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SpringerBriefs in Plant Science SpringerBriefs present concise summaries of cutting-edge research and practi- cal applications across a wide spectrum of fields. Featuring compact volumes of 50–125 pages, the series covers a range of content from professional to academic. Typical topics might include: • A timely report of state-of-the-art analytical techniques •  A bridge between new research results, as published in journal articles, and a  contextual literature review • A snapshot of a hot or emerging topic • An in-depth case study or clinical example •  A presentation of core concepts that students must understand in order to make  independent contributions. SpringerBriefs in Plant Sciences showcase emerging theory, original research, re- view material, and practical application in plant genetics and genomics, agronomy, forestry, plant breeding and biotechnology, botany, and related fields, from a global  author community. Briefs are characterized by fast, global electronic dissemination,  standard publishing contracts, standardized manuscript preparation and formatting  guidelines, and expedited production schedules. http://www.springer.com/series/10080 Gideon Ladizinsky • Shahal Abbo The Search for Wild Relatives of Cool Season Legumes 1 3 Gideon Ladizinsky Shahal Abbo The RH Smith Faculty of Agriculture, The RH Smith Faculty of Agriculture, Food & Environment Food & Environment The Hebrew University of Jerusalem The Hebrew University of Jerusalem Rehovot Rehovot Israel Israel ISSN 2192-1229 ISSN 2192-1210 (electronic) SpringerBriefs in Plant Science ISBN 978-3-319-14504-4 ISBN 978-3-319-14505-1 (eBook) DOI 10.1007/978-3-319-14505-1 Library of Congress Control Number: 2015933148 Springer Cham Heidelberg New York Dordrecht London © Springer International Publishing Switzerland 2015 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recita- tion, broadcasting, reproduction on microfilms or in any other physical way, and transmission or in- formation 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 publica- tion 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, express or implied, with respect to the material contained herein or for any errors or omissions that may have been made. Printed on acid-free paper Springer is part of Springer Science+Business Media (www.springer.com) Preface The origin and domestication of legumes described in this book began to be seri- ously studied when it became apparent that, while this kind of information had  been quite extensively researched with regard to cereals, there was little knowledge  available on pulses. At the end of the 1960s, a study on the wild relatives of chick- pea was initiated by the senior author, and this was followed by similar studies on  broad bean, fenugreek, common vetch, bitter vetch, and lentil, with a moderate  emphasis on the pea. The junior author joined the project as a PhD student in the  late 1980s to study the genetics of interspecific hybrid embryo abortion in lentil,  and later developed his own research programs on chickpea domestication and its  evolution under domestication, and on wild peas. While this book describes mainly  the findings of our research, pertinent results obtained by others are mentioned and  evaluated. Studies of the wild relatives of each of the abovementioned legumes included  evaluation of their taxonomic status, morphological variations, ecological require- ments, exploration of their distribution, and seed collection in their natural habitats.  Protein profiles of the seeds were examined to obtain preliminary hints of their  affinity with the cultigens. Plants grown from these seeds were used to establish  their karyotypes, produce intra- and interspecific hybrids, and analyze their chro- mosome pairing at meiosis and fertility. The aim of these investigations was to identify the potential wild gene pool of the domesticated forms. Genetic variations among accessions, particularly in the genus Lens, were assessed by the studies of  isozymes and chloroplast DNA. The main findings of such research were discov- ery of the chickpea wild progenitor Cicer reticulatum and the arrangement of an- nual wild chickpea species in three crossability groups, one of which includes the  domesticated form and its wild progenitor, as well as C. echinospermum, which can be used to improve chickpea cultivars. Chickpea domestication appears to be  a remarkable achievement of prehistoric farmers. It became a crop only after the  seed dormancy of the wild progenitor was overcome and a vernalization-insensitive type was selected. This enabled spring sowing, thereby minimizing damage by as- cochyta blight. v vi Preface The wild species of lentil were found to form three crossability groups. One  contains the domesticated form, its wild progenitor, and the wild Lens odemensis. From this group, L. tomentosus was isolated by hybrid embryo abortion and hybrid  sterility. Both L. odemensis and L. tomentosus were delimited as new species during the course of that study. While the idea that L. culinaris ssp. orientalis is the wild progenitor of lentil had already been put forward by Barulina in the 1930s, we were  able, with the aid of characteristics that are monomorphic in the cultigens but vari- able in the wild form, to identify the wild genetic stock that gave rise to the domesti- cated form. All of the wild lentil species exhibit strong seed dormancy, making their  sowing unprofitable. It seems that selection of dormancy-free types by prehistoric  humans was done in wild stands. Our efforts with wild peas have been only modest, and have proceeded along  with two lines: study of amphicarpy in Pisum fulvum, and assessment of the seed yield of different wild pea species under natural conditions and under cultivation. Faba bean and fenugreek are two legumes whose wild progenitors and any other  genetically related species have not yet been identified. The suggestion that the faba  bean originated from the Vicia narbonensis group of species was refuted on the basis of differences in chromosome numbers and shape, seed protein profile, and  cross-incompatibility. Fenugreek is a minor crop with no known genetically related  wild species. The wild Trigonella berythea is the only species known to be cross- compatible with the cultigens, but the resulting hybrid seeds either do not germinate  or they give rise to the albino seedlings that soon die. Common vetch and bitter vetch are fodder plants. The common vetch and its  related form are traditionally divided into a number of species, but are treated here  as an aggregate under the epithet Vicia sativa. Morphologically and karyotypically  this aggregate is remarkably variable, but generally there is no direct relationship  between a given karyotype and a specific morphological species. The V. sativa ag- gregate is in a stage of incipient speciation, although interkaryotypic hybrids still  can be obtained, most of which are partially fertile. This indicates that gene flow  between members of different karyotypes is still possible. Bitter vetch is a minor crop that is utilized to feed ruminants but is poisonous to  monogastric animals. Its wild progenitor is found mainly in Turkey, and relation- ships between the wild and the domesticated form were established by breeding  experiments during the course of this study. Because the wild bitter vetch grows in  nature in small populations and sparse stands, it is not a classical forage plant. Why and how it has been adopted as such remains a mystery. Contents 1 The Lens Genus ......................................................................................... 1 1.1 Morphology and Taxonomy ............................................................... 2 1.1.1 Lens Morphology .................................................................... 3 1.1.2 Key to the Lens Species .......................................................... 3 1.2   Description, Ecology and Distribution  ............................................... 4 1.2.1 Lens culinaris ssp. culinaris ................................................... 4 1.2.2 Lens culinaris ssp. orientalis .................................................. 4 1.2.3 Lens ervoides .......................................................................... 5 1.2.4 Lens ervoides in Ethiopia and Uganda  ...................................   6 1.2.5 Lens lamottei ........................................................................... 7 1.2.6   Lens nigricans ......................................................................... 8 1.2.7 Lens odemensis ....................................................................... 9 1.2.8 Lens tomentosus ...................................................................... 10 1.3 Genetics of Morphological Variation in Wild Lentils ........................ 10 1.4 Dwarfing Genes in the Genus Lens .................................................... 12 1.5   Variation in Molecular Markers  ......................................................... 13 1.6   Chromosome Variation  ....................................................................... 13 1.7   Crossability and Cytogenetic Relationships  ....................................... 15 1.7.1    Genetic Nature of Hybrid Embryo Breakdown in  the Genus Lens ........................................................................ 17 1.8 The Wild Gene Pool of Lentil ............................................................ 17 1.8.1 Potential of the Wild Genetic Resources ................................ 18 1.9 The Species Concept .......................................................................... 19 1.10 Lentil Domestication .......................................................................... 21 1.10.1 W ild Lentil as a Food Source for Prehistoric Humans ........... 21 1.10.2 T he Mystery of Early Lentil Cultivation ................................ 22 1.11  Wild Genetic Stock of the Domesticated Lentil  ................................. 27 vii viii Contents 2 The Annual Species of the Cicer Genus ��������������������������������������������������� 29 2�1 T axonomy of Annual Cicer Species ��������������������������������������������������� 29 2�1�1 K ey to the Annual Cicer Species ������������������������������������������� 30 2�2 Morphology of Annual Cicer Species ������������������������������������������������ 30 2�2�1 Cicer arietinum ssp. arietinum ����������������������������������������������� 30 2�2�2 Cicer arietinum ssp. reticulatum �������������������������������������������� 31 2�2�3 Cicer bijugum ������������������������������������������������������������������������� 31 2�2�4 Cicer chorassanicum �������������������������������������������������������������� 31 2�2�5 Cicer cuneatum ���������������������������������������������������������������������� 31 2�2�6 Cicer echinospermum ������������������������������������������������������������� 31 2�2�7 Cicer judaicum ����������������������������������������������������������������������� 32 2�2�8 Cicer pinnatifidum ����������������������������������������������������������������� 33 2�2�9 Cicer yamashitae �������������������������������������������������������������������� 33 2�3 The Search for the Wild Progenitor of Chickpea ������������������������������� 34 2�4 D istribution and Ecology of Other Annual Wild Chickpea Species ������������������������������������������������������������������������������� 37 2�4�1 Cicer bijugum ������������������������������������������������������������������������� 37 2�4�2 Cicer chorassanicum �������������������������������������������������������������� 37 2�4�3 Cicer cuneatum ���������������������������������������������������������������������� 37 2�4�4 Cicer judaicum ����������������������������������������������������������������������� 37 2�4�5 Cicer pinnatifidum ����������������������������������������������������������������� 39 2�4�6 Cicer yamashitae �������������������������������������������������������������������� 40 2�5 C rossability Relationships Among the Annual Chickpea Species ����� 40 2�6 Economic Potential of Wild Genetic Resources of Chickpea ������������ 42 2�7 The Mystery of Cicer canariense ������������������������������������������������������� 44 2�8 Pathosystems of Ascochyta Blight in Chickpea and Its Wild Relatives ������������������������������������������������������������������������������������ 45 2�9 Chickpea Domestication �������������������������������������������������������������������� 49 2�9�1 Evidence for Introgression from Domesticated to Wild Chickpea ������������������������������������������������������������������������ 52 3 The Pisum Genus ��������������������������������������������������������������������������������������� 55 3�1 Morphology and Taxonomy ��������������������������������������������������������������� 55 3�1�1 Key to Pisum Species and Subspecies ����������������������������������� 56 3�2 D istribution and Ecology ������������������������������������������������������������������� 57 3�2�1 Pisum fulvum �������������������������������������������������������������������������� 57 3�2�2 Pisum sativum ssp� elatius ������������������������������������������������������ 59 3�2�3 Pisum sativum ssp� humile ����������������������������������������������������� 60 3�3 C rossability and Species Relationship ����������������������������������������������� 62 3�4 Species Relationships in Pisum as Inferred from DNA Sequence Comparisons ���������������������������������������������������������������������� 64 Contents ix 3.5 Utilization of Wild Pisum Germplasm for Pea Improvement ............ 66 3.5.1 Pisum fulvum .......................................................................... 66 3.5.2 Pisum sativum ssp. humile ..................................................... 67 3.5.3 Pisum sativum ssp. elatius ...................................................... 67 3.6 Pea Domestication .............................................................................. 68 4 Legumes with No Documented Genetic Relatives .................................. 71 4.1 Faba Bean ........................................................................................... 71 4.1.1 C hromosome Numbers and Karyotypes ................................ 72 4.1.2 S eed Protein Profiles .............................................................. 73 4.1.3 Crossability Experiments ....................................................... 73 4.1.4 Characterization of the Hypothetical Wild Progenitor of Faba Bean ................................................ 74 4.2 Fenugreek ........................................................................................... 75 5 Fodder Crops ............................................................................................. 77 5.1 Vicia Sativa, Aggregate of Common Vetch ........................................ 77 5.1.1 C hromosomal Variation and the Nature of Interkaryotypic Hybrids ..................................................... 78 5.1.2 Evolution of Vicia Sativa Aggregate ...................................... 84 5.1.3 E conomic Potential of Vicia Sativa Wild Genetic Resources ......................................................... 85 5.1.4 P olyploidy in the Vicia Sativa Aggregate ............................... 86 5.2 Vicia ervilia: Bitter Vetch ................................................................... 87 Glossary ........................................................................................................... 91 References ........................................................................................................ 95

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