Biology of Populus and its Implications for Management and Conservation

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1 A PUBLICATION OF THE NATIONAL RESEARCH COUNCIL OF CANADA MONOGRAPH PUBLISHING PROGRAM Biology of Populus and its Implications for Management and Conservation Edited by R.F. Stettler University of Washington, College of Forest Resources, Seattle, WA 98195, USA H.D. Bradshaw, Jr. Center for Urban Horticulture, Box , University of Washington, Seattle, WA , USA P.E. Heilman Washington State University, WSU Puyallup Research & Extension Center, Puyallup, WA 98371, USA and T.M. Hinckley University of Washington, College of Forest Resources, Seattle, WA 98195, USA NRC MCCNtC RESEARCH PRESS Ottawa 1996

2 Table of Contents Abstract/Resume Preface (R.F. Stettler, H.D. Bradshaw, Jr., P.E. Heilman, and T.M. Hinckley) viii ix PART I. Evolution, genetics, and genetic manipulation. Overview (R.F. Stettler and H.D. Bradshaw, Jr.) 1 CHAPTER 1. Systematics and evolution of Populus (J.E. Eckenwalder)... 7 Introduction 7 Definition of the genus Populus 7 Relationships of Populus 12 Classification of Populus 14 Evolution 19 Conclusions 30 References 30 CHAPTER 2. The genecology of Populus (R.E. Farmer, Jr.) 33 Introduction 33 Adaptation to the growing period 35 Adaptation to climate 39 Morphology. 41 Reproductive characteristics 43 Diseases and insects 44 Molecular genetics 44 Natural variation and wood production 47 Conclusions 48 Topics deserving further study 50 Acknowledgements 50 References 50 CHAPTER 3. Life history, ecology, and conservation of riparian cottonwoods in North America (J.H. Braatne, S.B. Rood, and P.E. Heilman) 57 Introduction 57 Environmental characteristics of riverine systems 58 Life history and ecological properties of riparian cottonwoods 61 Sexual reproduction and establishment 63 Asexual reproduction 70 Establishment of riparian cottonwoods in nonalluvial habitats 70 Growth and maturation 71

3 Conservation and restoration of riparian cottonwood forests 73 Causes of decline in riparian cottonwood populations 74 Conservation and restoration strategies 79 Acknowledgements 80 References 80 CHAPTER 4. The role of hybridization in the genetic manipulation of Populus (R.F. Stettler, L. Zsuffa, and R. Wu) 87 Introduction Natural hybridization Crossability Hybridization for increased productivity Hybridization as analytical tool Conclusions and implications 106 Questions deserving study 107 Acknowledgements 108 References 108 CHAPTER 5. Controlled reproduction of Populus (B.J, Stanton and M. Villar) 113 Introduction 113 Collection of reproductive material 114 Pollen management 119 Seed production 121 Progeny propagation 128 Concluding comments 131 References 132 CHAPTER 6. Poplar breeding and selection strategies (S. Bisoffi and U. Gullberg) 139 Introduction 139 Features of poplar breeding 140 Multiple generation breeding 142 Clonal selection 147 Concluding remarks 153 References 155 CHAPTER 7. Quantitative genetics of poplars and poplar hybrids (D.E. Riemenschneider, H.E. Stelzer, and G.S. Foster) 159 Introduction 159 A short summary of estimation methods 160 A short summary of prediction methods Quantitative genetics and poplars 163 Conclusions and recommendations 176 References 178 IV

4 CHAPTER 8. Molecular genetics of Populus (H.D. Bradshaw, Jr.) Introduction 183 Quantitative genetics at the molecular level 185 The genetic architecture of growth, development, and adaptation 189 The future of molecular genetics in Populus 193 Acknowledgements 197 References 197 CHAPTER 9. Cellular and molecular biology of Agrobacterium-mediated transformation of plants and its application to genetic transformation of Populus (K.-H. Han, M.P. Gordon, and S.H. Strauss) 201 Introduction 201 Tissue culture and regeneration 201 Gene transfer methods 203 Major components of Agrobacterium-mediated transformation system Future prospects 216 Acknowledgements 216 References 216 CHAPTER 10. The specificity of fungal pathogens of Populus (G. Newcombe) 223 Introduction 223 Foliage diseases 226 Stem diseases 233 Host-pathogen interactions 236 Concluding remarks 240 Acknowledgements 241 References 241 CHAPTER 11. Ecological and evolutionary implications of hybridization: Popw/ws-herbivore interactions (T.G. Whitham, K.D. Floate, G.D. Martinsen, E.M. Driebe, and P. Keim) 247 Introduction 247 Defensive chemistry 249 Resistance and introgression in hybrid swarms 254 Herbivore responses to hybrids 258 Conservation of cottonwood hybrid zones 264 Conclusions and implications 269 Acknowledgements 270 References 270

5 PART II. Physiology of growth, productivity, and stress response Overview (T.M. Hinckley) 277 References 281 CHAPTER 12. Leaf growth physiology (E. Van Volkenburgh and G. Taylor) 283 Introduction 283 Leaf development in Populus 283 Physiology of leaf cell growth 288 Environmental effects on leaf expansion 290 Solving questions using Populus 295 References 297 CHAPTER 13. Physiology of secondary tissues of Populus (F.W. Telewski, R. Aloni, and J.J. Sauter) 301 Introduction 301 The vascular cambium and control of vascular differentiation 302 The xylem 307 Wounding responses and compartmentalization 313 Xylem translocation and phloem transport 315 Bark structure and rhytidome development 316 Storage in secondary tissues 317 Concluding remarks 321 References 323 CHAPTER 14. The structure and function of Populus root systems "(K.S. Pregitzer and A.L. Friend) 331 Introduction 331 Root system structure 332 Root growth and mortality 337 Root physiology 341 Conclusions 349 Acknowledgements 350 References 350 CHAPTER 15. Carbon acquisition and allocation (R. Ceulemans and J.G. Isebrands) 355 Introduction 355 Carbon assimilation 357 Carbon allocation and partitioning 373 Models 384 Summary: future needs and direction 391 References 392 VI

6 CHAPTER 16. Water relations (T.J. Blake, J.S. Sperry, T.J. Tschaplinski, and S.S. Wang) 401 Introduction 401 Xylem transport 403 Water relations adjustments 408 Conclusions 417 Acknowledgements 418 References 419 CHAPTER 17. Stress physiology abiotic (D.S. Neuman, M. Wagner, J.H. Braatne, and J. Howe) 423 Introduction 423 Flooding stress 424 Salinity stress 431 Freezing stress 437 Ozone stress 443 Perspectives 448 References 449 CHAPTER 18. Production physiology (P.E. Heilman, T.M. Hinckley, D.A. Roberts, and R. Ceulemans) 459 Introduction 459 Aboveground components and production 460 Belowground components and production 475 Environmental controls of production 479 Summary and conclusions 484 References 485 CHAPTER 19. Linking physiology, molecular genetics, and the Populus ideotype (D.I. Dickmann and D.E. Keathley) 491 Introduction 491 The ideotype concept 493 Achieving the ideotype: can physiology and molecular genetics play a part? 498 Summary and conclusions 510 References 511 CHAPTER 20. Trends in poplar culture: some global and regional perspectives (L. Zsuffa, E. Giordano, L.D. Pryor, and R.F. Stettler) 515 Introduction 515 General overview 515 Some regional perspectives 520 Global outlook 536 Acknowledgements 537 References 537 vii