Advances in Haploid Production in Higher Plants Alisher Touraev (cid:129) Brian P. Forster (cid:129) S. Mohan Jain Editors Advances in Haploid Production in Higher Plants Editors Dr. Alisher Touraev Dr. S. Mohan Jain Max F. Perutz Laboratories University of Helsinki University of Vienna Department of Applied Biology Dr Bohrgasse 9/4 FI-00014 Helsinki 1030 Vienna Latokartanonkaari 5 Austria Finland Dr. Brian P. Forster BioHybrids International Ltd. Earley, Reading RG6 5FY United Kingdom ISBN 978-1-4020-8853-7 e-ISBN 978-1-4020-8854-4 Library of Congress Control Number: 2008831064 © 2009 Springer Science + Business Media B.V. No part of this work may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfilming, recording or otherwise, without written permission from the Publisher, with the exception of any material supplied specifically for the purpose of being entered and executed on a computer system, for exclusive use by the purchaser of the work. Printed on acid-free paper 9 8 7 6 5 4 3 2 1 springer.com Preface The importance of haploids is well known to geneticists and plant breeders. The discovery of anther-derived haploid Datura plants in 1964 initiated great excitement in the plant breeding and genetics communities as it offered shortcuts in producing highly desirable homozygous plants. Unfortunately, the expected revolution was slow to materialise due to problems in extending methods to other species, including genotypic dependence, recalcitrance, slow development of tissue culture technologies and a lack of knowledge of the underlying processes. Recent years have witnessed great strides in the research and application of haploids in higher plants. After a lull in activities, drivers for the resurgence have been: (1) development of effective tissue culture protocols, (2) identification of genes con- trolling embryogenesis, and (3) large scale and wide spread commercial up-take in plant breeding and plant biotechnology arenas. The first major international symposium on “Haploids in Higher Plants” took place in Guelph, Canada in 1974. At that time there was much excitement about the potential benefits, but in his opening address Sir Ralph Riley offered the following words of caution: “I believe that it is quite likely that haploid research will contrib- ute cultivars to agriculture in several crops in the future. However, the more extreme claims of the enthusiasts for haploid breeding must be treated with proper caution. Plant breeding is subject from time to time to sweeping claims from enthu- siastic proponents of new procedures. Mention may be made of induced mutations and induced polyploids. The new techniques usually put an additional weapon in the armoury of the breeder but they rarely provide the total defence initially sug- gested. Let us study and consider haploids fully but let us be cautious and not take an unrealistic view of the contributions haploids can make to plant breeding real though I expect these to be.” It has taken several years to “study and consider hap- loids” to a point where these concerns can be addressed and understood, but thanks to advances in recent years we are now at this point. Today there are published tissue culture protocols in over 200 plant species, ranging from Aconitum to Zingiber. These have been collated in the book, “Doubled Haploid production in crop plants: a manual” (Maluszynski et al. 2003). This manual is a valuable contribution to the practical application of our topic and we do v vi Preface not wish to repeat a list of protocols here. Rather we would like to complement the manual by emphasizing the scientific basis of haploid and doubled haploid produc- tion in higher plants and to showcase the impact in plant breeding and biotechnol- ogy. Similarly we have striven not to duplicate the contents of the recently published book: “Haploids in crop improvement II”, Springer (Palmer et al. 2005), which contains information on various aspects of haploidy technology and limitations of haploidy utilisation in breeding programmes. Here we have focused on major advances in understanding, such as the genetic control of embryogenesis. Success in identifying the controlling genes has propelled further investigations in func- tional genomics in unravelling the mechanisms involved especially in respect to regulation and their manipulation. These studies are now linking in with gene expression, metabolism studies as well as changes in cell ultra-structure. The take up of doubled haploidy by plant breeders has been no less impressive. Doubled haploidy provides the fastest route to the production of pure breeding lines and the coupling of doubled haploidy with genetic markers provides a rapid and commer- cially attractive means of cultivar production. Doubled haploidy has now become an essential biotechnology in the development of most major crop species and many vegetables, and is rapidly expanding into other species, particularly those of high value, e.g. medicinal, aromatic and ornamental plants. The widespread activity in haploid research has also resulted in unexpected findings and novel applications, which have prompted studies in scientific understanding and patent claims to pro- tect and capitalise on innovative applications. In February 2006 Vienna hosted a major international symposium on “Haploidy in Higher Plants III”, which in itself marked the renaissance in haploidy and dou- bled haploidy. The symposium attracted top experts from around the world and it is their work plus invited contributions that has formed the basis of this book. The book is composed of five sections. In the first, ‘Technology improvement’ reviews are provided that describe advances in the various methods used to produce haploids both generally and specifically for model and major crop species. In the past, many breakthroughs in haploid and doubled haploid production have been achieved through empirical methods. Today the emerging disciplines of functional genomics, proteomics and metabalomics along with increasingly sophisticated methods in cell biology are fuelling the drive for a more scientific basis aimed at understanding the biological processes involved; these approaches are described in section 2 (Biological mechanisms). The third section, ‘Breeding and genetics’ cov- ers the application of the haploid/doubled haploid technologies to what is tradition- ally considered to be the primary end users – plant geneticists and plant breeders. In section 4, the expansion of these technologies is aptly demonstrated in ‘New species’. Finally, Novel applications are considered. We, the editors, have striven to provide a current and balanced view of the sci- ence and application of haploidy/doubled haploidy in plants, in what is a topic undergoing rapid advancement by scientist and take up by end users. The book, although confined to a relatively restricted topic, has widespread appeal ranging from cell biologists, embryologists, research in plant genetics, biotechnology and genetic engineering, and molecular biologists and with considerable commercial end user relevance. Preface vii References Maluszynski et al. (2003). Doubled haploid production in crop plants: a manual. Kluwer, Dordrecht Palmer CE et al. (2005). Haploids in crop improvement II”, Springer, Berlin Riley R. (1974). The status of haploid research. In: Kasha K.J. (ed.), Proceeding of the First International Symposium on Haploids in Higher Plants: Advances and Potential. Univ. Guelph, Guelph, Canada, pp 3–9 Contents Preface ............................................................................................................. v 1 Progress in Doubled Haploid Technology in Higher Plants ................ 1 . M. Wędzony, B.P. Forster, I. Zur, E. Golemiec, M. Szechyńska-Hebda, E. Dubas, and G. Gotębiowska 2 Doubled Haploids via Gynogenesis ........................................................ 35 B. Bohanec 3 Overview of Barley Doubled Haploid Production ................................ 47 P. Devaux and K.J. Kasha 4 Production of Doubled Haploids in Brassica ........................................ 65 J. Gil-Humanes and F. Barro 5 An Overview on Tobacco Doubled Haploids ........................................ 75 K. Belogradova, I. Lewicka, E. Heberle-Bors, and A. Touraev 6 An Overview of Triticale Doubled Haploids ......................................... 87 F. Eudes and A. Chugh 7 Patents and Haploid Plants ..................................................................... 97 J.M. Dunwell 8 Gene Expression Profiling of Microspore Embryogenesis in Brassica napus ...................................................................................... 115 M.R. Malik and J.E. Krochko 9 Expression Profiles in Barley Microspore Embryogenesis .................. 127 M. Muñoz-Amatriaín, J.T. Svensson, A.M. Castillo, L. Cistué, T.J. Close, and M.P. Vallés ix x Contents 10 Proteomics in Rapeseed Microspore Embryogenesis ......................... 135 J. Cordewener, F. van der Wal, R. Joosen, K. Boutilier, and T. America 11 Programmed Cell Death and Microspore Embryogenesis ................ 147 A.L. Varnier, C. Jacquard, and C. Clément 12 Albinism in Microspore Culture .......................................................... 155 A.M. Torp and S.B. Andersen 13 Doubled Haploids in Breeding Winter Oilseed Rape ......................... 161 C. Möllers and M.C.M. Iqbal 14 Anther Culture Derived Doubled Haploids in Oat ............................ 171 E.M. Kiviharju 15 Barley and Wheat Doubled Haploids in Breeding ............................. 179 J. Weyen 16 Rice Doubled Haploids and Breeding .................................................. 189 J. Pauk, M. Jancsó, and I. Simon-Kiss 17 Potato Haploids and Breeding .............................................................. 199 V.-M. Rokka 18 Current Status of Doubled Haploids in Medicinal Plants ................. 209 A.M.R. Ferrie 19 Microspore Embryogenesis in Selected Medicinal and Ornamental Species of the Asteraceae ................................................. 219 U. Bal and A. Touraev 20 Carrot Doubled Haploids ...................................................................... 231 K. Górecka, D. Krzyżanowska, W. Kiszczak, U. Kowalska, and R. Górecki 21 Haploids and Doubled Haploids in Fruit Trees .................................. 241 M.A. Germanà 22 Haploidy in Tef ....................................................................................... 265 L. Gugsa and J. Kumlehn 23 A Novel and Reversible Male Sterility System Using Targeted Inactivation of Glutamine Synthetase and Doubled Haploidy .......... 285 A. Ribarits, A.N.K. Mamun, S. Li, T. Resch, M. Fiers, E. Heberle-Bors, C.-M. Liu, and A. Touraev Contents xi 24 Embryogenic Pollen Culture: A Promising Target for Genetic Transformation ........................................................................ 295 J. Kumlehn 25 Immature Pollen as a Target for Gene Targeting ............................... 307 T. Resch, E. Ankele, R. Badur, B. Reiss, E. Herberle-Bors, and A. Touraev 26 Induction of Semi-Dwarf, Salt Tolerant Rice Mutants from a Tall Salt Tolerant Indica Landrace .......................................... 319 R. Afza, M. Guzmann, F.J. Zapata, E. Tumimbang-Raiz, G. Gregorio, B.P. Forster, and C. Mba 27 Chromosome Doubling in Monocots .................................................... 329 A.M. Castillo, L. Cistué, M.P. Vallés, and M. Soriano 28 Tracking Gene and Protein Expression During Microspore Embryogenesis by Confocal Laser Scanning Microscopy ................. 339 P.S. Testillano and M.C. Risueño Chapter 1 Progress in Doubled Haploid Technology in Higher Plants . M. Wędzony, B.P. Forster, I. Zur, E. Golemiec, M. Szechyńska-Hebda, E. Dubas, and G. Gotębiowska Abstract In the early 1990s, many basic protocols were developed for haploidy and doubled haploidy, but most were inefficient. During the last decade, progress in technology has been achieved mainly by empirical, time and cost consuming testing of protocols; as a consequence success was proportional to the number of laboratories involved. In the most frequently studied crops (barley, wheat, triticale, maize, rice and rapeseed) improved protocols are now used routinely in breeding and although several problems remain the benefits make doubled haploidy well worthwhile. Significant advances have also been achieved in vegetable, fruit, ornamental, woody and medicinal species, though responses in many remain low with legumes being particularly recal- citrant. There has been resurgence in doubled haploids over the last few years with protocols published for almost 200 plant species. The present review aims to show the recent progress in haploid and doubled haploid technology of higher plants. Keywords Technology, androgenesis, gynogenesis, media composition, wide crossing, regeneration, albinism Introduction Doubled haploidy is the fastest route to homozygosity in plants. The core motivation for developing broadly applicable protocols to raise doubled haploids (DHs) are at least three fold: (1) DHs are useful in fixing traits rapidly in desirable combinations M. Wędzony((cid:2)) Institute of Plant Physiology Polish Academy of Sciences, Niezapominajek 21, 30-239, Kraków, Poland and Pedagogical University of Kraków, Pochorążych 2, 30-084, Kraków, Poland E-mail: [email protected] B.P. Forster Biohybrids International Ltd, Po Box, 2411, Earley Reading RG6 5FY, UK . I. Zur, E. Golemiec, M. Szechyńska-Hebda, E. Dubas, and G. Gotębiowska Institute of Plant Physiology Polish Academy of Sciences, Niezapominajek 21, 30-239, Kraków, Poland A. Touraev et al. (eds.) Advances in Haploid Production in Higher Plants, 1 © Springer Science + Business Media B.V. 2009