Cellulose : Molecular and Structural Biology.

Preface ....................................................................................................... xiii Chapter 1: Many Paths up the Mountain: Tracking the Evolution of Cellulose Biosynthesis ........................................ 1 David R. Nobles, Jr. and R. Malcolm Brown, Jr. 1. Introduction ........................................................................................ 1 2. Sequence Comparisons ....................................................................... 3 3. Eukaryotic Cellulose Synthases ........................................................... 4 3.1. The case for a cyanobacterial origin of plant cellulose synthases ..................................................................... 4 3.2. Lateral transfer of cellulose synthase in the urochordates .......... 4 3.3. The cellulose synthase of Dictyostelium discoideum ................... 5 4. Bacterial Gene Clusters ....................................................................... 6 4.1. Introduction .............................................................................. 6 4.2. Characterized gene clusters ........................................................ 6 5. Novel Gene Clusters ............................................................................ 8 5.1. Introduction .............................................................................. 8 5.2. Group III ................................................................................... 8 5.3. Group IV ................................................................................... 10 6. Concluding Remarks ........................................................................... 12 References ........................................................................................... 12 Chapter 2: Evolution of the Cellulose Synthase (CesA) Gene Family: Insights from Green Algae and Seedless Plants ........................................... 17 Alison W. Roberts and Eric Roberts 1. Overview ............................................................................................. 18 2. The Prokaryotic Ancestry of Eukaryotic CesAs .................................. 21 3. Green Algal CesAs and the Evolution of Terminal Complexes ........... 23 4. CesA Diversification and the Evolution of Land Plants ...................... 25 4.1. Evolution of tracheary elements ................................................ 25 4.2. Functional specialization of CesA proteins ............................... 26 v vi Table of Contents 4.3. Tip growth and the function of Cellulose synthase-like type D (CslD) Genes ............................................................... 26 4.4. CesA and CslD genes of the moss Physcomitrella patens .......... 27 5. Analysis of CesA Function by Targeted Transformation in P. patens ......................................................................................... 28 Acknowledgments ............................................................................. 28 References ......................................................................................... 29 Chapter 3: The Cellulose Synthase Superfamily ....................................... 35 Heather L. Youngs, Thorsten Hamann, Erin Osborne and Chris Somerville 1. Introduction ...................................................................................... 35 2. Identification of Cellulose Synthase .................................................. 37 3. Toward a Functional Analysis of Cellulose Synthase ........................ 38 4. Identification of the Cellulose Synthase-like Genes ........................... 40 Acknowledgments ............................................................................. 45 References ......................................................................................... 46 Chapter 4: Cellulose Synthesis in the Arabidopsis Secondary Cell Wall ................................................................................... 49 Neil G. Taylor and Simon R. Turner 1. Introduction ...................................................................................... 50 2. irx Mutant Isolation and Characterization ........................................ 50 3. Three CesAs Are Required for Secondary Cell Wall Cellulose Synthesis ............................................................................ 51 4. Function of Multiple CesA Proteins during Cellulose Synthesis ....... 52 5. Localization of CesA Proteins ........................................................... 54 6. Conservation of CesA Protein Function in other Species ................. 55 7. Other irx Genes Required for Secondary Cell Wall Formation .......... 55 8. Identifying Novel Genes Required for Secondary Cell Wall Formation Using Expression Profiling .............................................. 57 9. Alternative Approaches to Studying Cellulose Synthesis in the Secondary Cell Wall ..................................................................... 58 10. Conclusions ....................................................................................... 59 References ......................................................................................... 59 Chapter 5: From Cellulose to Mechanical Strength: Relationship of the Cellulose Synthase Genes to Dry Matter Accumulation in Maize ............................................................................... 63 Roberto Barreiro and Kanwarpal S. Dhugga 1. Introduction ...................................................................................... 64 2. Role of Cellulose in Stalk Strength .................................................... 65 3. Carbon Flux through Cellulose Synthase .......................................... 65 4. Alteration of Cellulose Formation in Plants ...................................... 66 5. Mass Action and Metabolic Control ................................................. 68 Table of Contents vii 6. The Cellulose Synthase Gene Family ................................................. 71 7. Expression Analysis of the ZmCesA Gene Family ............................ 73 8. Rationale for Future Transgenic Work .............................................. 76 9. Summary ........................................................................................... 77 References .......................................................................................... 78 Chapter 6: Cellulose Biosynthesis in Forest Trees ..................................... 85 Kristina Blomqvist, Soraya Djerbi, Henrik Aspeborg, and Tuula T. Teeri 1. The Properties of Wood .................................................................... 86 1.1. Formation of wood cells .......................................................... 86 1.2. Reaction wood ......................................................................... 88 2. Cellulose Synthesis ............................................................................ 89 2.1. Rosettes: the machinery of cellulose synthesis .......................... 90 2.2. CesA and Csl ............................................................................ 90 2.3. Other enzymes and proteins involved in cellulose synthesis ................................................................. 96 2.4. Other metabolic processes involved in cell wall biosynthesis ...................................................................... 98 3. In Vitro Cellulose Synthesis ............................................................... 99 Acknowledgments ............................................................................. 100 References .......................................................................................... 100 Chapter 7: Cellulose Biosynthesis in Enterobacteriaceae ........................... 107 Ute Römling 1. Introduction ...................................................................................... 107 2. The Cellulose Biosynthesis Operon in Salmonella typhimurium and Escherichia coli ........................................................................... 109 3. Regulation of the Expression of the bcsABZC Operon ..................... 112 4. Regulation of Cellulose Biosynthesis ................................................. 112 5. Regulation of csgD Expression .......................................................... 114 6. Function of AdrA ............................................................................. 115 7. Occurrence of the Cellulose Biosynthesis Operon among Enterobacterial Species ...................................................................... 116 8. Differential Expression of Cellulose among Enterobacteriaceae ........ 118 9. Coexpression of Cellulose with Curli Fimbriae ................................. 118 10. Conclusions ....................................................................................... 119 Acknowledgments ............................................................................. 119 References .......................................................................................... 120 Chapter 8: In Vitro Synthesis and Analysis of Plant (1Æ3)-b-D-glucans and Cellulose: A Key Step Towards the Characterization of Glucan Synthases ................................................... 123 Vincent Bulone 1. Introduction ...................................................................................... 124 2. In Vitro Approaches for the Study of β-glucan Synthesis ................... 127 viii Table of Contents 2.1. Optimization of the conditions for callose and cellulose synthesis .................................................................... 127 2.2. Structural characterization of in vitro products ........................ 132 2.3. Purification of callose and cellulose synthases ......................... 140 References .......................................................................................... 142 Chapter 9: Substrate Supply for Cellulose Synthesis and its Stress Sensitivity in the Cotton Fiber ............................................... 147 Candace H. Haigler 1. Introduction ........................................................................................ 148 2. Overview of Cotton Fiber Cellulose Biogenesis .................................. 149 2.1. The role of cellulose biogenesis in cotton fiber development ...................................................................... 149 2.2. Changes in cellulose characteristics throughout cotton fiber development ...................................................................... 151 2.3. The role of the microtubules in cotton fiber cellulose synthesis ...................................................................... 152 2.4. Molecular biology of cotton fiber cellulose biogenesis ............... 152 2.5. Biochemistry of cotton fiber cellulose biogenesis ....................... 153 3. Substrate Supply for Cotton Fiber Cellulose Biogenesis ...................... 154 3.1. A role for sucrose synthase ......................................................... 154 4. Intrafiber Sucrose Synthesis as a Source of Carbon for Secondary Wall Cellulose Synthesis .................................................... 158 5. A Role for Sucrose Phosphate Synthase in IntraFiber Cellulose Synthesis .............................................................................. 160 6. Stress Sensitivity of Cellulose Synthesis .............................................. 161 Acknowledgments ............................................................................... 163 References ........................................................................................... 163 Chapter 10: A Perspective on the Assembly of Cellulose-Synthesizing Complexes: Possible Role of KORRIGAN and Microtubules in Cellulose Synthesis in Plants ....................................... 169 Inder M. Saxena and R. Malcolm Brown, Jr. 1. Introduction ........................................................................................ 170 2. Structure and Composition of Cellulose-Synthesizing Complexes ........................................................................................... 171 3. Stages in the Assembly of the Rosette Terminal Complex in Plants .............................................................................................. 172 4. Possible Role of KORRIGAN in the Digestion of Glucan Chains and in the Second Stage of the Assembly of the Terminal Complex .................................................................... 174 5. Role of Microtubules in Cellulose Biosynthesis ................................... 177 Table of Contents ix 6. Summary ............................................................................................. 178 Acknowledgments ............................................................................... 179 References ........................................................................................... 179 Chapter 11: How Cellulose Synthase Density in the Plasma Membrane may Dictate Cell Wall Texture .................................................. 183 Anne Mie Emons, Miriam Akkerman, Michel Ebskamp, Jan Schel and Bela Mulder. 1. Textures of Cellulose Microfibrils ....................................................... 183 2. Hypotheses about Cellulose Microfibril Ordering Mechanisms .......... 184 2.1. Microtubule-directed microfibril orientation ............................. 184 2.2. The liquid crystalline self-assembly hypothesis .......................... 186 2.3. Templated incorporation hypothesis .......................................... 187 3. The Geometrical Model for Cellulose Microfibril Orientation ............ 188 4. A role for Cortical Microtubules in Localizing Cell Wall Deposition ........................................................................... 191 5. Criticism on the Geometrical Model ................................................... 192 6. Outlook on the Verification/Falsification of the Geometrical Theory .................................................................. 194 References ........................................................................................... 195 Chapter 12: Cellulose-Synthesizing Complexes of a Dinoflagellate and other Unique Algae .............................................................................. 199 Kazuo Okuda and Satoko Sekida 1. Introduction ........................................................................................ 199 2. Assembly of Cellulose Microfibrils in Dinoflagellates ......................... 200 3. Occurrence of Distinct TCs in the Heterokontophyta ......................... 205 4. Diversification in Cellulose Microfibril Assembly ............................... 210 References ........................................................................................... 212 Chapter 13: Biogenesis and Function of Cellulose in the Tunicates ............ 217 Satoshi Kimura and Takao Itoh 1. Introduction ........................................................................................ 218 2. Texture of the Tunic in the Ascidians .................................................. 219 3. Cellulose-Synthesizing Terminal Complexes in the Ascidians ............. 220 4. A Novel Cellulose-Synthesizing Site in the Tunicates .......................... 225 5. Occurrence of a Cellulose Network in the Hemocoel of Ascidians ........................................................................................ 227 6. Structure and Function of the Tunic Cord in the Ascidians ................ 230 7. Occurrence of Highly Crystalline Cellulose in the Most Primitive Tunicate, the Appendicularians ......................... 231 8. Origin of Cellulose Synthase in the Tunicates ..................................... 233 9. Summary ............................................................................................. 233 References ........................................................................................... 234 x Table of Contents Chapter 14: Immunogold Labeling of Cellulose-Synthesizing Terminal Complexes ................................................................................... 237 Takao Itoh, Satoshi Kimura, and R. Malcolm Brown, Jr. 1. Introduction ........................................................................................ 238 2. The Cellulose-Synthesizing Machinery (Terminal Complexes) ............ 238 3. Advances in the Understanding of Cellulose Synthases ...................... 241 4. How to Prove if the Rosette or Linear TC is the Cellulose-Synthesizing Machinery? ............................................ 242 5. Labeling of Freeze Fracture Replicas .................................................. 243 6. Specific Labeling of Rosette TCs ......................................................... 247 7. Specific Labeling of Linear TCs .......................................................... 249 8. The Mechanism of Labeling of Cellulose Synthases ........................... 249 9. Future Perspectives on SDS-FRL and Research in Cellulose Biosynthesis ..................................................................... 250 Acknowledgments ............................................................................... 252 References ........................................................................................... 252 Chapter 15: Cellulose Shapes ................................................................... 257 Alfred D. French and Glenn P. Johnson 1. Introduction ........................................................................................ 257 2. Cellulose Polymorphy and Crystal Structures ..................................... 258 2.1. The polymorphs ......................................................................... 259 2.2. High-resolution structure determinations .................................. 260 2.3. The dominant twofold shape in crystals ..................................... 260 2.4. Topological nightmare ............................................................... 262 2.5. Interdigitation ............................................................................ 263 3. Other Cellulosic Polymers ................................................................... 264 4. Information from Small Molecules in Self-Crystals and Protein-Carbohydrate Complexes ................................................. 264 5. The φ,ψ to n,h Conversion Map .......................................................... 266 6. Crystal Structures in φ,ψ Space ........................................................... 268 6.1. Cellulose and its oligomers ........................................................ 268 6.2. Small molecules ......................................................................... 269 6.3. Protein-cellodextrin complexes .................................................. 270 6.4. Lactose-protein complexes ......................................................... 272 7. Computerized Energy Calculations Based on Molecular Models .......................................................................... 273 8. Summary ............................................................................................. 278 Acknowledgments ............................................................................... 282 References ........................................................................................... 282 Table of Contents xi Chapter 16: Nematic Ordered Cellulose: Its Structure and Properties ............................................................................................ 285 Tetsuo Kondo 1. Introduction ........................................................................................ 285 2. Structure of Nematic Ordered Cellulose ............................................. 287 2.1. What is nematic ordered cellulose? ............................................. 287 2.2. Nematic ordered α-chitin and cellulose/α-chitin blends (Kondo et al. 2004) .................................................................... 294 2.3. Another type of nematic ordered cellulose: Honeycomb-patterned cellulose (18) .......................................... 297 3. Properties of Nematic Ordered Cellulose ........................................... 297 3.1. The exclusive surface property of NOC and its unique application ................................................................................. 297 4. The Future .......................................................................................... 301 5. Materials and Methods ....................................................................... 302 5.1. Materials .................................................................................... 302 5.2. Water-swollen cellulose film from the DMAc/LiCl solution ....... 302 5.3. Preparation of NOC from water-swollen cellulose films. ............ 303 5.4. Preparation of NOC template in Schramm-Hestrin (SH) medium ................................................ 303 Acknowledgments ............................................................................... 304 References ........................................................................................... 304 Chapter 17: Biomedical Applications of Microbial Cellulose in Burn Wound Recovery ............................................................................. 307 Wojciech Czaja, Alina Krystynowicz, Marek Kawecki, Krzysztof Wysota, Stanisław Sakiel, Piotr Wróblewski, Justyna Glik, Mariusz Nowak and Stanisław Bielecki 1. Introduction ........................................................................................ 308 2. Experimental Design ........................................................................... 309 2.1. Never-dried MC membranes preparation .................................. 309 2.2. Clinical trials ............................................................................. 310 3. Clinical Outcomes ............................................................................... 311 3.1. High conformability, moisture donation, and faster healing ...... 311 3.2. MC is particularly useful in the treatment of facial burns .......... 315 4. Conclusions ......................................................................................... 319 Acknowledgments ............................................................................... 319 References ........................................................................................... 319 Chapter 18: Cellulose as a Smart Material ............................................... 323 Jaehwan Kim 1. Introduction ........................................................................................ 324 2. Experiments ........................................................................................ 327 xii Table of Contents 2.1. EAPap sample preparation ........................................................ 327 2.2. EAPap actuator performance .................................................... 329 2.3. EAPap actuation principle ......................................................... 331 2.4. Mechanical test of EAPap ......................................................... 336 3. Potential Applications ......................................................................... 339 4. Summary ............................................................................................. 341 Acknowledgment ................................................................................ 342 References ........................................................................................... 342 Index .......................................................................................................... 345 Color Plates ...............................................................................................