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Manual on Mutation Breeding Second Edition EMS D E S N M U TECHNICAL REPORTS SERIES No. 119 Manual on Mutation Breeding Second Edition JOINT FAO/IAEA DIVISION OF ATOMIC ENERGY IN FOOD AND AGRICULTURE It^I INTERNATIONAL ATOMIC ENERGY AGENCY, VIENNA, 1 977 MANUAL ON MUTATION BREEDING SECOND EDITION TECHNICAL REPORTS SERIES No.119 MANUAL ON MUTATION BREEDING SECOND EDITION A JOINT UNDERTAKING BY THE FOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONS AND THE INTERNATIONAL ATOMIC ENERGY AGENCY INTERNATIONAL ATOMIC ENERGY AGENCY VIENNA, 1977 MANUAL ON MUTATION BREEDING SECOND EDITION IAEA, VIENNA, 1977 STI/DOC/lO/119 ISBN 92-0-115077-6 Printed by the IAEA in Austria April 1977 FOREWORD The idea of compiling a Manual on Mutation Breeding goes back to the year 1963. Plans for the manual were developed in 1966 by a Panel on Co-ordination of Research on the Production and Use of Induced Mutations in Plant Breeding held in Vienna. It was felt that such a manual would be very useful for advanced university studies, for special courses and plant breeders. Most of the members of that panel volunteered to contribute to the book, and other authors were also brought in, the manual being finally compiled and edited in 1970 by the Plant Breeding and Genetics Section of the Joint FAO/1AEA Division of Atomic Energy in Food and Agri- culture. The first edition went out of print in 1975 and progress in various fields made it desirable not simply to reprint the book but to prepare an updated and revised second edition. The second edition is slightly larger owing to some additional chapters and new literature references. A key- word index has been added too, for easier reference. We are grateful for all the critical comments and suggestions received from readers of the first edition and hope that they find the second edition of greater value for both students and professional plant breeders. We are greatly indebted to the many experts who supplied various sections to the manual. Their contributions are indicated in the List of. Contents. Acknow- ledgement for valuable assistance is due particularly to Bjorn Sigurbjornsson, who compiled the manuscript for the second edition. We should again be happy to receive critical comments, notification of any errors, and suggestions. CONTENTS 1. INTRODUCTION: MUTATIONS IN PLANT-BREEDING PROGRAMMES (B. Sigurbjornsson) 2. MUTAGENIC RADIATION 2.1. Radiation types and radiation sources (R. W. Briggs, M.J. Constantin) 7 2.2. Radio biology (G. Ahnstrom) 21 2.3. Dosimetry (H. Eisenlohr) 28 2.4. Objects and methods of treatment (R. W. Briggs, C.F. Konzak) 33 2.5. Radiation sensitivity and modifying factors (B. V. Conger, C.F. Konzak, R.A. Nilan) 40 2.6. Methods of applying pre- and post-treatments (B. V. Conger, C.F. Konzak, R.A. Nilan) 43 3. CHEMICAL MUTAGENS 3.1. Review of main mutagenic compounds (H. Heslot) 51 3.2. Mode of action (O.P. Kamra, H. Brunner) 59 3.3. Objects and methods of treatment (O.P. Kamra, H. Brunner) 65 3.4. Dose (O.P. Kamra, H. Brunner) 66 3.5. Modifying factors (O.P. Kamra, H. Brunner) 69 3.6. Methods of pre- and post-treatment in chemical mutagenesis (C.F. Konzak, K.R. Narayanan) 71 3.7. Handling and disposal of chemical mutagens (K.R. Narayanan, C.F. Konzak, C. Sander) 74 3.8. Examples of treatment procedures (K. Mikaelsen, C.F. Konzak) 78 4. OTHER CAUSES OF MUTATIONS (F. D'Amato) 4.1. Genetic constitution 81 4.2. Physiological conditions 84 4.3. Environmental causes 84 5. MUTAGEN EFFECTS IN THE FIRST GENERATION AFTER SEED TREATMENT 5.1. Plant injury and lethality (H. Gaul) 87 5.2. Cytological effects (H. Gaul) 91 5.3. Sterility (H. Gaul) 96 5.4. Chimeras (G. Eriksson, D. Lindgren) 98 5.5. Tester stocks for early detection of mutations (R. W. Briggs) 103 6. TYPES OF M UTATIONS 6.1. Genome mutations (A. Gustafsson, I. Ekberg) 107 6.2. Chromosome mutations: Changes in chromosomes, generally without alteration of chromosome number (K. Gustafsson, I. Ekberg) 108 6.3. Extranuclear mutation (A. Gustafsson, R. W. Briggs) 114 6.4. Mutations in characters with continuous variation (R.E. Scossiroli) 118 7. INDUCED-MUTATION TECHNIQUES IN BREEDING SEED-PROPAGATED SPECIES 7.1. Selecting parents and handling the Mj-M3 generations for the selection of mutants (C.F. Konzak, K. Mikaelsen) 125 7.2. The detection of induced mutations (M.S. Swaminathan) 138 7.3. Identification, evaluation and documentation of mutants (H. Hansel, C.F. Konzak) 142 7.4. Factors influencing the mutant spectrum and the quality of mutants (K. Borojevic, W. Gottschalk, A. Micke) 146 7.5. How to breed improved varieties by using induced mutations (A. Micke) 150 8. INDUCED-MUTANT TECHNIQUES IN BREEDING ASEXUALLY PROPAGATED PLANTS 8.1. General considerations, breeding methods and selection of parents (C. Broertjes) 159 8.2. Mutagen treatment and handling of treated material (C. Broertjes) 160 8.3. Cultivars of vegetatively propagated plants developed through mutation induction (A. Micke) 166 9. PLANT CHARACTERS TO BE IMPROVED BY MUTATION BREEDING 9.1. Yielding ability (K. Aastveit) 169 9.2. Flowering and ripening time (T. Kawai) 171 9.3. Adaptability (B. Sigurbjornsson) 172 9.4. Plant type and growth habit <T. Kawai) 173 9.5. Resistance to lodging and stem breakage (G. T. Scarascia-Mugnozza) 174 9.6. Shattering and shedding resistance (W. Gottschalk) 177 9.7. Tolerance to low temperature, drought, heat and salinity (W. Gottschalk) 178 9.8. Disease and pest resistance(E.A. Favret, C.F. Konzak, A. Micke) 180 9.9. Quality (R. Rabson, B. Sigurbjornsson, A. Micke) 188 10. UTILIZATION OF MUTAGEN EFFECTS FOR SOLVING PARTICULAR PLANT- BREEDING PROBLEMS 10.1. Chromosomal reconstruction (A. Hagberg, C.R. Bhatia, C.J. Driscoll) 193 10.2. Alteration of the reproductive system (M.S. Swaminathan, C.J. Driscoll) 195 10.3. The use of mutagens for facilitating recombination (C.R. Bhatia) 199 11. EXPERIENCE WITH INDUCED MUTATIONS IN CROSS-BREEDING (A. Micke).... 201 12. SOMATIC CELL GENETICS AND INDUCED MUTATIONS (P. S. Carlson, H.H. Smith) 205 13. WHEN TO USE MUTATIONS IN PLANT BREEDING (R.D. Brock) 213 BIBLIOGRAPHY : 221 SUBJECT INDEX 281 LIST OF AUTHORS 285 1. INTRODUCTION: MUTATIONS IN PLANT-BREEDING PROGRAMMES Mutations are the ultimate source of all variability in organisms. Variability caused by induced mutations is not essentially different from variability caused by spontaneous mutations during evolution. Methods of using induced mutations in plant breeding are basically the same as those used for newly arisen spontaneous mutations and will be dealt with simultaneously in this book. Mutations can be used for plant breeding in many different ways as shown in Table I, which is based on a classification by Gaul (1963). 1.1. DIRECT USE OF MUTATIONS The direct use of mutations is a very valuable supplementary approach to plant breeding, particularly when it is desired to improve one or two easily identifiable characters in an otherwise well-adapted variety. The main advantages are that the basic genotype of the variety is usually only slightly altered (as compared with hybridization of two distinct varieties), while the improved character(s) is (are) added, and that the time required to breed the improved variety can be shorter than when hybridization is used to achieve the same result. Several cultivars derived from direct utilization of induced mutants have shown that, for example, short straw, earliness, and resistance to certain diseases can be introduced in otherwise well-adapted varieties without significantly altering their other attributes (Sigurbjornsson and Micke, 1974). However, one should watch for pleiotropic effects of the mutated'gene(s) and other unwanted induced mutations (see section 7.4). If undesirable characters are observed, it is advisable to cross the mutant back to the parent line and select for individuals containing the desired mutation(s) that are free of other undesirable changes. Because of the nearly identical genotypes of the parents, one or two backcrosses should normally be sufficient to rid the mutant of such undesirable characters as have been caused by independently induced mutations. In the case of close linkage or severe cases of pleiotropism, the mutagenic treatment should be repeated. One can repeat the treatments with the parent strain or re-treat the mutant line. A desired mutation can be recovered in a homozygous stage already in the M2 or M3 generation as compared with the F6 or F7 generation in the case of hybridization. In the case of plants that flower within the year of seeding this method has resulted in the release of cultivars after three and a half to six years from initial mutagen treatment. The time required for breeding has thus been cut considerably. For species that take several years to reach flowering stage the time-saving could be much greater. Fruit trees are good examples of this (see section 8). When using the mutation-breeding technique, one should pay particular attention to the selection of the parent variety. It should, in general, be the best adapted variety for the location in question. The successful performance of a mutant line does not only depend on the newly discovered beneficial mutation. The genotype of which it is a part determines its other agronomic attributes such as adaptibility, resistance, quality and yield. It is, therefore, essential that the general performance of the mutant line be carefully tested under field conditions. This can be started already in the M3 or M4 generations. If agronomic performance is not satisfactory and if the mutation is a worthwhile improvement in a given character, it is advisable to cross the mutant to a well-adapted and high yielding variety (Table I).
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