Monographs on 18 Theoretical and Applied Genetics Edited by R. Frankel (Coordinating Editor), Bet-Dagan M. Grossman, Urbana· H. F. Linskens, Nijmegen P. Maliga, Piscataway· R. Riley, London Monographs on Theoretical and Applied Genetics Volume 1 Meiotic Configurations A Source of Information for Estimating Genetic Parameters By 1. Sybenga (1975) Volume 2 Pollination Mechanisms, Reproduction and Plant Breeding By R. Frankel and E. Galun (1977) Volume 3 Incompatibility in Angiosperms By D. de Nettancourt (1977) Volume 4 Gene Interactions in Development By L.1. Korochkin (1981) Volume 5 The Molecular Theory of Radiation Biology By K.H. Chadwick and H.P. Leenhouts (1981) Volume 6 Heterosis Reappraisal of Theory and Practice Editor: R. Frankel (1983) Volume 7 Induced Mutations in Plant Breeding By W. Gottschalk and G. Wolff (1983) Volume 8 Protoplast Fusion Genetic Engineering in Higher Plants By Y.Y. Gleba and K.M. Sytnik (1984) Volume 9 Petunia Editor: K.C. Sink (1984) Volume 10 Male Sterility in Higher Plants By M.L.H. Kaul (1988) Volume 11 Tree Breeding: Principles and Strategies By G. Namkoong, H.C. Kang, and 1.S. Brouard (1988) Volume 12 The Wheat Rusts - Breeding for Resistance By D.R. Knott (1989) Volume 13 Population Dynamics and the Tribolism Model Genetics and Demography By R.F. Costantino and R.A. Desharnais (1991) Volume 14 Genetic Improvement of Tomato Editor: G. Kalloo (1991) Volume 15 Evolution and Function of Heterostyly Editor: S.C.H. Barrett (1992) Volume 16 Distant Hybridization of Crop Plants Editors: G. Kalloo and 1.B. Chowdhury (1992) Volume 17 Cytogenetics in Plant Breeding By J. Sybenga (1992) Volume 18 Cytogenetics of the Festuca-Lolium Complex By P.P. lauhar (1993) p. P. lauhar Cytogenetics of the Festuca-Lolium Complex Relevance to Breeding With 53 Figures Springer-Verlag Berlin Heidelberg New York London Paris Tokyo Hong Kong Barcelona Budapest Prof. Dr. Prem P. lauhar USDA-ARS, Red River Valley Agricultural Research Center Northern Crop Science Laboratory State University Station Fargo, North Dakota 58105-5677 U.S.A. Library of Congress Cataloging-in-Publication Data. lauhar, Prem P. Cytogenetics of the Festuca-Lolium complex: relevance to breeding/P.P. lauhar. p. cm.-(Monographs on theoretical and applied genetics: 18) Includes bibliographical references and index. ISBN-13: 978-3-642-84088-3 e-ISBN-t3: 978-3-642-84086-9 DO I: to. t 007/ 978-3-642-84086-9 1. Fescue-Breeding. 2. Lolium-Breeding. 3. Fescue-Cytogenetics. 4. Lolium-Cytogenetics. I. Title. II. Series. SB201.F48J38 1992 633.2'37-dc20 92-15946 This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilm or in any other way, and storage in data banks. Duplication of this publication or parts thereof is permitted only under the provisions of the German Copyright Law of September 9, 1965, in its current version, and permission for use must always be obtained from Springer-Verlag. Violations are liable for prosecution under the German Copyright Law. © Springer-Verlag Berlin Heidelberg 1993 Softcover reprint of the hardcover lst edition 1993 The use of general descriptive names, registered names, trademarks, 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. Typesetting: Best-set Typesetter Ltd, Hong Kong 3113145-5 4 3 2 1 0 - Printed on acid-free paper To the memory of my father, whose life was a constant struggle for social justice and rights for all. Preface Considering the widespread occurrence and ecological diversity of grasses, their dominance over vast regions of our globe, and their prime importance to humankind, we, the experts, may congratulate ourselves on having become authorities on the most important single family of organisms in the world of life, rivaled only by the human family itself. G. Ledyard Stebbins 1987 From times immemorial man has relied heavily on grasses, directly and indirectly, for most of his basic needs. Grasses have had a profound impact on the development of human societies and influenced civilization - perhaps in more ways than any other crop. A paramount factor in the evolution of civilizations is a steady supply of food. Grasses were indeed essential to the "evolution" of wild and domestic grazing animals, which, in turn, also catalyzed the "evolution" of humankind. Thus, grasses and human civiliza­ tion were inseparably associated, the association marked by intimate coevolution of grasses, herbivores, and man. Even today, on a world-wide basis, acreage under grasslands is es­ timated to be twice that of cropland. However, grasslands are still one of the largest underdeveloped resources in the world. Knowledge acquired from an array of scientific disciplines should help tap their enormous potential for increasing agricultural productivity. It should also help preserve the vast global germplasm resources, the rich reservoirs of desirable genes. Festuca and Lalium are valuable genera of temperate grasslands. The two genera are closely related. They contain some very productive, nutritious, and well-adapted grasses widely used for agricultural and recreational purposes. They are an important source of feed for the livestock industry that provides us with wholesome meat, milk, wool, etc. They also play an important role in conserving and stabilizing soils. These grasses are widely distributed in temperate and cool climates in Europe, North Africa, South America, Asia, South and East Africa, Australia, and New Zealand; and they have been spread by man to several other parts of the world. Because of their importance in temperate agriculture, the Festuca­ Lalium group of grasses have been extensively studied by agronomists, animal scientists, plant breeders, taxonomists, and cytogeneticists. However, no integrated information has been published on them. The results of various studies remain scattered in numerous journals and periodicals in different languages, some of which are not easily accessible to a common researcher. I have attempted, therefore, to compile and synthesize import­ ant information on certain aspects of these grasses into this volume. The ultimate objective would be to use information from different fields of enquiry to improve the quantity and quality of forage and to stop or reduce VIII Preface genetic erosion. This book is, however, not intended to be an encyclopedic review of different topics. This book deals mainly with the cytogenetics of agronomically important grasses of the Festuca-Lolium complex and discusses their basic importance and breeding relevance. The perennial nature of these grasses makes them particularly amenable to cytogenetic and breeding manipulation. Modern biotechnological developments also hold considerable promise for their improvement. While the current available knowledge on the Festuca-Lolium group will be synthesized and discussed, what we do not know about them will also be highlighted. The book consists of 15 chapters, which deal with various facets of cytogenetics of the Festuca-Lolium group in relation to breeding. An appropriate introduction to each chapter and to several sections in different chapters puts the information in proper perspective to help the reader. Each chapter can be read independently, an arrangement that has inevitably led to some duplication between chapters. However, repetition has been kept to a minimum by giving cross-references. Some speculations have been made to stir ideas and interest among researchers. After all, science can and does build on speculates. Chapter 1 describes the general importance of the Festuca-Lolium group of grasses, their complementary attributes, and the promise they hold for agricultural and recreational purposes. Chapter 2 deals with various taxonomic treatments given to this fascinating group. Certain taxonomic controversies are highlighted and the multidisciplinary approach to revealing phylogeny emphasized. Karyotypic features in relation to species evolution and divergence are discussed in Chapter 3. How species divergence in diploids is accompanied by massive changes in the amount of nuclear DNA is described. Discussion of genetic control of diploid-like meiosis in polyploid species of Festuca forms the main body of Chapter 4, the cyto­ genetic basis of reproductive isolation among ecotypes of hexaploid tall fescue is described in Chapter 5, and the phylogenetic and breeding impli­ cations of these phenomena are discussed in Chapter 6, where possibilities of manipulating the diploidizing mechanism to effect intergeneric gene transfers are outlined. Chapter 7 deals with B chromosomes and their controversial role in stabilizing meiosis. Polyploidy and its value in plant improvement are described in Chapter 8, while Chapter 9 outlines the natural occurrence and artificial induction of haploidy in species of Lolium and Festuca, particularly F. arundinacea. Pairing in haploids shows the haplo-insufficiency of the genetic control of pairing; production of homo­ zygous inbred lines is also briefly described. In Chapter 10, aneuploids are described and their use in elucidating cytogenetic architecture of hexaploid tall fescue is briefly stated. While Chapters 11 and 12 deal with intra- and inter-specific, and intergeneric hybridizations, genome relationships, and the development of