Amaranthus: A Promising Crop of Future

Saubhik Das

Amaranthus: A Promising Crop of

Future Saubhik Das Department of Botany Taki Government College West Bengal , India

ISBN 978-981-10-1468-0 ISBN 978-981-10-1469-7 (eBook) DOI 10.1007/978-981-10-1469-7

Library of Congress Control Number: 2016945253

© Springer Science+Business Media Singapore 2016 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifi cally the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfi lms 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 specifi c 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, express or implied, with respect to the material contained herein or for any errors or omissions that may have been made.

Printed on acid-free paper

This Springer imprint is published by Springer Nature The registered company is Springer Science+Business Media Singapore Pte Ltd. Pref ace

Pseudocereals are promising crops for coming decades keeping in view the global food security. Amaranths are a leading group of plants among the pseudocereals that have the great potentiality to prevent malnutrition espe- cially in the low-income food-defi cient countries. Though its cultivation and use as food grains have great antiquity, but later its cultivation was ignored by a larger section of people of the world. It lagged behind the conventional cereal crops in spite of being nutritionally very much competitive to those. In the last few decades, the nutritive potentialities and other unique features of amaranths have been rediscovered in different corners of the world. Research done so far on amaranths has surfaced its unique and unparallel nutritive value, wide adaptability, herbicide resistance property and vast germplasm variability. Its germplasm variability has opened up a new avenue to evolve much improved varieties or cultivars. Little efforts have been devoted to improve its genetic background by applying biotechnological methods. But that yielded signifi cant achievements. Keeping in view its tremendous poten- tiality to become a super crop of coming decades, much more attention is to be given to it especially when the conventional crops are overburdened with a task to feed the huge world population. To increase the food production at a global level and at a sustainable basis, the importance of amaranths cannot be ignored. In this book, attempts have been made to accumulate updated information, signifi cant research achievements and knowledge about amaranths also to revive interest about amaranths and to explore them comprehensively. The author expresses his sincere gratitude to Dr. David Brenner (curator of amaranth, North Central Regional Plant Introduction Station, Iowa State University, Ames, Iowa), Dr. Duilio Iamonico (Department of PDTA, Section of Environment and Landscape, University of Rome Sapienza, Italy) and Dr. J. C. Rana (head, Division of Germplasm Evaluation, ICAR-NBPGR, New Delhi, India), having pioneering contribution in amaranth research, for their help and cooperation. The author is also thankful to Dr. G. Nessom (Flora of North America Association); Dr. K. N. Gandhi (Harvard University Herbaria); the director of the Central National Herbarium, Howrah, Shibpur,

v vi Preface

West Bengal, India; and the head of the Department of Vegetable Crop, TNAU, Coimbatore for their necessary Cooperation. The effort devoted in this book will be successful if the book gets recogni- tion and appreciation. The author welcomes relevant comments and sugges- tions for future improvement of this book.

Taki , West Bengal , India Saubhik Das Contents

1 Introduction ...... 1 1.1 General ...... 1 1.2 Agricultural Development and Global Food Security ...... 1 1.3 Underutilised Crop ...... 3 2 Pseudocereals: An Efficient Food Supplement ...... 5 2.1 General ...... 5 2.2 Floristic Composition of India ...... 5 2.3 Pseudocereals: An Alternative Food Resource ...... 6 2.4 Nutritive Value of Pseudocereals ...... 7 3 Amaranths: The Crop of Great Prospect ...... 13 3.1 General ...... 13 3.2 Grain Amaranths: A Nutritive Supplement to Major Cereals ...... 14 3.3 Different Species of Grain Amaranths ...... 15 3.3.1 Amaranthus hypochondriacus L...... 18 3.3.2 Amaranthus cruentus L...... 20 3.3.3 Amaranthus caudatus L...... 21 3.4 Potentiality of Grain Amaranth as Food ...... 23 3.5 Vegetable Amaranths ...... 25 3.6 Species of Vegetable Amaranths ...... 27 3.6.1 Amaranthus tricolor L...... 27 3.6.2 Amaranthus blitum L...... 29 3.6.3 Amaranthus dubius L...... 30 3.6.4 Amaranthus cruentus L...... 30 3.6.5 Amaranthus viridis L...... 32 3.6.6 Amaranthus graecizans L...... 33 3.7 Nutritive Value of Vegetable Amaranths ...... 34 3.8 Uses of Amaranths ...... 35 3.8.1 Uses of Vegetable Amaranths ...... 35 3.8.2 Uses of Grain Amaranths ...... 36 3.9 Culinary Properties of Amaranths ...... 39 3.10 Bioactive Components and Medicinal Properties ...... 39 3.11 Weed Amaranths ...... 42

vii viii Contents

4 Infrageneric Classification of Amaranths ...... 49 4.1 General ...... 49 4.2 Synopsis of Infrageneric Classifi cations of Amaranths ...... 49 4.2.1 Amaranthus Subgen. Acnida Aellen ex K.R. Robertson ...... 50 4.2.2 Amaranthus Subgen. Amaranthus ...... 50 4.2.3 Amaranthus Subgen. Albersia Grenier & Godron ...... 51 4.3 Infrageneric Classifi cation After the Modifi cation by Mosyakin and Robertson (1996) ...... 51 4.3.1 Amaranthus Subgen. Acnida (L.) Aellen ex K.R. Robertson ...... 52 4.3.2 Amaranthus Subgen. Amaranthus ...... 53 4.3.3 Amaranthus Subgen. Albersia ...... 54 4.4 Provisional Key to Some Edible Species of Amaranthus...... 55 5 Taxonomy and Phylogeny of Grain Amaranths ...... 57 5.1 General ...... 57 5.2 Morphological Approach in Solving Taxonomic Disputes ...... 58 5.3 Biochemical and Molecular Approach in Understanding Systematics and Taxonomy ...... 65 5.4 Cytogenetical Approach in Understanding Species Relationship ...... 75 5.5 Taxonomic Delimitation in Vegetable Amaranths ...... 80 5.6 Key to the Varieties of Amaranthus tricolor L...... 80 5.7 Taxonomic Delimitation in Weed Amaranths ...... 88 5.8 Different Phylogenetic Concepts on Grain Amaranths ...... 88 6 Weed and Herbicide Resistance ...... 95 6.1 General ...... 95 6.2 Resistance to Different Types of Herbicides...... 95 6.2.1 Resistance to Photosystem II Inhibitors ...... 96 6.2.2 Resistance to Acetolactate Synthase (ALS) Inhibitors ...... 96 6.2.3 Resistance to Protoporphyrinogen Oxidase (PPO) Inhibitors ...... 96 7 Distribution and Maintenance of Amaranth Germplasm Worldwide ...... 99 7.1 General ...... 99 7.2 Germplasm Collection Centres in the World ...... 99 7.3 Genome Resource Development of Amaranthus ...... 104 8 Breeding of Amaranths ...... 107 8.1 General ...... 107 8.2 Breeding Behaviour in Amaranths ...... 108 8.3 Objectives in Amaranth Breeding ...... 112 Contents ix

8.4 Conventional Breeding by Selection ...... 116 8.5 Hybridisation and Heterosis in Amaranths ...... 118 8.6 Male Sterility in Amaranth Breeding ...... 125 8.7 Wild Gene Pool, Landraces and Heritability ...... 126 8.8 Mutation Breeding ...... 132 8.9 Polyploidy in Amaranth Breeding ...... 136 8.10 Genetic Engineering and Biotechnological Approach in Genetic Improvement ...... 137 8.11 Genetic Transformation in Breeding Amaranths ...... 138 8.12 Molecular Markers and Genome Research for Using Genetic Resources in Plant Breeding ...... 142 8.13 Molecular Breeding in Amaranths ...... 143 8.14 Plant Tissue Culture Techniques in Amaranth Breeding .... 144 9 Evolution of Sexuality in Amaranths ...... 147 9.1 General ...... 147 9.2 Trend of Sexuality in Plants ...... 147 9.3 Sexuality in Amaranths ...... 149 10 Cultivation of Amaranths ...... 153 10.1 General ...... 153 10.2 Seed Dormancy and Seed Viability of Amaranthus ...... 154 10.3 Agroclimatic Condition for Cultivation of Amaranths ...... 159 10.3.1 Climate ...... 159 10.3.2 Soil ...... 160 10.3.3 Temperature ...... 160 10.3.4 Rainfall ...... 161 10.3.5 Day Length ...... 161 10.4 Cultivation Practices ...... 161 10.4.1 Sowing of Seeds ...... 161 10.4.2 Seedling Growth and Interculture ...... 162 10.4.3 Manuring ...... 163 10.4.4 Obstacles in Productivity in Grain Amaranths ..... 163 10.4.5 Harvesting ...... 164 10.4.6 Pests and Diseases ...... 165 11 Future Prospects in Amaranth Research ...... 167

Appendices ...... 173 Appendix I ...... 173 Appendix II ...... 174

References ...... 189