Induced Mutation Breeding 9811994889, 9789811994883

Table of contents :
Foreword
Preface
Contents
About the Author
Chapter 1: Introduction/Review
1.1 Mutagenesis and Radiobiological Aspects
1.2 Acute and Chronic Irradiations
Chapter 2: Impact of Mutation Technique
Chapter 3: Definition
3.1 Types of Mutations
Chapter 4: Mutagens
4.1 Treatment Procedure
Chapter 5: Experiment and Data Collection
5.1 Experiment
5.2 Data Collection
5.2.1 Common Characters: Vegetative
5.3 Floral Characters
5.3.1 Analysis of Data and Interpretation
5.3.2 Mitotic Index Calculation
Chapter 6: Radiosensitivity
6.1 Split Dose, Storage
6.2 Acute and Chronic Radiation
6.3 Experimental Materials
6.4 Mutagen, Propagules, and Treatment Dose
6.5 Enzyme-Related Radioprotection
6.6 Experiment: Bulb Propagated Crop (Allium cepa)
6.7 Experiment: Seed Propagated Crop (Lens culinaris)
Chapter 7: LD50 Dose
7.1 Experiment: Seed Propagated Crops: X-Irradiation
7.2 Makhana (Euryale ferox Salisb.): Gamma Irradiation
7.3 Experiment: Vegetatively Propagated Crops: Rose: Gamma Irradiation
7.4 Chrysanthemum: Gamma Irradiation
Chapter 8: Chromosomal Aberrations
8.1 Mitotic Chromosomes
Chapter 9: Morphological Abnormalities
9.1 Experiment: Seed Propagated Crops
9.2 Experiment: Vegetatively Propagated Crops: Chrysanthemum and Rose
Chapter 10: Histology
10.1 Micromorphology
Chapter 11: Effects on Pollen Grain
11.1 Experiment: Seed Propagated Crops
11.2 Vegetatively Propagated Crops
Chapter 12: Colchi-Mutation
12.1 Experiment
Chapter 13: Combined Treatment
Chapter 14: Recurrent Irradiation
14.1 Chrysanthemum
14.2 Rose
Chapter 15: Detection of Mutations
15.1 Seed Propagated Crops
15.2 Vegetatively Propagated Crops: Chrysanthemum
15.3 Detection of Mutants in Later Generations
Chapter 16: Management of Chimera and In Vitro Mutagenesis
16.1 Experiment
Chapter 17: Induced Mutagenesis in Cross-Breeding
Chapter 18: Adventitious Bud Technique
Chapter 19: Present Status of Mutation Technology
Chapter 20: Spontaneous Mutation (Bud Sport)
Chapter 21: Somaclonal Variation
Chapter 22: Nanoparticles
References

Citation preview

Subodh Kumar Datta

Induced Mutation Breeding

Induced Mutation Breeding

Subodh Kumar Datta

Induced Mutation Breeding

Subodh Kumar Datta Floriculture Laboratory National Botanical Research Institute Lucknow, India

ISBN 978-981-19-9488-3 ISBN 978-981-19-9489-0 https://doi.org/10.1007/978-981-19-9489-0

(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

Mutation plays an essential role in evolution. It is the ultimate source of new alleles in plants that creates new genotypes. Mutation-induction techniques are a valuable tool, competitive to more expensive and complicated transgenesis and other molecular approaches. In the past 80 years, plant breeders have made extensive use of induced genetic diversity to select for desirable traits and in crop improvement. Mutation breeding has played a major role in floriculture, horticulture, and agronomy. Vast knowledge developed on induced mutation, worldwide, is spread in the form of scientific journals, books, bulletins, and catalogues. However, to date, there was no comprehensive reference book on mutation breeding. It is difficult for researchers, students, and breeders to get an overview of earlier and recent developments covering the whole spectrum of activities on induced mutation. This book is an exhaustive account on mutation breeding, covering all important crops. It has been prepared based on published information as a reference document. This practical guide of approximately 400 pages covers an impressive number of 50 pages of useful references listed in a tabular form. The book contains 22 synthetic chapters on various important basic and practical aspects of mutation that are very informative and the reader can effortlessly follow the references on the topic-wise manner. The book provides an authoritative review account of many aspects related to crop improvement through induced mutations. It caters not only to classical mutation breeding (colchicine mutation, irradiation) but also to relevant modern aspects, such as somaclonal variation and the utilization of nanoparticles. This compilation will be a valuable source of information for next-generation mutation breeding scientists. The editor of this book—Dr. Subodh K. Datta—is a distinguished researcher with 35 years of experience in induced mutagenesis. He has begun his scientific career in 1971 at Bose Institute, Kolkata, India, and obtained his Ph.D. degree in 1976 from Calcutta University. Dr. Datta continued his research activities at the National Botanical Research Institute (NBRI) and received D.Sc. degree in Botany from Kanpur University in 1994 for extensive induced mutagenesis work on vegetatively propagated ornamentals. He retired from NBRI in July 2007 as Scientist “G” v

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(Director’s Grade Scientist), Coordinator and Head, Botanic Garden and Floriculture Division. Nonetheless, Dr. Datta successfully continued his research as the Council of Scientific and Industrial Research (CSIR) Emeritus Scientist at Bose Institute, Kolkata until 2013. In 2015, he has completed a Visiting Research Professorship at the Department of Life Science and Bioinformatics, Assam University. Recently, Dr. Datta has published a number of highly informative papers in scientific journals and in the form of book chapters. The main research field of the editor included various aspects related to cytomorphology, conventional breeding followed by selection, in vivo and in vitro mutagenesis, tissue culture, management of chimera, plant conservation, molecular characterization, dehydration of flowers, and foliage and floral craft on different ornamentals. In the course of his professional career, Dr. S.K. Datta has created over 100 new cultivars of different ornamentals for the floriculture trade. Moreover, medicinal plants, vegetable crops, essential oil-bearing plants, and non-edible oil-bearing crops were included in his mutation breeding work. The editor has broad experience with both chemical and physical mutagens applied on seed and vegetatively propagated crops of various genera and botanical families. The target of his studies covers amaryllis, Asiatic hybrid lily, bougainvillea, chrysanthemum, dahlia, gerbera, gold lantana, marigold, gladiolus, hibiscus, rose, tuberose, orchids, and others that are internationally competitive. Dr. Datta continues to be a prolific writer and an author or co-author of over 250 research papers (both original and reviews) on induced mutagenesis and has published an impressive number of books, book chapters, and bulletins. A sincere attempt has been made by Dr. S.K. Datta to highlight a complete and comprehensive scenario on induced mutation activities worldwide in the form of a reference book so that innovations made so far can be used judiciously for this sector. This practical guide provides an illustrated mutation breeding work, cropand topic-wise. It is an important treatise that contains a large volume of information on different aspects of induced mutations. The book includes suggestions for future need-based planning, considering modern developments of ongoing and oncoming research areas. It will be an invaluable asset to both mutation breeding scientists and commercial companies interested in creating novel cultivars of various crops. Faculty of Agriculture and Biotechnology, Bydgoszcz University of Science and Technology, Bydgoszcz, Poland

Dariusz Kulus Ph.D.

Preface

To increase genetic variability within a crop, several plant breeding methods are followed. One of the main methods is cross breeding with which the breeder tries to combine the beneficial characters from different sources into one genotype. From such pooled genotypes, sometimes it is possible to directly select a particular genotype which is superior to the existing cultivars. Mutation breeding, on the other hand, makes use of possibilities of altering genes by exposing seeds or other plant parts to physical and/or chemical mutagens. Mutation breeding is an established method for crop improvement. My involvement with induced mutations goes back to 1971 when I started my research career at Bose Institute, Kolkata, India on induced mutagenesis for crop improvement for my Ph.D. work. I was associated with Council of Scientific and Industrial Research-National Botanical Research Institute (CSIR-NBRI), Lucknow, India for almost 30 years (1977–2007) and was engaged in a multidisciplinary research program on induced mutagenesis on various crops with special reference to ornamental crops. National Botanical Research Institute Lucknow is one of the pioneer institutions where commendable work has been done on induced mutagenesis. My main research activities were development of new and novel varieties through in vivo and in vitro mutagenesis. My long association with mutation breeding scientists has immensely benefited my comprehension on mutation. The benefits of induced mutagenesis are well known and the promises it has are still unfathomable. Induced mutagenesis has generated a vast amount of genetic variability and has played a significant role in plant breeding and genetic studies. Vast literature/knowledge generated on induced mutation, worldwide, is spread in the form of books, bulletins, catalogues, scientific journals, newsletters, newspapers, popular magazines, etc. It is difficult for all researchers, students, and breeders to get an overview of earlier and recent developments covering the whole spectrum of activities on induced mutation. An attempt has been made in this book to put together all information to develop a complete documentation of the results of the research conducted by different researchers over the last more than 80 years as a reference book. The main objective of the book is to give a coherent and concise vii

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account of all work done on induced mutagenesis with an emphasis on recent developments. The document has been prepared only from published information as a reference document. All important publications of scientists on different crops have been cited. The publication of this book is planned to reveal multifarious activities done on different aspects of mutation so that achievements made so far can be used judiciously for designing new experiments in future. It is my sincere endeavor to present a complete and comprehensive scenario on mutation breeding research carried out in different countries through this book. The book also provides a comprehensive review account of many aspects of current interest and progress on mutation research. The number of papers/review articles/books, etc. published on mutation breeding, worldwide, is so voluminous that it is difficult for one scientist to collect all. All the references have been cited for technical support and interpretation. Crop improvement is a never-ending project and there is always search for new techniques. Spontaneous bud mutations, somaclonal variations, and application of nanoparticles have played a very important role in developing different types of changes in different crop plants including ornamentals. Genetic changes in plants through somaclonal variation caused by mutation are regenerated from tissue culture. Spontaneous mutations are the most significant source of genetic variation and their characterization provides important information about evolutionary processes and induced mutation. The literature on application of genotoxic effects of nanoparticles in agriculture is increasing. Important references on these topics have been cited in the book. I sincerely acknowledge and profusely thank all my professional colleagues of mutation breeding for publishing a good amount of information on mutation activities. I have tried to put together all references to develop a complete documentation of the results of the research conducted by different scientists all over the world last about 80 years. The book deals with all the important and relevant aspects of mutation breeding. The book provides authoritative review account of many aspects related to induce mutagenesis in the field of crop improvement. The references have been arranged crop wise and important topic wise in the book which caters not only classical mutation breeding but also relevant modern aspects. I am sure that the information of the book will be of great help to researchers, teachers, students, and breeders for planning future strategies for the development of new varieties. I thank all scientific societies, book publishers, and journal’s editors who have published research work on induced mutations. I sincerely acknowledge all of them from where I collected all references. Taking all aspects together, it is an excellent reference book of updated information on the mutation breeding. The book will be an excellent informative document for researchers and students for understanding application of induced mutations in crop improvement and biological research. The book will provide an up-to-date comprehensive reference manual for proper application of mutagenesis. Important and useful reference lists have been cited in all important crops and on basic aspects of mutagenesis. Though meticulous care has been taken in reviewing but while dealing with such a voluminous work, some

Preface

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mistakes/errors might have crept in, despite my best efforts and non-availability of publications. I wish to record my sincere thanks to all the Directors, CSIR-NBRI for providing all working facilities, favorable suggestions, and constant encouragement. I sincerely thank my teacher, Late Prof. Dr. R.K. Basu, D.Sc. Cytogenetic Laboratory, Department of Botany, Bose Institute, Kolkata, India under whose guidance I did my Ph.D. thesis work. I sincerely acknowledge my long association at National Botanical Research Institute, Lucknow, India where I did all my research activities on different ornamental crops on multidisciplinary aspects. I specially thank my own students and convey my deepest feelings of gratitude to all of them. I shall be failing my duty if I do not record the blessings of my parents, whose unfathomable love and blessings kept me ever up and doing. It would be a remiss, if I don’t acknowledge the warmth of my wife who always stood by me steadfast. She is a charming, understanding, sociable lady who took care of me and family with effusive affection. She admirably adjusted me to pursue my research interest with unison devotion. I record my sincere gratitude to my wife Dr. Kalyani Datta and son Shouvik Dutt who extended full cooperation in many invisible ways during my research. I warmly appreciate cooperation and approval of Springer Nature Singapore Pte. Ltd., 152 Beach Road, #21-01/04 Gateway East, Singapore 189721, Singapore for publishing the book in excellent manner. I am confident that the book will be widely accepted by the students, teachers, and researchers in the field of crop improvement. I hope that this book would provide valuable data and also be a reference material for future research activities in this area. Lucknow and Kolkata, India

Subodh Kumar Datta

Contents

1

Introduction/Review . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.1 Mutagenesis and Radiobiological Aspects . . . . . . . . . . . . . . . . . 1.2 Acute and Chronic Irradiations . . . . . . . . . . . . . . . . . . . . . . . . .

1 9 14

2

Impact of Mutation Technique . . . . . . . . . . . . . . . . . . . . . . . . . . . .

75

3

Definition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.1 Types of Mutations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

81 81

4

Mutagens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.1 Treatment Procedure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

85 88

5

Experiment and Data Collection . . . . . . . . . . . . . . . . . . . . . . . . . . 5.1 Experiment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.2 Data Collection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.2.1 Common Characters: Vegetative . . . . . . . . . . . . . . . . . 5.3 Floral Characters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.3.1 Analysis of Data and Interpretation . . . . . . . . . . . . . . . 5.3.2 Mitotic Index Calculation . . . . . . . . . . . . . . . . . . . . . .

. . . . . . .

91 91 92 93 94 94 94

6

Radiosensitivity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.1 Split Dose, Storage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.2 Acute and Chronic Radiation . . . . . . . . . . . . . . . . . . . . . . . . . . 6.3 Experimental Materials . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.4 Mutagen, Propagules, and Treatment Dose . . . . . . . . . . . . . . . . 6.5 Enzyme-Related Radioprotection . . . . . . . . . . . . . . . . . . . . . . . 6.6 Experiment: Bulb Propagated Crop (Allium cepa) . . . . . . . . . . . 6.7 Experiment: Seed Propagated Crop (Lens culinaris) . . . . . . . . .

95 102 102 103 103 105 106 107

7

LD50 Dose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109 7.1 Experiment: Seed Propagated Crops: X-Irradiation . . . . . . . . . . 110 7.2 Makhana (Euryale ferox Salisb.): Gamma Irradiation . . . . . . . . . 110

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7.3 7.4

Experiment: Vegetatively Propagated Crops: Rose: Gamma Irradiation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111 Chrysanthemum: Gamma Irradiation . . . . . . . . . . . . . . . . . . . . 111

8

Chromosomal Aberrations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113 8.1 Mitotic Chromosomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114

9

Morphological Abnormalities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117 9.1 Experiment: Seed Propagated Crops . . . . . . . . . . . . . . . . . . . . . 118 9.2 Experiment: Vegetatively Propagated Crops: Chrysanthemum and Rose . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122

10

Histology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 123 10.1 Micromorphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124

11

Effects on Pollen Grain . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125 11.1 Experiment: Seed Propagated Crops . . . . . . . . . . . . . . . . . . . . . 125 11.2 Vegetatively Propagated Crops . . . . . . . . . . . . . . . . . . . . . . . . 127

12

Colchi-Mutation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131 12.1 Experiment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132

13

Combined Treatment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135

14

Recurrent Irradiation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139 14.1 Chrysanthemum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140 14.2 Rose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140

15

Detection of Mutations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15.1 Seed Propagated Crops . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15.2 Vegetatively Propagated Crops: Chrysanthemum . . . . . . . . . . 15.3 Detection of Mutants in Later Generations . . . . . . . . . . . . . . .

16

Management of Chimera and In Vitro Mutagenesis . . . . . . . . . . . . 149 16.1 Experiment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157

17

Induced Mutagenesis in Cross-Breeding . . . . . . . . . . . . . . . . . . . . . 161

18

Adventitious Bud Technique . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163

19

Present Status of Mutation Technology . . . . . . . . . . . . . . . . . . . . . . 165

20

Spontaneous Mutation (Bud Sport) . . . . . . . . . . . . . . . . . . . . . . . . . 171

21

Somaclonal Variation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173

22

Nanoparticles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177

. . . .

143 143 144 144

References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 179

About the Author

S. K. Datta, Ph.D., D.Sc. is an internationally acclaimed expert in floriculture and mutation breeding. Dr. Datta was engaged in both basic and applied research for the improvement of floriculture for over more than 35 years at CSIR-NBRI, Lucknow, India. His main field of research was induced mutations and improvement of ornamental plants. For mutation breeding work, he applied both physical (X-rays, Gamma rays) and chemical (EMS, MMS, Colchicine) mutagens and developed more than 100 new varieties in different ornamentals. He has published series of research papers (approx. 338) on different aspects related to floriculture and induced mutagenesis. He has published three books, edited five books and eight bulletins on different aspects related to induced mutagenesis and floriculture. Dr. Datta visited Humboldt Universitat Zu Berlin, Berlin for 2 months (under CSIR-DAAD Scientists Exchange Programme) and Korea for 1 month (INSA-KOSEF Scientists Exchange Programme). International Atomic Energy Agency (IAEA), Vienna selected Dr. Data as “Expert on Mission” for evaluation of mutation breeding projects sponsored by IAEA to Philippines, Jakarta, Indonesia for project evaluation mission. Dr. Datta organized international training program as supervisor on induced mutagenesis sponsored by IAEA, Vienna at Central Research Institute for Horticulture, Cipanas, Indonesia. Dr. Datta presented papers in different international symposium—at VIth International Congress of SABRAO held at Tsukuba, Japan; International Nuclear Conference held at Putra World Trade Centre, Kuala Lumpur, Malaysia; International Symposium on “Underutilized Plant Species for food, nutrition, income and sustainable development” held at Arusha, Tanzania; FAO/IAEA International Symposium on the contribution of Plant Mutation Breeding to Crop Improvement, held at International Atomic Energy Agency, Vienna, Austria—four times.

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Chapter 1

Introduction/Review

Discovery of induced mutation is one of the most important breakthroughs in the history of genetics. Hermann J. Muller got Nobel Prize in Physiology or Medicine in 1946 for his discovery of induction of mutations using X-ray irradiation. Muller did experiments with fruit flies in 1927. Stadler (1928a, b) demonstrated that radiation can induce genetic variability in plants. Physical mutagens, mostly ionizing radiations, have been used widely for creating genetic variability and maximum mutant varieties have been developed. Since 1960, gamma rays have become the most commonly used mutagenic radiation in plant breeding. Ion beam radiation has also been tested as an effective and unique mutagen. Then α- and β-particles, fast neutrons, UV light, and even space radiation have also been demonstrated as useful mutagens. Induced mutagenesis is now an established method for crop improvement. Mutation breeding is considered as the obvious means to perfect the breeding products of conventional plant breeding and a possible shortcut for inducing desired genetic alterations in outstanding cultivars. Mutation techniques by using ionizing radiations and other mutagens have successfully produced quite a large number of new promising varieties in different plant species. Since beginning, there is a stepby-step improvement in technical procedure for the application of induced mutation for crop improvement and voluminous knowledge have developed for the successful and accurate application of the technique. The mutation breeding research began around 1930 but large-scale practical applications started in most countries after 1945. The foregoing pages contain a bird’s eye view of the prospects, procedures, possibilities, and problems of mutation breeding. The concept of induced mutation for crop improvement developed dates back to the beginning of the twentieth century. First observation on artificial induction of genetic changes was reported by De Vries (1905), Gager (1908), Muller (1927), and Stadler (1928a, b, 1932). A study of induced mutations commenced in 1927 by Muller who discovered the mutagenic property of X-ray in Drosophila. Stadler (1928a, b) demonstrated that similar mutation could be induced in plants (Maize and Barley) by X-ray. In the same year, mutation was also induced in Nicotiana by Goodspeed and Olson (1928). © The Author(s), under exclusive license to Springer Nature Singapore Pte Ltd. 2023 S. K. Datta, Induced Mutation Breeding, https://doi.org/10.1007/978-981-19-9489-0_1

1

2

1 Introduction/Review

Practical mutation breeding procedure by use of mutagenesis strictly for plant breeding developed after publications of Friesleben and Lein (1942). Hoffmann (1959), Gustafsson (1947a, b), and Mackey (1956) made systematic studies and generated information on optimal treatment doses, treatment conditions, and mutation frequency and mutation spectra. Nilan et al. (1965) concentrated on radiation treatment conditions and combined treatment for specific mutation. New varieties of rice appeared in the market as a result of mutation breeding work in developing countries (Sigurbjornsson and Micke 1969, 1974). In the beginning, X-irradiation on dry seed was applied as a standard method for studying the mutation process. Soon other types of irradiation such as gamma rays (acute and chronic), neutrons (fast and thermal), electrons, photons, alpha-rays from radon, and beta rays from phosphorus32 and Sulfur35 were included in the experiments. Mutation breeding experiment was based primarily on X-rays. But now gamma rays are mainly used for mutation and to a smaller extent fast or thermal neutrons are also used. A number of chemical mutagens were identified for their mutagenic properties (Robbelen 1959; Ehrenberg 1960, 1971; Auerbach 1946, 1961; Auerbagh 1974; Auerbach and Robson 1946, 1947; Fischbein et al. 1970; Rapoport 1956; Rapoport et al. 1966; Schwochau and Hadwiger 1996; Konzak et al. 1965; Kuleshov et al. 1974; Mee et al. 1969; Veleminsky and Gichner 1970; Veleminsky et al. 1970; Westergaard 1957, 1960; von Wettstein 1961). Several groups of chemical compounds have been distinguished as mutagens, such as alkylation agents, antibiotics, base analogs, and others. Alkylation agents are among the chemicals more frequently used. Most commonly used chemical mutagens are ethyl methanesulfonate (EMS), methyl methanesulfonate (MMS), diethyl sulfate (dES), ethyleneimine (EI), ethyl nitroso urethane (ENU), ethyl nitroso urea (ENH), methyl nitroso urea (MNH), Azides, etc. Various chemical mutagens give a higher mutation rate; some produce a higher ratio of gene mutations vs. deletions or other chromosome mutations. But there are several practical problems with chemical mutagens (soaking of seeds, penetration to the relevant target cells, safety of handling and disposal, poor reproducibility, persistence of the mutagen or its metabolites). Slowly use of physical mutagens was widespread and use of chemical mutagens was limited. Over the years many different mutagenic agents have been applied. In the beginning, mostly X-ray, 32P and, to a smaller extent, gamma rays were used, in a wide range of doses, dose rates, and concentrations. Food and Agriculture Organization of the United Nations (FAO) and the International Atomic Energy Agency (IAEA) created the joint FAO/IAEA Division of Nuclear Techniques in Food and Agriculture on October 1, 1964. IAEA and FAO of the United Nations through the joint FAO/IAEA Division of Nuclear Techniques in Food and Agriculture, Vienna serves as the global focal point for nuclear cooperation, mobilizing peaceful applications of nuclear science and technology for critical needs in developing countries. The joint FAO/IAEA Division has been promoting the efficient use of mutation techniques since the late 1960s. In 1969, the Joint FAO/IAEA Division started to organize training courses for plant breeders on the induction and use of induced mutations. This was a period when many students received fellowship training in “peaceful applications of nuclear techniques”

1

Introduction/Review

3

including the genetic effects of radiation. The dawn of the “Atomic Age” following World War II saw a boom of interest in utilizing ionizing radiation for peaceful purposes. By 1950, mutation induction research began to flourish in several countries using mainly 60Co gamma rays or neutrons obtainable from various newly established nuclear research centers. The IAEA Technical Cooperation Projects (TC Projects) and Coordinated Research projects (CRPs) are playing a unique role in capacity building, promoting research, and efficient use of nuclear techniques for plant breeding and networking among scientists of both developing and developed countries. The joint FAO/IAEA Programmers have been promoting the research, development, and application of nuclear techniques in food and agriculture in the Member States for more than 50 years. FAO/IAEA published the “Manual on Mutation Breeding” in 1970, which was the first book of its kind in the world. FAO/IAEA generated extensive knowledge and literature on almost all basic and applied aspects of induced mutagenesis. Details of mutant varieties officially developed and released worldwide are available at Mutant Variety Database (FAO/IAEA Mutant Variety Database, MVD, http://www-mvd.iaea.org/MVD/default.htm) (FAO/IAEA 1965, 1969a, b, 1970a, b, 1971a, b, 1973, 1975, 1976, 1977a, b, 1979, 1981, 1982, 1983, 1984a, b, 1986, 1988, 1989, 1991; Burkart 2009; Pandey 2009; Ishige 2009; Cetto 2009; Lagoda et al. 2009; Maluszynski et al. 2000; Micke 1991a, b; Shu 2009a, b). Author (Dr. S.K. Datta) started his research career on induced mutagenesis in 1971 and did voluminous work till his retirement in 2007. An appreciable amount of literature has been generated on crop improvement using classical and modern induced mutagenesis techniques on different aspects like radiosensitivity, selection of material, methods of exposure to mutagens, determination of suitable dose of mutagen, combined treatment, recurrent irradiation, split dose, colchi-mutation, detection of mutation, mutation frequency and spectrum of mutations, nature of chimerism, classical and modern methods for management of chimera, in vitro mutagenesis, isolation of mutants, cytological—biochemical and molecular characterization of mutants, commercial exploitation of mutant varieties, etc. (Datta 2012, 2014, 2015a, b, 2017). The following crops were selected under improvement programme: vegetable (Trichosanthes anguina L, T. cucumarina, Cucurbita maxima L, Cephalandra indica, Luffa acutangula Roxb., Lagenaria ciceraria), medicinal (Trigonella foenum-graecum L, Mentha citrate Ehrh), pulse (Winged Bean—Psophocarpus tetragonolobus L. D.C.), oil-bearing (Jatropha curcas L, Rosa damascena, Cymbopogon flexuosus (Nees) Wats), and ornamentals (Amaryllis, Asiatic Hybrid Lily, Bougainvillea, Canna, Chrysanthemum, Dahlia, Gerbera, Gladiolus, Hibiscus, Lantana depressa Naud, Tagetes erecta, Rose, Tuberose, Mesembryanthemum criniflorum L.F., Narcissus tazetta, etc.). Both physical (X-ray and Gamma rays) and chemical (EMS, MMS, Colchicine) mutagens were used as mutagens. Promising mutants have been isolated in Jatropha curcas (both tall and dwarf, high branching, high fruit, and oil yielding and high biomass yielding—Datta and Pandey 1992, 1993, 1996, 2005, 2013; Datta et al. 1998a, b; Pandey 1995; Pandey and Datta 1995); Cymbopogon flexuous (morphological and agronomical characters—Banerji et al. 1987a, b); Rosa damascena or Damask rose

4

1 Introduction/Review

(morphological characters, flower color—Gupta et al. 1988); Winged Bean (dwarf mutant with determinate growth habit, early fruiting—Jugran et al. 1986, 2001); Mentha citrata (hairy mutant and mutant with higher herbage yield—Gupta et al. 1979); Trigonella foenum-graecum (small seeds, bold seeds, green and chocolate seed coat color against normal green, dwarf and high branching, etc.—Laxmi et al. 1982, 1983a, b; Laxmi and Datta 1986, 1987, 1989; Datta and Laxmi 1992; Datta and Goel 2005); and Trichosanthes anguina (early flowering, short thick fruit, increased fruit weight, yellow fruit, high oil and punicic acid yielding, crinkled leaf, etc.—Basu and Datta 1977, 1982; Datta and Basu 1977; Datta 1986a, 1987a, 1989a, 1989b, 1991a, 1992b, 1994a). More than 76 new mutant varieties have been developed using gamma radiation in different ornamental plants (Bougainvillea-5, Perennial portulaca-6, Chrysanthemum-43, Hibiscus-1, Rose-16, Tuberose-2, Lantana depressa-3). The mutations were mainly in flower color/shape and chlorophyll variegation in leaves (Datta 1988a, 1989c, 1992a, 1994b, 1997a, b, 2000, 2004, 2005a, b, 2009a, b, c, 2015a, b). It is clear from these reports that the author from his induced mutagenesis work, successfully developed mutants in vegetable, medicinal, pulse, and oil-bearing crops. But the success was very prominent in ornamental crops. At this stage, there is no need to review the work done on induced mutagenesis. Series of books, research and review papers have already been published in detail on prospects, utilization of induced mutations, list of released, commercialized and approved mutant varieties in different crop plants (Abo-Hegazi 1991; Ahloowalia and Maluszynski 2001; Ahloowalia et al. 2004; Alkema 1971a, b; Amano 1981, 1985, 1986, 2001, 2006; Anonymous 1973, 1982a, b, 1991a, b, 1992, 1994a, b, 1995; Antonyuk 1991; Ashri 1982; Awan 1991, 1999, 2005; Bhatia 1991; Blakeslee et al. 1922; Blixt and Gottschalk 1975; Brar and Jain 1998; Brock 1965a, b, c, 1971; Broertjes 1968a*; Broertjes 1969a, 1969b, 1969c, 1969d, 1972a, 1972b, 1974a, 1974b, 1977a, 1977b, 1977c, 1977d, 2012; Breider 1963, 1964a, b; Broertjes and Van Harten 1988; Brunner 1991, 1992, 1995a, b; Buzzati-Traverso 1913; BuzzatiTraverso and Scossiroli 1958; Caldecott 1953; Caldecott et al. 1959; Casarett 1968; Chandramouli et al. 1989; Chen et al. 2006; Clayton and Robertson 1955, Clayton and Robertson 1964; Darlington and McLeish 1951a, b; D’Amato 1965, 1986, 1990; Datta 1988a, 1997a, b, 2001, 2002, 2004, 2005a, b, 2009a, b, c, 2012, 2014, 2015a, b; Datta and Chakrabarty 2005; Davies et al. 1963; De Vries 1905; Doll and Sandfaer 1969; Donini et al. 1984; Drake et al. 1983; Dryagina et al. 1966, 1967; Dryagina and Kazarinov 1966; Dryagina and Limberger 1974; Dubinin 1964; Enken 1966a, b; Etter 1965; FAO/IAEA 1965, 1969a, b, 1970a, b, 1971a, b, 1977a, b, 1981, 1982, 1984a, b, 1986, 1989, 1991; Friesleben and Lein 1942; Frey 1965; Gottschalk and Wolf 1983; Gamma Field Symposia (Japan) (1962, 1963, 1964, 1965, 1966), Gardner 1968, Gaul et al. 1971, 1972; Gaul 1958, 1960, 1961a, b, 1962, 1963, 1964a, b, 1965a, b, 1977a, b; Gichner and Veleminsky 1970; Gottschalk 1967; Granhall 1953; Gustafsson 1938, 1942, 1947a, b, 1954, 1960a, b, 1969a, b, 1975, 1986; Gustafsson and Teden 1954; Gustafsson et al. 1950, 1969; Gustafsson and Lundqvist 1980; Gustafsson and Mackey 1948; Gustafsson and von Wettstein 1958; Hagberg 1953, 1961; Hagberg et al. 1952, 1958; Harney 1976; van Harten 1998; Hine and Brownell 1956; Hoffmann 1959; Hoffmann and Walther 1961; Holst and Nagel 1997; Hough et al. 1965; Howland and Hart 1977; Institute of

1

Introduction/Review

5

Radiation Breeding 1962, 1963, 1964; Heinze and Schmidt 1998; IAEA 1961, 1965, 1971a, b, 1981, 1984a, b, 1985, 1986a, b, 1987, 1992, 1997, 2011; Jain 1998a, b, 2002, 2005, 2006a, b, 2010; Jorgensen 1974; Kahl and Meksem 2010; Kawai and Amano 1991a, Kawai and Amano 1991b; Kharkwal 2012a, b; Khvostova 1966; Kiyosawa and Nomura 1988a, b; Kodym et al. 2012; Konzak 1956a, b, 1957, 1959a, b, 1984; Konzak et al. 1976; Konzak and Mikaelsen 1977a, b; Kivi et al. 1974; Kuzin 1956; Lapins 1983; Lawrence 1965; Liamngee et al. 2021; Liu et al. 2006a, b; L€onnig 2005; Linqing 1991; McClintock 1948, 1956; MacKey 1956, 1981; Mba et al. 2009; Lonning 2005; Levy and Ashri 1975; Malepszy et al. 1973; Maluszynski 1995, 1999; Maluszyuski et al. 1992, 1995, 2000; Mahamad et al. 2006; Mandal and Datta 2005a; Marie 1967; Matsuo and Yamaguchi 1962; Mee et al. 1969; Mohd. Nazir et al. 1998; Micke 1974, 1979a, b, 1984a, b, c, 1987, 1988a, b, 1989, 1991a, b, 1992, 1996; Micke and Donini 1982, 1993; Micke et al. 1980, Micke et al. 1987, 1990a, b; Murty et al. 2004; Muller 1928a, b, 1954a, b; Nakagawa 2008; National Bureau of Standards Handbook (NBSH), No. 73 1960; de Nettancourt 1969; Nichterlein et al. 2000; Nilan 1966, 1967, 1972b; Nilan and Konzak 1961; Nilan et al. 1962a, b; Novak 1991a, b; Novak and Brunner 1992; Nybom 1961; Nybom and Koch 1965; Ohba 1971; Olmo 1960a, b; Parry et al. 2009; Patterson and Muller 1930; Privalov 1967; Rao et al. 1975a, b; Redei 1974; RivielloFlores et al. 2022; Robbelen 1957, 1990; Robbelen and Heun 1991; Roby 1972a, b, c; Ross et al. 1997; Roux 2004; Royani et al. 2021; Rutger 1992, 2006; Saccardo and Monti 1984; Sakinah 1998; Scarascia-Mungnozza 1965; Schum and Preil 1998; Scholz 1957; Schulz 1971; Scossiroli 1962, 1965; Serebrovsky 1929; Shapiro 1956a, b; Shu et al. 1997; Siddiqui and Khan 1999; Sigurbjornssen 1977; Sikora et al. 2011; Silvy and Mitteau 1986a, b; Somsri et al. 2008; Sparrow 1966a, b; Sparrow and Konzak 1958; Sparrow and Cuany 1959; Smith 1958, 1961a, b, 1971, 1972; Spiegel-Roy 1990; Swaminathan 1961, 1963a, b, 1965a, b, c, 1969, 1971, 1972; Swaminathan et al. 1968; Stadler 1930a, b; Stubbe 1929, 1940, 1942, 1958, 1959, 1967; Sigurbjornsson and Micke 1969; Scheibe and Micke 1967; Shamel and Pomeroy 1936; Song and Kang 2003; Shu et al. 2012a, 2012b; Strauss 1958; Szarejko and Maluszynski 1980; Szarejko et al. 1991; Timofeeff-Ressovsk 1932; Tomlekova 2010; Yamagata 1981; von Wettstein 1961; Vasline et al. 2005; Van Harten 1998; Van Harten and Broertjes 1986; Veleminsky 1965; Veronica et al. 2011; Vinh et al. 2009; von Wettstein et al. 1959; de Vries 1901; Wang 1986, 1985, 1991, 1992, 1998; Wange and Li 2005; Waugh et al. 2006; von Wettstein et al. 1959; Wu et al. 2000; Xiao 1983; Yamaguchi 1983, 1991, 1994, 2001, 2005; Yonezawa and Yamagata 1977; Zacharias 1956a, b; Zhu et al. 1991; Zheng 1995; Zhukov 1966, 1967; Zschege and Haaring 1962; Zwintzscher 1966a, b, etc.). These review articles have covered almost all aspects of induced mutagenesis. The widespread use of induced mutants in plant breeding programs throughout the world has led to date to the official release of close to 3218 mutant varieties from more than 170 different plant species. As mentioned, there are huge publications (both research papers and review articles) by many workers highlighting their success and failure on induced mutagenesis. But unfortunately, the background information for the success/failure of

6

1 Introduction/Review

mutation has not been adequately documented. Therefore, the author prepared a few review articles to highlight different technical information which are most important for the successful application of induced mutagenesis in ornamental crops. An attempt has been made to highlight different important basic aspects (radiosensitivity, experimental procedure, treatment, LD50 dose, material and time for treatment, data collection, analysis of data, detection and isolation of mutant/s, role of combined and recurrent treatment, sensitivity of mutant genotype, acute and chronic irradiation, colchi-mutation, domestication, management of chimera, in vitro mutagenesis, etc.) which may be helpful as a guideline for large scale mutagenesis work on any ornamental crop (Datta 2012, 2014, 2015a, b). Before starting mutation breeding work one should have up-to-date knowledge about all practical approaches and their limitations. Clear breeding objectives have to be set and a proper strategy must be outlined before any program is started. Researcher should select the best genotype and should have a solid knowledge of the genetics of the material. Selection of mutagens depends mainly upon the material and to some extent on the objective. Next is to find out the optimum working dose range and tolerance to mutagens from pilot experiment. Determination of population size and optimum time of treatment are needed for effective work. Different factors such as the type of mutagen, dose and dose rate, genotype, and growth conditions play a very important role in the outcome of any mutation experiment. Application of dose depends on the radiosensitivity of the material, plant part, and state of development of the experimental plant. Vegetatively propagated plants are not suitable for chemical mutagenesis due to poor uptake and penetration. Vegetatively propagated ornamentals are very suitable for mutation work. All plant parts can be irradiated by one method or another. Seeds, pollen grains, whole plants, cuttings, tubers, corms, bulbs, stolons, tissues, or organs in artificial cultures, etc., can be treated. For any experimental work, a given dose of radiation (dose rate) has a significant effect, both qualitatively and quantitatively. For this reason, dose rate should be carefully chosen and recorded in all experiments. At an early stage, a series of papers have been published on basic aspects of induced mutagenesis. It covers type of mutagen, mutagen efficiency, working dose, sensitivity to mutagen, acute and chronic irradiation, storage effect, split dose, etc. Different parameters suitable for determining the biological effects of mutagens have been mentioned. The main question is what mutagen is to be used—chemicals or ionizing radiations? A wide range of chemicals are available, many of which induce, after seed treatment, a high mutation frequency, and a favorable mutation spectrum. The treatment of plant parts however is not as successful, the main reason being the poor penetration into the material and the deleterious effect of hydrolysis products. Vast literature/ knowledge generated by researchers at an early stage is spread in the form of books, bulletins, catalogs, scientific journals, etc. It is difficult for all researchers, teachers, and students to get an overview of earlier and recent developments covering the whole spectrum of activities on induced mutagenesis. The author tried to put together all available information on mutagens and dose to develop a complete documentation of the results of the research conducted by different scientists over the last 80 years. The primary objective of the write-up is to give a coherent and concise account of earlier work with an emphasis on recent developments. The document has been prepared only from published information

1

Introduction/Review

7

accessible to the author. All important and relevant publications on basic classical mutation have been cited. It is a sincere endeavor of the author to present a complete and comprehensive scenario on mutagens and dose recommended by the earlier worker. This book also provides an authoritative review account of many recent interests and progress in the field of mutation. The number of papers published on mutation, worldwide, is so voluminous that it is difficult to cite all in this book. The author tried to compile the worldwide mutation work alone. The main emphasis has been given to highlight some randomly selected publications of common nature. This will provide all the necessary information on the induced mutation technique. Though meticulous care has been taken in reviewing while dealing with such a voluminous work, some mistakes/errors might have crept in, despite the author’s best efforts. The shortcoming is mainly due to the author’s limitations and nonavailability of literature. The literature survey makes some confusion. It is not clear from some publications whether they have used X-ray or Gamma rays. There is also confusion on unit of radiation doses. Therefore, an attempt has been made in this book to mention the units of different radiations so that uniformity will be maintained by future researchers. Before starting the experiment, one should check thoroughly all available basic information like mutagen, dose, exposure of material, radiosensitivity, biological effectiveness of mutagens, etc. One should go through these early publications for the proper designing of experiments. Differences in relative biological effectiveness (RBE) of X-rays, gamma rays, or neutrons or differences in radiosensitivity of related plants and within polyploidy series are very interesting to radiobiologists. Radiosensitivity of dry dormant seeds is different according to the species or genus, and is also dependent on physiological conditions. The induction of bud mutation in apples by several types of radiation was studied by Granhall (1953). Scions of apples were exposed to X-, gamma, beta rays and neutrons and grafted on rootstocks, and the RBE between these radiations were discussed. The primary biological effects on fruit trees decreased in the order of neutrons>X-rays>32P beta rays and 60Co gamma rays. The gamma ray effect was closely related to that of X-rays in their general biological effect as pointed out by Gray (1954), Spencer and Blackeslee (1954). Matsumura and Fujii (1958a, b) reported that the effect of X-ray was more than that of gamma rays with camphor tree and lemon tree, strawberry, and sweet potato. These facts might be due to the time factor or irradiation, namely gamma irradiation needed a longer time than the X-rays. On the other hand, differences in the primary effect between X- and gamma- rays might be due to the difference in ionization density or quantum energy. Radiosensitivity of plants under the same physiological conditions varies according to the species or variety (Fujii and Matsumura 1958a, b). On the other hand, Smith (1942) reported high radiation susceptibility of a recessive homozygote in X1 generation concerning germination rate and plant height, but he obtained no apparent difference in the mutation rate in the X2 generation. A similar examination was also carried out by Gregory (1956a, b, c) with several closely related lines of peanuts, and variation in sensitivity to X-rays was observed among certain lines. Similar investigations were done by Lamprecht (1958) and Gelin et al. (1958) with pea varieties which were different in 1–3 major genes. Lamprecht pointed out that genotypical constitution

8

1 Introduction/Review

has a great influence on the sensitivity to X-rays in the X1 generation, but Gellin et al. (1959) concluded that variation in this respect is mostly due to the physiological condition of the materials. The difference between gamma rays and X-rays is in the frequency of electromagnetic radiation. Gamma rays and X-rays are both types of electromagnetic radiation, and gamma rays are higher on the frequency spectrum and have more energy than X-rays. There are a few more differences between X-rays and Gamma Rays. The key difference is the source: X-rays are emitted by the electrons outside the nucleus and gamma rays are emitted by the excited nucleus itself. Moreover, gamma rays are highly penetrating and highly energetic ionizing radiation. Gamma rays have shorter wavelengths than X-rays. X-rays have a wavelength ranging from 0.01 to 10 nm and energies in the range of 100–100 keV. X-ray wavelengths are shorter than those of UV rays and typically longer than those of gamma rays. X-rays would have energies in the range of 1–100 keV, while gamma rays would have energies of about 1 MeV and above. Gamma rays have about 1000 times more energy than X-rays (Adlienė and Adlytė 2017; Draganić and Gupta 1973; Mehta and Parker 2011; IAEA 2004; ICRU 1998; Legrand et al. 2012; Podgorsak et al. 2005). Different units which are used in the radiation studies are Roentgen (R): The roentgen is a unit used to measure a quantity called exposure. Rad (radiation absorbed dose): The unit rad is used to measure the quantity of absorbed dose. This relates to the amount of energy actually absorbed in some material and is used for any type of radiation and any material. One rad is defined as the absorption of 100 ergs per gram of material. Gray (Gy): The gray is also a unit like Rad used to measure a quantity called absorbed dose. One gray is equal to one joule of energy deposited in one kg of material. Absorbed dose is often expressed in terms of hundredths of a gray, or centigrays. One gray is equivalent to 100 rads. Absorbed dose is the amount of energy that ionizing radiation imparts to a given mass of matter. The dose rate is a measure of how fast a radiation dose is being received. Different materials that receive the same exposure may not absorb the same amount of energy. If we look at the literature, the source of radiations and their units are not uniform. A good amount of literature mentions the use of source of radiation treatment in the same paper gamma rays or X-rays. Similarly, doses are mentioned as rad, krad, Krad, Gy, R, kR, Kr, etc. for both Gamma rays and X-rays. Scientifically it is correct. But it creates confusion. To avoid such confusion and considering abovementioned energy level of X- and Gamma rays, it is proposed to follow a uniform pattern of unit. The unit rad may be used for any type of radiation, but it does not describe the biological effects of the different radiations. The rad is a deprecated unit of absorbed radiation dose, defined as 1 rad = 0.01 Gy = 0.01 J/kg. The Rontgen or roentgen (R) has mainly been used for the calibration of X-ray machines. The Rontgen X-radiation is referred to with terms meaning Röntgen radiation, after Wilhelm Röntgen, who is usually credited as its discoverer. rad is short for radiation while roentgen is the unit for X-ray radiation. It is wise to use rad for gamma rays and R for X-rays. Following the International System (SI), this basic radiation unit for absorbed dose is defined as the Gray (Gy). The Gray is equivalent

1.1

Mutagenesis and Radiobiological Aspects

9

to the absorption of 1 Joule (J) of energy per kilogram (kg) of irradiated material. The SI unit for absorbed dose is the gray (Gy), but the “rad” (Radiation Absorbed Dose) is commonly used. 1 rad is equivalent to 0.01 Gy. For X-rays the unit may be roentgen “R” (1 roentgen = R), in plural kilo roentgens kR, a unit equal to 1000 roentgens [1 kR = 1000 roentgen]. The gamma radiation unit may be “rad,” 1000 rad = 1 Krad (1 kilorad). The rad may be replaced by the gray (Gy): 100 rads = 1 Gray; 1 kilorad (krad) = 10 gray (Gy). In radiation biology for any biological effect, a specific amount of radiation is responsible. The dose of radiation is most important for any biological effect. The absorbed dose for any ionizing radiation is defined as the amount of energy absorbed per mass of irradiated matter at the point of interest (ICRU 1964, 1971a, b, 1993, 1998; Adlienė and Adlytė 2017; Boudou et al. 2004; Brunner 1995a; Fricke and Hart 1966; Haninger and Henniger 2016; Harrison 2013; Klassen et al. 1999; Mehta and Parker 2011; Podgorsak et al. 2005; Schreiner 2004; Tang et al. 1984). The special unit of absorbed dose is the rad, where 1 rad = 100 erg/g = 10-2 joule/kg. The dose rate is usually expressed as rad/h, rad/min, and rad/s. In many cases, absorbed dose is calculated from the measured number of ions produced in air by the ionizing radiations (X-ray, gamma radiation) which is called “exposure.” The special unit of the exposure is the roentgen, where 1 R = 2.85 × 10-4 coulomb/kg air. The usual unit of exposure rate are R/h, R/min, or R/s. The effects of different dose rates have seldom been studied although it appears that especially the rate of survival and the amount of chimerism may be affected by dose rate (McCrory and Grun 1969; van Harten et al. 1972).

1.1

Mutagenesis and Radiobiological Aspects

For any radiobiological work, Cobalt-60 and Caesium-137 are used as the main sources of gamma rays (Cowan and Meinhold 1962; D’Amato et al. 1962a, b, 1964a, b; Singleton et al. 1961). The half-life of 60Co is 5.3 years and the same for 137Cs is 30 years. For routine classical mutagenesis work, the first approach is what is the best mutagen, what is the appropriate dose, what is the best object of mutagen application (e.g., Seeds, pollen, whole plant), what is the best treatment conditions (e.g., acute vs. chronic irradiation, concentration vs. duration, dry vs. presoaked seeds), etc. Determination of radiosensitivity in plants is very important in radiobiological studies and in plant breeding by radiation. It has been demonstrated that many biological, physical, and physiological factors can alter the radiosensitivity of higher plants (Ahnstroem 1989; Albokari et al. 2012; Altenbung 1934; Attix 1966; Brunner 1992, 1995a, b; Stadler 1929; Lea 1947a, b; Basu and Datta 1982; Bishop 1952; Bostract and Sparrow 1970; Bowen 1962; Brock 1965a, b, 1971, 1977, 1979; Brock and Micke 1979; Bacq and Alaxander 1955; Broertjes 1977a, b, c, d; Caldecott et al. 1954, 1957; Caldecott 1955, 1961; Carlson and Smith 1977; Chopra 2005; Conger and Johnston 1956; Croteau and Bohr 2013; Datta 1984, 1992b, 2001; Davis 1962a, b; Doring and Stubbe 1938; Eguchi-Kasai and Cox 1995; El-Metainy

10

1 Introduction/Review

et al. 1973; Freisleben and Lain 1942a, Freisleben and Lain 1942b, Freisleben and Lain 1943a, 1943b; Froier and Gustafsson 1944a, b; Froier et al. 1941, 1942; Fujii 1960a, b; Fujii and Matsumura 1958a, b, 1959; Gill et al. 2015; Gray 1954; Gustafsson 1940, 1944, 1947a, b; Johns 1961; Joshi and Sharma 1975; Iqbal et al. 1991; Kimball 1987; Kondo and Matsumura 1958; Konzak and Singleton 1952; Maliga et al. 1981; Robbelen 1982; Mandal and Basu 1977, 1978, 1980; Matsumura 1960; Matsumura and Fujii 1955a, b, 1958a, b, 1959a, b; MacKey 1954a, b; Micke 1984a, b; Miksche and Rudolph 1968a, b; Mericle and Mericle 1957; Mee et al. 1969; Metainy et al. 1971; Muller 1954a, b; Nishiyama et al. 1959; Notani 1961; Novak and Micke 1988; Ostergren et al. 1958; Robbelen 1959; Sax 1940, 1942, 1955, 1963; Singleton et al. 1956; Smith 1943, 1946, 1950; Sparrow 1954; Sparrow et al. 1961a, b; Sparrow and Evans 1961; Sparrow and Forro 1953; Sparrow and Gunckel 1955; Sparrow and Gunkel 1956; Sparrow and Konzak 1958; Sparrow and Miksche 1960; Sparrow et al. 1952a, b, 1965a, b, 1967, 1968; Sparrow an Schwemmer 1974; Sparrow and Sparrow 1965; Sparrow and Schairer 1980; Steffensen 1957, 1958; Sherman 1952; Nirula 1963; Nishiyama and Amano 1963; Nishiyama and Ichikawa 1961; Woodwell and Rabuck 1967; Nilan 1956; Sybenga 1964; Stoilov et al. 1966; Smith 1942; Lawrence 1963; Ohba and Sinaka 1961; Wu et al. 2012a, b; Yamaguchi 1956, 1969a, b, 1973, 1979; Ukai 1967, 1970; Underbrink et al. 1968; Yonezawa and Yamagata 1977). The aspect of radiosensitivity has been mentioned in a separate chapter. Treatment with physical and chemical mutagens can induce cytological, morphological, physiological, and genetical changes in cells and tissues (Beal and Scully 1950; Bowen and Cawse 1962; Chadwick and Leenbouts 1981; Davis and Wall 1961; Etter 1965; Evans 1962, 1966; Ford 1948; Gunckel 1957; Kihlman 1966). These in turn can result in modifications in the growth and development of roots, stems, leaves, and flowers. Manifestation of aberrant plant growth is a very striking effect of treatment of propagules with mutagens. Development of different types of chromosomal aberrations is also noticed during root tip mitosis. Abnormal plant growth is an immediate effect after mutagen treatment. Early studies and their results have enriched the basic knowledge on the subject of radiosensitivity. This subject has been further enriched step by step through advanced studies. Reduction in plant height after exposure to ionizing radiations has earlier been reported by a number of workers (MacKey 1956; Gunckel and Sparrow 1954; Yamaguchi 1958a, b; Basu 1960, 1962a, b; Datta and Basu 1977). Quastler and Baser (1950), Gnanamurthy et al. (2012a, b), and Russel and Martin (1952) are of opinion that physiological effects are responsible for radiation damage as measured by growth reduction. Skoog (1935) and Gordon and Weber (1950, 1953) have suggested that destruction and inactivation of auxin by X-rays bring about radiation damage. Gordon (1957) has demonstrated inactivation of auxin and decrease in auxin content with increase in radiation doses. Inactivation of IAA (Mika 1952), inactivation of rate of assimilation, altered nutritional level (Went and Theimen 1937; Gunkel and Theimann 1949; Ehrenberg 1955), inactivation of enzyme system (Barron 1952), and changes in oxidation and reduction system (Frossberg and Nybom 1953a, b; Nybom 1953) after treatment with ionizing radiations are probably responsible for reduction in

1.1

Mutagenesis and Radiobiological Aspects

11

seedling height. Sax (1942) and Lea (1947a, b) have shown that the survival of plants to maturity and reduction in seedling height depends on the nature and extent of chromosomal damage. Increasing the frequency of chromosomal damage with increase in dose may be responsible for less germinability and reduction in plant growth. Sparrow and Gunckel (1955) have suggested that chromosomal damage associated with non-chromosomal damage plays an important role in growth reduction. Radiation-induced abnormal plant growth has been reported by a number of workers and this aspect has been reviewed earlier in detail by Gunkel and Sparrow (1954). Growth rate analysis showed stimulation of growth in some cultivars at initial stage. Stimulation of growth after treatment with different ionizing radiations has been reported in different crops (Sax 1942, 1963; Kersten et al. 1943; Shull and Mitchell 1933; Sparrow 1954; Charles and John 2016). The author and his colleagues have made a series of experiments to explore the correlation among mutagens, morphological and cytological aberrations, and its biochemical status. Datta and Basu (1977) and Datta (1988b) treated seeds of four members of cucurbitaceous (Cucurbita maxima L., Cephalandra indica Naud, Trichisanthes cucumarina L., and Trichosanthes anguina L.) with X-rays and colchicines. All the materials showed reduction in plant height and different forms of morphological abnormalities irrespective of X-ray or colchicine treatment. Chromosomal breakage as one of the factors for the development of abnormal morphology of plants has been suggested earlier (Thoday 1951; Sparrow et al. 1952a, b). Materials treated with X-rays and colchicine resulted in the formation of chromosomal aberrations and polyploidy cells, respectively and the plants developed from the same treatment showed aberrant plant growth (Datta and Basu 1977; Datta 1988b). Different forms of morphological abnormalities were common to both X1 and C1 generations. Differential sensitivity of the materials to X-rays and colchicine was recorded. If chromosomal aberrations due to X-irradiation are responsible for morphological abnormalities and reduction in plant height, there must have been some correlation between chromosomal aberrations and morphological changes. The material with maximum induced chromosomal aberrations should show maximum morphological abnormalities and reduction in plant height. The material which was most sensitive to X-rays and colchicine with respect to the formation of chromosome aberrations and polyploidy did not show either maximum morphological abnormalities or reduction in growth. Therefore, radiation and colchicine-induced cytological effects cannot be considered as the factors which directly lead to the development of abnormal plant parts and reduction in seedling height. It has been presumed that either any change in the chromosome level (aberrations or polyploidy) disturbs the pathway of some chemical reactions or mutagens directly hit on these pathways which in turn lead to abnormal plant growth. It is possible that both the pathways (one for morphology of leaves and another for growth of plant) are differentially sensitive to X-rays and colchicine and the degree of sensitivity varies from material to material. Different plant materials are differentially sensitive to external physical and/or chemical stress. Not only different genus and species but also different cultivars, different parts of the same plant, and even different tissues are sensitive differentially (Mandal and Basu 1980). This sensitivity is not only applicable to plants but also to all other biological

12

1 Introduction/Review

samples. Datta (1984) has reviewed the subject radiosensitivity of cells. Datta (1988c) reported external stress not only changes the chromosomal behavior but also disturbs the pathway of some chemical reactions related to abnormal plant growth and defense mechanism of cells. Datta (1992b) reported that radiosensitivity is a genotype-dependent mechanism in the damage or repair/protection of stressinduced damage within biological materials. Dry and water-soaked germinated seeds of Lens culinaris were treated with 5, 10, and 20 Krad gamma rays. Chromotoxic effects and enzyme-related defense mechanisms have been studied (Datta et al. 2011; Hossain et al. 2003a, b). Gamma ray stressed condition was found to have a lethal effect on cell division and induced chromosomal abnormalities during root tip mitosis in both the seed materials (dry and wet). The severity of chromotoxic effects increased with increased stress due to higher doses. Both the enzymes (peroxidase and superoxide dismutase) were increased several fold over the control in germinated seeds when treated with gamma rays. After gamma treatment, the level of free radical formation was high in germinating seeds in comparison to dry seeds. These free radicals interacted with each other to produce organic peroxide which were highly reactive and believed to be responsible for a good portion of biological and genetic effects of radiation (Singh 1996). Dry seeds were metabolically inactive and thus antioxidant enzymes failed to protect cells from free radical damage. As the half-life of free radicals is a few micro seconds, e.g., 2–4 μs for 1O2 (singlet oxygen) in water (Foyer and Harbinson 1994), they exerted their effect on chromosomes immediately. When gamma ray treatments were applied to germinated seeds, antioxidant enzymes remained in the active stage. To scavenge the excess free radicals they exhibited greater activity in comparison to dry seeds with an increase in dose. Different pathways are differentially sensitive and the severity of sensitivity varies from material to material (Datta 1988c). Different enzyme-mediated defense systems against ROS are differentially sensitive to stress (gamma rays) or any change at the chromosomal level may stimulate their activities (Hossain et al. 2003a, b). Sensitivity of Allium cepa to gamma rays has been determined on the basis of cytological and enzymatic changes. Datta et al. (2011) treated bulbs of Allium cepa with different doses of gamma rays (2, 4, and 6 Krad or in another form 0, 20, 40, and 60 Gray) and studied the chromosomal behavior and antioxidant enzymes (SOD, APX, G-POD, GR, and catalase (CAT)) status on third and 30th day respectively to understand the level of radiosensitivity. The role of oxidative stress due to gamma ray treatment in the form of chromosomal aberrations and the protective enzymatic systems in relation to root tip mitosis were studied. Abnormal chromosomal behavior and enzyme-related defense mechanism studies due to gamma ray-induced stress condition is perhaps the first report in germinating roots of the bulbous plant. A positive correlation between chromosomal abnormalities and antioxidant enzymes related defense mechanism of cell has been established. Any plant propagule (seed, bulb, cuttings/suckers, etc.), when exposed to any external physical and/or chemical agents, develops stress on physiological, biochemical, and cytological activity at the cellular level. These in turn can result in the modification chromosomal behavior, changes in growth, and development of roots, stems, leaves, and flowers (Frossberg and Nybom 1953a, b; Gunkel 1962; Gunckel and Sparrow 1954, 1961; Gordon

1.1

Mutagenesis and Radiobiological Aspects

13

1957; Datta and Basu 1977; Datta 1988c; Hossain et al. 2003a, b). Allium cepa L. as a test system was introduced in 1938 by Levan to investigate the effects of several external agents on living cells. Allium test is a simple but very efficient method for rapid and extensive evaluation of toxicity and teratogenicity of such agents (Bellani et al. 1991). The results obtained by the Allium test were found similar to those obtained with different chemicals tested in other organisms (Fiskesjo 1998). Allium test has been successfully applied to study the cytotoxic/genotoxic effects of chemicals and radiations and the results have been reported from time to time (Pandey et al. 1994; Adam et al. 1990; Ahmad and Yasmin 1962; Bellani et al. 1991; Kumar and Sinha 1991; Datta et al. 1998a, b; Shukla and Datta 1999, 2000, 2002; Liang 1983; Hossain et al. 2002; Yildiz and Evrim 2008; Majewska et al. 2008; Bulbul et al. 2008). Clastogenic effects and enzyme-related defense mechanisms have been studied on dry and water-soaked germinated seeds of Lens culinaris after treatment with different doses of gamma rays (Hossain et al. 2003a, b). Gamma ray-induced chromosomal aberrations were very high on third day which sharply decreased after 1 month. Deleterious effects of radiation in form of abnormal chromosomal behavior are found up to a certain period of growth, after which the activities are mostly normal. Datta and Basu (1977) reported that the percentage of plants with X-ray-induced morphological abnormalities decreased with aging. Diplontic selection gradually eliminated the abnormal cells and plants appeared to be slowly returning to near normalcy. With an increase in the dose of gamma rays, there was a decrease in mitotic index and an increase in chromosomal aberrations, SOD-, APX-, GR-, and Guaiacol peroxidase activity and MDA content. SOD-, APX-, GR-, and Guaiacol peroxidase activity increased to its maximum after gamma irradiation on third day and thereafter decreased. Increased positive significant changes of antioxidant enzymes with an increase in gamma ray doses were prerequisite to protect living cells from the stress effect of gamma rays. Antioxidant enzyme content was almost the same or slightly increased in control and gamma ray-treated bulbs on the 30th day. Increased chromosomal aberrations during initial root tip growth (third day) in A. cepa is due to gamma ray-induced stress. Since antioxidant metabolism has been shown to be important in determining the ability of the plant to survive in oxidative stress, an upregulation of these enzymes would help to reduce the build up of ROS (Levitt 1980; Bowler et al. 1992, 1994). An increase in the activities of SOD and GR in roots of A. cepa indicated enhanced production of free radicals under gamma ray treatment stress conditions. The level of free radical formation was high immediately after gamma irradiation which is reflected through a higher percentage of chromosomal abnormalities. Radiation-induced free radicals produce highly reactive organic peroxide which is believed to be responsible for biological and genetic damage (Singh 1996). Half-life of free radicals is a few microseconds in water (Foyer and Harbinson 1994) which affects chromosomal behavior immediately, and therefore maximum chromosomal abnormalities were recorded at the initial stage. Antioxidant enzymes remain in an active stage in developing root and help to scavenge the excess free radicals in dividing cells. During the root growth period, the MDA content in developing roots decreased as compared to initial gamma irradiation, indicating a decrease in oxidative stress.

14

1 Introduction/Review

MDA, a decomposition product of polyunsaturated fatty acids hydroperoxides, has been utilized very often as a suitable biomarker for lipid peroxidation (Bailley et al. 1996), which is an effect of oxidative stress/damage. Living cells under stress condition develop a protective mechanism to scavenge the same stress-induced free radicals through the formation of antioxidant enzymes (Larson 1988; Noctor and Foyer 1998; Halliwell and Gutteridge 1989). An increase in SOD-, APX-, GR- and Guaiacol peroxidase activity and MDA content in the root of A. cepa indicate enhanced production of free radicals under gamma ray-induced stress condition. Sensitivity of A. cepa to gamma rays has been determined on the basis of cytological and enzymatic changes. Datta (1992d) reported earlier that radiosensitivity is a genotype-dependent mechanism in the damage or repair/protection of stress-induced damage within biological materials. Gamma radiation effects on all these parameters can be utilized as test protocols for routine testing of radiosensitivity of any plant. It will enhance our present understanding of the mechanism of radiation effects on plants and will promote interdisciplinary research. Different pathways related to antioxidant enzymes are differentially sensitive to gamma ray doses. Since antioxidant metabolism has been shown to be important in determining the ability of plants to survive in oxidative stress, an upregulation of these enzymes would help to reduce the build up of ROS. This factor could be a key to design adequate methods to improve radiosensitivity in the crop improvement program.

1.2

Acute and Chronic Irradiations

Radiation doses are grouped into two categories: acute and chronic dose. Exposures that are continued over long periods of time (usually weeks, months, or years) are referred as chronic (low dose rate). Exposures delivered in minutes or a few hours are referred to as acute (high dose rate). Gamma sources can be used both in acute and prolonged (chronic) form. Experimental results indicate that plants are differentially sensitive to acute (gamma chamber—gamma room) and chronic (gamma field) radiation separately and in combination with both methods (Asghar and Khan 1988; Bari 1971; Davis and Wall 1960a, b, 1961; Donini 1967; Eriksson et al. 1966; Kovalchuk et al. 2007; Miksche et al. 1962; Moghaddam et al. 2011; Sparrow and Christensen 1953; Nishida et al. 1967; Sparrow and Singleton 1953; Sparrow et al. 1961a, b; Taheri et al. 2014; Tangpong et al. 2009). Chronic irradiation results in maximum mutation frequency and spectrum in chrysanthemum over acute irradiation. A combination of the two yielded ten times more mutation rate and non-chimerical mutants (Nagatomi and Degi 2009). The optimum dose of chronic irradiation has been determined to be almost 2.5 times more than acute irradiation for chrysanthemum cuttings, petal, and/or bud culture. This technique has been successfully applied and developed a number of useful mutant varieties in ornamentals and other crops (Bauer 1957; Brock and Franklin 1966; Broertjes 1971a, b; Contin et al. 1963; Conger et al. 1973a, b; Cuany et al. 1958a, b; Stadler 1931; Donini and Scarascia-Mugnozza 1968; Fujii 1960a, b; Fujii and Matsumura 1967; Kang et al.

1.2

Acute and Chronic Irradiations

15

2010; Kovalchuk et al. 2000; Natarajan and Maric 1961; Matsumura 1964; Mabuchi and Matsumura 1964; Mergen and Thielges 1966; Miksche et al. 1962; Nagatomi 1991a, b, 1992, 2002; Nagatomi et al. 1993a, b, 1996a, b, c, d; de Nettancourt 1969; Nayar 1969; Nybom et al. 1956; Richter and Singleton 1995; Singleton 1965; Sparrow and Konzak 1958; Sparrow et al. 1961a, b; Sparrow 1965; Ukai and Yamashita 1969; Yamashita 1964, 1967). The decision to be made by the investigator concerning the most appropriate plant part or stage to expose requires a thorough knowledge of the organism and clear objectives of the experiment. The procedures and techniques for irradiating the whole plant in a gamma field or a gamma room and treatment of seedling or small plants in X-ray machines or gamma source in a greenhouse have been reported (Sparrow (1966a, b). Dose level vary with many factors like—dry and weight seed, seed size, seed coat, seed moisture, postirradiation storage, etc. (Froese-Gertzen 1962; Froese-Gertzen et al. 1963, 1964; Briggs 1966; Konzak et al. 1972; Gaul et al. 1972). Duration of the treatment period should be calculated so that there is a thorough infusion of mutagen to the target tissue. Presoaked seeds shorten the duration of treatment. Presoaking, short treatment with high concentrations at relatively high temperature, long duration treatment, buffering, and periodical renewal of fresh mutagenic solution for the highest mutagenic effectiveness have been studied (Heslot et al. 1961a, b; Konzak et al. 1965; Ehrenberg et al. 1966; Brunner and Mikaelsen 1971). The temperature of a mutagenic solution has a great influence on mutagenic chemicals. Knowledge of hydrolysis half-lives of the chemical mutagens at various temperatures is very important. Hydrolysis half-lives, temperatures, duration of treatment, and rate of reaction efficiency of chemicals have been worked out (Mikaelsen et al. 1968; Brunner and Mikaelsen 1971). They have reported hydrolysis half-lives of dES and EMS at 0, 5, 10, 20, 30, and 40 °C are 59.2, 27.6, 13.1, 3.34, 1.00, and 0.32 h, and 1716, 796, 378, 93.1, 25.9, and 7.92 h, respectively. A number of other modifying factors like presoaking, pH of the mutagenic solution, metallic ions, carrier agents, etc., can influence the effect of chemical mutagens before, during, or after the treatment (Savin et al. 1968; Mikaelsen et al. 1968; Brunner et al. 1968; Brunner and Mikaelsen 1971; Ramanna and Natarajan 1965; Mikaelsen 1956, 1967, 1968a, b, 1969; Wagner et al. 1968; McCalla et al. 1968; Ehrenberg and Gichner 1967; Kamra and Brunner 1970, 1977; Kamra et al. 1960; MoutschenDahmen and Degraeve 1965; Moutschen-Dahmen and Moutschen-Dahmen 1963; Moutschen-Dahmen et al. 1959; Swiecicki 1987; Bhatia and Narayanan 1965; Patrick and Haynes 1964; Bhatia 1967; Siddiq et al. 1968a, b; Kaul 1969; Bender and Gaul 1966, 1967). Treatment procedures are very important in chemical mutagens. Seeds and buds may be treated either in a dormant state or in the actively metabolizing and synthesizing stage. These may be dipped or soaked in a mutagenic solution of the appropriate concentration. The dose of chemical mutagen is decided on the basis of concentration, duration of treatment, and temperature during treatment. For chemical mutagen, volume of the mutagen is very important. Volume should be sufficiently large so that each material gets the opportunity to absorb the same number of moles of mutagen. The role of high concentration and low concentration treatments for

16

1 Introduction/Review

long periods on physiological and genetical characters have been worked out (Konzak et al. 1965; Mikaelsen et al. 1968; Savin et al. 1968; Brunner and Mikaelsen 1971). There are early reports regarding the recommended concentrations for various mutagens like EMS (0.05–0.3 M or 0.3–1.5%, Mikaelsen et al. 1968, Konzak et al. 1965, Savin et al. 1968); dES (0.015 M–0.02 M or 0.1–0.6%, Mikaelsen et al. 1968, Konzak et al. 1965); EI (0.85–9.00 mM or 0.05–0.15%, Wagner et al. 1968); ENH and MNH (1.2–14.0 mM or 0.01–0.03%) and NaN3 (0.001–0.004 M, Gichner et al. 1968, Savin et al. 1968). Pollen grains were treated with UV radiation, X-rays, and chemical mutagens and their comparative sensitivity have been studied (Emmerling 1955; Neuffer 1957; Brewbaker and Emery 1962; Brewbaker et al. 1965a, b; Ikenga and Mbuchi 1966; Kaplan 1949; Loh and Cooper 1966a, b; Brock and Franklin 1966; Rudolph 1965a, b; Stadler 1928a, b; Bianchi and Contin 1963; Briggs and Smith 1965; Sparrow and Konzak 1958; Li et al. 1989; Taner et al. 2000, 2003; Yanmaz et al. 1999). Ultra-violet radiation has limited use for treating spores or pollen grains. The biological effectiveness of light varies with the wavelength. Wavelength in the range 2500–2900 nm are biologically most effective (Altenbung 1934; Brewbaker et al. 1965a, b; Ito 1964; Castronuovo et al. 2014; Emmerling 1955; Jardim et al. 2015; Neuffer 1957; Kovacs and Keresztes 2002; Kirby-Smith 1963; Liu et al. 2015; Lois 1994; Mba 2013; Mba et al. 2012; Johns et al. 1965; Cline and Salisbury 1966; Ikenga and Mbuchi 1966). Pollen grains are irradiated at a distance of 5.5 cm from the lower surface of the tube at a dose rate of 3,112,400 erg/cm2 min. The penetrating ability of beta particles is lower than that of X- or gamma rays. Experiments were mostly conducted with 32P, 35S, 131I, 90Sr, 90Y, and 14C (Amaldi 2000; Beal and Scully 1950; Faegri and Deure 1960; Kawai 1954, 1962, 1963a, b; Michaelis and Kaplan 1953; Mackie et al. 1952; Hungate and Marinelli 1952; L’Annunziata 2016; McQuade and Friedkin 1960; Kharkwal et al. 2004; Matsumura 1962; Mba et al. 2012; McKelvie 1963; Sparrow and Konzak 1958; Stein and Sparrow 1966; Bell 1970; van Harten 1998). The dosages ranged from 2.5 to 40 μCi per seed. 32P has a half-life of 14.5 day and 35S, 87 day. Neutron energy is obtained from a reactor due to 235U fuel nuclear fission. The energy spectrum of the majority of neutrons between 0.5 and 2.0 MeV are termed fast neutrons. By reaction with moderators, such as carbon and hydrogen, fast neutrons are reduced in energy to about 0.025 eV and are then called thermal neutrons. Series of biological experiments have been conducted using various neutron sources (An and Wang 1985; Smith 1961a, b; Burtscher and Casta 1967; Casta 1968a, b; Johnson and Poston 1967; Goodman 1972; Sparrow et al. 1972; Underbrink et al. 1971; Smith et al. 1964; Harle 1965; Rossi 1966; Glubrecht and Hamann 1968). Neutrons at low fluxes obtained from radioisotopic sources such as Californium-252, 210P-Be, 238P-Be, or 241Am-Be have been used in mutation induction (c.f. Matsumura 1966, Reinig 1968, Karelin et al. 1997). Fast and thermal neutron irradiation has been applied in mutation work (Byrne 2013; Solomko 1965a, b; van Harten et al. 1972; Gomez Cuervo and Nelson Estrada 1972; Miedema 1973a, b). Doses and dose rates used vary greatly and are dependent on the reactor use. High-energy protons, used by Tarasenko (1977a, b),

1.2

Acute and Chronic Irradiations

17

were found to be as effective as fast neutrons. When comparing the two irradiation types it can be summarized as follows: the injurious action of neutrons differs from that of X-ray in several respects. The seeds are 20–30 times more sensitive to neutrons than to X-rays and germinating seeds are two to three times more sensitive to neutrons than dormant seeds. Neutrons are approximately 10 times as effective as X-rays in producing chromosome disturbances and about 50–100 times more effective in increasing the mutation rate in the second generation. Neutrons produce relatively more chlorophyll mutations than X-rays. Many types of accelerators have been constructed, e.g., the Cockroft-Walton, Van de Graaff, betatron, cyclotron, synchrocyclotron, synchrotron, and linear accelerator (Amaldi 2000). In general practice, they are used to accelerate protons, deuterons, and electrons. Few reports are available on the radiobiological effect of such high-energy particles. Caldecott (1955) gave information on experiments in which he used 2-MeV electrons obtained with a Van de Graaff electrostatic generator. Smith et al. (1965) have used the Alternating Gradient Synchrotron (AGS) at Brookhaven National Laboratory, a proton synchrotron operating at energies up to 33 GeV. Micke et al. (1964a, b) used the AGS to study the genetic effects of almost pure pi mesons and muons. No particular advantage of using particulate radiation from accelerators for the induction of mutations has yet been found. Treatment conditions play a very important role in mutation induction. The first is dose rate of radiation which should be very carefully chosen and recorded in all experiments. An important question is the selection of mutagens and optimum dose—radiation (type, dose, and dose rate) or chemicals (type, concentration, length of treatment, pretreatment, post washing). The best practice is the empirical one: the starting material should first be exposed to a range of doses, centered around the optimum one, either that found in the literature or a calculated optimum (Sparrow et al. 1967). Mutation frequency increases with increasing dose (linearly with X- and gamma rays, more exponentially with neutrons), but survival and capacity to regenerate decrease with increasing dose. One must therefore choose some point between a low dose (100% survival, low mutation frequency) and a high dose (low survival, high mutation frequency). At high dose, too many mutational events per cell may be induced, with an increased risk of a favorable mutation being accompanied by one or more unfavorable genetic changes. Somatic mutation frequency has been reported with high-intensity and low-intensity radiation exposure (Sax 1939; Sparrow et al. 1961a, b; Nishiyama et al. 1962a, b, 1964, 1966; Ichikawa and Nishiyama 1963; Matsumura 1964; Bianchi and Giaccetta 1964; Smith and Rossi 1966; Yanesawa and Yamagata 1977). The dose to be used obviously depends upon the radiosensitivity of the material involved. However, even when one knows the dose-response curve, it is difficult to decide which dose is to be preferred—a light one which only slightly interferes with the subsequent development of the material but induces a low mutation frequency, a heavy dose which retards normal development but induces a high(er) mutation frequency. It is not wise to recommend simply applying a dose that is mentioned in the literature. One pilot experiment should be carried out by exposing the starting material to a range of doses. To determine the LD50 or optimum working dose a number of parameters may be selected like Germination/sprouting

18

1 Introduction/Review

percentage, 15-day seedling/sprout height, morphological abnormalities at the age of 30 days, chromosomal aberrations during root/shoot tip mitosis, meiotic abnormalities, pollen grain sterility/fertility, survival, mutation frequency, etc. (Datta 1997). To cite an example, the suitable radiation dose for the induction of somatic mutations in chrysanthemum has been reported by many workers. Datta and his group detected LD50 of chrysanthemum cultivars varies between 1.5 and 2.5 krad. Previous workers have reported that some of the cultivars withstood 3000r X-rays and the optimum dose lay between 2000 and 4000r (Jank 1957a, b; Sheenan and Sagawa 1959). Fujii and Mabuchi (1961) found that 2–4 Krad gamma rays showed the optimum number of survival while Bowen et al. (1962) found only 50% lethality after 4.3 Krad. Dowrick and El-Bayoumi (1966a, b) reported that 14 Krad of gamma rays was the suitable dose. Some authors however used higher doses like 25 Krad gamma rays (Cawse 1965, 1966), 10–12 kR gamma rays (Yamakawa and Sekiguchi 1968), and 8 Krad gamma rays (Broertjes 1966a, b). The use of such higher doses was probably the result of low dose rate application (1 kr/days’ 125–150 rad/ha). On the basis of the earlier and present experiments the optimum dose of gamma rays for inducing mutations is reported to be 1.5–2.5 Krad for chrysanthemum. As such a wide range of doses have been reported for a single species by many workers, it is not wise to start the mutation experiment on the basis of available literature. It is wise to do a pilot experiment to determine the optimum dose (Datta 2001). For X-rays and gamma rays, doses range from 4 to 100 Gy for treatment of vegetative parts in vivo, 20–30 Gy being the doses preferred by most workers. Doses of 80–100 Gy are usually lethal. One must choose some point between a low dose (100% survival, low mutation frequency) and a high dose (low survival, high mutation frequency). Induced mutagenesis has played a major role in the development of superior mutant plant varieties all over the world. The global impact of mutation-derived varieties in major crops has been reviewed. Research papers, review papers, and books convincingly documented the contribution of induced mutations to the increase of agricultural production (Anonymous 1973, 1977a, b, 1979, 1982a, b; Ahloowalia 1997, 1998; Ahloowalia et al. 2004; Ali 2008; Ahmad et al. 2010; Akerberg and Hagberg 1963; Al-Qurainy and Khan 2009; Alexander and Stacey 1958; Alexander et al. 1961; Al-Safadi et al. 2002; Al-Enezi and Al-Khoyri 2012; Amano and Yamaguchi 2001; Atienzar and Jha 2006; Aurigue 2018; Avemanco et al. 1989; Azad et al. 2018a, b; Bado et al. 2015, 2016a, b, c; Bajpay and Dwivedi 2016; Bakri et al. 2005; Banerji 2009, 2014; Batista et al. 2008; Bhatia et al. 1991a, b, 2001; Blixt 1972; Bokhari Khera 2015; Bradshaw 2016; Brenner et al. 1961; Bressan et al. 1981; Brock 1976, 1977; Brock and Micke 1979; Brock and Rochford 1963; Broertjes 1965; Broertjes 1968a; Broertjes 1968b, c, d, e, f, g, 1969a, b, c, d, e, 1972a, b, c, d; Broertjes and Alkema 1970, 1971; Broertjes and van Harten 1978; Brown 2013; Buiatti 1989; Bull et al. 2007; Burnner and Mikaelsen 1971; Cagirgan 2018; Casarett 1968; Cassels 1998; Chadwick and Leenbouts 1981; Chadwick and Leenhouts 2005; Chaudhary and Chaudhary 2014; Chikelu 2013, Chopra and Sharma 1985; Cooke 1953; Coretchi et al. 2018; Cuevas et al. 2009, 2015; Dale 1966; D’Amato and Gustafsson 1948; D’Amato et al. 1964a, b; D’Souza 2008; D’Souza et al. 2009; Dahiya et al. 2013; Danso et al. 2008;

1.2

Acute and Chronic Irradiations

19

Das et al. 1974, 1977a, b; Datta 1990c, 2002, 2015a, b, 2018a, b; Datta and Datta 2018; Datta et al. 1993; Davis 1971; Decourtye 1969, 1978, 1979; Dellaert 1979; Danso et al. 1990; De Loose 1964a, b; De Micco et al. 2010, 2011; Devreux et al. 1986; Diamantis 1974; Do et al. 2009; Díaz-López et al. 2017; Dighton et al. 2008; Dommergues 1961, 1962a, b, c; Dommergues et al. 1966a, b; Dong et al. 1985, 2016a, b; Donini 1974, 1982; Donini and Micke 1984; Donini and Rossi 1979; Donini and Sonnino 1988, 1998; Draganić and Gupta 1973; Drake 1991; Drake et al. 1998; Dubinina and Dubinin 1968; Dulieu 1970; Ehrenberg and Granhall 1952; Elazazi et al. 2018; Enken 1967; Esnault et al. 2010; FAO/IAEA 1961, 1965, 1973, 1988; Favret 1959, 1960a, b, 1962, 1965, 1971; Favret and Ryan 1957; Ferrary 1965; Filali et al. 1997; Fisher and Lapins 1966; Forster and Shu 2012; Freese 1961; Freidman 1985; Freisleben and Lain 1942; Friedt 2003; Froese-Gertzen et al. 1963; Froier and Gustafsson 1944a, b; Fujii 1981; Gady et al. 2009; Jagadeesan and Ramabhatta 2021; Gago et al. 2009; Gaul 1962; Geras’kin et al. 2018; Ghanim et al. 2018a, b; Granhall 1954; Geier 2012; Geraskin et al. 2013; Germana 2012; González-Jiménez 2004; Gressel and Levy 2006; Jankowicz-Cieslak et al. 2017; Jankowicz-Cieslak and Till 2015, 2016, 2017; Goldstein et al. 2015; Gottschalk 1986; Gottschalk and Wolff 2012; Gomez-Pando et al. 2009; Gonzalez 2004; Gonzalez et al. 2008a, b; Grant 1994; Gustafsson 1940, Gustafsson et al. 1967; Gvozdenovic et al. 2009a, b; Granhall, et al. 1949; Gregory 1961, 1965; Gustafsson and Gadd 1962, 1965a, b, 1966; Grant and Salamone 1994; Gruszka et al. 2012; Gupta 1998a, b; Hadzim et al. 1994; Hagmann and Schultheiss 2018; Harney 1976; Hartwell et al. 2008; Haspolat et al. 2014; Haq 2008, 2009; Hawliczek et al. 2020; Hazama 1967a, b; Hernández-Muñoz et al. 2020; Hirochika et al. 2004; Harle 1972, 1974; Heinze and Schmidt 1998; Henikoff and Comai 2003; Henry et al. 2014a, b; Heslot 1962, 1964; Heslot et al. 1959, 1961a, b, 1966; Hitoshi 2008; Hoffmann 1959; Hoffmann et al. 1982; Hoffinger et al. 2016; Holme et al. 2019; Holst and Nagel 1997; Hough et al. 1965; Hough and Weaver 1959; Huy and Xuan 2018; IAEA 1967, 1968, 1971a, b, 1974, 1976, 1977; IAEA-TECDOC-1426 2004; Ibrahim 2018; Ibrahim et al. 2018; Ichikawa 1992; Ilieva-Stameva 1971; Iqbal et al. 1991; Ismail and Horie 2017; Iwamasa 1983; Jacobsen and Schouten 2007; Jain 1998a, b, 2006a, b, 2010a, 2012a, b; Jain and Spencer 2006a, b; Jain et al. 2010; Jankowicz-Ciesla et al. 2018; Jan et al. 2012; Jankowicz-Cieslak et al. 2016a, b; Jiang and Ramachandran 2010; Jin 1994; Joiner et al. 2001; Jompuk et al. 2008; Jo and Kim 2019; Joshua 1983, 2000; Joshua and Jambhulkar 2003; Kalchenko et al. 1991; Kamile and Ayse 2015; Kapoor 1981; Karelin et al. 1997; Karmous et al. 2018; Kaul 1973; Kawai 1967, 1975, 1980, 1983, 1986; Kawai and Amano 1991a, b; Keightley and Halligan 2009; Kennedy et al. 2014; Kharkwall 2017; Kharkwal et al. 2008; Kharkwal and Shu 2009; Klekowski 2011; Knuth 1974; Knuth and Kaufmann 1974; Kodym and Afza 2003; Konzak et al. 1984; Kovacs and Keresztes 2002; Kozhevnikov et al. 2018; Kukimura 1976, 1982; Kukimura et al. 1975a, b, 1976a, b; Kharkwal 1983; Khalatkar and Bhargava 1982; Kharkwal et al. 2004, 2010; Kharkwal 2012a, b; Kleinhofs et al. 1974; Korystov and Narimanov 1997; Kozgar 2014; Kulus 2018; Kuzin et al. 1984; Lagoda et al. 2012; Linqing 1991; Lunden 1964; Lupins 1983; Kojima et al. 2011; Konzak

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et al. 1973; Kool 1982; Kuckuck 1965; Kudr and Heger 2015; Kunzel and Scholz 1972; Kurowska et al. 2016; Kuzin et al. 1975; Kwasniewska and Kwasniewski 2013; Lacey 1977a, b; Lamo et al. 2017; Lagoda 2012; Latado et al. 2001; Lee et al. 2009a, b; Leung et al. 2001; Liu et al. 2006a, b, 2018; Lundqvist 1986, 2005, 2014; Leitao 2012; Leonard et al. 1983; Li et al. 2002; Lineberger et al. 1993; Liu 2021; LoSchiavo et al. 1989; Lönnig 2005; Loveless 1966; Lundquist 2008, 2009; Lundqvist 2018; Lundqvist et al. 2012; Luo et al. 2013, 2016; Luxiang et al. 2018; Lynn 1967; Lysikov 1968; Mai et al. 2008; MacKey 1981; Mba 2013, 2018; Mba et al. 2005, 2010, 2012; Maluszynski and Kasha 2002; Maluszynski and Szarejko 2003; Maluszynski et al. 1994, 2001a, b, 2009a, b; Mandakh et al. 2018; Mergen 1963; Miksche and Shapiro 1964; Medina et al. 2004; Mericle and Mericle 1962; Murty et al. 2007; Micke 1968a, 1985, 1988a, b; Micke et al. 1990a, b; Manjaya 2007; Maruyama et al. 1991; Mashkin et al. 1974; Marton et al. 2010; Mergen 1963; Micco et al. 2011; Miksche and Shapiro 1964; Mikaelsen 1961; Miller and Miller 1987; Miller et al. 1984; Mohajer et al. 2014; Monson-Miller et al. 2012; Nazir et al. 1998; Mokobia and Anomohanran 2005; Mugiono and Hendratno 1995; Mukherjee 1974; Mukherjee and Basu 1976; Nayar 1969; Nakagawa 2008, 2009a, b, 2018; Nishida 1973; Nazir et al. 1998, 2003; Necas 1975; Neuffer 1994; Neuffer et al. 2009; Newton et al. 1989; Nielen 2002; Nilan et al. 1973; Nur et al. 2018; Nybom et al. 1952; Ogor and Odatola 1975; Okamura et al. 2006a, b; Oldach 2011; Oladosu et al. 2015, 2016; Olsen and Dineva 2017; Owais and Kleinhofs 1988; Pacher and Puchta 2017; Pan et al. 2015; Parasuraman and Weerasinghe 2018; Pathirana 1985, 2011; Pathirana et al. 2011; Patterson and Muller 1930; Penna and Jain 2017; Petruccioli et al. 1976; Piluek and Wongpiyasatid 2010; Piri et al. 2011; Perry 1985; Phommalath et al. 2018; Pimonrat and Suraninpong 2009; Pollard 1964; Privalov 1963, 1968a, b; Pohlheim 1981; Pawar et al. 1991; Pokou and Koffi 2018; Potsch 1966b, 1967; Powell and Murray 1978; Pratt 1983; Predieri 2000, 2001; Predieri and Divrgilio 2007; Prina et al. 2012; Przybyla 1994; Purnamaningsih and Hutami 2016; Qüesta et al. 2013; Rai et al. 2011; Raina et al. 2016; Ramnath 2018; Rapoport 1966a, b; Relichova 1984a, b; Reyes-Castro et al. 2018; Rice et al. 2000, Ries et al. 2000; Rizwan et al. 2017; Roux 2004; Roest 1977; Ro and At 2000; Rutger 2006; Rudolph 1965a, b, 1972; Russell et al. 1958; Saccardo et al. 1993; Sacerdot et al. 2005; Sadiq and Owais 2000; Saito 2016; Sakinah 1998; Saliskova and Rapopout 1993; Sanada and Amano 1998; Sanjay 2007; Sanjuán and DomingoCalap 2016; Sarsu et al. 2018a, b, c, 2021; Scheibe 1959; Schneeberger et al. 2009; Schwarzacher and Heslop-Harrison 2018; Sega 1984; Selvarasu and Kandhasamy 2013; Sharma 1986; Shimpei et al. 2010; Shu 2009a, b, 2018; Shu et al. 2009, 2011, 2012a, b; Sitch and Maluszynski 1992; Schum 2003a, b; Sharma 1995; Sharma and Kharkwal 1982; Shouming 1994; Shoba Sivasankar et al. 2021; Siagian et al. 1981; Shapiro and Broertjes 1961; Shu and Lagoda 2007; Singh and Verma 2015; Siddiqui et al. 1991; Sigurbjornsson 1975, 1983; Sigurbjornsson and Micke 1974; Sikora et al. 2011; Sivasankar et al. 2020; Soedjono 2003; Solanki and Waldia 1997; Skirvin 1978; Slota et al. 2017; Smith 1961a, b; Smith and Kersten 1942; Soeranto et al. 2001; Sparrow et al. 1952a, b, 1971; Spencer 1955; Srisombun et al. 2009; Stadler 1928c; Stankovic 1962; Suganthy et al. 1994; Suprasanna and Rao 1997; Suprasanna and Nakagawa 2012; Suprasanna et al. 2012a, b, 2014; Suprasanna and

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Jain 2022; Swaminathan 1964; Swaminathan and Kamra 1961; Swanson et al. 1989; Szarejko 2012; Szarejko and Forster 2007; Tadele et al. 2018; Tai 2007; Tawfik et al. 2011; Tester and Langridge 2010; Tomlekova 2010; Tedoradze 1961; Tham et al. 2001; Till 2016; Till et al. 2016a, b, 2018; Torres and De 2008; Thinh et al. 2007; Toker 2009; Toker et al. 2007, 2012a, b; Tomlekova et al. 2014a, b, c; Torikoshi et al. 2007; Tramontano et al. 2018; Tulmann et al. 2011; Tullman et al. 1998; Turelli 1984; Udage 2021; Udensi et al. 2012; Ukai 1982; Ukai and Yamashita 1974; Vaugh and Wilson 1980; Van Harten 1982, 2002; Van Harten 1988; Van Heck 1962; Velasquez et al. 2018; Vollmann and Jankowicz-Cieslak 2016; Vough and Wilson 1980; Wilde et al. 2012; Wani et al. 2014a, b, c; Walles 1967, 1973; Walbot 1988; Wang et al. 2004; Weider 1956; Whitmore 1995; Wi et al. 2007; Williams 2003; Yamaguchi 1994, 2001, 2018; Yuan et al. 2014; Zaichkina et al. 2004; Zakir 2018; Zhang and Fan 2018; Zhukov and Osipova 1976, Zhu 2008; Zhao 2013). Induced mutations will continue to play a significant role in improving world food security in the coming years and decades. There are many success stories of induced mutations. Mutation breeding has been most successful in ornamental plants. The heterozygous nature of many of the cultivars offers high mutation frequency. The capability of gamma rays in inducing desirable mutations in ornamental plants is well understood from a significant number of new varieties developed via direct mutation breeding. Mutation breeding work especially vegetatively propagated plants have proved to be a valid tool, supplementary to sexual breeding, and sometimes, a unique approach to the genetic improvement of specific characteristics in commercial and promising varieties. Mutations may take place in somatic cells as well as in germinal cells. Germinal cell mutation gives rise to mutated seeds and mutated progenies. Somatic cell mutation gives rise to patches of tissues with mutated cells. Such tissues may develop buds and shoots and bud sports or bud mutations arise in this way. When vegetatively propagated, these buds may give rise to clones of mutated plants. Today, a much deeper and more comprehensive understanding of the principles of mutagenesis and the genetic-implications of induced mutations is possible. The main stumbling-block for practical mutation breeding in vegetatively propagated plants is the chimera formation after the treatment of buds. When an entire branch is mutated, isolation of mutant tissue is possible through conventional propagation methods while a small sectorial mutation in the floret cannot be isolated using existing conventional propagation techniques. Therefore, a large number of new flower color/shape mutants are lost every year. Hence, to isolate and establish the mutated chimeric tissues in a pure form, an appropriate tissue culture method of plant regeneration directly from florets was necessary. The protocol has already been standardized for in vitro regeneration of different crops and recommended somaclonal variation and in vitro mutagenesis for crop improvement (Bajaj 1971; Balkema 1971, 1972; Duron and Decourtye 1986a, b; Hill 1969; Ben-Jaacov and Langhans 1972; Earle and Langhans 1974; Evans and Sharp 1983; Heath-Pagliuso et al. 1988; Nikaido and Onozawa 1989; Malaure et al. 1991a, b; Novak 1991a, b; Nagatomi et al. 1993a, b; Kaul et al. 1990; Lu et al. 1990; Duncan 1997; Remotti 1998a, b; Jain 2007). The main advantage of this technique is to overcome chimera

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formation. In vitro mutagenesis experiments can be conducted with a large population, within limited space, and at any time of the year. The chances of getting solid mutant are more in in vitro mutagenesis. The main advantage of this method is that it helps to avoid chimera formation in the M1V1 (Maliga 1984; Ahloowalia 1995; Jain 1997b; Chai et al. 2004; Jain and Maluszynski 2004; Maluszynski et al. 1995; Kuksova et al. 1997; Predieri 2000, 2001). Both conventional (Datta 1990a, 1992b, c, 1995, 1997a, b) and in vitro techniques have been standardized by the author and his team for the management of chimera. In vitro technique has been discussed in a separate chapter. Disease resistance: Considering high crop losses from diseases and pests, efforts were made in many countries to apply mutagenesis work to develop disease resistance characteristics on many plant species (Abdel-Hafez and Robbelen 1979, 1981; Anderson et al. 1996; Badigannavar et al. 2005; Bhagwat and Duncan 1998a, b; Bolik et al. 1986; Bouter 1977; Caldecott et al. 1959; Chapman et al. 1959; Chen et al. 1986, 2013; Cornu et al. 1977; De Jong et al. 1995; Darmodjo and Sastrowijono 1977; FAO/IAEA 1971a, b, 1977a, b, 1983, 1991; Fadl 1983; Joergensen 1975; Robbelen et al. 1977; IAEA-TECDOC-728 1991). A number of desirable mutants with improved resistance have been reported in different crop species (Aziz et al. 1980; Cepero et al. 2018; Cherukupalli et al. 2018; Damasco et al. 2007; Das and Rahman 1989; De Jong et al. 1995; Effmert and Tellhelm 1984; FAO/IAEA 1971a, b, 1977a, b; Favret 1960a, b, 1965, 1971; Friebe et al. 1991; Fujita 1974; Gomez-Pando 2018a, b; Green Jr. and Path 1975; Haq and Hassan 1980; Haq et al. 1983, 1999, 2001; Hamada et al. 1999; Jankowicz-Ciesla et al. 2018; Jorgensen 1975, 1988, 1992, 1996; Kantoglu et al. 2010; Kiyosawa and Nomura 1988a, b; Konzak 1956b, 1984; Koo and Myers 1955; Lebeda and Svabova 2010; Lee et al. 2014; Lin and Lin 1960; Lestari et al. 2006; Matthews and Dow 1975, 1976; Micke 1974, 1977, 1983, 1992; Murthy 1973; Murray and Todd 1972; Masuda et al. 1997; Mejlhede et al. 2006; Micke et al. 1990a, b; Nabors et al. 1975; Nakai et al. 1988, 1990a, b; Neale 1976; Nakai 1991; Ohta 1974; Liu et al. 2003; Nambisan et al. 1980; Nilan 1973; Oldach et al. 2008; Pathirana 1992; Pham and Nguyen 2018; Pophaly et al. 2006; Rianawati et al. 2013; Robbelen and Heun 1991; Robbelen et al. 1977; Sairam et al. 1997, 1998; Sanada et al. 1988; Saxena et al. 2008; Saunders et al. 1992; Shah et al. 2009a, b; Shapiro 1956a, b; Si et al. 2008; Shirnina et al. 1977; Skou 1985; Sohoo et al. 1985; Sparnaaij 1991; Sreekantappa and Marulasiddappa 2018; Tabira et al. 1993; Tarasenko 1977a, b; Tellhelm and Stelter 1978, 1984; Timmis et al. 1990; To et al. 1989; Todd 1990; Tonnemaker et al. 1992; Torp and Jorgensen 1986; van de Werken 1988; Verghese and Robbelen 1984; Wagner et al. 2018; Wallace 1965; Williams et al. 1992; Worland and Law 1991; Yonli et al. 2018; Zaman et al. 2007). Quality improvement: Considerable work have been carried out on different crops for improving their chemical compositions like starch quality, enzyme activities, protein quality, protein composition, beta carotene, phenolic compounds, oil content and quality, palmitic acid, etc. (Aney 2013; Auld et al. 1992a, b; Azeez et al. 2017; Badigannavar et al. 2007a, b; Bhatia et al. 1991a, b; Brock 1965a, b, c, 1979; Brunner and Keppl 1991a, b; Caoyrgan and Yyldyrym 1989; Chatterjee et al. 2000;

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Cheng 2014; Clayton and Robertson 1955, 1964; Daskalov and Baralieva 1992a, b; Datta 1987b, 1989a; De Jong et al. 1991; Down and Andersen 1956; Duitama et al. 2017; FAO/IAEA 1969a, b, 1970a, b, 1979; Fernández-Martínez et al. 1997; Green and Dribnenki 1996; Green and Marshall 1984; Habben and Larkins 1995; Jaisi et al. 2013; Leij et al. 1991; Iida et al. 1993; Liu et al. 1998; Ismail et al. 1977; Kaul 1973; Khan et al. 2009; Kaul 1980a, b; Kelestanie et al. 2018; Kuzin et al. 1975, 1976; Kozgar et al. 2012; Kumamaru et al. 1997; Liu et al. 2007a, b, c, d; Miklashevichs and Walden 1997; Miu 1973; Miu et al. 1973a, b; Mondal et al. 2011a, b, Muller 1984; Malepszy et al. 1977; Nagatomi et al. 1998a, b; Nair 1992; Odipio et al. 2017; Oladosu et al. 2014; Olsen 1958; Preil et al. 1983, Preil et al. 1988, 1991; Rahman et al. 1994; Rennie 1981; Robbelen 1990; Rowland 1991; Sakai and Suzuki 1964; Scarascia-Mungnozza 1965; Sianipar and Purnamaningsih 2018; Suzuki et al. 2008a, b; Takagi and Rahman 1995; Takagi et al. 1989; Tanaka et al. 2002; Thompson et al. 2019; Tomlekova et al. 2017; Vardhan and Shukla 2017; Vollmann et al. 1997; von Wettstein 2009; Von Wettstein 1995; Farkash et al. 2006; Yang et al. 2016). Phylogenetic Significance: The induced mutations play an important role in understanding the phylogenetic history of several plant species and in studying the phylogenetic pathway and affinities in many plant genera (Goldschmidt 1955; Swaminathan 1963a, b; Gottschalk 1961, Gottschalk 1969; Stube 1966; Nerker 1973, 1976; Goldenberg 1965; Laxmi and Datta 1989). Ion beam, as a new mutation technique, has been widely used in mutation breeding. Effects of ion beams have been investigated on several plants including a few ornamentals. Different kinds of energies and ions such as laser beam irradiation (helium-neon laser, He-Ne), helium (He), carbon (C), neon (Ne), and argon (Ar) with 220 MeV C ions being the most common are used in these studies. Mutation frequency and spectrum induced by C ions were more. C ion-induced mutations with complex patterns of colorations, novel mutant phenotype, wide spectrum of flower color and shape have been reported in ornamental crops like chrysanthemum, carnation, cyclamen, dahlia, delphinium, limonium, petunia, rose, Torenia, Lotus, osteospermum, verbena, etc., and also in other crops (Arabidopsis, banana, tobacco, rice, etc.). Ion beams have the potential to be used in induced mutagenesis work on ornamental crop breeding and also the new basic knowledge generated from these studies can be exploited in biotechnology and molecular biology (Abe et al. 2000, 2002, 2007, 2018; An and Wang 1985; Azad et al. 2018a, b; Bae et al. 2004; Blakely 1992; Chen et al. 1992, 2008a, b, c, 2010; Chinone et al. 2008; Choi et al. 2018; Cui and Luo 1999; Davies and Bateman 1963; Dong and Li 2012; Dong et al. 2007, 2008, 2015, 2016a, b, 2017; Drake 1970; Du et al. 2014; Durante and Cucinotta 2008; Dudley 1999; Ewa et al. 2004; Feng et al. 2006, 2009; Fukuda et al. 1995; Furutani et al. 2008; Fujii et al. 1966; Goodhead 1995; Granhall et al. 1949; Guo et al. 2007; Hamada et al. 1999; Hamatani et al. 2001; Han et al. 2006; Hara et al. 2003; Hase et al. 1999a, b, 2002, 2012a, b, 2018; Hayashi et al. 2007; He et al. 2011; Hirono et al. 1970; Hou et al. 2008a, b; Huang et al. 1996; Ichida et al. 2007; Ikegami et al. 2005; Ishii et al. 2003; Iizuka et al. 2008; Jankowicz-Cieslak and Till 2015, 2016, 2017; Kalimullah

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et al. 2003; Kanaya et al. 2008; Kazama et al. 2007, 2008a, b; Kiefer 2002; Kishinami et al. 1994, 1996; Kondo et al. 2009; Krasaechai et al. 2009; Li et al. 1996, 2018a, b; Ling et al. 2013; Liu et al. 2005a, b, 2006a, b, 2007a, b, 2008, 2012, 2013, 2014; Mei et al. 1994a, b, 1995; Morishita et al. 2003; Maekawa et al. 2003; Matsumura et al. 2010a, b; Miyazaki et al. 2002, 2006; Nagatomi et al. 1995, 1996a, b, c, d, 1997, 1998a, b, 2003; Nakai et al. 1995; Nakayama et al. 2012; Nguyen and Huy 2018; Okamura et al. 2003, 2006a, b, 2015; Pathirana 2012; Phanchaisri et al. 2007; Promnart et al. 2017; Puripunyavanich et al. 2018; Qiu et al. 1991; Qian et al. 2007; Reyes-Borja et al. 2007; Roychowdhury and Tah 2013; Rybianski 2000; Saraye et al. 2018; Sasaki et al. 2008; Sakashita et al. 2002; Schlatholter et al. 2005; Shikazono et al. 2002a, b, 2003a, b, 2005; Shimono 2000; Shirao et al. 2007, 2013; Shu et al. 2012b; Sugiyama et al. 2008a, b; Sun et al. 1996; Suzuki et al. 2005; Tanaka 1999, 2009, 2012; Tanaka et al. 1996a, b, 1997a, b, 2010; Takahashi et al. 2005; Ueno et al. 2002, 2004, 2005; Ukai and Yamashita 2010; Vilaithong et al. 2004; Wang 2006; Wang et al. 1992, 2003a, b, c, d, 2012a, b; Wakita et al. 2008; Wang and Zhang 2007; Watanabe 2001; Watanabe et al. 2008; Wei 1989; Wei et al. 1998, 2003; Wu et al. 1999, 2009; Xie et al. 2007, 2008, 2018; Xin et al. 2007; Yokota et al. 2007a, b; Yamaguchi et al. 2003, 2009; Yang and Tobias 1979; Yang et al. 1997; Yano et al. 2000; Yu et al. 1991a, b, 2007a, b, 2014; Zengquan et al. 2003; Zhang et al. 2008; Zhao et al. 2006; Zhou et al. 2006a, b, c). A new technique and method of mutation induction, i.e., mutagenesis in space (spaceflight-induced mutation technique or “space-breeding”) has been set up to enhance genetic diversity to breed new crop varieties (Dutcher et al. 1994; Liu and Zheng 1997) as the space flight experiments showed that the space conditions are mutagenic (Halstead and Dutcher 1987; Mei et al. 1998). Combined effects of cosmic radiation and microgravity are the main causes of the genetic changes in plants induced by space conditions (Halstead and Dutcher 1987; Gu and Shen 1989). Space-induced mutation breeding of crops (ornamentals—yet to start) may be a novel and effective way to create distinctive genetic resources due to its wide mutation spectrum, high frequency of useful genetic variation, and short breeding period. More than 2000 accessions belonging to 133 species (e.g., rice, wheat, cotton, rapeseed, sesame, pepper, tomato, alfalfa, Spirulina (Arthrospira platensis), etc.) have been reported. But the chance of space flight experimentation is very limited due to the requirement of major investment and technological support (Chen et al. 1994; Cheng et al. 2018; Cucinotta and Durante 2006; Fukuda et al. 1995; Gao et al. 2000, 2008; Guo et al. 2010a, b, 2018a, b, c; Gu et al. 2018; Hou et al. 2009; Hu et al. 2010; Liu et al. 1996, 2000a, b, c, 2002, 2004, 2005a, b, c, d, 2007a, b, c, d, 2009; Luo et al. 2006; Magori et al. 2010; Mei et al. 1994a, b; Qiu et al. 1998; Ren et al. 2010; Shi et al. 2000; Wen and Liu 2001; Wang and Cheng 2018a, b; Xiao et al. 2008; Xu et al. 1995; Yu et al. 2006, 2013; Zhang et al. 2006). There has been substantial technological development—transposable elements for mutation induction, the use of restriction endonuclease for site-directed, homologous recombination, DNA markers linked to mutated genes for marker-assisted selection and tracing of the gene, and target-induced local lesions in genomes (TILLING), as well as different variation versions for high throughput-screening of mutated alleles. There is a

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revolution in plant genomics since improvements in technologies and adoption of novel approaches which have greatly reduced the cost of sequencing the genome of an organism. Reverse-genetics methodologies allow the quick identification of plants carrying mutations in known genes (Wang et al. 2009a, b; Ostergaard and Yanofsky 2004; Alonso and Ecker 2006). Targeting Induced Local Lesions in Genomes (TILLING) is one of the high-throughput, non-transgenic reverse-genetic approaches technique combining chemical mutagenesis with a sensitive DNA screening technique that enables the recovery of individuals carrying allelic variants at candidate genes. TILLING combines traditional mutagenesis followed by high-throughput mutation discovery which can improve the efficiency of using induced mutations to develop crops with improved traits (Amri et al. 2018; McCallum et al. 2000a, b; Coe et al. 2018; Colbert et al. 2001; Datta et al. 2016; Eliot et al. 2008; Greene et al. 2003; HawliczekStrulak et al. 2017; Henikoff et al. 2004; Ishikawa et al. 2010; Caldwell et al. 2004; Chawade et al. 2010a, b; Stemple 2004; Slade and Knauf 2005; Perry et al. 2003; Fernandes 2004; Till et al. 2003, 2004, 2006, 2007a, b, 2009; Ritchie and Nielsen 2006a, b; Xin et al. 2008; Gady et al. 2009; Gilchrist and Haughn 2005; Gilchrist et al. 2006a, b; Kurowska et al. 2011; Sato et al. 2006; Bradley et al. 2007; Cooper et al. 2008; Suzuki et al. 2008a, b; Talame et al. 2008a, b, 2009; Tadele et al. 2009; Wang et al. 2008a, b, c; McCallum et al. 2000a, b; Mejlhede et al. 2006; Martin et al. 2009; Minoia et al. 2010a, b; Chawade et al. 2010a, b; Dahmani-Mardas et al. 2010; Dong et al. 2009a, b; Gilchrist and Haughn 2010; Gottwald et al. 2009; Guo et al. 2018a, b, c; Henry et al. 2014a, b; Laouar et al. 2018; Li et al. 2001a, b; Ritchie and Nielsen 2006a, b; Nieto et al. 2007; Rashid et al. 2011; Schmitt et al. 2012; Simsek and Kacar 2010; Slade et al. 2005; Stephenson et al. 2010; Szarejko and Maluszynski 2011; Rigola et al. 2009; Porch et al. 2009; Szurman-Zubrzycka and Szarejko 2018; Szarejko et al. 2017; Tadele et al. 2010; Tsai et al. 2011; Knoll et al. 2011; Uauy et al. 2009; Wang et al. 2007a, b; Kurowska et al. 2018a, b). TILLMore, a TILLING resource recently developed in Morex barley, is suitable for both forward- and reverse-genetics screening (Talame et al. 2008a, b; Martin et al. 2009; Yi et al. 2009). An improved technique “Endonucleolytic Mutation Analysis by Internal Labeling” (EMAIL) has been developed using capillary electrophoresis which offers an increased degree of sensitivity in the detection of rare mutations in specific genes in pooled samples. EMAIL technique introduces a significant improvement over TILLING approach and offers the plant breeder a new tool for efficient screening of induced point mutation at an early stage for variants in genes of specific interest before taking plants to field trial (Oleykowsky et al. 1998; Comai and Henikoff 2006; Caldwell et al. 2004; Henikoff and Comai 2003; Comai et al. 2004; Slade and Knauf 2005; Cordeiro et al. 2006; Gilchrist et al. 2006a, b; Mejlhede et al. 2006; Sato et al. 2006; Nieto et al. 2007; Cross et al. 2008; Lee et al. 2009a, b; Till et al. 2010). An attempt has been made to summarize the mutation breeding work carried out throughout the world, i.e., crop, mutagen, dose, author, etc., have been shown in Table 1.1.

Chrysanthemum (C. morifolium Ramat) Rooted cuttings, unrooted cuttings, suckers, young plant, mature Plant, leaf explants, in vitro

Calathea crocata Young plant/cuttings Chionodoxa Bulbs

Material Drosophila Ornamental Crops Tuber and bulb crops Amaryllis/Hippeastrum Bulbs Anemone Anemone coronaria, seeds Blue Daisy (Brachycome multifida) Nodal segments Bougainvillea sp. Stem cuttings Anonymous (1989)

Abraham and Desai (1977a), Anonymous (1988, 1989, 1991a, 1991b), Banerji (2008), Banerji et al. (1987a, b), Chen et al. (2012a, b), Datta (1992a), Datta and Banerji (1990, 1994, 1997), Datta et al. (1995), Deng and Liu (1990), Gupta and Nath (1977), Gupta and Shukla (1974), Hong and Shaode (1990), Jayanthi et al. (1999), Judsri et al. (2016), Nath et al. (1983), Sharma et al. (2002), Srivastava et al. (2002), Swaroop et al. (2015), Wang and Chen (2014), Waroonyupa et al. (2016) Anonymous (1988) Alkema (1974a, b)

X-rays 10–50 Gy

γ rays 1 kR, 1.5 krad, 1–5 krad, 250–1250 rad Colchicine 0.05%

X-rays upto 75 Gy, survival upto 15 Gy., 150–175 Gy no survival γ rays 15, 20 Gy; 5–25 Gy [1.92 Gy/min]; 1.75 krad; 1.25–2.5 krad; 10–50 Gy; 30–50 Gy; 2000 rads; 1.5– 0.5 Krad; 1, 3, 5, 7, 15 kR; 1.5–2.5 Krad; acute 20– 100 Gy; Chronic 25–150 Gy for 100 days;

Ahloowalia (1992), Ahmad et al. (2012), Aisyah and Marwoto (2001), Anonymous (1985a, b, 1988, 1991a, b, 1996), Bajpay and Dwivedi (2017, 2019), Banerji and Datta (1993), Banerji et al. (1996), Barakat et al. (2010a, b), Bowen (1964), Bowen et al. (1962), Broertjes (1966a, b, c, 1967), Broertjes (1968a, Broertjes (1968b, 1969a, 1979b), Broertjes et al. (1976, 1983a, b), Broertjes and De Jong (1984), Broertjes and Lock (1985), Bush et al. (1974), Castillo-Martínez et al. (2015), Chen et al. (2007), Crandell et al. (1966), Dao et al. (2006), Datta (1990a, 1992d), Datta and Banerji (1995a, b, 1986a, b, c, d, e, f, g, 1991a, b, c, d), Dalsou and Short (1987), Dash et al. (2000), Datta and Gupta (1980, 1981a, 1983a), De Jong and Custers (1986), Dwivedi et al.

Alkema (1974a, b)

X-rays 100–150 Gy

X-rays 9 Gy

Author Muller (1930, 1946), Sadiq and Owais (2000) Kaicker and Singh (1979)

Mutagens/dose X-rays, sodium azide γ (Gamma) rays 5 and 10 Gy

Table 1.1 Material, dose and author of mutation work

26 1 Introduction/Review

Broertjes et al. (1980), Datta (1991b) Chakrabarty et al. (1999, 2000), Datta et al. (2001), Hossain et al. (2006a, b), Mandal et al. (2000a, b), Misra et al. (2003), Misra and Datta (2007), Tymoszuk and Zalewska (2014), Zalewska et al. (2010, 2011), Wei et al. (2014) Kapoor et al. (2015) Jain et al. (1961), Rana (1964a, b, c, d, 1965a, b) Qiang et al. (2005)

γ rays 1.5–2.5 Krad

γ rays 1.0–2.0 Krad

γ rays 0.5 and 1 Gy; EMS (0.026 and 0.050%); X-irradiation

γ rays γ rays

γ ray γ rays 10 and 15 Gy

Mutant genotype. Rooted cuttings Recurrent irradiation rooted cuttings Management of chimera Rooted cuttings; Ray florets; Leaf explants C. paludosum Poir Annual Chrysanthemum Celosia cristata

(continued)

(2009), Goo (2000), Gupta and Datta (1978), Gupta and Jugran (1978), Gupta and Shukla (1971), Haspolat et al. (2020), Huitema et al. (1986a, b, 1987, 1989, 1991), Hu et al. (1989), Ichikawa et al. (1970), Jank (1955), Jerzy and Zalewska (1996a, b, 1997), Jiang et al. (2004), Jompuk et al. (2001), Jung-Heiliger (1979), Kang et al. (2007a, b), Kapadiya et al. (2016), Kaul et al. (2011), Kim et al. (2016, 2019a, b, c), Kumar et al. (2006, 2012a, b, 2017a, b), Kumari et al. (2013), Lamseejan et al. (2000), Latado et al. (2004), Lee et al. (2008), Lema-Rumińska and Zalewska (2002, 2005), Lema-Ruminska and Zalewska (2004), Liu et al. (2000a, b, c), Machin (1971), Matsumura et al. (1961), Miler et al. (2020, 2021), Miler and Kulus 2018, Miñano et al. (2009), Nagatomi 1991b, Nagatomi et al. (1996a, b, c, d), Nagatomi and Degi (2009), Nakajima and Kawara (1967), Okamura et al. (2015), Park et al. (2007a, b), Patil et al. (2015), Pillai and Abraham (1996a, b), Prasad et al. (2008), Priya et al. (2009), Renu and Lal (2016), Salleh et al. (2015), Satory (1975), Shant and Bhardwaj (2016), Shukla and Datta (1993a), Shimotsuma and Sakurai (1962), Singh and Bala (2015), Sun et al. (2007), Soliman et al. (2014), Tank (1957), Telem et al. (2015), Tulmann and Latado (1996), Ueno et al. (2002, 2004), Verma and Prasad (2019), Verma et al. (2012), Wang et al. (1996, 2011), Weaver (1963), Widiarsih and Dwimahyani (2018), Yamaguchi et al. (2010), Ruprechet (1961), Yamakawa (1970), Yamaguchi et al. (2008), Zalewska et al. (2007), Anne and Lim (2021), Datta and Gupta (1984b, 1987), Datta (1990a, b), Datta and Banerji (1988) Datta (1985a, 1996)

X-rays 1.75 krad, 18 Gy; 18 Gy; 5–25 Gy [0.92 Gy/ min]; Colchicine 0.0625%; Ion beam Colchicine 0.0625 and 0.125%

culture, calli, regenerated plantlets

1.2 Acute and Chronic Irradiations 27

Verbena hortensis, Carthamus tinctorius

Dianthus chinensis Endymion (E. hispanicus) leaves Gladiolus Dormant corms, cormels, seeds

Material Cosmos sulphureus Crocus Tubers Dahlia variabilis Dormant tubers, tubers Irradiation experiments should be carried out immediately after harvest when no visible eyes can be detected Delphinium Dianthus caryophyllus Unrooted cuttings

Table 1.1 (continued)

Heavy-ion beam X-rays 4 Gy Colchicines 1% 7 h γ rays 25–150 Gy; 1–12.5 Krad; X-rays 40–75 Gy; 4 Gy optimum; 10–40 Krad; Chemical mutagen Colhicine 1% 7 h Fast neutrons 142–994 rads NMU, NEU, MMS, EMS, DS, Formalin, Ethylene etc. γ rays

γ rays X-rays 80 Gy γ rays 50 Gy

X-rays 10–40 Gy γ rays 20–30 Gy

Mutagens/dose

Anonymous (1967), Banerji and Datta (1987), Datta 1987b, Banerji et al. (1981, 1994), Broertjes and Bakker (1984), Bhajantri and Patil (2013), Buiatti and Tesi (1968), Buiatti et al. (1965a, b, 1967, 1969, 1970), Dogra et al. (2017), Dryagina (1964, 1975a, b), Drygina and Kazarinov (1966), Gupta and Banerji (1977, 1984), Grabowska (1972, 1975), Grabowska and Mynett (1970a, b, 1974), Iba et al. (1964, 1965), Isaev et al. (1960), Jenkins (1961), Kaicker and Singh (1983), Kumari and Sarkar (2018), Marck (1954), Meshitsuka et al. (1963a, b), Misra (1976, 1977, 1978, 1982, 1983), Misra and Bajpai (1978, 1983a, b), Misra and Choudhary (1979), Moes (1966, 1969), Negi et al. (1983), Raghava et al. (1988), Rahi et al. (1998), Sedelnikova (1988), Spencer (1955), Sheenan and Sagawa (1959), Shukla et al. (2018), Usenbaev and Nazarenko (1970), Yamamoto et al. (1958), Zhakote and Murin (1994) Wu et al. (2005a, b)

Kolar et al. (2020) Anonymous (1988, 1989), Bhattacharya (2003), Dommergues et al. (1966a, b), Hemalatha (1998), Roychowdhury et al. (2012a, b), Roychowdhury and Tah (2011e), Sagawa and Mehlquist (1956) Sugiyama et al. (2008a, b) Alkema (1974a, b), Broertjes and Alkema (1970)

Author Broertjes and Van Harten (1988), Estilai (1978), Gupta and Samata (1967), Mitsukuri and Arai (1965), Syakudo (1964) Anonymous (1977a, b, 1988), Asahira et al. (1975), Broertjes (1968g), Broertjes and Ballego (1967, 1968, 1969), Das et al. (1975, 1977a, b, 1978), Dube et al. (1980), Dong et al. (2007), Grabowska and Mynett (1964), Lantin and Decourtye (1970), Lawrence (1931)

28 1 Introduction/Review

Lilium longiflorum Manita Unrooted cuttings Manolito Unrooted cuttings Muscari armeniacum Leaf pieces

Lilium leichtlinii, seed

Iris Dormant bulbs Lilium, Bulb scales

Kenaf (Hibiscus cannabinus) Hyacinthus

Ornamental hibiscus (Hibiscus moscheutos), seeds Rose of Sharon (Hibiscus syricus syriacus L.), cuttings Hibiscus rosa-sinensis cv. ‘Alipur Beauty’ stem cutting Srivastava and Mishra (2005), Das et al. (1974, 1977a, b), Song et al. (1999), Banerji and Datta (1986, 1988)

γ rays 200, 400 Gy; X-rays 2.5–3.5 Gy; 10 Gy; Fast neutrons 1–6 Gy; DES γ rays 200, 400 Gy

Colchicine, 10, 20, 30 mg/ L X-rays γ rays 250–1000rad X-rays 3–10 Gy Fast neutrons 1.6–5 Gy γ rays 20 Gy X-rays, Fast neutrons Colchicine 0.2–0.4% 7 h

X-rays 2–5 Gy γ rays

X-rays 2–5 Gy, fast neutrons 1–6 Gy X-rays 10 Gy

Hong et al. (1980)

γ rays 1–4 Krad

Broertjes and Alkema (1970), Roest and Bokelmann (1981)

Anonymous (1988)

Hopper and Peloquin (1968) Anonymous (1988)

(continued)

Anonymous (1991a, b), Bowen and Sparrow (1954), Brown and Cave (1953, 1954a, b), Broertjes (1969b), Broertjes and Alkema (1970), Chiba et al. (2005), Chinone et al. (2005), Cuany et al. (1958a, b), Grassotti et al. (1987), Hopper and Peloquin (1968), Lizuka and Ikeda (1963), Loh and Cooper (1966a, b), Lyakh and Lagron 2005a, Lagron and Lyakh (2002), Mitra (1958), Van Groenestijn and Van Tuyl (1983), Wang et al. (1989) Taek et al. (2005)

Dryagina (1989), Hekstra and Broertjes (1968), Konzak and Randolph (1956)

De Mol (1926, 1931, 1933, 1934, 1937a, b, 1940, 1953)

Balogun et al. (2009)

Anonymous (1989), Kang et al. (2007a, b)

γ rays 5, 10, 20, 30, 50, 70, 100 Gy

1.2 Acute and Chronic Irradiations 29

Scilla (S. sibirica), Bulbs Tradescantia Mesembryanthemum Seedlings Marigold (Tagetes erecta L., White marigold In vitro raised seed and rooted cuttings Glebionis segetum (corn marigold)

Mutant genotype Ranunculus, Bulbs

Polyanthes (P. tuberose) Bulbs

Material Narcissus jonquille Bulbs Narcissus tazetta cv. Cicily white Bulb Ornithogalum Leaf parts

Table 1.1 (continued)

Broertjes and van Harten (1988)

X-rays 3–10 Gy; fast neutron 1.6–5 Gy; colchine 0.1–0.4% γ rays 2–4 Krad

Agarwal et al. (2002), Rahi et al. (1998), Singh et al. (2008a, b), Zhou et al. (2006a, b, c)

Kapoor et al. (2014)

γ rays

Neumann et al. (1976) Chaturvedi et al. (1997)

Alkema (1974a, b), Broertjes and Van Harten (1988), Chakravarty and Sen (2001)

γ rays 2–4 Krad; 500–2000 rad

γ rays 500–2000 rad X-rays 100 Gy optimum dose X-ray 3.5 Gy optimum dose γ rays, Chemical γ rays 5–50 Krad

Datta et al. (2003a, b), Rahi et al. (1998)

γ rays 5–50 Krad

Abhangrao et al. (2020), Abraham and Desai (1976), Estrada-Basaldua et al. (2011), Gupta et al. (1974), Kumari and Sarkar (2018), Koley and Gantait (2018), Kaintura et al. (2016), Kainthura and Srivastava (2015), Kayalvizhi et al. (2017), Navabi et al. (2016), Pohare et al. (2013), Pooja et al. (2016), Salvana (2004), Shukla and Datta (1993b), Younis and Borham (1975) Datta and Shukla (1996) Alkema (1974a, b)

Author Alkema (1974a, b), De Mol (1926), Lu et al. (2007)

Mutagens/dose X-rays 4–5 Gy

30 1 Introduction/Review

Guzmania peacockii Seeds Aechmea fasciata Aechmea bromelifolia Seeds, Seedlings Cactus Rhipsalidopsis (Easter cactus) and Zygocactus (Z. truncatus—

Begonia In vitro adventitious bud

De Loose (1966, 1969, 1973), IAEA (1972), Michelssen (1984)

c.f. Broertjes and van Harten (1985)

X-/γ rays 15 Gy

De Loose (1966, 1969, 1973)

(continued)

Pratiwi (2010), Venkatachalam and Jayabalan (1991, 1992, 1994a, b) Bin et al. (2006), Broertjes and van Harten (1988), Chen (2009), Cuany et al. (1985), Doring and Stubbe (1938), Puchooa (2005), Puchooa and Sookun (2003), Sekiguchi et al. (1971), Stubbe and Doring (1938a, b), Sekiguchi et al. (1971), Sheela and Sheena (2014), Sparrow and Pond (1956a, b), Te-Chato and Susanon (2005) Anonymous (1977a, b), Broertjes (1982), Broertjes and van Harten (1988), Benetka (1987), Brown and Harney (1974), Doorenbos (1973), Doorenbos and Karper (1975), Harney (1976), IAEA (1972), Lin and Molnar (1983), Linderman (1968), Matsubara (1982), Matsubara et al. (1971, 1974, 1975), Mikkelson et al. (1975), Molnar (1976), Roest et al. (1981), Shigematsu and Matsubara (1972), Soedjono (1988)

X-rays 300–400 Gy EMS 0.5%, 1%

X-rays 20–30 Gy X-rays 300–400 Gy EMS 0.5, 1% X-rays/Gamma rays 1, 2 Gy, Fast neutrons X- or γ rays 33 Gy

EMS 0.01–0.5% γ rays 7.5 Gy X-/γ rays/ 12–50 Gy, colchicine γ rays γ rays 2.5–10 Gy

Zinnia elegans Flowering pot plants Anthurium. Callus Antirrhinum majus

Anonymous (1988, 1991a, b)

γ rays

Philodendron erubescens ‘Gold’ Weigela Thunb. In vitro plantlets Zephyranthes Buds Banerjee (1967), Spencer (1955), Tisch (1974)

Anonymous (1988), Custers et al. (1977), De Mol (1949), Grabowska and Mynett (1970a, b), Matsubara et al. (1963, 1965), Matsuda (1960), Meshitsuka et al. (1962), de Mol van Oud Loosdrecht (1956), Myodo (1942a, b), Nezu (1962, 1963a, b, 1964, 1965, 1967), Nezu and Obata (1964a, b), Van Eijk and Eikelboom (1981a, b), Li et al. (2022) Karunananda et al. (2018)

X-rays 2.5–8 Gy γ rays

Tulip (Tulipa sp) Dormant, Small and large bulbs

1.2 Acute and Chronic Irradiations 31

Christmas cactus), Freshly detached segments Calathea (syn. Maranta) Plants/cuttings Crossandra (C. infundibuliformis) rooted/unrooted cuttings Cyclamen Young tubers, seeds Euphorbia (E. pulcherrima) Adventitious buds, Suspension culture E. splendens rooted cuttings Gentiana Fuchsia Hoya carnosa (decorative pot plants) rooted cuttings Saintpaulia (African violet) Detached leaves (Adventitious Buds)

Material

Table 1.1 (continued)

Nakatsuka et al. (2005) c.f. Broertjes and van Harten (1988), Bouharmont and Dabin (1986) Anonymous (1988), Van Raalte (1980)

Ando et al. (1986), Anonymous (1988), Broertjes (1968b, c, 1971a, 1972a), Craig and Hampson (1979), Espino and Vazquez (1981), Eyerdom (1981), Grunewaldt (1980, 1983, 1988), Hentrich and Beger (1974), Jungnickel (1977), Kelly and Lineberger (1981a, b), Lineberger and Druckenbrod (1985), Plummer and Leopold (1957), Polheim (1974, 1980, 1981), Pohlheim and Beger (1974) (c.f. Pohlheim and Pohlheim (1976)), Relichova (1984a, b), Seneviratne and

γ rays (20 Gy), acute, chronic or fractionated irradiation 2 × 20 Gy

Spontaneous mutation X-rays 25 Gy X or γ rays 50 Gy

X-ray 5–40 Gy Fast neutrons 2.5–20 Gy, repeated/chronic irradiation, EMS, thermal neutron

c.f. Broertjes and van Harten (1988), Lalitha et al. (1992)

c.f. Broertjes and van Harten (1988)

Author

Breider (1959), c.f. Broertjes and van Harten (1988), Ishizaka (2018), Kameari et al. (2012), Sugiyama et al. (2008a, b) Bergann (1967a, b), Canul-Ku et al. (2012), Kleffel et al. (1986), Koo and Cuevas-Ruiz (1974), Love (1966, 1972), Preil and Engelhardt (1982), Preil et al. (1983), Robinson and Darrow (1929), Stewart (1965)

X-rays 90–100 Gy

X-/γ rays 11 Gy Optimum dose X- or γ rays, optimum dose approx. 20 Gy

Mutagens/dose

32 1 Introduction/Review

Van Raalte (1980) Love (1966, 1972) Alston and Sparrow (1962), Arisumi (1978), Weigle and Butler (1983)

Broertjes and Leffring (1972), Horn (1984), Johnson (1948), Nakronthap (1974), Shama Rao and Singh (1976), Sharma Rao (1977), Schwemmle and Robbelen (1962), Stein and Sparrow (1963, 1966), Stein and Steffensen (1959a), Van Dordrecht (1984)

c.f. Broertjes and van Harten (1985) Bergann (1967b), Craig (1963), Grunewaldt (1983), Jadrná et al. (2011), Kameya (1975), Pohlheim (1977), Pohlheim et al. (1972, 1976), Potsch (1964, 1969), Skirvin and Janick (1976), Stewart et al. (1974), Shigematsu (1973)

X-/γ rays 50 Gy Fast neutron X-/γ rays ±30 Gy, EMS

X-rays 10–30 Gy, γ rays 15–30 Gy

X-/ γ rays Gy 10–15 Gy X-rays 5–12.5 Gy NMU or γ rays

Acute and Chronic Irradiations (continued)

Ahmad et al. (2018) Anonymous (1988, 1991b), Datta (1995)

X-rays 10–40 Gy, fast neutrons 10–20 Gy X-/γ rays 20 Gy, acute dose, NMU γ rays γ rays 10–30 Gy

Achimenes Freshly detached leaves Kohleria Internode explants H. cannabinus Wild sage (Lantana depressa) Stem cuttings Hoya, rooted cuttings Poinsettia Impatiens (I. platypetala) rooted cuttings Kalanchoe K. laciniata, K. daigremontiana, cv. Singapur Adventitious buds, Leaf explants, callus or Cell suspension, In vitro adventitious buds Nertera Pelargonium (P. zonale) Geier (1983, 1988, 1989, 1994), Parliman and Stushnoff (1979)

Colchicines, X-/γ rays 30 Gy

Streptocarpus Leaf segments, leaf cuttings

Wijesundara (2004), Wijesundara (2007), Sparrow and Schairer (1980), Sparrow et al. (1960), Zhou et al. (2006a, b, c), Warfield (1973) Anonymous (1975a, 1991a, b), Broertjes (1968a, 1969c, 1970, 1973, 1982), Broertjes et al. (1969), Brown (1971, 1973, 1974), Brown and Davies (1971), Choudhary (1976), Davies (1971), Davies and Hedley (1975), Nishiyama et al. (1959), Van Raatle and Van Raatle-Wichers (1974), Zeven (1972, 1973) Broertjes (1971b, 1972b, 1973, 1974a, b, 1976, 1977a), Broertjes et al. (1983a, b), IAEA (1972)

1.2 33

Atak et al. (2011), c.f. Broertjes and van Harten (1974), De Loose (1966, 1971a, b, 1974a, b), IAEA (1972), de Loose (1971), Preil and Walther (1983), Streitberg (1966a, b, 1967a, b)

Nagatomi et al. (1993a, b) Das et al. (1977a, b)

c.f. Broertjes and Van Harten (1988) Abraham and Radhakrishnan (2009), Love and Mullenax (1964), Love and Constantin (1965, 1966), Love and Malone (1967)

c.f. Broertjes and van Harten (1988)

γ rays γ rays, chronic 24–88 Gy [0.1–0.37 Gy/h], X-rays 5–10 Gy

X-/γ rays 5–10 Gy (1–2 Gy/min) Fast neutrons 2–10 Gy γ rays 2–10 Gy; 1–100 Gy

X- or γ rays 5–10 Gy

Boonbongkarn et al. (2013), Jiranapapan et al. (2011), Miyazaki et al. (2006), Sawangmee et al. (2011a, b), Sasaki et al. (2008), Suwanseree et al. (2011), Suzuki et al. (2000), Taychasinpitak et al. (2016), Tandon and Bhutani (1965) Bergann and Bergann (1982)

Author

X-rays 20–60 Gy, X- or γ rays 20–60 Gy, chronic γ rays, Recurrent irradiation

X-rays