Photosynthesis: Structures, Mechanisms, and Applications (eBook)

Harvey J.M. Hou, born in 1962 in China, is a Professor in the Department of Physical Sciences at Alabama State University, Montgomery, Alabama, USA. He received his B.Sc. in Physical Chemistry in 1984 from Wuhan University, and completed his Ph.D. in Analytical Chemistry in 1993 at Peking University (Beijing, China) with Xiaoxia Gao. He had his postdoctoral training at Chinese Academy of Sciences with Peisung Tang and Tingyun Kuang, at Iowa State University with Parag Chitnis, and at the Rockefeller University with David Mauzerall. Since 1995, he has served as a faculty member at the Chinese Academy of Sciences, Gonzaga University, the University of Massachusetts at Dartmouth, and the Alabama State University. He began his research career in photosynthesis in 1993, working on photosystem II (PS II). In 1996, he visited the laboratory of Jacque Breton in France and studied the orientation of pigments in PS II. In the laboratory of Parag Chitnis, he examined the organization of PS I. Working with David Mauzerall, he systematically investigated the thermodynamics of electron transfer reactions in photosynthesis using pulsed photoacoustics. His work has uncovered a significant entropy change of reaction in PS I; further, he has demonstrated that the entropy change in PS I is dramatically different from that in PS II. Since he established his laboratory in 2002, he has maintained his long-term collaboration with David Mauzerall on the thermodynamics in cyanobacterial PS I and in Heliobacteria. In 2006, he began collaboration with Gary Brudvig at Yale University and Dunwei Wang at Boston College on artificial photosynthesis, and has developed a manganese/semiconductor system for solar energy storage. His research group has also investigated the responses of cyanobacteria and cranberry plants to environment. He co-chaired a symposium at the 15th International Congress of Photosynthesis Research, chaired the 28th Annual Eastern Regional Photosynthesis Conference, and he co-organized the 38th Annual Midwest/Southeast Photosynthesis Meeting.

Foreword 5
Preface 7
Contents 10
Contributors 13
About the Editors 17
1: Photosynthesis: Natural Nanomachines Toward Energy and Food Production 20
1.1 Introduction 20
1.2 Organization and Function of Photosynthetic Nanomachines 21
1.3 Structures and Mechanisms of the Water Oxidation Nanomachines 24
1.4 Applications of Artificial Photosynthetic Nanomechines 26
References 27
2: Structure and Function of the Reaction Centre - Light Harvesting 1 Core Complexes from Purple Photosynthetic Bacteria 29
2.1 Introduction 30
2.2 The Purple Photosynthetic Bacteria 30
2.3 Mechanism of Photosynthesis in the Purple Photosynthetic Bacteria 31
2.4 Building Block of the Core Complex of Purple Photosynthetic Bacteria 32
2.4.1 Pigment Molecules 32
2.4.2 LH1 Subunit, ?/beta-Polypeptide Pair 35
2.4.3 Reaction Centre 37
2.4.4 PufX Protein 37
2.5 Structural Diversity of the RC-LH1 Complexes 39
2.5.1 X-Ray Crystal Structure of the Core Complex from Rps. palustris 40
2.5.2 X-Ray Crystal Structure of the Core Complex from Rba. sphaeroides 40
2.5.3 X-Ray Crystal Structure of Core Complex from Tch. tepidum 43
References 45
3: Recombinant Light Harvesting Complexes: Views and Perspectives 50
3.1 Pigment-Protein Reconstitution: The Technique 51
3.2 Achievements Obtained by Using Recombinant Complexes 54
3.3 Recombinant LHCII Proteins for the Study of NPQ 56
3.4 Future of NPQ Research 59
3.5 Biology Serving Nanotechnology: Applications of Reconstitution Technique 61
3.6 Concluding Summary 63
References 63
4: Alternative Electron Acceptors for Photosystem II 67
4.1 Introduction 68
4.1.1 The Structure and Function of Photosystem II 68
4.2 Investigating the Reductase Activity of Photosystem II 70
4.2.1 The Hill Reaction 70
4.2.2 Native Electron Acceptors in the Plastoquinone B Pocket 70
4.2.2.1 Quinone Analogs: Artificial Electron Acceptors from the QB Site 70
4.2.2.2 Herbicides that Inhibit at the QB Binding Site 71
4.3 DCMU-Insensitive Electron Acceptors 71
4.3.1 Ferricyanide 71
4.3.2 Hg2+ 72
4.3.3 Silicomolybdate 73
4.3.4 PSII Mutagenesis to Redirect Electron Flow 73
4.3.5 Designer Electron Acceptors - Co[(terpy)2]3+ 73
4.3.6 Electrodes 73
4.4 Factors Influencing PSII Reductase Activity and Redox Potential of QA 74
4.4.1 Herbicides 74
4.4.2 Quinone Pocket Hydrogen-Bonding Interactions 74
4.4.3 D1 Isoforms 75
4.4.4 OEC Perturbations 75
4.4.4.1 Challenging the ``Long-Range Effect´´ of OEC Perturbations 76
4.5 Factors Influencing Electron Transfer from QA to QB 76
4.5.1 Non-Heme Iron and Bicarbonate 76
4.5.2 Additional Quinones and Cytochrome b559 77
4.5.3 Stromal Side Lipids 77
4.5.3.1 Phosphatidylglycerol 77
4.5.4 Small Transmembrane Polypeptides 78
4.6 Conclusion 78
References 79
5: The Cl- Requirement for Oxygen Evolution by Photosystem II Explored Using Enzyme Kinetics and EPR Spectroscopy 83
5.1 Introduction 83
5.2 Chloride in PSII 84
5.2.1 Early Studies of the Cl- Dependence in O2 Evolution 84
5.2.2 Cl- Depletion of PSII Preparations from Higher Plants 85
5.2.3 S2 State EPR Signals and Chloride 86
5.2.4 XRD and EXAFS Structural Characterizations of Cl- Sites 88
5.3 Activators 90
5.3.1 Studies of Cl- Activation in Cyanobacteria 90
5.3.2 Quantifying the Affinity of Cl- Binding Sites in Higher Plant PSII 92
5.3.3 Bromide and Nitrate Activators 95
5.4 Inhibitors 96
5.4.1 Ammonia 96
5.4.2 Fluoride 98
5.4.3 Acetate 99
5.4.4 Azide 101
5.4.5 Iodide 102
5.4.6 Nitrite 104
5.5 Concluding Remarks 105
References 105
6: Vectorial Charge Transfer Reactions in the Protein-Pigment Complex of Photosystem II 112
6.1 Structural Peculiarities of Photosystem II 113
6.2 Functioning of Photosystem II 113
6.2.1 Photochemical Processes 114
6.2.2 The Catalytic Cycle of Water Oxidation 115
6.2.3 Reduction of Tyrosine Cation Radical YZ 118
6.2.4 Lipophilic Electron Donors 118
6.2.5 Synthetic Mn-Containing Complex 119
6.2.6 Hydrophilic Electron Donors 119
6.2.7 Mechanism of Electron Transfer 119
6.2.8 Vectorial Charge Transfer Reactions in Photoactivated Apo-OEC-PS II Core Complexes 121
6.3 Concluding Remarks 122
References 122
7: Function and Structure of Cyanobacterial Photosystem I 125
7.1 Introduction 126
7.2 Peripheral Subunits of PS I 128
7.2.1 PsaC 129
7.2.2 PsaD 130
7.2.3 PsaE 133
7.3 Integral Membrane Subunits 135
7.3.1 PsaA and PsaB 135
7.3.1.1 Introduction 135
7.3.1.2 Antenna Systems 139
Antenna Chlorophylls 139
Antenna Carotenoids 143
7.3.1.3 Lipids 144
7.3.1.4 Electron Transfer Chain 145
P700, The Primary Electron Donor 146
Accessory Chlorophylls 150
A0, The Primary Electron Acceptor 152
A1, The Second Electron Acceptor 154
FX, The First FeS Cluster 156
7.3.1.5 Docking of Plastocyanin/Cytochrome c6 158
7.3.2 PsaF 159
7.3.3 PsaI 162
7.3.4 PsaJ 164
7.3.5 PsaK 165
7.3.6 PsaL 167
7.3.7 PsaM 169
7.3.8 PsaU or PsaX 170
7.4 Concluding Remarks 172
References 173
8: How Light-Harvesting and Energy-Transfer Processes Are Modified Under Different Light Conditions: STUDIES by Time-Resolved ... 183
8.1 Introduction 183
8.2 Method 186
8.2.1 Time-Resolved Fluorescence Spectroscopy 186
8.2.2 Global Fitting Analysis 186
8.3 Modification of Energy Transfer 186
8.3.1 Excitation Energy Regulation in Phycobilisomes 186
8.3.2 Modification of Energy Transfer from Phycobilisomes in the Cyanobacterium Arthrospira platensis 189
8.3.3 Excitation Balance in Photochemical Reaction Centers in Red Algae 190
8.3.4 Light Harvesting by Carotenoids in the Higher Plant Arabidopsis thaliana and Green Algae Codium fragile 192
8.4 Conclusions 194
References 195
9: Interaction of Glycine Betaine and Plant Hormones: Protection of the Photosynthetic Apparatus During Abiotic Stress 199
9.1 Introduction 200
9.2 Biosynthesis and Accumulation of Glycine Betaine in Response to Abiotic Stress 201
9.3 Glycine Betaine Alleviates Abiotic Stress Effects on Crop Growth and Yield via an Interaction with Photosynthesis 202
9.4 Glycine Betaine Protects the Photosynthetic Apparatus from Abiotic Stress 205
9.5 Glycine Betaine Interaction with Reactive Oxygen Species Aids in the Protection of the Photosynthetic Apparatus 206
9.6 Glycine Betaine Interaction with Plant ``Stress´´ Hormones and Photosynthesis 206
9.7 Glycine Betaine Interactions with Plant ``Growth´´ Hormones 209
9.8 Modelling the Interactions Between Glycine Betaine, Plant Hormones, Photosynthesis and Reactive Oxygen Species 209
References 210
10: Photosynthetic Responses Under Harmful and Changing Environment: Practical Aspects in Crop Research 217
10.1 Introduction 218
10.2 Photosynthesis and Productivity in Crop Plants 218
10.2.1 Gas Exchange Analyses of Photosynthetic Limitations 221
10.2.2 Mesophyll Limitations to Photosynthesis 223
10.2.3 The Chlorophyll Fluorescence Methods in Crop Research 224
10.2.3.1 Saturation Pulse Method 225
10.2.3.2 Analysis of Fast Chlorophyll Fluorescence Induction 227
10.2.3.3 Chlorophyll Fluorescence Imaging 229
10.2.4 Photosynthetic Data in Context of Plant and Canopy 230
10.3 The Photosynthesis and Abiotic Stress Factors 232
10.3.1 Limitation of Photosynthesis by Drought 232
10.3.2 Limitation of Photosynthesis by High Temperature 238
10.4 Future Challenges: The Central Role of Plant Phenotyping 244
10.5 Concluding Remarks 247
References 248
11: Effects of Environmental Pollutants Polycyclic Aromatic Hydrocarbons (PAH) on Photosynthetic Processes 263
11.1 Introduction 263
11.2 Structure and Properties of PAH 264
11.3 Effects of PAHs on Plant Growth 265
11.4 Effects of PAHs on Photosynthesis 266
11.5 Effects of PAHs on Photosystem II Heterogeneity 267
11.6 Photo-Toxicity of PAH 267
11.7 Phytoremediation Methods 269
11.8 Conclusions and Future Prospective 270
References 271
12: Chlorophyll Fluorescence for High-Throughput Screening of Plants During Abiotic Stress, Aging, and Genetic Perturbation 274
12.1 Introduction 274
12.2 The Basic Principle of Chl Fluorescence 275
12.3 Chlorophyll Fluorescence as a Tool to Study Abiotic Stresses 276
12.4 Chlorophyll Fluorescence as an Approach to Investigate the Aging Process 279
12.5 Chlorophyll Fluorescence as a Biomarker in Large-Scale Phenomics Studies 281
12.6 The Application of Chl Fluorescence in Forestry and Crop Management 282
12.7 Concluding Remarks 282
References 283
13: Adaptation to Low Temperature in a Photoautotrophic Antarctic Psychrophile, Chlamydomonas sp. UWO 241 287
13.1 Introduction 288
13.2 Ultrastructure and Cell Morphology 289
13.3 Membrane Fatty Acid Composition 289
13.4 Cold-Adapted Enzymes 290
13.5 Energy Metabolism 292
13.6 Photosynthesis at Low Temperatures 293
13.7 Acclimation to Light and Low Temperatures 294
13.8 Chlamydomonas sp. UWO241 297
13.8.1 Natural Habitat of UWO241 297
13.8.2 Growth of UWO241 297
13.8.2.1 Temperature 297
13.8.2.2 Salt 298
13.9 Adaptation of UWO241 to Low Temperature 298
13.10 Photosynthetic Electron Transport 300
13.10.1 Structure of PETC 300
13.10.2 Function of PETC 300
13.10.3 Acclimation to Temperature and Irradiance 301
13.10.4 Acclimation to Light Quality 302
13.10.5 State Transitions 303
13.10.6 Thylakoid Polypeptide Phosphorylation Profile 303
13.10.7 PsbP-Like Proteins 304
13.11 A Model of Cyclic Electron Flow in Chlamydomonas sp. UWO241 305
13.12 Conclusions 307
References 308
14: Nitric Oxide Mediated Effects on Chloroplasts 316
14.1 Introduction 317
14.2 Sources of Nitric Oxide in Plants 318
14.3 Effect of Nitric Oxide on Photosynthetic Pigment Dynamics 318
14.4 Interaction of Nitric Oxide with Oxygen Evolving Complex 319
14.5 Effect of Nitric Oxide on Photosynthetic Electron Transport 320
14.6 Effect of Nitric Oxide on the Photophosphorylation in Chloroplasts 322
14.7 Ameliorating Effect of Nitric Oxide on Photosynthetic Stress Responses 322
14.7.1 Nitric Oxide Under Osmotic Stress 322
14.7.2 Nitric Oxide Under Temperature Stress 323
14.7.3 Nitric Oxide Under High Light Stress and UV Radiation 323
14.7.4 Nitric Oxide Under Heavy Metals 324
14.7.5 Nitric Oxide Under Herbicides 325
14.8 Conclusion 326
References 327
15: Nanostructured Mn Oxide/Carboxylic Acid or Amine Functionalized Carbon Nanotubes as Water-Oxidizing Composites in Artifici... 332
15.1 Introduction 332
15.2 Experiments 333
15.3 Results and Discussion 335
15.4 Conclusions 340
References 341
16: Self-Healing in Nano-sized Manganese-Based Water-Oxidizing Catalysts 343
16.1 Introduction 343
16.2 Self-Healing in Water-Oxidizing Catalysts 344
16.3 Manganese Based Water-Oxidizing Catalyst 346
References 350
17: A Robust PS II Mimic: Using Manganese/Tungsten Oxide Nanostructures for Photo Water Splitting 352
17.1 Introduction 352
17.2 Manganese Oxo Dimer and Manganese Oxo Oligomer 353
17.3 Design, Synthesis, and Structural Characterization of Manganese/Tungsten Oxide Nanostructure 357
17.4 Catalytic Activity of Manganese/Tungsten Oxide Nanostructure 359
17.5 Mechanism of the Manganese/Tungsten Oxide System 362
17.6 Conclusions 364
References 365
18: Time-Resolved EPR in Artificial Photosynthesis 368
18.1 Introduction 369
18.1.1 Transient Paramagnetic Species in Natural Photosynthesis 369
18.1.2 Differences Between Natural and Artificial Photosynthesis 369
18.2 Time-Resolved EPR Methods 371
18.2.1 Transient EPR 371
18.2.2 Pulsed EPR 371
18.3 Quasi-Static Polarization Patterns 372
18.3.1 Spin-Polarized, Weakly-Coupled Radical Pairs 373
18.3.2 Strongly-Coupled Radical Pairs 377
18.3.3 Sequential Radical Pairs 379
18.3.4 Triplet States 381
18.4 Time Dependent Effects 383
18.4.1 Electron Transfer 383
18.4.2 Quantum Beats 384
18.4.3 Out of Phase Echo Modulation 384
18.5 Recent Results 385
18.5.1 Early Results on Donor-Acceptor Complexes 385
18.5.2 Sequential Electron Transfer in Triads 386
18.5.3 Quantum Beats 387
18.5.4 Echo Modulations 388
18.5.5 Polymer-Fullerene Blends 389
18.6 Concluding Remarks 391
References 391
19: Artificial Photosynthesis Based on 1,10-Phenanthroline Complexes 397
19.1 Introduction 397
19.1.1 Aim and Scope 397
19.1.2 Dyes in DSSCs 400
19.1.3 Electrolytes in DSSCs 401
19.2 1,10-Phenanthroline-Based Electrolytes in DSSCs 403
19.3 Concluding Remarks 409
References 411
20: Concluding Remarks and Future Perspectives: Looking Back and Moving Forward 414
20.1 Looking Back and Moving Forward 414
References 418
Index 422