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Perennial Cereal Crops for the Cold Temperate Zone: Agronomy, Physiology, Sink Regulation and Disease Resistance

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Perennial Cereal Crops for the Cold Temperate Zone: Agronomy, Physiology, Sink Regulation and Disease Resistance PERENNIAL CEREAL CROPS FOR THE COLD TEMPERATE ZONE: AGRONOMY, PHYSIOLOGY, SINK REGULATION AND DISEASE RESISTANCE By Nikhil Sankara Jaikumar A DISSERTATION Submitted to Michigan State University in partial fulfillment of the requirements for the degree of Crop and Soil Sciences --- Doctor of Philosophy Ecology, Evolution and Behavioral Biology --- Dual Major 2013 ABSTRACT PERENNIAL CEREAL CROPS FOR THE COLD TEMPERATE ZONE: AGRONOMY, PHYSIOLOGY, SINK REGULATION AND DISEASE RESISTANCE By Nikhil Sankara Jaikumar Within the last few decade, renewed attention has been devoted to the development of high yielding, herbaceous perennial plants that can meet the demand for food, forage and fiber in environmentally sustainable ways. While woody plants and annual herbaceous plants have been successfully selected to achieve high levels of reproductive allocation, herbaceous perennial plants with very high levels of sexual reproduction do not presently exist. The lack of high yielding herbaceous perennials is of intense interest to agronomists, horticulturalists, evolutionary biologists, population ecologists, ecophysiologists, and breeders. However, perennial cereal crops face challenges if they are to be economically feasible, including genetic incompatibilities, resource tradeoffs between sexual reproduction and multi- year survival, and potential for disease buildup. In a series of studies between 2008 and 2012, I explore these issues within the context of three perennial cereal species: perennial wheat (Triticum aestivum x Thinopyrum spp.), perennial rye (Secale cereale x montanum) and intermediate wheatgrass (Thinopyrum intermedium). Chapter 1 of this dissertation is a literature review dealing with the history of perennial cereal breeding, potential environmental benefits, and challenges that perennial cereals face. Chapter 2 reports on a two-year agronomic study of four accessions of perennial wheat and one accession of perennial rye. Chapter 3 describes a series of experiments exploring differences in photosynthetic rates, and other traits affecting resource accumulation, between perennial and annual cereals. Chapter 4 explores the extent to which photosynthesis and metabolism in perennial cereals is responsive to changes in carbohydrate supply / demand ratio and whether this responsiveness differs between perennial and annual cereals. Finally, Chapter 5 explores the resistance of three perennial cereal species to the fungal disease Fusarium head blight (FHB). In brief, we find the following. Perennial wheat achieves grain yields of approximately 50% of annual wheat, while perennial rye achieves 75% of annual rye. Both species show an ability to maintain these yields into their second year, and show later flowering than annuals. Perennial wheat, perennial rye and intermediate wheatgrass show 10-60% higher photosynthetic rates than their annual analogues. Intermediate wheatgrass shows declining photosynthetic rates with increasing plant age. These differences are driven primarily by biochemical rather than hydraulic changes. Perennial wheat and rye are more sink-limited than their annual analogues, and show an ability to mostly compensate for moderate source/sink changes. The perennial species appear to show a more conservative reproductive strategy than their annual relatives. Intermediate wheatgrass shows high resistance to FHB, while perennial rye is moderately susceptible and perennial wheat accessions vary in susceptibility. Thus our studies provide novel contributions to the growing literature on perennial cereals, illustrating some physiological traits of perennial cereals as well as some of the problems they face. We hope that our results can contribute to advancing efforts to achieve high yielding herbaceous perennials, as well as to improving our understanding of how life history, source-sink balance and whole-plant age interact to affect resource acquisition rates. Copyright by NIKHIL SANKARA JAIKUMAR 2013 ACKNOWLEDGEMENTS Sincere acknowledgements and thanks are due to all of the people who have helped me with this research over the last several years. In particular I would like to acknowledge the following: My committee members (Sieg Snapp, James Flore, Jen Lau, Doug Landis and Janet Lewis) for invaluable guidance; John Green, and Mark Freeman for invaluable technical assistance and project management; Lee Siler and Randy Laurenz for planting help, disease monitoring assistance and technical advice; Arianna Pikus, Lacey Culbertson, Dan Kane, Emily May and Iman Sylvain for field assistance and help with data collection and maintenance of experiments; Randy Laurenz and Sue Hammar for helping me with my greenhouse experiment; Wayne Loescher for help in designing protocols for carbohydrate measurements and for training me in carbohydrate analysis; Wayne Loescher and Bert Cregg for the loan of critical gas exchange equipment; Alex Eilts and Tom Sharkey for helping train me in gas exchange protocols and help me design my gas exchange experiments; and a variety of people including Jeannine Cavender- Bares, N. Michelle Holbrook, Lawren Sack, Christian Körner, Peter Reich, Deborah Roach, Richard Shefferson, Lee DeHaan, David Van Tassel and others for critical advice and comments on my research topics and methods as they developed. I would like to express my deepest gratitude to all of you. v TABLE OF CONTENTS LIST OF TABLES x LIST OF FIGURES xiii CHAPTER 1. POTENTIAL BENEFITS AND CHALLENGES FACING PERENNIAL CEREAL CROPS 1 Overview 2 Intermediate Wheatgrass: A Case Study in Domestication 5 Hybridization of Grain Crops (Poaceae) with Perennial Wild Relatives 7 Perennial Wheat and Rye: Case Studies in Grass x Cereal Hybridization 9 Potential Environmental Benefits of Perennial Agroecosystems 15 Nutrient Usage and Leaching 15 Improved Carbon Sequestration 19 Improved Soil Quality 22 Resistance and Tolerance to Herbivory and Insects 23 Challenges to Perennial Cereal Crops 25 Genetic Incompatibility 26 Disease Pressure 30 Perenniality vs. Productivity Tradeoffs 33 Reproductive Photosynthesis 36 Annual vs. Perennial Differences in Resource Acquisition 38 Source vs. Sink Limitation of Metabolism 41 Conclusion 55 REFERENCES 57 CHAPTER 2. AGRONOMY OF PERENNIAL WHEAT AND RYE 71 Introduction 72 Materials and Methods 75 Site Characteristics 75 Accession Selection 75 Experimental Design 76 Management 77 Measurements 78 Data Analysis 79 Results 80 Environment 80 Yield of First-year Perennials and Annuals 81 Effect of Plant Age on Yield and Yield Components 83 Plant Growth, PSCR and Phenology 84 Discussion 86 Conclusion 92 vi Acknowledgements 93 APPENDIX 94 REFERENCES 106 CHAPTER 3. RESOURCE ACQUISITION TRAITS IN PERENNIAL AND ANNUAL CEREALS 111 Introduction 112 Materials and Methods 116 Site and Goals 116 Selection of Accessions 118 Design and Layout 122 Mid-season Photosynthetic Rates 123 Diurnal Variation in Photosynthesis 123 Conductance, Transpiration and Ci / Ca 123 Triose Phosphate Utilization, Carboxylation, Electron Transport and Stomatal Limitation 123 LMR and LMA 126 Experiment 3: Characterizing the Intermediate Wheatgrass Population 126 Experiments 4 and 5 127 Statistical Analysis 127 Results 129 Photosynthetic Rates (Experiment 1) 129 Photosynthetic Rates (Experiment 2) 129 Biochemical Traits 130 Hydraulic Traits 131 Diurnal Variation in Photosynthesis 132 Morphological Traits 132 Experiment 3: Sampling the Intermediate Wheatgrass Population 132 Experiment 4: Replicating Wheat vs. Wheatgrass Comparisons 133 Experiment 5: Post-Sexual Cycle Regrowth and Photosynthetic Rate 133 Discussion 134 Conclusion 140 Acknowledgements 140 APPENDICES 142 APPENDIX A. Figures and Tables for Chapter 3 143 APPENDIX B. Modeling the A / Ci Curve 156 REFERENCES 163 CHAPTER 4. SOURCE VS. SINK REGULATION IN PERENNIAL AND ANNUAL CEREALS 169 Introduction 170 Materials & Methods 173 Site and Plant Materials 173 vii Experimental Design 173 Source / Sink Manipulations 175 Survival 176 Photosynthetic Rates 176 Nonstructural Carbohydrates 177 Seed Mass and PSCR 180 Fall Flowering and Fall Photosynthesis 180 Pilot Experiment 181 Statistical Analysis 181 Results 182 Survival 182 Spring Photosynthetic Rates 183 Nonstructural Carbohydrates 184 Seed Mass and Regrowth 186 Fall Flowering and Photosynthesis 187 Pilot Experiment 187 Discussion 188 Conclusion 194 Acknowledgements 195 APPENDICES 196 APPENDIX A. Tables and Figures for Chapter 4 197 APPENDIX B. Pictures of Perennial Cereal Experiments 214 REFERENCES 221 CHAPTER 5. RESISTANCE TO FUSARIUM HEAD BLIGHT IN PERENNIAL CEREALS 227 Introduction 228 Materials & Methods 230 Experimental Design 230 Experiment 1: Plant Materials 231 Experiment 1: Planting Protocol 233 Experiment 1: Inoculation and Observation of Response 234 Experiment 1: Post-sexual Cycle Regrowth 235 Experiment 2: FHB Symptoms on Naturally Infected Plants in the Field 236 Statistical Analysis 237 Results 238 Experiment 1: Overall ANOVA 238 Experiment 1: Resistant and Susceptible Lines 238 Experiment 1: Regrowth 239 Experiment 2: Natural Occurrence of FHB in the Field 240 Discussion 240 Acknowledgments 244 APPENDIX 245 REFERENCES 253 viii CHAPTER 6: CONCLUSION. BROADER IMPACTS OF PERENNIAL CEREALS 258 ix LIST OF TABLES Table 1. Accessions and planting year / age in 2008-2010 agronomy study 100 Table 2. Dates of field operations in 2008-2010 agronomy study 101 Table 3. Means, Fisher’s LSD values, and F-values for yield and yield components, averaged over 2009 and 2010 102 Table 4. Means, Fisher’s LSD values, and F-values for yield and yield components
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