Lipids in Photosynthesis

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1 Lipids in Photosynthesis Essential and Regulatory Functions Edited by Hajime Wada Department of Life Sciences Graduate School of Arts and Science University of Tokyo Japan and Norio Murata National Institute for Basic Biology Okazaki Japan 21 Springer

2 Contents From the Series Editor Contents Preface Contributors The Editors Author Index Color Plates vii xv xxi xxiii xxvii xxix CP1-CP16 1 Lipids in Thylakoid Membranes and Photosynthetic Cells 1-9 Hajime Wada and Norio Murata Summary 1 I. Introduction 2 II. Genome Sequences Reveal Genes Involved in Lipid Synthesis 2 III. Mutagenesis and Transgenic Techniques 4 IV. Crystallographic Analysis of Soluble and Membrane-Bound Proteins 5 V. Electron Microscopy 6 VI. Improvements in Analytical Techniques 6 VII. Physical Methods for Assessing the Fluidity of Membrane Lipids 6 VIII. DNA Microarrays 6 IX. Perspective 7 Acknowledgements 7 References 7 2 Fatty Acid Biosynthesis in Plants - Metabolic Pathways, Structure and Organization Adrian P Brown, Antoni R. Slabas and John B. Rafferty Summary 12 I. Introduction 12 II. Biosynthesis of Fatty Acids 12 III. Export of Fatty Acids from the Plastid 15 IV. Biosynthesis of Polyunsaturated Fatty Acids, Unusual Fatty Acids and Chain Elongation 16 V. Acylation Reactions and the Biosynthesis of Triacylglycerol 17 VI. Structural Models for Fatty Acid Biosynthetic Pathway Components 20 VII. Integration of Fatty Acid Biosynthesis with Other Metabolic Machinery 27 VIII. Conclusions 28 xv

3 Acknowledgements 28 References 28 3 Biosynthesis and Function of Chloroplast Lipids Mie Shimojima, Hiroyuki Ohta and Yuki Nakamura Summary 35 I. Introduction 36 II. Structures of Chloroplast Lipids 36 III. Biosynthesis of Chloroplast Lipids 38 IV. Regulation of Lipid Biosynthesis 44 V. Concluding Remarks 49 Acknowledgements 50 References 50 4 Lipids in Plant Mitochondria Radin Sadre and Margrit Frentzen Summary 57 I. Introduction 58 II. Lipid Composition of Mitochondrial Membranes 59 III. Lipid Biosynthetic Pathways within Mitochondria 60 IV. Concluding Remarks 70 Acknowledgements 70 References 70 5 Plant Sphingolipids: Structure, Synthesis and Function Ming Chen, Edgar B. Cahoon, Mariana Saucedo-Garcia, Javier Plasencia and Marina Gavilanes-Rulz Summary 78 I. Introduction 78 II. Sphingolipid Structure 79 III. Synthesis of Sphingolipids 82 IV. Sphingolipid Turnover 96 V. Sphingolipid Function 97 VI. Concluding Remarks 107 Acknowledgements 107 References Lipids in Algae, Lichens and Mosses Reimund Goss and Christian Wilhelm Summary 117 I. Introduction 118 II. Lipids in Algae 118 III. Lipids in Lichens 126 IV. Lipids in Mosses 129 V. Conclusions 131 Acknowledgements 132 References 132 xvi

4 7 Molecular Genetics of Lipid Metabolism in the Model Green Alga Chlamydomonas reinhardtii Eric R. Moellering, Rachel Miller and Christoph Benning Summary 139 I. Introduction 140 II. General Differences in Lipid Metabolism between Chlamydomonas and Seed Plants 140 III. Membrane Glycerolipid Biosynthesis 142 IV. Fatty Acid Desaturation 147 V. Neutral Lipid Metabolism 148 VI. Perspectives 149 Acknowledgements 150 References Lipid Biosynthesis and its Regulation in Cyanobacteria Norihiro Sato and Hajime Wada Summary 157 I. Introduction 158 II. Characteristics of Cyanobacterial Lipids 159 III. Biosynthesis of Lipids 161 IV. Regulation of Lipid Biosynthesis 170 V. Conclusions and Future Perspectives 172 Acknowledgements 173 References Heterocyst Envelope Glycolipids Koichiro Awai, Sigal Lechno-Yossef and C. Peter Wolk Summary 179 I. Introduction 180 II. Chemical Structure of the Heterocyst Envelope Glycolipids 180 III. Physiological Role of the Heterocyst Envelope Glycolipids 181 IV. Deposition of the Heterocyst Envelope Glycolipids 183 V. Biosynthetic hgl Genes; Predicted Heterocyst Envelope Glycolipid Biosynthetic Pathway 184 VI. Regulation of Heterocyst Envelope Glycolipid Biosynthesis 186 VII. The Organization of hgl Genes Resembles that of Polyunsaturated Fatty Acid-Biosynthetic Genes and Has Been Found in Only One Cyanobacterium that Does Not Form Heterocysts 188 VIII. Are Heterocyst Envelope Glycolipid Biosynthetic Domains Distinguishable from those of Other Polyketide Synthases? 190 IX. Perspectives 194 Acknowledgements 198 References 198 xvii

5 10 Lipids in the Structure of Photosystem 1, Photosystem Hand the Cytochrome b 6f Complex Jan Kern, Athina Zouni, Albert Guskov and Norbert Krau ß Summary Introduction 204 II. Lipids Localized in the Structural Models 208 III. Functions of Lipids 225 IV. Conclusions and Perspectives 237 Acknowledgements 237 References The Role of Phosphatidylglycerol in Photosynthesis Hajime Wada and Naoki Mizusawa Summary Introduction 244 II. Biosynthesis of Phosphatidylglycerol 245 III. Phosphatidylglycerol in Photosynthetic Membranes 247 IV. The Role of Phosphatidylglycerol in Photosynthesis 250 V. The Role of Phosphatidylglycerol in Non-Photosynthetic Processes 256 VI. Conclusions and Future Perspectives 258 Acknowledgements 258 References The Role of Glycolipids in Photosynthesis Peter Dörmann and Georg Hölzl Summary 265 I. Introduction 266 II. Glycoglycerolipids in Photosynthetic Membranes 266 III. Enzymes Involved in Glycoglycerolipid Biosynthesis 267 IV. The Role of MGDG, DGDG and SQDG in Oxygenic Photosynthesis 270 V. The Role of Glycoglycerolipids in Anoxygenic Photosynthesis 277 VI. Conclusions 278 References Role of Lipids in the Dynamics of Thylakoid Membranes Conrad W Mullineaux and Helmut Kirchhoff Summary Introduction 284 II. Methods for Measuring Thylakoid Membrane Dynamics 284 III. Lipid and Protein Mobility in Thylakoid Membranes 287 IV. Effects of Lipid Composition an Membrane Fluidity and Tolerance of Low Temperatures 291 V. Importance of Lipids for Creating Diffusion Space for Proteins and Plastoquinone 292 Acknowledgements 293 References 293 xviii

6 14 Architecture of Thylakoid Membrane Networks Reinat Nevo, Silvia G. Chuartzman, Onie Tsabari, Ziv Reich, Dana Charuvi and Eyal Shimoni Summary 295 I. Introduction 296 II. Techniques in Electron Microscopy of Plant Samples 297 III. Thylakoid Network Organization in Cyanobacteria and Algae 300 IV. Thylakoid Networks of Higher Plants 307 V. Concluding Remarks 319 Acknowledgements 320 References Regulatory Role of Membrane Fluidity in Gene Expression Dmitry A. Los and Vladislav V Zinchenko Summary 329 I. Introduction 330 II. Modulation of Membrane Fluidity 330 III. Perception of Membrane Rigidification 334 IV. Perception of Membrane Fluidization 339 V. Multifunctional Sensors 341 VI. Conclusions and Perspectives 342 Acknowledgements 343 References Lipid Trafficking in Plant Photosynthetic Cells Juliette Jouhet, Emmanuelle Dubots, Eric MareChal and Maryse A. Block Summary 349 I. Introduction 350 II. Lipid Trafficking lnvolved in Formation of Chloroplast Lipids 351 III. Lipid Trafficking Induced by Phosphate Deprivation 355 IV. Molecular Mechanisms Involved in Lipid Trafficking 357 V. Conclusions 367 Acknowledgements 367 References Regulatory Roles in Photosynthesis of Unsaturated Fatty Acids in Membrane Lipids Suleyman 1. Allakhverdiev, Dmitry A. Los and Norio Murata Summary 373 I. Introduction 374 II. Engineered Decreases in Unsaturation of Fatty Acids in Membrane Lipids in Synechocystis 375 III. Engineered Increases in Unsaturation of Fatty Acids in Membrane Lipids in Synechococcus 379 IV. Modulation of Unsaturation of Fatty Acids in Membrane Lipids in Higher Plants 380 xix

7 V. Conclusions 384 Acknowledgements 385 References Oxidation of Membrane Lipids and Functions of Oxylipins John Browse Summary 389 I. Introduction 390 II. Chemical and Enzymatic Pathways of Oxylipin Synthesis 390 III. The Synthesis and Function of Jasmonate in Higher Plants 393 IV. Conclusions and Perspectives 401 Acknowledgements 401 References Biosynthesis and Biotechnology of Seed Lipids Including Sterols, Carotenoids and Tocochromanols Knut Meyer and Anthony J. Kinney Summary 407 I. Introduction 408 II. Sterols 409 HI. Carotenoids 412 IV. Tocochromanols 418 V. Acyl Lipids 422 References Advanced Mass Spectrometry Methods for Analysis of Lipids from Photosynthetic Organisms Bettina Seiwert, Patrick Giavalisco and Lothar Willmitzer Summary 445 I. Introduction 446 II. Lipid Extraction 447 III. Chromatographic Methods for Lipid-Class Separation 447 IV. Mass-Spectrometry-Based Lipid Analysis 451 V. Qualification and Quantification 457 VI. Outlook 458 References 459 Index 463 xx

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