, Bengaluru, India G. A. Rajanna ICAR - Directorate of Groundnut Research, Anantapur, AP, India V. K. Wasnik ICAR - Indian Grassland and Fodder Research Institute, Jhansi, India # The Author(s), under exclusive license to Springer Nature Singapore Pte Ltd. 2023 R. K. Singhal et al. (eds.), Molecular Interventions for Developing Climate-Smart Crops: A Forage Perspective, https://doi.org/10.1007/978-981-99-1858-4_11

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crucial (Dutta et al. 2022). In India, paddy is grown on 43.8 million hectares (Mha), yielding 118.43 million tons (Mt) and 165.8 Mt of grain and straw, respectively. Due to its easy method of cultivation, short window for growing wheat, increasing mechanised harvesting and insufficient farm mechanisation for incorporation, low acceptance of paddy straw as fodder and lack of its practical utility, burning is the most popular strategy for handling paddy crop wastes. Each year, 50 Mt of paddy straw is burned with around half taking place in North-West India in the months of October and November. Residue burning produces particulate matter (1.5 Mt), carbon dioxide (150 Mt) as well as other GHGs and volatile organic compounds. This pollution causes a variety of respiratory illnesses in people, reduces soil nutrient and carbon status and disturbs soil biological activity (Kaur et al. 2022). Seasonal variations affect the accessibility of good fodder for livestock grazing. Generally, fodder is in plentiful supply during the wet season, whereas it is scarce during the dry season. Maiorella (1985) and Doyle et al. (1986) suggested that quantities of paddy grains and straw that can be harvested are the same for each hectare is the foundation for the anticipated amount of rice straw production. Paddy straw and other crop residues are readily accessible in huge numbers, shortly following the harvest seasons in many agricultural nations. These farm wastes are used for a variety of purposes, including the manufacture of mushrooms, board or paper, organic fertiliser, fuel and feed for livestock. To feed buffalo, cattle, goats and sheep, paddy straw is a commonly accessible, useful and affordable feed resource. Rice straw from a field is typically hauled and stacked by livestock farmers, who then use it to provide a reserve feed during the time of scarcity. It has been demonstrated that feeding livestock pure rice straw throughout their rapid growth and early lactation phases has an impact on the animal health and performance. High silica and lignin content is another factor for low accessibility of paddy straw as fodder along with low digestion of nutrients. Therefore, pre-treatment is required to maximise its impact on boosting the production of milk and meat. With a notable improvement in intake of feed, digestibility and performance of animals, technology based solutions have been innovated to maximise rice straw’s nutritional and feeding qualities.

11.2

Crop Residue Generation

Around 682.6 Mt of dry residual biomass are produced in India each year and 58.6% of this biomass is produced during Kharif season, 38.9% during the Rabi and 2.5% during summer season. Uttar Pradesh tops in residue generation with 19.1% during the Kharif followed by Maharashtra, Gujarat and Punjab (16.2%, 8.8% and 7.5%, respectively). Likewise, Uttar Pradesh remains first in Rabi season residue generation with 17% followed by Madhya Pradesh, Rajasthan and Bihar (12.7%, 10.7% and 8.5%, respectively) (Jain et al. 2018). Cereal crops and sugarcane account for the highest share of all residue formation in India. In Indo-Gangetic Plain (IGP), including the states of Uttar Pradesh, Punjab and Haryana contributes to national rice and wheat production at an extent of 27.13% and 57.66%, respectively, which

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Table 11.1 Crop residue production and burning estimation in India and world

India

Crops All crops

Paddy Asia

World

Total biomass All crops Paddy Paddy

Residue production (Mt) 350 682.6 683 178.5 225.5 –

Residue burning (Mt) – – 87 – – 730

– 826

250 –

980

–

References Mandal et al. (2004) Jain et al. (2018) Datta et al. (2020) Mandal et al. (2004) Jain et al. (2018) Streets et al. (2003)

Goswami et al. (2019) Goswami et al. (2019)

created crop residue formation from rice becomes a significant problem. In India, paddy is grown in 43.78 Mha, producing 118.43 Mt of grains (Ministry of Agriculture and Farmers Welfare 2020). According to Satpathy and Pradhan (2023), each Mega gram (Mg) of harvested rice grain yields about 1.4 Mg of straw during 2019–2020. In North-West India, paddy generates biomass up to 33% followed by wheat and sugarcane (22% and 17%, respectively) and around 84% of the world’s total production of paddy residue is generated in Asia (Jain et al. 2018; Goswami et al. 2019). The crop residue production in India and world is estimated as per Table 11.1.

11.3

Crop Residue Burning and Its Implication

Agricultural straw or crop residues leftover is purposefully fired up in the field after harvest and this method of residue management is known as stubble burning or crop residue burning. Burning crop residues is becoming less common worldwide particularly in developed countries because of strict regulations and market driven strategies. However, the scenario is different in South-East Asia, because there are not any practical and affordable alternatives or technologies to dispose of large amounts of biomass and their economic gains are likewise questionable (Shyamsundar et al. 2019). Majority of crop residue burning is done by farmers of South-East Asian countries and this practice has risen over time (Cassou et al. 2018). Each year, Asia burns about 730 Mt of biomass, primarily burning forests (45%), crop residues (34%) and grasslands (20%). In spite of financial support to manage residues, China tops in the total crop residue burning followed by India (Streets et al. 2003). Each year, India produces 683 Mt of crop residue, out of which 80% is employed as fuel, fodder and industrial raw materials. According to Datta et al. (2020), approximately 87 Mt surplus residues fired up in the crop fields. Out of which 60% was paddy straw and its burning estimated to 50 Mt each year

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Table 11.2 Major pollutants emission (Gg/year) from major crop residue burning in India (source: Ravindra et al. 2019) Pollutants Particulate matter (PM2.5) Particulate matter (PM10) Sulfur dioxide Carbon dioxide Carbon monoxide Nitrous oxide Nitrous oxides Ammonia Volatile organic compounds Elemental carbon Organic carbon Polycyclic aromatic hydrocarbons

Rice 418 458.29 9.07 59,275.74 4683.64 24.17 114.82 206.48 352.53 25.68 150.58 0.026

Wheat 264.57 175.35 12.31 54,974.27 816.38 22.76 52.30 39.99 215.34 4.92 8.92 0.04

Sugarcane 42.11 44.33 2.33 12,523.52 384.57 8.20 28.82 11.08 24.38 7.76 36.57 0.02

Cotton 11.66 13.01 0.63 4022.31 316.46 2.21 7.45 3.89 20.93 2.45 5.47 0.01

(Bhattacharya et al. 2021). In India, farmers practise residue burning of different crops including 8–80% of rice residues, 25% of sugarcane residue, 10–23% of wheat residues and 10% of maize, cotton, millets, jute and rapeseed-mustard (Jain et al. 2014). Table 11.1 depicts the estimation of crop residue burning in India and world. In North-West India, burning of paddy straw is estimated to around 23 Mt which was half of the total paddy residue burning in the country in order to make their agricultural land ready for following wheat sowing (NAAS 2017). According to the total number of burning incidents throughout India’s three largest states, Punjab has the highest followed by Haryana and UP. In the three states combined, there were around 83,000 fire accidents occurred during October-November 2021, which was 35.6% less than the number of events reported in 2016 and decreased by 30.6%, 55.8% and 56.6% in Punjab, Haryana and UP, respectively (CREAMS-IARI 2021). In Punjab, 20 Mt of paddy residue was produced during 2019 from which 9.8 Mt being fired up over 37.4% of area under paddy cultivation. Haryana produced 7.0 Mt of rice straws, of which 1.24 Mt was burned over 17.7% of the entire area used for paddy cultivation. About 16 Mt of rice straws were produced in Uttar Pradesh, but only 0.0042 Mt of that total were burned across 0.2% of the land used to grow paddy (EPCA 2020). Paddy residue burning takes place in the months of October and November, when there is more particulate matter (PM) in atmosphere because of extreme humidity, lower temperature and little breeze. So, in North-West India, air pollution and its negative effects still exist. The burning paddy stubble raises the PM concentration which results in a thick smog and every living thing in this area suffers greatly from the residual burning aftereffects. The harmful effects of this process on soil health include the loss of organic matter in the form of carbon dioxide (CO2) and nitrogen (N) in the form of nitrate (NO3-) (Kumar et al. 2019). The Punjab Agricultural University reported that residue burning resulted in 0.824 Mt losses of major nutrients from the soil annually. In addition, it has a negative effect on the biological

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activity of soil especially in the topsoil (top 2.5 cm) (Turmel et al. 2015). Stubble burning emits GHGs such as CO2, carbon monoxide (CO), nitrous oxides (NO, NOx), sulfur dioxide (SO2), methane (CH4) and hydrocarbons as well as other harmful radioactive chemicals as depicted in Table 11.2. Burning also releases suspended particulate matter (SPM) and volatile organic compounds (VOCs) into air, which can lead to persistent disruptive lung disorders and raise the conditions like eyes irritation, cataracts, corneal opacities and in due course sightlessness (Saxena et al. 2021). According to Abdurrahman et al. (2020) and Adam et al. (2021), polluting air brought on by burning of residue increases the chance of respiratory infections and heart illnesses. Burning of residue degrades environment and soil quality (Lohan et al. 2018), decreases the amount of nutrients available to plants (Ademe 2015), negatively impacts the soil’s microbiological and physicochemical structure and increases the likelihood of accelerated erosion (Kumar et al. 2015).

11.4

Paddy Straw as Livestock Feed Resource

In many agricultural nations across the world, rice producers create a lot of paddy straw and more than 90% of the livestock population in South-East Asia including China and Mongolia is typically fed with paddy straw which was roughly 30–40% of total region supply (Devendra and Thomas 2002). When the supply of high-quality forages is insufficient, paddy straw is the substance of the lean months for livestock. This was supplied as the main source of food to satisfy dietary requirement without meeting all the vital nutrients all alone. To increase palatability, intake, protein content and digestibility of the feed by the animals, paddy straw can be added to an extent of 60% in combination with remaining feed materials. Following rice harvest, livestock producers gather and stockpile rice straw in an uncomplicated shelter that is typically constructed from materials found nearby or stored in outside stacks. During the fodder scarcity period from January to May, the preserved residue is typically used as animal feed. The livestock population as well as the availability of forage gardens determines how much rice straw is used as fodder. Given that, the average farmer has just 1–2 ha of land, the estimated yearly production of rice straw is only 10–15 tonnes, from which it is possible to maintain 3–4 animal units either of buffalo or cattle. In order to achieve normal animal performance, if a farmer is willing to maintain more animals, he has to go for cultivation of additional green fodder, silage and hay making and other crop residues along with feed supplements.

11.5

Nutritional Quality of Paddy Straw

The protein level of rice straw ranges from 3% to 6% and it contains thick cell wall which is made up of degradable carbohydrate components like starch, cellulose, hemicellulose, lignin, acid detergent fibre (ADF) and neutral detergent fibre (NDF). Rice straw is typically utilised bulky feed to fulfil the stomach requirements and 80%

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Table 11.3 Proximate and mineral composition of paddy straw (source: Babu et al. 2022) Parameters Dry matter content Organic matter content Crude protein Crude fibre Acid detergent fibre Neutral detergent fibre Nitrogen free extract Cellulose Hemicellulose Ash Acid-insoluble ash Acid detergent lignin

Concentration (%) 88–96.87 83–90 2–6.5 30–40 49–73.01 39.83–85 40–46 31.96–60 13–32.24 11–16 3–5 4.63–13.2

Parameters Nitrogen Phosphorus Potassium Calcium Magnesium Sulfur Zinc Iron Copper Manganese Boron Silicone

Concentration (%) 0.65 0.10 1.40 0.52–2.24 0.91–2.21 0.075 0.02 0.04–0.45 0.005 0.07 0.001 4.25–13

of the materials in this are possibly biodegradable and act as an energy source. It contains more dry matter between 92% and 96%, but low crude protein (CP) (3–7%) concentrations (Shen et al. 1998). During the developing and fruiting stages, rice plant structure is maintained by silica and lignin composition, but when consumed by mammals, these ingredients take on an indigestible form. Due to low digestible nutrient contents, low palatability, changeable nutritional composition, high oxalates and silica content along with adulterants if improperly handled, it has a limited use. However, rice straw is still a useful source of animal feed, especially during El Nino events or other critical times when fresh animal feed is scarce. In addition, paddy straw is a better feed material if processed appropriately mainly during the unproductive period of milch animals. The proximate and mineral composition is depicted in Table 11.3. Paddy straw typically contains NPK (0.7%, 0.23% and 1.75%, respectively) in terms of nutrients. About 26.26 and 22.13 Mt of total nutrients are included in rice residue annually throughout Asia and the rest of the world (Goswami et al. 2019). Calcium, magnesium and potassium (0.53%, 0.24% and 1.58% of DM, respectively) are all present in higher concentrations in rice straw. However, it has low levels of sodium, iron, phosphorus and manganese (0.13%, 0.07%, 0.12% and 0.07%, respectively) (Shen et al. 1998). The amount of phosphorus comprised of 0.02–0.16% in paddy straw is insufficient to provide the necessary requirement for normal animal growth and fertility (Jackson 1977). Although its calcium concentration (0.4%) is thought to be sufficient to cover the daily needs of animals, is not always the case. Negative calcium balance was recorded with cattle fed with paddy residue even though straw contains enough calcium and making the calcium bioavailable from paddy straw is important as much the calcium composition (Nath et al. 1969). Similar studies by Joshi and Talapatra (1968) found that animals fed diets including wheat straw and sorghum stover had larger positive calcium balances than those fed diets containing rice straw, despite the latter’s higher calcium intake. The authors claim that adding calcium supplements to the animal’s diets of rice straw is safe. Since

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majority of these minerals are interrelated to one another in the form of acidinsoluble ash, further research is needed to determine their bioavailability.

11.6

Dietary Intake and Digestibility of Paddy Straw

Generally, an animal can consume paddy straw only less than 2% of its body weight every day. According to Devendra (1997), livestock can take up to 1.2 kg paddy straw per 100 kg of its weight on daily basis. The amount of rice straw that each animal consumes varies and controlled by its amount that is included in the dietary ratio and the way that paddy straw is produced and processed affects how much of it they consume. The amount of rice straw that an animal consumes is significantly increased by chopping, application of chemicals or microorganisms. Since it is heavy or takes up more room in the rumen when fed, its intake is reduced and the digestibility of the straw is impacted by sluggish transit rate of straw fed and subsequent fermentation in the stomach. By breaking up the straw, more microorganisms are able to enter the rumen and ferment its biodegradable components. For proper digestibility of cereal straws, the leaf to stem ratio is crucial. Comparatively, rice straw contains a higher leaf percentage (60%) than others like barley and oats (35% and 43%, respectively) (Sarnklong et al. 2010; Theander and Aman 1984). Because there are more leaves than stems, the leaves in vitro dry matter digestibility (IVDMD) is lower (50–51%) as compared to stems (61%) (Vadiveloo 2000). These findings were corroborated by Phang and Vadiveloo (1992) that IVDMD in goats for paddy stems and leaves was 68.5% and 56.2%, respectively. The leaves can be pre-treated by chemicals to boost their propensity to decompose. The IVDMD of the leaves was significantly higher than that of the stems after 21 days of treatment with 4% urea solution (Vadiveloo 2000). To maximise digestibility, one should take into account this increase in rice straw’s feeding value. Offering rice straw to livestock resulted in DM digestibility ranging from 45% to 50%. Glucanase, cellulase and hemicellulase are a few of the enzymes released in the reticulo-rumen that may be able to break down the rice straw’s cell wall constituents (Schiere and Ibrahim 1989). The above mentioned enzymes were secreted by rumen bacteria rather than being generated by the animals themselves. The amount of lignification’s or higher lignin content directly impacts how digestible the nutrients are in rice straw. The silica impacted directly on cell wall and its digestibility because it creates a barrier which inhibits its breakdown by microbes and leads to inadequate hydrolysis of paddy straw by enzymes (Dohnani et al. 2003).

11.7

Paddy Straw Pre-treatment for Livestock Feed

Farmers now have access to developed technology that can assist them to improve the nutritional quality of paddy straw for animal feeding and increase its utility. Different physical, chemical and biological processing as well as combinations of

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these are included in this approach (Ibrahim 1983). However, because they demand additional inputs and farmers must observe progress before adopting these technologies, implementation takes time.

11.7.1 Physical Pre-treatment When rice straw is used as livestock feed, the physical process is an efficient and affordable way to improve nutrient utilisation and recycling. The physical pre-treatment of paddy residue is done to increase intake, increase delectableness and enhance livestock’s potential digestibility. The physical process includes drenching, chopping, grinding, pelleting, baking under pressure and irradiate with gamma rays. These methods encourage the transformation of paddy straw physically including the reduction of particle size which shortens the animal’s ruminative period and the enriching softness to fibrous components of the straw makes the straw more edible to the animal and to improve the digestibility. These procedures made straw’s particles smaller, making it easier for rumen microorganism activity. When employing these methods, it is important to correctly balance the size of particles and passing rate of the treated straw ingested. The lower particle size of rice straw encourages voluntary intake and increases feed passage rate, but this has a detrimental impact that it makes the nutrients in the straw less digestible due to lesser exposure of feed components to rumination and other fermentation process in the rumen by microbes. An economical and popular method of treating paddy residue is soaking. The fibrous components like lignin and cellulose are softened by treating the paddy straw in water overnight. The animal will consume more of the straw and the nutrients will be easier for it to digest. Soaking and steaming both have a straight impact on delignification of cell wall. The effects of steaming or introduction of lignocellulosic components of paddy to extreme pressure create a favourable environmental condition for various processes like enzymatic activity, improving digestibility and availability of nutrients from paddy straw (Walker 1984). According to Milstein et al. (1987), heat pre-treatment raises the digestibility of cellulose from 20% to 40%. Another technique to think about is pressure steaming rice straw. Farmers may incur more costs as a result of the process demands more energy. Pre-treating using steam for various roughages and rice straw at higher pressure of 15 bars for 5 min at a moisture content varied from 30% to 70% resulted that steam pressure separated the various rice straw parts including hemicellulose, cellulose, lignin and sugars (Rangnekar et al. 1982; Liu et al. 1999). Similar finding was also recorded by Ooshima et al. (1984), who used conserved glass jars with handy partitions into lignocellulosic materials and increased rice straw nutrition by microwave irradiation technique at a frequency of 2450 MHz.

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11.7.2 Chemical Pre-treatment Since more than a century ago, scientists have used chemicals to enhance the nutritional content of rice straw in an effort to improve animal feed intake and digestibility (Kamstra et al. 1958). Sodium hydroxide (NaOH), ammonia (NH3) and urea are frequently researched and utilised in paddy straw chemical processing to progress its flavour, intake and digestibility. These chemicals degrade the lignocellulose bonds that are delicate in an acidic or alkaline environment. Alkali agents are a widely researched and used in agricultural purpose. Alkaline reagents like NaOH, NH3 or urea are engrossed into the cell wall of paddy straw and react with lignocellulosic components which break the ester bonds among lignin, cellulose and hemicellulose. The structural fibre in the paddy straw expands as a result of the alkali being absorbed into them and making them available for fermentation by various microbes (Chenost and Kayouli 1997; Lam et al. 2001).

11.7.2.1 Pre-treatment with Sodium Hydroxide Since the 1940s, cereal straw has been treated with NaOH to prevent mould (Mcanally 1942). NaOH (1.5%) is applied to the straw in a container for 24 h to treat it. After being cleaned with cold water, the treated straw is tested for in vitro digestibility. According to the research findings, the NaOH pre-treated straw is 28% more palatable and digestible than the untreated ones. FAO (2012) has advised using the Beckman procedure, which is comparable to Mcanally’s (1942) instructions for treating straw with NaOH. The Beckman procedure likewise employs 1.5% NaOH, but the treatment time is only 18–20 h before tap water is rinsed out. By breaking the bond between the straw’s lignin and cellulose components, the NaOH has an effect by slowing down protein breakdown and speeding up lignin breakdown. This allows more time to degradation by enzyme activity. The chemical processing of NaOH on the contents of paddy straw cell walls is beneficial for the disintegration of ester bonds between the phenols and cellulose fractions of straw and favouring enzymatic hydrolysis (Jackson 1977; Berger et al. 1994; Arieli 1997; Wang et al. 2004). The straw treated with NaOH performed better than ammonia treated straw in terms of cattle performance, increased intake and palatability as well as the improved digestibility. Treating rice straw with NaOH is not a common practice among farmers. This is due to the fact that NaOH is more expensive and occasionally unavailable than urea treatment (Chaudhry and Miller 1996; Vadiveloo 2000). Additionally, greater usage of NaOH, more than 10 g of the daily salt requirement for adult animals is harmful (Sundstol and Coxworth 1984). 11.7.2.2 Pre-treatment with Ammonia It has been determined that treating paddy straw with ammonia (anhydrous and aqueous) and other compounds which release ammonia improves the straw propensity to degrade (Fadel-Elseed et al. 2003; Selim et al. 2004). NH3 is comparable to NaOH treatment to paddy straw. Due to its availability and ability to be produced by the hydrolysis of urea, NH3 has been found to be preferable over using NaOH. In addition to improving rice straw’s ability to break down, NH3 treatment also adds

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nutrients mainly nitrogen, raising the crude protein of the straw (Abou-EL-Enin et al. 1999). Since NH3 prevents mould from growing on the treated straw, it can also be used as a preservative (Calzado and Rolz 1990). Other advantages of NH3 treatment include lowering the expense of purchasing high protein supplements and improving livestock acceptance and ingestion rate. NaOH rather than NH3 treatment is more effective in increasing the energy composition of straw. However, since NH3 supplies extra nitrogen to the straw, employing it is typically more commercial for farmers than NaOH (Liu et al. 2002). Sheep fed gaseous NH3 (3 g per 100 g of dry matter) treated straw for 4 weeks and noticed a comparable increase in CP from 51 to 115 g/kg and N content from 8.16 to 18.4 g/kg. The treated straw’s NDF slightly decreased from 571 to 551 g/kg, but ADF increased from 303 to 327 g/kg (Selim et al. 2004).

11.7.2.3 Pre-treatment with Urea The most practical and popular chemical approach for treating rice straws is urea treatment. Both large and small-scale livestock farms can adapt to it. Primary role of urea in the fermentation process is to improve the crude protein content of the paddy straw being treated. Urea or NH3 (urea-molasses solution) is utilised most efficiently at optimum moisture (30%) of the straw. To create ammonia-nitrogen, urea must first be hydrolysed or subjected to ureolysis (Sahnounea et al. 1991). Molasses played as an energy source, so that the fermentation of cellulose from the straw can proceed more quickly. Hence, rice straw can be made into a comprehensive and secure base feed for livestock by adding urea or urea in combination with molasses (Langar et al. 1985). A substance called urea is a source of non-protein nitrogen for livestock as well as nitrogen for crops. It is a crystalline substance that is available locally in markets, easy to handle. Urea can be used to efficiently cure rice straw at various concentrations, ranging from 1% to 5%. The necessary amount of urea should first be dissolved in water before being sprayed onto the rice straw. A silo, an empty drum or a plastic bag can be used to store the treated straw. Farmers can readily accept this treatment method because it is useful and practical (Sundstol and Coxworth 1984). Urea is less expensive over pure NH3 or NaOH. Different low degradable carbohydrates rice varieties on being treated respond better with urea treatments than good quality rice varieties as evidenced by a raise in IVDMD from 45% to 58% (Vadiveloo 2003). More number of studies were conducted on paddy straw with alone urea treatment or combining with various chemicals in laboratory (Reddy 1996; Shen et al. 1998) or in field condition (Prasad et al. 1998; Vu et al. 1999; Akter et al. 2004) and the results clearly showed advancement in biological value or nutritive composition of treated paddy residue. 11.7.2.4 Pre-treatment with Lime It is anticipated that treating paddy straw with lime (CaO or Ca(OH)2) may improve fibre degradability similar to NaOH. Lime is more soluble in water than urea or NaOH, but it is nevertheless a calcium source for livestock on low calcium diets. Straw can be treated with lime in different ways (soaking and ensiling). When combined with urea, the lime therapy has complementing effects and work well

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together to improve the treated straw’s capacity to degrade and increase its calcium and nitrogen contents (Nguyen 2000). Paddy straw treated with 4–6% Ca(OH)2 had a greater IVDMD value after ensiling and also its combination with urea produced superior effects than a single component (Pradhan et al. 1997). Sirohi and Rai (1995) employed 3% urea and 4% Ca(OH)2 at 50% moisture level and discovered that this was the best method for treating rice straw to brought treated straw’s increased digestibility and biodegradable nutrients. The use of Ca(OH)2 and other alkali had synergistic effects on rice straw being treated to be more safer and practical to employ, according to Saadulah et al. (1981) and Hadjipanayiotou (1984). Treated paddy straw with pure Ca(OH)2 provided conflicting outcomes in terms of its impact on the lignin breakdown or deterioration (Trach et al. 2001). Treated straw had a palatability issue, which caused animals to consume less dry matter. Although lime treatment had no effect on the amount of nitrogen, it seemed to be more effective at delignifying or reducing the amounts of hemicellulose and NDF in the straw. There were some undesirable interactions between the two chemicals when lime and urea were raised during the treating process. The lime at 6% seemed to be high enough for rumen cellulose synthesis, whereas 2% urea all alone appeared to be very less for successful therapy.

11.7.3 Biological Pre-treatment Processing of paddy straw biologically uses a variety of microorganisms, including bacteria and fungus as well as enzymes. Various fungal strains have the ability to affect the contents of the straw’s cell walls, enhancing the pace of breakdown and availing additional nutrients to the livestock. Biological agents are used to treat paddy straw to increase its nutrient content by the delignifying action. These enzymes released by fungi had a great affinity to metabolise lingocelluloses. However, due to technical skill gaps and a lack of assets to generate and manage great amounts of fungi, its current use in impoverished nations is still very much in doubt (Jalc 2002). The biological treatment of straw raises several issues and challenges that need to be resolved (Schiere and Ibrahim 1989). For instance, certain fungus species produce toxic compounds that are poisonous to both humans and animals yet are not edible. In order to develop and reproduce, fungi also need certain environmental conditions before, after and during the treatment process (pH, temperature, pressure, O2 and CO2). As mycology advances, it is possible now to culture and produce fungus or purify enzymes for the treatment of rice straw using straightforward procedures or instructions. Livestock farmers can employ commercially available enzyme inoculants or additives to boost the efficiency of their production and their farm income while also lowering the cost of purchasing these substrates (Beauchemin et al. 2004).

11.7.3.1 Pre-treatment with Fungi White-rot fungi are lignocellulolytic components of agricultural wastes, particularly wood and are recognised to have degrading or rotting qualities. According to

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Eriksson et al. (1990), they acquired the ability of degrading and metabolising cell wall components in suitable settings through enzymes. These species have capability to solubilise the lignin to be referred as ‘lignin degraders’ and can advance the nutritional worth of feed by providing more carbohydrates in degradable nature for fermentation in rumen (Yamakava and Okamnto 1992; Howard et al. 2003). These fungi secrete many extracellular enzymes which alter lignin that includes lignin peroxidase, manganese dependent peroxidase, phenol oxidase, aryl-alcohol oxidase, glyoxal oxidase, etc. (Arora et al. 2002; Novotny et al. 2004; Arora and Gill 2005; Lechner and Papinutti 2006). Few fungal groups have the ability to dissolve the bindings or bonding between lignin and polysaccharides of straw by decomposing or acting directly on free phenolic monomers (Chen et al. 1996). The IVDMD of treated straw was improved by other fungal species. Incubation of straw with three white-rot fungus species for 30 days resulted that IVDMD in rice stems and leaves was improved by Pleurotus sajor-caju. However, compared to other fungal species, the results utilising Cyathus stercoreus showed the higher IVDMD (Karunanandaa et al. 1992, 1995; Fazaeli et al. 2006). The rate of reaction of white-rot fungi with their substrates depends on the type of fungus. There were some species of fungi that easily digest carbohydrates (simple sugars, cellulose and hemicellulose) before degrading lignin, which causes livestock to have less energy available (Karunanadaa and Varga 1996; Jalc 2002). Depending on the kind of white-rot fungi, the time required to incubate straw varies. Mycelial development was expected to cause some energy crisis during the early stages of incubation, but a while after, few fungal species prefer to hit lignin over other cell components which provide more energy for the livestock. These days, it is critical to conduct research in mycology using fungal species that favour lignin degradation of rice straw. Mycologists can create better strains of fungi once these species have been identified (Rodrigues et al. 2008). Paddy straw can also be used for growing edible mushrooms. Pleurotus ostreatus and Volvariella sp. are two examples of edible fungi and these are simple to grow and leftover mycelia of mushroom bedding can boost the rice straw’s crude protein and digestible carbohydrate content.

11.7.3.2 Pre-treatment with Enzymes Enzymatic reactions cause the metabolic breakdown of any complex substance into simplest form. The microorganisms mediated enzymes help in breakdown of paddy straw and they work very specifically on the substrates that need to be broken down. Degradable enzymes are commercially accessible in the market as cellulase, hemicellulase, glucanase and xylanase. However, many factors including animal body temperature and particle size, processing and packaging of enzyme products have an impact on their stability and efficacy. The enzymes utilised in the livestock feed commercially are of either bacterial origin (Lactobacillus and Staphylococcus sp.) or fungal origin (Aspergillus niger, Trichoderma longibrachiatum and Aspergillus oryzae) (Colombatto et al. 2003). Intensified treatment of enzymes degrades paddy residue very quickly. Furthermore, employing enzymes had shown to increase milk production in dairy cows, weight gain and production of wool in sheep (Wang et al. 2004; Jafari et al. 2005; Rodrigues et al. 2008). Due to additional input costs

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and the limited knowledge on enzymatic products, the enzymatic treatment of paddy straw is yet not commercialised popularly among livestock farmers.

11.8

Limitations of Paddy Straw as Livestock Feed

Paddy straw as livestock feed is restricted by a number of variables comprising poor digestion, a low intake of animals, very low protein content, etc. For the preparation of straws before feeding to animals, technologies have been developed to overcome the mentioned challenges. When employed by farmers, though it’s practicable, issues with health and environment taking into account and varies depending on their capacity and capability. The primary restriction on the physical treatment of straw is the need to grind it into smaller particle sizes. Reduced particle size has the advantage of encouraging increased intake because it speeds up the animal’s transit of the ingested feed. But it reduces the time period required for rumination and degradation by microbes as a drawback and subsequently slows down the breakdown and digestion of the straw constituent. According to Uden (1988), grinding and pelleting reduced the DM digestibility in cattle from 73% to 67% due to lower fermentation rate (9.4–5.1%/h), shorter retention time of the solids (73–54 h) and increased intake (Stensig et al. 1994). Due to their limited ability to purchase equipment and the potential for poor or even adverse advantages for the farmers, small-scale farms are also unable to use machines for physical treatment and processing of crop leftovers (Schiere and Ibrahim 1989). One of the problems limiting the use of chemical treatment of straw may be the associated expenses. The treatment of rice straw has considerable benefits in improving nutritional content, animal activity and income, but farmers should still weigh all of their options before deciding whether or not to use treated straw. Chemical treatment of straw has some drawbacks including safety concerns like toxicity and environmental contamination. The main disadvantages of using a particular strain of fungi to treat rice straw through microbiology is that it can break down lignin and also other straw constituents like cellulose and hemicellulose. Another restriction on its practical use in treating straws is the incubation period. It is necessary to investigate these fungi to determine their ideal incubation period in order to maximise the biological value of straw. Some fungi species have ability to decompose lignocellulosic straw material within 1 or 2 weeks of incubation period. Additionally, some fungus can create toxins that can harm both people and animals. Therefore caution should be taken while utilising them to treat with rice straw.

11.9

Conclusion

Among agricultural by-products, rice straw is the most plentiful, least expensive and useful source of livestock food. Due of difficulties in collection, transportation and storage, it is only occasionally used as cattle feed. Because of the fact that rice straw is not nutritious as grasses, low in protein and difficult to digest, it cannot provide

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nutritional requirements needed by high milk yielding cattle. Through physical processing, chemical pre-treatment or biological treatment, it is possible to improve the nutritional composition, digestibility and utility of paddy straw. Additionally, mechanisation is crucial for facilitating the gathering, transportation, stacking and processing of rice straw from the field in order to capitalise use of rice straw as livestock fodder on a larger scale. Because of the limited knowledge and resources, uncertainty regarding the suitability of technologies for use on farms and their potential advantages for livestock producers and animals, the implementation of technologies is relatively low. However, these technologies are useful in generating extra revenue for livestock farmers through the sale of milk or live animals. So, rice straw continues to be the practical, plentiful and affordable source of fodder for livestock in nations where there is a shortage of feed or a lack of high-quality forages.

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Engineering Interventions for Climate-Resilient Forage Production

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Amit Kumar Patil, Naseeb Singh, Partha Sarathi Singha, Monika Satankar, Sheshrao Kautkar, S. K. Singh, and P. K. Pathak

12.1

Introduction

India faces the issue of combating poverty through the implementation of a development strategy that, in addition to bolstering the country’s climatic resilience, also improves the nation’s food and water security (Satankar et al. 2020). Overusing both fertilizer and water results in a kind of waste that not only harms the local ecosystem and our planet’s climate but also hurts the farmer’s business line. Fertilizer that is applied in excess causes soil bacteria to produce nitrous oxide, a greenhouse gas with a heat-trapping effect that is staggeringly more than 260 times greater than that of carbon dioxide. Farmers around the world are already seeing their livelihoods put in jeopardy as a direct result of climate changes (Campbell et al. 2014). Dryland agriculture is especially susceptible to the effects of drought in arid and semi-arid regions, which are home to more than 40% of the world’s population and include 650 million of the world’s most impoverished and food-insecure people (Kritee et al. 2019). Agriculture, animal husbandry, and dairying are all activities that have continued to be an essential component of human existence ever since the beginning of the civilizing process. These efforts have not only added to the food basket and

A. K. Patil (*) · S. K. Singh · P. K. Pathak ICAR - Indian Grassland and Fodder Research Institute, Jhansi, India N. Singh ICAR - Research Complex for NEH Region, Umiam, India P. S. Singha Assam University, Silchar, Assam, India M. Satankar IARI, New Delhi, India S. Kautkar ICAR - Central Institute for Research on Cotton Technology (CIRCOT), Mumbai, India # The Author(s), under exclusive license to Springer Nature Singapore Pte Ltd. 2023 R. K. Singhal et al. (eds.), Molecular Interventions for Developing Climate-Smart Crops: A Forage Perspective, https://doi.org/10.1007/978-981-99-1858-4_12

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Table 12.1 Estimates of demand and supply of fodder resources in India (mt) Year 1995 2000 2005 2010 2015 2020 2025

Supply (mt) Green 379.3 384.5 389.9 395.2 400.6 405.9 411.3

Dry 421 428 443 451 466 473 488

Demand (mt) Green 947 988 1025 1061 1097 1134 1170

Dry 526 549 569 589 609 630 650

Deficit of demand (%) Green Dry 59.95 19.95 61.10 21.93 61.96 22.08 62.76 23.46 63.50 23.56 64.21 24.81 64.87 24.92

Source: Khanna (2014)

increased the power of draught animals, but they have also helped to the preservation of ecological equilibrium. Because of India’s accommodating climate and topography, the animal husbandry and dairying industries have been able to play a significant role in the country’s socioeconomic development. They also play a vital role in the generation of gainful employment in the rural sector, notably among the landless, small and marginal farmers, and women, in addition to providing food that is inexpensive and nutritious to millions of people. India is home to 56.7% of the world’s buffalo population, 12.5% of the world’s cow population, and 20.4% of the world’s small ruminant population (MoFAD, GoI 2021b). In addition to playing a significant part in the economy as a whole, livestock production is also an essential source of income for landless and marginal farmers. In the fiscal year 2020–2021, agricultural production (the crop sector) contributed 8.96% (at constant prices) to total gross value added, while the livestock sector contributed 4.90% (at constant prices) to total GVA. Animal husbandry is one of the key drivers of growth in rural incomes, and increased public investment in the livestock sector is required to achieve the goal of doubling farmers’ incomes. Agricultural diversification is accomplished through animal husbandry. Both livestock production and dairying are highly dependent on the quantity and quality of fodder available. It is becoming increasingly difficult to make green fodder available to all dairy farmers these days (Table 12.1). As the years go by, Table 12.1 data demonstrates the growing shortage of green and dry fodder supplies to satisfy the necessary demand. The cost of feed accounts for around 70–75% of the entire milk cost, whereas green fodder contributes 35% of the total input feed. Green fodder offers the necessary nutrients or minerals for milk production and the health of dairy animals. Due to this situation, dairy farmers were compelled to look for alternative approaches of generating premium green fodder (MoFAD, GoI 2021b). In the current scenario, when competing demands on land expansion for food and cash crops, the possibility of extending the area under fodder crops is extremely low, and the likelihood of doing so is almost impossible. It is therefore imperative to develop a multi-pronged strategy for the appropriate availability of fodder in order to offer a buffer for the farmer even in times of climate unpredictability. This is because it is essential to establish a multi-pronged strategy for adequate availability of fodder.

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This strategy interiliac envisions the supply of quality seeds, the promotion of production of fodder crops, the extension of fodder cultivation to currently fallow and unutilized lands, the promotion of dual-purpose varieties of crops that have the potential to meet fodder requirements both during the growing season and during the off-growing season, the promotion of non-traditional fodder, post-harvest technologies for the preservation of fodder, and other such things. In addition, adopting smart and climate-friendly agricultural machinery which increases field performance with the timeliness of operation and better fuel efficiency to reduce the time taken for operation and saving fuel would not only benefit monetarily to the farmer but also saves the degradation of climate and the environment. Hence the inclusion of this smart and climate-friendly machinery would be a promising approach to increase fodder production yield. This would open up the possibility of increasing the availability of fodder derived from a variety of forage crops like berseem, pearl millet, sorghum, maize, and oat (AFDP, MoA 2011).

12.2

Climate-Smart Technology

According to the Food and Agriculture Organization (FAO), low carbon farming methods, also known as climate-smart farming, are those that (a) sustainably boost productivity and income, (b) help in adapting to climate change and creating resilience, and (c) take whatever steps are necessary to cut or eliminate emissions of greenhouse gases. In the context of Indian agriculture, “smart farming” has recently emerged as a pressing necessity. It is far more productive than the more conventional approaches to farming. The concept of “smart farming,” which includes the use of sensors and automated irrigation methods, can assist in the monitoring of a variety of agricultural factors, including temperature, soil moisture, and more. This would make it possible for farmers to check on their crops from any location. In addition, digital and physical infrastructures may be integrated more easily with the help of smart farming, which is something that is beneficial to small farmers. The inability of India’s small and marginal farmers to successfully integrate digital and physical infrastructures, which impedes the increase of their earnings, is a significant barrier (Das et al. 2022). The most vulnerable people to the effects of climate change are low-income and subsistence farmers whose livelihoods depend on the land. Savy practices and technology can encourage alleviation of hazards produced by a variety of factors, including climate change. Recently, India is frequently putting up endeavor to develop and put into standard rules that would make agriculture more sustainable. Artificial intelligence has the potential to completely transform the current farming and agricultural practices that are prevalent. The dynamic corporate structure of India exhibits an opportunity for collaborations between private companies and the Government of India to contribute to the progress of a refined agricultural sector (IBEF 2021a). Stagnation in agricultural yields, an increase in soil degradation, an increase in the shortage of water, a reduction in biodiversity, and an increase in the frequency of natural disasters are all factors that are making it difficult for farmers to sustain. Additionally, India’s

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agricultural sector is accountable for approximately 14% of the country’s overall GHG emissions. Agriculture that is climate-smart (CSA) is a system of farming that aims to concurrently produce food and energy apparently that is both environmentally and socially responsible (Aryal et al. 2016). It can assist in the transformation of agri-food systems into one that is more responsive to the effects of climate change. Farmers in India are beginning to understand the value of climate-smart agriculture (CSA). The CSA model is an integrated strategy for managing agricultural land, livestock, forests, and fisheries. It also addresses the intertwined problems of food insecurity and accelerating climate change. The CSA can assist India in accomplishing the following goals:

12.2.1 Increased Productivity CSAs can help farmers produce more food while maintaining quality. This will improve farmers’ nutrition security and help them earn more money, particularly those who are impoverished or on the verge of going bankrupt (Patil et al. 2021a, b).

12.2.2 Improved Resilience People who practice CSA may be less susceptible to diseases brought on by drought, pests, and other climate change-related shocks. A long-stressed and unfavorable environment can also be taken care of and improved by farmers.

12.2.3 Reduced Emissions This is likely to be one of the most important benefits of the CSA. Automation makes jobs less labor-intensive, which helps cut down on emissions per calorie of food produced, stops trees from being cut down, and lowers the amount of greenhouse gases like carbon dioxide released into the air. This will cause people to use less power from sources that are not good for the environment. Smart farming lets farmers grow different kinds of crops, which helps them depend less on the monsoon.

12.3

Forage Production and Conservation Machinery

The cultivation and consumption of fodder include operations comparable to those of any other crop, such as seedbed preparation, sowing, weeding, harvesting, etc. Before feeding fodder to animals, it is necessary to undertake initial processing on the collected fodder crop. The production, processing, and consumption of fodder are labor-intensive, time-consuming, and energy-intensive processes. The optimal production and exploitation of fodder crops necessitate minimal and timely

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operation (Patil et al. 2021a, b). Delay in the operation of fodder production frequently results in quick loss of moisture content and deterioration of fodder quality. There are special mechanization needs associated with fodder crops. At the time of harvest, a number of fodder crops are multi-cut and create a volume of green and dry matter. Their production relies on the timely harvesting of materials to allow growth for the subsequent crop (Mohammadi and Omid 2010). Large volume and mass manipulation necessitate the operation of suitable apparatus. Under Indian conditions, the majority of farming communities have small (1–3 hectare) land holdings. A small farmer with typically two to ten animals devotes a tiny percentage (up to 10%) of his cultivated land to fodder production. Accordingly, Indian conditions necessitate machinery and their magnitude to meet their farming demand. Conservation involves size reduction and drying. When agricultural leftovers are wet or green fodder needs to be dried for various products, drying, size reduction, baling/ densification/pelleting, etc. are used. Material handling and transport can be done manually or mechanically to reduce labor. After harvesting, agricultural commodities like grasses are transported for drying, size reduction, storage, feeding, etc. After harvesting and threshing wheat, wheat straw is stored, sold, or fed. Thus, material handling and transport should ease labor.

12.4

Engineering Interventions

12.4.1 AI-Based Crop Yield Prediction Model This facilitates timely agricultural guidance to farmers. Microsoft and the Indian government have partnered to support the success of Indian farmers. This alliance aims to help farmers earn more money by providing them with more harvests and better price control via the use of AI tools (Kashyap and Panda 2003), The collaboration would increase the likelihood that farming would adopt AI in farming.

12.4.2 Artificial Intelligence for Fodder Production Artificial intelligence is founded on the idea that human intelligence can be described in a way that makes it simple for a computer to imitate it and carry out tasks of any complexity. Artificial intelligence has three main objectives: learning, reasoning, and perception. In every industry, including education, entertainment, finance, robotics, agriculture, e-commerce, etc., artificial intelligence has emerged as one of the most significant technological advancements. In multiple directions, it is revolutionizing agriculture and performing a very important part in the agricultural sector (Azimi et al. 2021). AI protects the agriculture industry from several concerns, including food safety, population increase, and climate change. Because of AI, modern agriculture has advanced in a greater direction. Crop yields, real-time inspection, harvesting, processing, and selling have all been enhanced by artificial intelligence. To identify numerous crucial criteria, including weed identification,

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yield detection, crop quality, and many more, various high-tech computer-based systems have been developed. Similar approaches using AI can be used for fodder crops to mitigate the effects of climate change on fodder production. In the following sections, various systems are explained that can be implemented for climate-resilient forage production and its conservation.

12.4.3 Nutrient Management Using Artificial Intelligence (AI) Plant nutrients are substances that are necessary for a plant’s life processes and can be found in soil, air, or water (for example, nitrogen, manganese, sulfur, potassium, carbon, hydrogen, oxygen, etc.). Nutrient management is the process of utilizing crop nutrients as efficiently as possible to increase productivity while safeguarding the environment. The fundamental premise of nutrient management is to balance soil nutrient inputs with crop demand. To manage plant nutrients, AI can be very useful. In past, many researchers used this technique to predict nutrient deficiency and manage the same. Jose et al. (2021) in their study detect and classify the nutrient deficiencies in tomato plants using a mobile application for which they used artificial intelligence. Aleksandrov (2022) proposed an AI-based tool for predicting nutrient deficiencies in bean plants, whereas Wulandhari et al. (2019) used a convolutional neural network which is the most recent technique in AI, to predict plant nutrient deficiency. Lee et al. (2022) proposed an AI-based intelligent precision nutrient analysis model to manage the nutrient precisely.

12.4.4 Water Stress Management Using Artificial Intelligence (AI) Water stress, often known as drought stress, is a type of plant abiotic stress that, if prolonged, poses a danger to plant productivity. Due to a scarcity of water, most major crop plant yields can be reduced by more than half (Kashyap and Panda 2003). Accurate water stress evaluation will boost agricultural output by optimizing plant water usage, maximizing plant breeding techniques, and increasing fodder yield. Azimi et al. (2021) proposed a deep learning network for the temporal analysis of stress-induced visual changes in plants and applied it to the specific water stress identification case in Chickpea plant shoot photos. Duarte-Carvajalino et al. (2021) estimate water stress using artificial intelligence in potato plants, while Soffer et al. (2020) detect water stress in corn using deep learning in real-time.

12.4.5 Predicting the Best Time to Sow and Harvest Using Artificial Intelligence (AI) The difference between a productive year and a failed crop is timely advice on seeding and harvesting crops. Sowing as well as harvesting critically depends on climatic factors like temperature, humidity, rainfall, sunlight, soil moisture,

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evapotranspiration, etc. Artificial intelligence can be utilized as an advanced analytical technique to figure out the best period for seeding and harvesting to maximize yield. This area has yet to be investigated, and if artificial intelligence is used in this aspect, fodder output is likely to improve.

12.4.6 AI-Powered Pest Detection System Pests are one of the biggest adversaries of farmers, causing agricultural loss including fodder crops (Ivelina 2018). AI-powered devices can determine whether or not an insect is present in a crop, as well as the type of insect (Liu and Wang 2021). The AI-powered device then sends alerts to farmers’ smartphones, allowing them to take necessary precautions and apply necessary pest control, so assisting farmers in their fight against pests in feed. Turkoglu et al. (2022) developed a convolutional neural network model to detect diseases and pests. In past, numerous researchers (Alves et al. 2020; Khanramaki et al. 2021; Rahman et al. 2020; Tassis et al. 2021) used artificial intelligence to detect pests in various crops which validates that a similar approach can be deployed to the fodder crops to alleviate the pests issues.

12.4.7 Intelligent Land Preparation For land preparation of fodders both tractors and implements are used in combination. To increase the productivity of the fodder land preparation, efficient operation of both tractors and implements is necessary. But it is difficult to identify the right combination of tractor and implements according to field condition. Hence decision support system (DSS) is useful in this regard. Wherein this system would guide the user about the recommended tractor to be used and correct matching of the implement for the same (Civelek and Say 2016; Sahu and Raheman 2008). Individually both tractor and implements need to be climate friendly and with use of intelligent systems both are expected to give better productivity for land preparation. In the upcoming subsections both intelligent tractors and machinery are taken up for discussion. Zero Emission Using Electric Tractor: The financial year of 2021 has seen 0.89 million units of tractor sold over India (www.tractorjunction.com). Most of them are internal combustion engine (ICE) tractors, hence it exhausts greenhouse gases (GHG) and unwanted particulate matter (PM) which degrades the climate and environment. To overcome this battery powered electric tractor was thought off (Ueka et al. 2013; Usinin et al. 2013; Mocera and Somà 2020). The battery powered electric tractors have zero emission with less noise and vibration compared to ICE and since it does not emit any GHG and PM hence it is climate friendly. With environmental benefits the initial capital expenditure of the battery powered vehicle is high, but the total cost of ownership (capital expenditure and running cost) will be low. Hence the return of investment for the farmer will be seen upon the more he/she uses the tractor and longer time they use it. The major cost of any battery electric

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vehicle is its battery ($140–$550/kW h) varying with its chemistry. But the electric tractors need to be evaluated in terms of total cost of ownership. With more improvement in the chemistry of the battery for the electric drive it is evident that battery powered electric tractor will become more and more cost competitive (Ueka et al. 2013; Usinin et al. 2013; Mocera and Somà 2020; Basma et al. 2021; Mousazadeh et al. 2010). The battery powered electric tractors need to be charged from the electric grid and in India still the major source of electric power is fossil fuels. Hence point of argument is that the battery powered electric tractor is indirectly making pollution. But with use of growing renewable energy source and development of renewable powered electric vehicle is the counter to this argument. The technology for the same is available by implementation on mass scale workout is the way forward (Basma et al. 2021; Mousazadeh et al. 2010). The electric tractor consists of battery which is the energy source with lithium iron phosphate (LFP), lithium nickel manganese cobalt (NMC), or lithium titanate (LTO) chemistries being generally used. The power they generate per kilograms of weight and the energy densities are important parameters to be considered for selecting a battery pack. The battery provides the required voltage and current to the power distributor which distributes the power to the inverter of the motor, DC– DC converter, the battery management system, and other auxiliary function if required. There are many configurations in which the motor can be mounted to the tractor, one is directly to the differential, directly to the power source hub or individually attaching motors to the wheels. There are many auxiliary power requirements by tractor to run the headlamps, electronic power control units, etc. This does not require a very high voltage system; hence the voltage is lowered down to accommodate these systems using a DC–DC power converters. Chargers are required to charge the electric tractor wherein combined charging system (CCS 2) chargers are used which as per Indian adopted charging standard.

12.4.8 Smart Rotavator One of the major machineries used for the land preparation is rotavator and it is also one of the most sold machineries in India (Singh 2014). There has also been recent development by introducing smart technologies to increase the productivity of operation by indicating the farmer with optimal range of operation. The trochoidal path of rotavator is dependent on the both the tractor forward speed and the peripheral speed of rotavator. Hence indicating the optimal range of these parameters would improve performance. One of the major developments in this regard is the development of Mahindra Tez-e rotavator (smart rotavator) system and another is a developed velocity ratio indicator system as discussed below. A smart rotavator consists mostly of three systems, as seen in Fig. 12.1. First, there is the integrated smart rotavator unit. The smart rotavator unit was made up of a sensor-integrated controller and cage. As depicted in Fig. 12.1, the smart rotavator unit was mounted on the input outer shaft of the side gear. To shield the controller with an integrated sensor system from the outside environment, a cage was created

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Fig. 12.1 Overview of smart rotavator

on the outer shaft. The battery management system, a communication module, and a Hall effect speed sensor make up the majority of the controller. The speed sensor detects this difference and feeds it to the controller, which then signals conditions it using the moving average algorithm and internally calculates the rotavator blade rpm by taking into account the side gear reduction ratio before sending it to the mobile app through the Bluetooth module. When the input shaft of the side gear rotates, it has grooves that cause a difference in the permeability effect. Additionally, the Bluetooth module utilizes an internal time clock to transmit the cumulative and current hours of operation. The second working method is a mobile app that notifies the user of the best operating zone via a green zone and flashes sound and vibration notifications when operating in any other zones. A change oil alert is one of the features of the mobile app, while a check oil alert is one that is sent after every 50 h of operation. The mobile application needs to be configured with the appropriate rotor gear for the excavator. The user interface of the mobile app includes a charge indicator for the internal battery and is available in multiple languages. The developed system’s user interface was a smartphone app. The mobile application was created for Android 5.0 and later. After the controller processes the speed and time data, the data are sent over Bluetooth to the mobile application. Other user interface parameters are treated internally. Figure 12.2 displays the user interface of a mobile app. The app features are described below: 1. Rotor Gears: The rotor gear of the rotavator is the set of gears that are used to change the speed of the rotavator blade depending on the usage and the state of the soil. The rotor 1:2:3:4 represents the gear sets 17/21:21/17:18/20:10/18, and the farmer selects one of them while the machine is running. The same gear set must be chosen by the farmer for this app as well. The ideal rpm range also varies depending on the rotor gear, according to the soil and farmer usage conditions. 2. Cumulative Hours: This function keeps track of the total number of operating hours. The rental hiring function required the addition of this feature.

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Fig. 12.2 Mobile app user interface

3. Trip Hours: This feature records the current operating hours. Resetting the time for the current action is possible by pressing the reset button. 4. Bluetooth ON/OFF: When a device is connected to a mobile app, the Bluetooth icon goes green to indicate ON and red to indicate OFF. 5. Serviceability: Every 400 h in a row, the app would suggest that it was time to change the oil, and every 50 h, it would signal that it was time to check the oil. 6. Display: A needle on a dial gauge in the app displays the rotavator blade’s rpm. Distinct portions of the dial gauge are distinguished by different colors. When a tractor is started, the rotavator blade rpm is displayed in a zone of white, which is the lowest rpm. The area of orange indicates where the farmer will be working at a lower rpm than ideal. Similar to this, the farmer will be running at a higher rpm than desirable in the red zone. While working in orange and red zones, there are sound and vibration alerts. The ideal working area is represented by the green zone

12.5

Hydroponic Fodder Production System

A hydroponic green fodder could be an option especially in hilly and arid regions. The hydroponic green fodder/sprouted feed rich in nutrients including protein, vitamins, and micronutrients could be fed as supplement for enhancing both quality and quantity of the milk (Singh et al. 2022). Hydroponics technique has proven useful and efficient for producing quality fodder/feed for livestock (Mobtaker et al. 2022). The technique is advantageous when compared with conventional agriculture because it controls the climatic conditions as well as plant nutrition. Hence, it is possible to get increased production, stable harvests of high quality fresh green fodder all the year round and can be produced on a commercial scale. The hydroponic fodder/feed is produced under completely controlled conditions (Singh et al. 2021) and is thus free from undesirable materials such as weeds, insects, dust,

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Fig. 12.3 Hydropic maize fodder production unit

insecticides, pesticides, germicides, and carcinogens. Hydroponics culture is probably the most intensive method of crop production in today’s agricultural industry (Anonymous, 2017). With the possibility of adjusting air and root temperature, light, water, plant nutrition and adverse climate, hydroponics can be made highly productive, conserving water and land, and protective of the environment (Fig. 12.3). An evaporative cool hydroponic (ECH) structure developed at IGFRI (12 ft × 12 ft × 10 ft) comprised of pipe framed structure made of iron pipe, covered with plastic sheet/shade net under which plants are grown under at least partially controlled environment without soil (Singh et al. 2022). The ECH structure was evaluated for maize crop. The maximum productivity of green biomass was 7.5 kg per kg of seed rate in 7 days. Water consumption was only 1.5–2 L/kg of green biomass. The fodder production capacity is about 1 quintal/day which is sufficient for ten animals. Farmers may construct the structure nearer to animal enclosure/shed by purchasing the required materials from the local market and with the help of skilled carpenters and mechanics. In summation, the artificial intelligence-based systems that researchers have already introduced to many crops can be expanded for fodder crops to increase productivity and offset the effects of climate change on fodder production.

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Conclusions

With adverse climatic situation due to growing greenhouse gases fodder production has been unable to meet the demand of market for past many years. Hence to increase the production without affecting climate through engineering solutions technologies are available and can be utilized for fodder production on a mass scale. The study shows that use of instrumentation, artificial intelligence for nutrient management, water stress management, predicting sowing and harvesting time as well as pest detection will help in optimal input for fodder production. To increase the productivity for land preparation since land size will not grow but the yield from the given land can be increased through use of smart implements, new advance technology based engineering intervention would help to have better field capacity, less fuel consumption, and better pulverization.

References Aleksandrov V (2022) Identification of nutrient deficiency in plants by artificial intelligence. Acta Physiol Plant 44:29. https://doi.org/10.1007/s11738-022-03363-0 Alves AN, Souza WSR, Borges DL (2020) Cotton pests classification in field-based images using deep residual networks. Comput Electron Agric 174:105488. https://doi.org/10.1016/j.compag. 2020.105488 Anonymous (2011) Accelerated Fodder Development Programme (AFDP) report. Ministry of Agriculture Department of Agriculture & Cooperation Krishi Bhavan, New Delhi Anonymous (2017) NAAS Policy Paper No. 85 Hydroponic fodder production in India. National Academy of Agricultural Sciences, New Delhi. 27 p Anonymous (2021a) Annual report-2020-21. India Brand Equity Foundation, New Delhi, p 23 Anonymous (2021b) Annual report-2021-22. Ministry of Fisheries, Animal Husbandry and Dairying, Govt. of India, New Delhi Aryal JP, Sapkota TB, Stirling CM, Jat M, Jat HS, Rai M, Mittal S, Sutaliya JM (2016) Conservation agriculture-based wheat production better copes with extreme climate events than conventional tillage-based systems: a case of untimely excess rainfall in Haryana, India. Agric Ecosyst Environ 233:325–335 Azimi S, Wadhawan R, Gandhi TK (2021) Intelligent monitoring of stress induced by water deficiency in plants using deep learning. IEEE Trans Instrum Meas 70:1–13. https://doi.org/ 10.1109/TIM.2021.3111994 Basma H, Beys Y, Rodríguez F (2021) Battery electric tractor-trailers in the European Union: a vehicle technology analysis (No. 2021-29). International Council on Clean Transportation, Washington, DC. https://theicct.org/publications/eu-tractor-trailers-analysis-aug21 Campbell BM, Thornton P, Zougmoré R, Van Asten P, Lipper L (2014) Sustainable intensification: what is its role in climate smart agriculture? Curr Opin Environ Sustain 8:39–43 Civelek Ç, Say SM (2016) A web-based decision support system to select proper machinery size and tractor power. Turkish J Agric Forestry 40(4):512–525 Das U, Ansari MA, Ghosh S (2022) Effectiveness and upscaling potential of climate smart agriculture interventions: farmers’ participatory prioritization and livelihood indicators as its determinants. Agric Syst 203:1–7 Duarte-Carvajalino JM, Silva-Arero EA, Góez-Vinasco GA, Torres-Delgado LM, Ocampo-Paez OD, Castaño-Marín AM (2021) Estimation of water stress in potato plants using hyperspectral imagery and machine learning algorithms. Horticulturae 7:176. https://doi.org/10.3390/ horticulturae7070176

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Ivelina N (2018) Insect pests in forage crops and integrated plant protection. ARTOAJ 17:1–14. https://doi.org/10.19080/ARTOAJ.2018.17.556038 Jose A, Nandagopalan S, Ubalanka V, Viswanath D (2021) Detection and classification of nutrient deficiencies in plants using machine learning. J Phys Conf Ser 1850:012050. https://doi.org/10. 1088/1742-6596/1850/1/012050 Kashyap PS, Panda RK (2003) Effect of irrigation scheduling on potato crop parameters under water stressed conditions. Agric Water Manag 59:49–66. https://doi.org/10.1016/S0378-3774 (02)00110-5 Khanna RS (2014) Fodder of the future: a review of the hydroponics technique. Indian Dairyman 66(5–6):1–5 Khanramaki M, Askari Asli-Ardeh E, Kozegar E (2021) Citrus pests classification using an ensemble of deep learning models. Comput Electron Agric 186:106192. https://doi.org/10. 1016/j.compag.2021.106192 Kritee K, Nair D, Araiza DZ, Reddy M, Proville J, Ahuja R (2019) Climate smart farming in India: a pathway to poverty alleviation, food security, and climate adaptation and mitigation. An online report with greenhouse gas flux data from rice and non-rice cropping systems from four agroecological regions in India. Environmental Defense Fund, New York, NY. 34 p Lee HA, Huang TT, Yen LH, Wu PH, Chen KW, Kung HH, Liu CY, Hsu CY (2022) Precision nutrient management using artificial intelligence based on digital data collection framework. Appl Sci 12:4167. https://doi.org/10.3390/app12094167 Liu J, Wang X (2021) Plant diseases and pests detection based on deep learning: a review. Plant Methods 17:22. https://doi.org/10.1186/s13007-021-00722-9 Mobtaker HG, Sharifi M, Shalmaei NT, Afrasiabi S (2022) A new method for green forage production: energy use efficiency and environmental sustainability. J Clean Prod 363:132562. https://doi.org/10.1016/j.jclepro.2022.132562 Mocera F, Somà A (2020) Analysis of a parallel hybrid electric tractor for agricultural applications. Energies 13(12):3055 Mohammadi A, Omid M (2010) Economical analysis and relation between energy inputs and yield of greenhouse cucumber production in Iran. Appl Energy 87(1):191–196 Mousazadeh H, Keyhani A, Javadi A, Mobli H, Abrinia K, Sharifi A (2010) Evaluation of alternative battery technologies for a solar assist plug-in hybrid electric tractor. Transp Res D Trans Environ 15(8):507–512 Patil AK, Singh SK, Sahay CS, Pathak PK, Dwivedi PN (2021a) Artificial Intelligence (AI): the future of innovation. Poster presented at ICAR-IGFRI. ICAR-IGFRI, Jhansi Patil AK, Sahay CS, Singh SK (2021b) Design and testing of sensor-based setup for precision seed treatment. Theme-conservation agricultural, smart mechanization and energy use. Extended summary: 5th International Agronomy Congress, PJTSAU, Hyderabad, Telangana, India, November 23–27, 2021, vol 1, pp 514–515 Rahman CR, Arko PS, Ali ME, Iqbal Khan MA, Apon SH, Nowrin F, Wasif A (2020) Identification and recognition of rice diseases and pests using convolutional neural networks. Biosyst Eng 194:112–120. https://doi.org/10.1016/j.biosystemseng.2020.03.020 Sahu RK, Raheman H (2008) A decision support system on matching and field performance prediction of tractor-implement system. Comput Electron Agric 60(1):76–86 Satankar M, Patil AK, Kautkar S, Kumar U (2020) Pearl millet: a fundamental review on underutilized source of nutrition. Multilogic Sci 34(10):1081–1084 Singh S (2014) Agricultural Machinery Manufacturers’ Association (AMMA-India) Singh SK, Patil AK, Kautkar S, Dwivedi PN (2021) Thermal performance for hydroponic maize fodder production. Bioscan 16(1):199–202 Singh SK, Patil AK, Kautkar S, Dwivedi PN, Singh AK (2022) Development and performance evaluation of evaporative cool hydroponic fodder production chamber. Range Manag Agroforest 43(1):132–138

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Soffer M, Lazarovitch N, Hadar O (2020) Real-time detection of water stress in corn using image processing and deep learning. EGU, Munich. https://doi.org/10.5194/egusphereegu2020-11370 Tassis LM, Tozzi de Souza JE, Krohling RA (2021) A deep learning approach combining instance and semantic segmentation to identify diseases and pests of coffee leaves from in-field images. Comput Electron Agric 186:106191. https://doi.org/10.1016/j.compag.2021.106191 Turkoglu M, Yanikoğlu B, Hanbay D (2022) Plant Disease Net: convolutional neural network ensemble for plant disease and pest detection. SIViP 16:301–309. https://doi.org/10.1007/ s11760-021-01909-2 Ueka Y, Yamashita J, Sato K, Doi Y (2013) Study on the development of the electric tractor: specifications and traveling and tilling performance of a prototype electric tractor. Eng Agric Environ Food 6(4):160–164 Usinin U, Gladyshev S, Grigoryev M, Shishkov A, Bychkov A, Belousov E (2013) Electric drive of an industrial tractor (No. 2013-01-2469). SAE technical paper. SAE, Warrendale, PA Wulandhari LA, Gunawan AAS, Qurania A, Harsani P, Triastinurmiatiningsih Tarawan F, Hermawan RF (2019) Plant nutrient deficiency detection using deep convolutional neural network. ICIC Express Lett 13:971. https://doi.org/10.24507/icicel.13.10.971

Promotion of Improved Forage Crop Production Technologies: Constraints and Strategies with Special Reference to Climate Change

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Ashish Kumar Gupta, M. L. Sharma, M. A. Khan, and P. K. Pandey

13.1

Introduction

Forage includes plant leaves and stems that may be fresh or preserved used for animals. Forage are edible parts of crops that not including the separated grain and basically utilized as a feed for grazing animals and also eaten by wildlife. The examples of forage crops are grasses, legumes, crucifers, etc. Forage crops are cultivated and may be used in the form of hay, pasture, fodder, and silage. The forage crops can be classified in various ways. But, in this chapter in order to create the simple understanding about classification the following two classification criteria has been used: (a) life cycle and (b) season of cultivation.

13.1.1 Life Cycle On the basis of life cycle forage crops can be divided into the following three categories: • Annual forage crops: In annual forage crop, establishment of seed is easier due to germination, growing, and maturation in one growing season. For example: cowpea, maize, berseem, sorghum, etc. • Biennial forage crops: Biennial forage crops require two growing seasons, at first season vegetative growth will occur and second growing season flowering

A. K. Gupta (✉) Rani Lakshmi Bai Central Agricultural University, Jhansi, Uttar Pradesh, India M. L. Sharma · M. A. Khan · P. K. Pandey Indira Gandhi Agricultural University, Raipur, Chhattisgarh, India # The Author(s), under exclusive license to Springer Nature Singapore Pte Ltd. 2023 R. K. Singhal et al. (eds.), Molecular Interventions for Developing Climate-Smart Crops: A Forage Perspective, https://doi.org/10.1007/978-981-99-1858-4_13

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will occur. For example: This category includes the crops of Brassica family, viz. turnips, rape, kale, etc. and some legumes like sweet clover. • Perennial forage crops: Perennial forage crops can sustain for more year and can go dormant for short time and then initiate germination from crowns, rhizomes, etc. For example: Guinea grass, Lucerne, Subabul (https://www.bighaat.com/ blogs/kb/forage-crops-and-its-importance-in-agriculture).

13.1.2 Season of Cultivation Similarly, on the basis of season of cultivation the forage crops can be also categorized into the following three categories: • Kharif forage crops: These crops are mainly grown between June and September. For example: Cowpea, Cluster bean, Field bean, Bajra, Sorghum, Maize. • Rabi forage crops: Such crops mainly grow between the month of October and December that may be extended up to the month of January. For example: Berseem, Lucerne, Oats, Barley. • Summer forage crops: The crops under this category are grown primarily between April and June. For example: Cowpea, Cluster bean, Field bean, Bajra, Sorghum, Maize (http://ecoursesonline.iasri.res.in/mod/page/view.php?id=5631 6) (Table 13.1).

13.2

Important Forage Crops Grown in India along with their Productivity

Among the kharif crops sorghum and rabi crops berseem cover almost 54% of the total cultivated forage area in the country.

13.2.1 Benefits of Forage Crops in Agriculture • Grass root systems of forage crops can improve the organic matter content of the soil, which ultimately improves the soil health as well as its fertility. • Forage crops provide the base to sustainable agriculture. • Forage crops contain vitamins, fiber, and proteins that enhance metabolic activity of the animals. • Forage crops are also an important feed material for poultry enterprises as it is rich in minerals which help in bone development, eggshell formation, fluid balance and in hormone production. • Forage crops reduce weed density and its incidence in the agricultural land. • It also plays an important role in checking the soil erosion.

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Table 13.1 The important forage crops grown in India and their green fodder productivity S. No. 1.

Vernacular name Berseem

2. 3. 4.

Lucerne Senji Shaftal

5.

Metha

English name Egyptian clover Alfalfa Sweet clover Persian clover Fenugreek

6. 7.

Lobia Guar

Cowpea Clusterbean

8. 9. 10. 11. 12. 13. 14. 15.

Rice bean Jai Jau Jowar/Chari Bajra Makka Makchari Chara sarson

Sutri Oat Barley Sorghum Pearl millet Maize Teosinte Chinese cabbage

Botanical name Trifolium alexandrinum Medicago sativa Melilotus indica Trifolium resupinatum Trigonella foenumgraecum Vigna unguiculata Cyamopsis tetragonoloba Vigna umbellata Avena sativa Hordeum vulgare Sorghum bicolor Pennisetum glaucum Zea mays Zea mexicana Brassica pekinensis

Green fodder productivity (tones/ha) 60–110 60–130 20–30 50–75 20–35 25–45 15–30 15–30 35–50 25–40 35–70 20–35 30–55 30–50 15–35

Source: http://agropedia.iitk.ac.in/content/area-under-fodder-production-india

• It also helps in enhancing crop diversity, wildlife habitat, and soil ecosystem services (https://www.bighaat.com/blogs/kb/forage-crops-and-its-importance-inagriculture; https://prepp.in/news/e-492-forage-crops-agriculture-notes).

13.3

Importance of Forage Crop in Livestock Sector

India occupies first rank in the production as well as in consumption of milk (Akshit et al. 2020). The quantity and quality of fodder crops is an essential factor that is directly responsible for augmenting productivity of livestock. Scarcity of quality fodder may result in gradual decrease in cattle population (Ekka et al. 2014). Forage crops serve a significant role in beef cattle industry and among them fodder alone contributes the two-third proportion of animal feed (Meena et al. 2017a). Addition of forage crops in the diet of dairy animal makes their feed cost much cheaper then concentrate ration (Pawar et al. 2019).

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Impact of Climate Change on Forage Crop

Climate change is a global phenomenon and its adverse impacts have been acknowledged and documented throughout the world. But the developing countries like India are more susceptible to climate change then the developed nations. It is so for the reason that agriculture is major contributor of their economy and livelihood source of their citizens (Parry et al. 2001; Kumar et al. 2017). Adverse effect of climate change is experienced in the form of changes in CO2 concentration and temperatures in the atmosphere, increased frequency of extreme precipitation events, and changes in weed, pathogen, insect, and pest incidences (Hopkins and Del Prado 2007; IPCC 2007; Akshit et al. 2020). All such changes in climate result in different environmental stresses on forage crops that consequently affect the development, growth, and production of forage crops negatively (Ziervogel et al. 2006; Tubiello Francesco et al. 2007). As per the IGFRI Vision 2050 (n.d.), at this moment in time India faces a net shortage of 44.00% of concentrate feed ingredients, 35.60% of green fodder, and 10.95% of dry crop residues. Experts considered climate change as one of the prime factors that affects such deficiency and it also impacts negatively on forage production and livestock management (Akshit et al. 2020).

13.5

Constraints in Forage Production, Its Availability, and Adoption of Improved Forage Production Practices

There are several factors that act as constraints in forage production, its availability and adoption of improved forage production practices. Among these some factors are very common that are faced by the dairy farmers and such common factors are described here. Regarding production constraints the major constraints at farm level are poor soil fertility, unavailability of regular grassland, lack of land availability for fodder cultivation, lack of timely availability of quality seed material, lack of need based training program on latest methods of forage cultivation, etc. (Suman et al. 2015; Banerjee and Biradar 2016; Meena et al. 2017b; Kumar et al. 2018; Worqlul et al. 2021). As far as constraints in forage availability are concerned the major ones are lack of knowledge of proper fodder conservation methods, poor fodder storage infrastructure facilities, damage and loss of stored fodder crops by termites or rats, lack of knowledge about sources of unconventional fodder (Pawar et al. 2019). While in case of major constraints in adoption of improved forage cultivation practices, the major restrictions are as follows: limited land size, scarcity of improved and quality forage seed, lack of awareness among the farmers about improved forage cultivation practices, poor reach out of existing extension services, erratic rain fall, lack of interest of farmers, lack of on farm trial (OFT) on latest methods of forage cultivation (Admassu 2008; Assefa et al. 2015; Suman et al. 2015; Yadessa 2015; Salo et al. 2017).

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On the basis of understanding gained through interaction with farmers, review of literatures, and discussion with experts working on this field the following strategies are suggested (Fig. 13.1).

13.6.1 Creating the Awareness Amongst the Dairy Farmers Extension functionaries should take some initiatives for creating awareness among the dairy farmers about the various forage crops that may fulfill their need for cattle feed. The forage crop promoted here should have the characteristics like nutrient rich, easily cultivable, economic, higher productivity, require minimal management efforts. Here efforts also made to aware the dairy farmers about the ill effects of climate change through climate change literacy, i.e. well coupled with the farmers’ own experience on climate change. In this stage extension method like conducting front line demonstration (FLD), exposure visit, and field day on proven climatesmart forage crops may prove wrathful in this stage.

Fig. 13.1 Conceptual framework of extension strategies for promoting forage crops and their climate resilient cultivation technologies

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13.6.2 Skill Enhancement and Up Gradation This stage mainly oriented toward imparting the need based and vocational training to dairy farmers to develop the skills among them about scientific cultivation of forage crops in order to ensure better quality along with higher productivity, which also ensures effective and efficient use of climate resilient technologies used for their cultivation. In order to facilitate experiential learning among the dairy farmers the extension agent approaches the farmers through farmers’ field school. It must be conducted at farmers’ field so that they can learn from what they have and where they live. Capacity building steps also incorporate the learning by doing principle of extension.

13.6.3 Dissemination of Relevant Technologies For this purpose on farm testing (OFT) may be proved as most suitable extension method because OFT is conducted at dairy farmers’ field that will facilitate the technology assessment and corresponding refinement in it. This method also motivates other dairy farmers to apply the same technology at their field. In order to facilitate smooth and effective dissemination a farmer centric transfer of technology (TOT) model should be developed. To make TOT model farmer centric the model should be specific to location where its implementation is planned and should be oriented to available resources of the dairy farmers specially the small and marginal one. For dissemination of climate resilient technologies for forage crop cultivation, the extension personnel should emphasize over mixed approach, i.e. combination of individual, group, and mass extension methods. In this way, by following such steps the extension personnel can ensure the easy and quick adoption of climate resilient technologies for forage crop cultivation.

13.6.4 Utilization of ICT Tools At present context, extension functionaries can use ICT tools as an alternate to conventional extension methods and it also supplements and facilitates the extension effort. ICT tools are mainly known for their wider coverage of farmers in shortest period of time. ICT tools especially the agri-app assist the extension efforts by providing the real time weather data and also provide weather based advisory services (WBAS) that is manly based on predicted weather data. Some successful examples on this lines are WBAS by District Agro-Meteorology Unit (DAMU), where farmers can get weather based advisory services, i.e. delivered twice in week, viz. Tuesday and Friday. Apart of this Meghdoot, Damini mobile apps were also launched by the Indian government institutions. Kisan mobile advisory services (KMAS) by KVK also play an important role in this context. In this way such apps help the farmers in the management of fodder crops in their field by considering the weather based data obtained from these ICT tools.

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13.6.5 Convergence: An Integrated Approach Convergence among different agricultural organizations involved in similar kind of area may enhance effective implementation of climate resilient technologies for forage crop production. An effective convergence may be established between: KVK, ATMA, ICAR institutes, Co-operative societies, NABARD, CAU/SAU, etc. Convergence mainly enhances the effective and efficient utilization of partner organizations’ resources and capabilities and also prevents from the duplicity of the work.

13.6.6 Development of Technology Capsule for Promoting Forage Crops and Their Climate Resilient Production Technologies and Ensuring Its Availability Here, the term technology capsule stands for aggregation of all relevant and appropriate techniques of forage crop cultivation in one place and then categorize the technologies as per farmer’s need, resources, and location. The dissemination of such capsule should be facilitated by developing farmer’s centric TOT model and by ensuring the effective coordination and active involvement of primary agriculture credit societies (PACS), office of village and block level extension functionaries, ATIC of CAU, SAU, and ICAR institutes. The selection of these centers is primarily based on their easy accessibility by the farmers.

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