Plant Breeding Principles and prospects EDITED BY M.D. Hayward AFRC IGER Welsh Plant Breeding Station, Aberystwyth, UK N.O. Bosemark Hilleshiig AB, Landskrona, Sweden I. Romagosa UdL-IRTA, Lleida, Spain Coordinating editor M. Cerezo, CIHEAM, Zaragoza, Spain ‘SPRINGER-SCIENCE*+BUSINESS MEDIA, B.V.First edition 1993 © 1993 Springer Science+Business Media Dordrecht Originally published by Chapman & Hall in 1993 ‘Typeset by Photoprint, Torquay ISBN 978.-94-010-4665-7 Apart from any fair dealing for the purposes of research or private study, or criticism or review, as permitted under the UK Copyright Designs and Patents Act, 1988, this publication may not be reproduced, stored, or transmitted, in any form or by any means, ‘without the prior permission in writing of the publishers, or in the «case of reprographie reproduction only in accordance with the terms of the licences issued by the Copyright Licensing Agency in the UK, oF in accordance with the terms of the licences issued by the appropriate Reproduction Rights Organization outside the UK. Enquiries concerning reproduction outside the terms stated here should be sent to the publishers at the London address printed on this page. ‘The publisher makes no representation, express or implied, with regard to the accuracy of the information contained in this book and cannot accept any legal responsibility or liability for any errors oF omissions that may be made, A catalogue record for this book is available from the British Library Library of Congress Catalogingsin-Publication data Plant breeding + principles and prospects / edited by M. Hayward, N. Bosemark, I. Romagosa ; coordinating editor, M. Cerezo. Pp. em. ~ (Plant breeding : v. 1) Includes bibliographical references and index. ISBN 978-94-010-4665-7 ISBN 978-94-011-1524-7 (eBook) DOI 10.1007/978.94-011-1524-7 1, Plant breeding. 2. Plant genetics. 1. Hayward, M.D. I, Bosemark, N._ Ill, Romagosa, I. IV. Series $B123,P547 1993 631.5°23-de20 9242654 cP © Printed on permanent acid-free text paper, manufactured in accordance with, the proposed ANSI/NISO Z.39,48-199X and ANSI Z. 39.48-19844 Induced mutations A. Micke and B. Donini 4.1 IS THERE A NEED FOR SUPPLEMENTING EXISTING GERMPLASM BY MUTAGENESIS? Any attempt at plant improvement requires genetic variability. Ancient ‘breeders’ resorted to the genetic variants provided by nature and selected among them those that more suited their desire in terms of fruit or grain size, taste, ease of harvest, suitability for storage, yield, etc. Resistance to biotic attacks and tolerance to abiotic stresses were also selected. This is how domestication began more than 10000 years ago, departing from the rigid requirement of fitness dominating natural evolution, and adapting certain suitable plant species more and more to the increasingly artificial conditions of man-made habitats (FAOMAEA, 1988). In this process of domestication, man relied ‘on spontaneously occurring variants coming from mutations (‘sports’). Outcrossing, leading to the combination and recombination of traits, has certainly aided crop plant evolution. Selec- tion favoured desired and eliminated undesired variants and led, in general, to a reduction of genetic variation. Conscious hybridization by man was performed rather late and only since the end of the 19th century have we understood the principles of hybridization, segregation and recombination. Varieties cultivated by farmers today comprise numerous desirable genes accu- mulated over centuries and may have reached an optimum plateau of a combination of good Plant Breeding: Principles and prospects. Edited by M.D. Hayward, N.O. Bosemark and I. Romagosa, Published in 1993 by Chapman & Hall, London. ISBN 0 412 43390 7. traits for a particular location and purpose. Cross-breeding among well adapted varieties is practised and selection of rare recombinants still gives some improvement. However, sub- stantial progress in plant breeding during the second half of the 20th century came from the courageous crossing of farmers’ varieties with exotic germplasm, where the term exotic may refer to varieties of exotic countries as well as to local landraces and weedy relatives of the crop species in question. Although geneticists estimate the number of genes possessed by a higher plant as being in the order of 100 000, it is assumed that breeders are actually dealing only with a few hundred genes. Asa consequence, when two established cultivars are crossed, the recombination that can be expected will be restricted to a relatively small fraction of the whole plant genome. Mutations, however, whether spontaneous or induced, may affect any of the 100 000 or more genes of a nuclear genome, and in addition also the ones located in cytoplasmic organelles Mutagenesis is a unique force in creating variation, as it may alter even those genes that are common to all the varieties of a species and therefore do not show any segregation after a cross within that species (‘house-keeping genes’). Therefore, when the environment (in a broad sense) imposes new fitness criteria upon a plant population, genetic variability from mutations is fundamental for the ability of the plant population to respond to the challenge (see Chapters 1 and 2). Without the capacity for undergoing mutation, the enforced selection might eventually lead to the extinction of a species. Application of mutagens can increase