LIFE SCIENCE Schlegel RYE Genetics, Breeding, RYE and Cultivation R Owing to its considerable winter hardiness, rye is a cereal that played a major Y role in the feeding of European populations throughout the Middle Ages. Recent data shows that rye is grown on about 5.4 million hectares, with a world E production of approximately 13 million tons. While still an important bread food in many countries, rye produced for bread making has decreased or stagnated, whereas production is increasing for other market segments. Particularly, rye G for feeding, ethanol processing, and biogas is promoted in Europe. e n The first comprehensive monograph on rye, Rye: Genetics, Breeding, and Cultivation e Genetics, Breeding, t gathers all the relevant and historic information from botany and genetics to utilization i c and sustainability of rye. The book covers taxonomy, morphology, and other botany- s related aspects of rye. It describes its physiology, cytology, and genetics, including , and Cultivation B use for genetic improvement of other cereals. The author addresses various types of r breeding such as population, hybrid, and molecular breeding. He also discusses rye e cropping, including seeding techniques, fungal and viral diseases, and predators. e d i The book examines the various uses for rye beyond bread making. This includes n feeding, biomass and biogas production, ethanol production, and other important g , characteristics such as phytosterol content and antioxidant activity. It also explores a the nutritional value of rye. Written by a leading expert in the field, this monograph n compiles the most important facets of rye research, past and present. d C u l t i v a t i o n K15939 RRoollff HH.. JJ.. SScchhlleeggeell 6000 Broken Sound Parkway, NW Suite 300, Boca Raton, FL 33487 711 Third Avenue New York, NY 10017 an informa business 2 Park Square, Milton Park www.crcpress.com Abingdon, Oxon OX14 4RN, UK www.crcpress.com K15939_cover.indd 1 8/16/13 12:57 PM RYE Genetics, Breeding, and Cultivation RYE Genetics, Breeding, and Cultivation Rolf H. J. Schlegel Boca Raton London New York CRC Press is an imprint of the Taylor & Francis Group, an informa business CCRRCC PPrreessss TTaayylloorr && FFrraanncciiss GGrroouupp 66000000 BBrrookkeenn SSoouunndd PPaarrkkwwaayy NNWW,, SSuuiittee 330000 BBooccaa RRaattoonn,, FFLL 3333448877--22774422 ©© 22001144 bbyy TTaayylloorr && FFrraanncciiss GGrroouupp,, LLLLCC CCRRCC PPrreessss iiss aann iimmpprriinntt ooff TTaayylloorr && FFrraanncciiss GGrroouupp,, aann IInnffoorrmmaa bbuussiinneessss NNoo ccllaaiimm ttoo oorriiggiinnaall UU..SS.. 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((CCCCCC)),, 222222 RRoosseewwoooodd DDrriivvee,, DDaannvveerrss,, MMAA 0011992233,, 997788--775500--88440000.. CCCCCC iiss aa nnoott--ffoorr--pprrooffiitt oorrggaanniizzaattiioonn tthhaatt pprroovviiddeess lliicceennsseess aanndd rreeggiissttrraattiioonn ffoorr aa vvaarriieettyy ooff uusseerrss.. FFoorr oorrggaanniizzaattiioonnss tthhaatt hhaavvee bbeeeenn ggrraanntteedd aa pphhoottooccooppyy lliicceennssee bbyy tthhee CCCCCC,, aa sseeppaarraattee ssyysstteemm ooff ppaayymmeenntt hhaass bbeeeenn aarrrraannggeedd.. TTrraaddeemmaarrkk NNoottiiccee:: PPrroodduucctt oorr ccoorrppoorraattee nnaammeess mmaayy bbee ttrraaddeemmaarrkkss oorr rreeggiisstteerreedd ttrraaddeemmaarrkkss,, aanndd aarree uusseedd oonnllyy ffoorr iiddeennttiiffiiccaattiioonn aanndd eexxppllaannaattiioonn wwiitthhoouutt iinntteenntt ttoo iinnffrriinnggee.. VViissiitt tthhee TTaayylloorr && FFrraanncciiss WWeebb ssiittee aatt hhttttpp::////wwwwww..ttaayylloorraannddffrraanncciiss..ccoomm aanndd tthhee CCRRCC PPrreessss WWeebb ssiittee aatt hhttttpp::////wwwwww..ccrrccpprreessss..ccoomm Contents List of Figures ...........................................................................................................xi List of Tables ............................................................................................................xv Preface.....................................................................................................................xix Acknowledgments ...................................................................................................xxi Author ..................................................................................................................xxiii Abbreviations .........................................................................................................xxv Chapter 1 Introduction ..........................................................................................1 Chapter 2 Botany ..................................................................................................9 2.1 Origin ........................................................................................9 2.2 Taxonomy and Cytotaxonomy .................................................11 2.3 Gross Morphology ...................................................................21 2.3.1 Plant Height ................................................................22 2.3.2 Spike ...........................................................................25 2.3.3 Spikelet .......................................................................26 2.3.4 Peduncle .....................................................................26 2.3.5 Rachis .........................................................................26 2.3.6 Node and Internode ....................................................27 2.3.7 Glume .........................................................................27 2.3.8 Lemma........................................................................28 2.3.9 Awn ............................................................................29 2.3.10 Palea ...........................................................................29 2.3.11 Flower .........................................................................30 2.3.12 Floret ..........................................................................30 2.3.13 Spike Differentiation ..................................................30 2.4 Root System .............................................................................31 2.5 Seeds ........................................................................................33 2.5.1 Seed Formation, Germination, and Xenia .................37 2.5.1.1 Seed Formation ...........................................37 2.5.1.2 Germination ................................................38 2.5.1.3 Sprouting .....................................................38 2.5.1.4 Xenia ...........................................................38 2.5.2 Long-Term Storage of Seed ........................................40 2.6 Flowering, Fertilization, and Apomixis ..................................43 2.6.1 Flowering ....................................................................43 2.6.2 Fertilization ................................................................44 2.6.3 Incompatibility ...........................................................44 2.6.4 Apomixis ....................................................................45 v © 2010 Taylor & Francis Group, LLC vi Contents 2.7 Rye Genebanks and Collections ..............................................47 2.7.1 Reproduction of Genebank Accessions ......................48 Chapter 3 Physiology ..........................................................................................51 3.1 Life Cycle ................................................................................51 3.1.1 Generative Features ....................................................51 3.1.2 Vegetative Behavior ....................................................56 3.2 Cold Tolerance .........................................................................57 3.2.1 Cytoplasm ...................................................................57 3.2.2 Hardening ...................................................................57 3.2.3 Inheritance ..................................................................58 3.2.4 Antifreeze Activity .....................................................59 3.3 Drought Tolerance ...................................................................60 3.4 Nutrition ..................................................................................63 3.4.1 Macronutrients ...........................................................63 3.4.2 Micronutrients ............................................................64 3.4.2.1 Iron ..............................................................64 3.4.2.2 Copper.........................................................66 3.4.2.3 Zinc .............................................................68 3.4.2.4 Manganese ..................................................71 3.4.2.5 Boron ..........................................................73 3.4.2.6 Molybdenum ...............................................73 3.4.2.7 Chloride ......................................................73 3.5 In Vitro Behavior .....................................................................73 3.5.1 Embryo Culture ..........................................................74 3.5.2 Tissue Culture ............................................................74 3.5.3 Protoplast Culture.......................................................75 3.5.4 Anther Culture and DHs ............................................77 3.5.4.1 Pollen Culture .............................................77 3.5.4.2 Doubled Haploids .......................................77 3.5.5 Somaclonal Variation .................................................79 3.6 Preharvest Sprouting ...............................................................80 3.7 Vernalization ...........................................................................81 Chapter 4 Cytology .............................................................................................85 4.1 Genome Structure ....................................................................85 4.2 Chromosome Number .............................................................89 4.3 Karyotype and Homeology .....................................................91 4.3.1 Karyotype ...................................................................91 4.3.2 Homeology .................................................................96 4.3.3 Neocentric Activity ....................................................97 4.4 Chromosome Pairing ...............................................................98 4.4.1 Meiotic Cycle .............................................................98 4.4.2 Prophase Pairing and Bouquet Stage .......................100 © 2010 Taylor & Francis Group, LLC Contents vii 4.4.3 Initiation of Synapsis ................................................103 4.4.4 Haploids and Chromosome Behavior .......................103 4.4.5 Diploids ....................................................................105 4.4.5.1 Variation of Chromosome Pairing ............105 4.4.6 Tetraploids ................................................................109 4.4.6.1 Chromosome Pairing in Egg Mother Cells .............................................109 4.4.7 Triploids ....................................................................110 4.4.7.1 Chromosome Pairing in Different Genetic Backgrounds ................................110 4.5 Sporogenesis ..........................................................................112 4.5.1 Microsporogenesis and Pollen Development ...........112 4.5.1.1 Unreduced Gametes ..................................114 4.5.1.2 Gametocides .............................................115 4.5.1.3 Pollination and Fertilization .....................116 4.5.1.4 Embryo Development ...............................116 4.5.1.5 Endosperm Development ..........................116 4.5.2 Macrosporogenesis ...................................................117 4.6 Primary Aneuploids ..............................................................117 4.6.1 Primary Trisomics and Telotrisomics ......................117 4.6.2 B Chromosomes .......................................................119 4.6.2.1 Midget Chromosomes ...............................123 4.7 Reciprocal Translocations .....................................................123 4.8 Genetic Donor for Other Crops .............................................124 4.8.1 α-Amylase Inhibitor .................................................125 4.8.2 Aluminum Tolerance ................................................125 4.8.3 Thaumatin ................................................................126 4.8.3.1 Genome Additions ....................................127 4.8.3.2 Chromosome Additions ............................127 4.8.3.3 Chromosome Substitutions .......................127 4.8.3.4 Chromosome Translocations ....................128 Chapter 5 Genetics ............................................................................................131 5.1 Nomenclature and Designation of Genes ..............................131 5.2 General ..................................................................................144 5.2.1 Microdissection ........................................................151 5.3 Chromosomal and/or Regional Localization of Genes and Markers ...........................................................................151 5.4 Linkages ................................................................................168 5.5 QTL Mapping ........................................................................177 5.6 Physical Mapping ..................................................................183 5.7 Comparative Mapping ...........................................................185 5.8 Gene Regulation ....................................................................189 5.9 DNA and Gene Transfer ........................................................191 5.9.1 Glutenin ....................................................................192 © 2010 Taylor & Francis Group, LLC viii Contents Chapter 6 Cytoplasm, Cytoplasmic Male Sterility, and Restorer .....................193 6.1 Cytoplasm ..............................................................................193 6.1.1 Chloroplast ...............................................................195 6.1.2 Chlorophyll Mutants.................................................195 6.1.3 Mitochondrial Genes ................................................196 6.2 Alloplasmic Rye ....................................................................196 Chapter 7 Breeding ...........................................................................................199 7.1 Diploid Rye ............................................................................204 7.1.1 Population Breeding .................................................205 7.1.1.1 Mass Selection ..........................................205 7.1.1.2 Family Selection .......................................205 7.1.1.3 Inbreeding .................................................207 7.1.1.4 Crossbreeding ...........................................209 7.1.1.5 Population Breeding .................................209 7.1.2 Synthetics .................................................................213 7.1.3 Hybrid Breeding .......................................................215 7.1.3.1 Production of Seed Parents .......................217 7.1.3.2 Pollen Parents ...........................................218 7.1.3.3 Restorer .....................................................218 7.1.3.4 Production of Hybrid Seed .......................220 7.1.4 Molecular Breeding ..................................................223 7.1.5 Mutation Breeding....................................................245 7.1.6 Interspecific and Intergeneric Crosses .....................250 7.1.6.1 Interspecific Crosses .................................250 7.1.6.2 Intergeneric Crosses..................................253 7.2 Tetraploid Rye ........................................................................257 7.2.1 Perennial Tetraploids ................................................260 7.3 Dual-Purpose Rye .................................................................261 7.4 Breeding Activities, Varieties, and Institutions Worldwide .........................................................................261 Chapter 8 Rye Cropping ....................................................................................265 8.1 No-Till Rye ............................................................................267 8.2 Seeding ..................................................................................268 8.2.1 Soil Type ...................................................................269 8.2.2 Water ........................................................................269 8.2.3 Nutrition and Dressing .............................................270 8.2.3.1 Micronutrients ..........................................271 8.2.4 Temperature ..............................................................271 8.2.4.1 Harvesting .................................................272 8.3 Diseases, Susceptibility, and Resistance ...............................272 8.3.1 Bacterial Diseases ....................................................272 © 2010 Taylor & Francis Group, LLC Contents ix 8.3.1.1 Bacterial Streak (Xanthomonas campestris) .......................272 8.3.1.2 Halo Blight (Pseudomonas coronafaciens pv. coronafaciens) .............274 8.3.2 Fungi .........................................................................274 8.3.2.1 Ergot (Claviceps purpurea) ......................274 8.3.2.2 Leaf Rust (Brown Rust) ............................277 8.3.2.3 Stem Rust (Puccinia graminis f. sp. secalis) ......................................................279 8.3.2.4 Powdery Mildew (Erysiphe graminis) ......280 8.3.2.5 Scab or Head Blight [Fusarium graminearum (anamorph) syn. Gibberella zeae] ........................................280 8.3.2.6 Fusarium Root Rot (Fusarium culmorum) .................................................281 8.3.2.7 Ramularia Leaf Spots (Ramularia collo-cygni) ...............................................282 8.3.2.8 Anthracnose [Colletotrichum graminicola syn. Glomerella graminicola (teleomorph), Alternaria sp., and Cladosporium herbarum] ............282 8.3.2.9 Black Head Molds [Alternaria sp., Cladosporium herbarum syn. Mycosphaerella tassiana (teleomorph), Epicoccum sp., Sporobolomyces sp., and Stemphylium sp.] ................................282 8.3.2.10 Pink Snow Mold, Fusarium Patch, or Snow Mold [Microdochium nivale syn. Fusarium nivale syn. Monographella nivalis (teleomorph)] .................................282 8.3.2.11 Leaf Spots (Rhynchosporium secalis) ......282 8.3.2.12 Strawbreaker, Eyespot, or Foot Rot [Pseudocercosporella herpotrichoides syn. Tapesia acuformis (teleomorph)] ......283 8.3.2.13 Take-All (Gaeumannomyces graminis) ....283 8.3.3 Viruses ......................................................................284 8.3.3.1 Barley Yellow Dwarf Virus ......................284 8.3.3.2 Tobacco Rattle Virus ................................284 8.3.3.3 Soil-Borne Viruses ...................................284 8.3.3.4 Wheat Spindle Streak Mosaic Virus ........285 8.3.3.5 Soil-Borne Cereal Mosaic Virus ..............285 8.3.4 Insects, Nematodes, and Other Predators ................287 8.3.4.1 Curl Mite (Aceria tosichella) ....................287 8.3.4.2 Fall Armyworm (Spodoptera frugiperda) .........................288 © 2010 Taylor & Francis Group, LLC