Genetics and Improvement of Barley Malt Quality (eBook)

Preface 7
Table of Contents 9
1 Barley Production and Consumption 14
1.1 Introduction 14
1.2 World Barley Production 15
1.3 World Barley Trade 19
1.4 World Barley Consumption 20
1.4.1 Feed Consumption 23
1.4.2 Malting Barley Consumption 23
1.4.3 Food Consumption 26
1.4.4 Other Industrial Uses 27
References 29
2 Barley Germplasm and Utilization 31
2.1 Origin and Evolution of Cultivated Barley 32
2.1.1 The Taxonomy of Barley 32
2.1.2 Origin of the Cultivated Barley 35
2.1.3 Evolution and Domestication of Cultivated Barley 38
2.2 Annual Wild Barley 38
2.2.1 Annual Wild Barley in the Near East Fertile Crescent 39
2.2 .1.1 Distribution 39
2.2.1.2 Diversity of target characters in H. spontaneum and potential use in breeding 40
2.2.2 The Annual Wild Barley in Qing-Tibetan Plateau 51
2.2.2.1 Genetic variation of some important characters 51
2.2.2.2 Utilization in barley breeding 52
2.2.3 Other Annual Wild Barley Species 53
2.2.4 Prospective 54
2.3 Perennial Wild.Barley Germplasm 54
2.3.1 The Variation of Target Traits 55
2.3.2 Approaches for Utilization of Perennial Wild Barleys 57
2.4 Cultivated Barleys 59
2.4.1 Primitive Barleys or Landraces 59
2.4.2 Commercial Varieties 60
2.4.3 Mutants 61
References 64
3 Chemical Composition in Barley Grains and Malt Quality 76
3.1 Introduction 76
3.2 Physical Structure 77
3.2.1 Grain Size 78
3.2.2 Dormancy 79
3.2.3 Grain Hardness 80
3.3 The Internal Structure 81
3.3.1 The Embryo 81
3.3.2 The Aleurone 83
3.3.3 The Barley Husk 84
3.4 The Barley Endosperm 84
3.4.1 Barley Carbohydrates 85
3.4.1.1 Starch 85
3.4.1.2 Non-starch polysaccharides 86
3.4.2 Grain Protein 87
3.4.2.1 Low protein barleys 88
3.4.2.2 Storage proteins 89
3.4.2.3 Hordeins, malting and brewing quality 89
3.5 Malt Quality 91
3.5.1 Diastatic Power 91
3.5.2 a-amylase 91
3.5.3 B-amylase 92
3.5.4 Limit Dextrinase 92
3.5.5 a-glucosidase 93
3.5.6 B-glucanase 93
3.5.7 Proteinase 94
3.5.8 Hot Water Extract 95
3.6 Feed Barley Quality Traits 96
3.7 Conclusion 97
References 98
4 Identification of Barley Varieties by Protein Profiling 112
4.1 Introduction 112
4.2 Extraction of Proteins for Protein Profiling 113
4.3 Characteristics of Barley Protein Profiles 114
4.3.1 Peak Size Calling 115
4.3.2 Location and Year 115
4.3.3 Phylogeny 115
4.4 Discussion 120
4.4.1 Specificity of Variety Identification 120
4.4.2 Location and Year 120
4.4.3 Malting versus Feed 121
4.4.4 Peak Inheritance 121
4.4.5 Contamination 122
4.4.6 Automation 123
4.5 Conclusion 124
Acknowledgement 124
References 124
5 B-glucans and Arabinoxylans 126
5.1 ß-glucan 126
5.1.1 Structure of B-glucan in Barley Grain and its Influence on Malting Quality 127
5.1.1.1 Structure and molecular weight of B-glucan 127
5.1.1.2 Effect of B-glucan on malt quality 128
5.1.2 Environmental and Genotypic Variation of B-glucan Content in Barley Grain 129
5.1.2.1 The variation of B-glucan content in barley grain 129
5.1.2.2 Genotypic variation of B-glucan content in barley grain 130
5.1.2.3 Environmental variation of B-glucan content in barley grain 131
5.1.3 Genetic Improvement of B-glucan Content in Barley Grain 133
5.1.4 The Relationships between B-glucan and Malt Quality and Some Agronomic Characters 136
5.1.5 Measurement of B-glucan Content 137
5.1.5.1 The enzymatic method 138
5.1.5.2 Calcofluor flow-Injection analysis (FIA) 138
5.1.5.3 High performance liquid chromatography method 139
5.1.5.4 Near-infrared reflectance spectroscopy method 139
5.2 Arabinoxylans 140
5.2.1 Structure of Arabinoxylans in Barley Grain and Its Influence on Malt Quality 140
5.2.1.1 Structure and molecular weight 141
5.2.1.2 Effects of arabinoxylan on malt quality 142
5.2.2 Environmental and Genotypic Variation of Arabinoxylans Content in Barley Grain 143
5.2.2.1 Amount and synthesis of arabinoxylans in barley brain 143
5.2.2.2 Genotypic and environmental variation of arabinoxylans 144
Acknowledgements 147
References 147
6 The Properties and Genetics of Barley Malt Starch Degrading Enzymes 156
6.1 Introduction 156
6.2 The Substrate: Starch 158
6.2.1 Starch Structure 158
6.2.2 Starch Gelatinisation 159
6.3 The Relationship between Malt DP Enzymes and Fermentability 160
6.3.1 Measurement of DP Enzymes 160
6.3.2 Measurement of Fermentability/ AAL 161
6.3.3 Prediction of Fermentability 164
6.4 The DP Enzymes 167
6.4.1 B-amylase 168
6.4.1.1 Biochemistry of B-amylase 169
6.4.1.2 Genetics of B-amylase 174
6.4.1.3 Amino acid substitutions and enzyme thermostability 175
6.4.1.4 Three dimensional structures of different isoforms 177
6.4.2 a-amylase 181
6.4.2.1 Biochemist ry of a-amylase 182
6.4.2.2 Measurement of a-amylase activity 183
6.4.2.3 Genetics of a-amylase 184
6.4.3 Limit Dextrinase Biochemistry and Genetics 184
6.4.3.1 Measurement of limit dextrinase activity 185
6.4.3.2 Biochemistry of limit dextrinase 186
6.4.3.3 Genetics of limit dextrinase 187
6.4.4 a-glucosidase Biochemistry and Genetics 188
6.5 Summary 190
Acknowledgements 191
References 191
7 The Role of Endosperm Cell Walls in Barley Malting Quality 203
7.1 Introduction 203
7.2 Composition of Walls of Barley Endosperm Cells 204
7.3 Properties of Major Wall Components 206
7.3.1 (1,3 1,4)-B-D-Glucans
7.3.2 Arabinoxylans 207
7.3.3 Glucomannans 209
7.3.4 Xyloglucans 209
7.3.5 Pectic Polysaccharides 210
7.3.6 Callose 210
7.3.7 Cellulose 210
7.4 Biosynthesis of Walls during Barley Grain Development 211
7.4.1 Morphology of Barley Grain Development 212
7.4.2 Enzymes Involved in Wall Polysaccharide Biosynthesis 214
7.4.2.1 (1,3 1
7.4.2.2 Arabinoxylan biosynthesis 217
7.4.2.3 Cellulose biosynthesis 218
7.4.2.4 Callose biosynthesis 219
7.4.2.5 Glucomannan biosynthesis 219
7.5 Germination 219
7.5.1 Hydrolysis of (1,3 1,4)-,B-D-glucan
7.5.1.1 (1,3 1,4)-B-D-glucan endohydrolases
7.5.1.2 (1-4)-B-D-glucan glucohydrolases 222
7.5.1.3 B-D-glucan exohydrolases 222
7.5.2 Hydrolysis of arabinoxylans 223
7.5.2.1 (1,4)-B-D-Xylan endohydrolases 224
7.5.2.2 Arabinoxylan arabinofuranohydrolases 226
7.5.2.3 a-L-Arabinofuranosidases and B-D-Xylosidases 227
7.5.3 Hydrolysis of Glucomannans 228
7.5.4 Hydrolysis of Callose 228
7.6 Effects of Wall Components on Malting Quality 229
7.6.1 Wall Components in Malting 229
7.6.1.1 (1,3 1,4)-,B-D-glucans
7.6.1.2 Arabinoxylans 232
7.6.2 Wall Components in Brewing 232
7.6.2.1 (1,3 1,4)-,B-D-glucans
7.6.2.2 Arabinoxylans 234
7.6.3 Effects of Wall Components on Other Malting Quality Parameters 235
7.7 Enhancing Malting Quality through Manipulation of Cell Wall Metabolism 236
7.7.1 Down-regulation of {1,3 1,4)-B-D-glucan Synthesis
7.7.2 Increasing (1,3 l,4)-B-D-glucanase Activity
7.8 Future Prospects 238
Acknowledgements 240
References 240
8 Barley Genomics and Malting Quality Improvement 251
8.1 Introduction 251
8.2 Platform Technologies for Barley 252
8.2.1 The Beadarray by Illumina 253
8.2.2 The GeneChip by Affymetrix 256
8.3 Linking Genotype and Phenotype via Component Traits 259
8.4 Identifying Genes Underpinning Malting Quality 260
8.4.1 Malting QTLs 261
8.4.2 Starch Synthesis and Degradation 263
8.4.3 Endosperm Cell Wall Degradation: Developmental Variation Adds Complexity 267
8.5 Future Developments 269
Acknowledgements 269
References 270
9 Genetic Improvement of Malting Quality through Conventional Breeding and Marker-assisted Selection 273
9.1 Conventional Approaches to Breeding for Malting Quality 273
9.1.1 Definition of Malting Quality 274
9.1.2 Breeding Methods for Malting Quality 275
9.1.3 Genetic Advance under Selection 277
9.1.4 Selection Strategies for Malting Quality 279
9.2 Marker-assisted Selection for Malting Quality 282
9.2.1 Marker and Mapping Technologies 282
9.2.1.1 Microsatellite or SSR markers 282
9.2.1.2 Diversity array technology (DArT) 284
9.2.1.3 Single nucleotide polymorphisms (SNPs) 284
9.2.1.4 Expressed sequence tags (EST) 285
9.2.1.5 Array-based technology and eQTL mapping 286
9.2.1.6 Association mapping 286
9.2.2 Mapping Quantitative Loci Controlling Malting Quality 287
9.3 Marker Assisted Selection for Major Genes Controlling Malting Quality 294
9.4 Marker-assisted Selection for QTLs Controlling Malting Quality 297
9.5 Summary 299
References 301
Index 306