Photosynthesis: Solar Energy for Life

205 Pages·2018·16.505 MB·English

by  Dmitry Shevela, Lars Olof Björn, Govindjee

10522_9789F81M32.i2n3d10d3 _tpP.inadgde 1i 11/10/18 4:42 PM
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World Scientific Publishing Co. Pte. Ltd.
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Library of Congress Cataloging-in-Publication Data
Names: Shevela, Dmitry, 1979– author. | Björn, Lars Olof, 1936– author. |
Govindjee, 1932– author.
Title: Photosynthesis : solar energy for life / by Dmitry Shevela (Umeå University, Sweden),
Lars Olof Björn (Lund University, Sweden),
Govindjee (University of Illinois at Urbana-Champaign, USA).
Description: New Jersey : World Scientific, 2018. | Includes bibliographical references and index.
Identifiers: LCCN 2018044556 | ISBN 9789813223103 (hardcover : alk. paper)
Subjects: LCSH: Photosynthesis.
Classification: LCC QK882 .S49488 2018 | DDC 572/.46--dc23
LC record available at https://lccn.loc.gov/2018044556
British Library Cataloguing-in-Publication Data
A catalogue record for this book is available from the British Library.
Cover design: Dmitry Shevela (Umeå University).
Front cover background image: Sunset Over the Gulf of Mexico. Image credit: NASA/Terry Virts;
Editor: Sarah Loff. Image credits for front cover foreground photos: Dr Joanna Porankiewcz Asplund
(Agrisera, Sweden) for the picture of the leaf; and Dr Natalia Voronkina (Kaluga State University,
Russia) for the microscopy picture of cells in the Sphagnum leaf.
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Contents
Preface ix
Acknowledgements xiii
About the Authors xv
Chapter 1: Introduction 1
1.1 Why Study Photosynthesis? 1
1.2 History 2
1.2.1 The discovery of photosynthesis 2
1.2.2 The concept of energy 5
1.2.3 Early research on photosynthesis 7
1.2.3.1 What absorbs the light for photosynthesis? 7
1.2.3.2 What goes in, and what comes out? 8
1.2.3.3 Which atoms go where? Is it possible
to answer this question? 10
1.2.3.4 How much light is required to drive
photosynthesis? 13
References 14
Chapter 2: The Photosynthetic Apparatus 17
2.1 Introduction 17
2.2 Chloroplasts 17
2.3 Thylakoid Membrane and Photosynthetic
Protein Complexes 20
2.4 Pigments 23
2.5 Lipids and Proteins 23
2.6 Evolution 26
References 26
Chapter 3: Basics of Photosynthesis: Light-Dependent Reactions 29
3.1 Overview: Harvesting Sunlight to Drive
Redox Chemistry 29
3.2 Capturing the Energy of Light 29
3.3 Conversion of Light Energy into Chemical Energy 36
v
Contents
3.3.1 Primary photochemistry 36
3.3.2 Electron-transfer pathways 41
3.3.2.1 The “Z-scheme” of photosynthesis 41
3.3.2.2 Photosynthetic water oxidation
(oxygen evolution) 43
3.3.2.3 Unique role of bicarbonate in light-induced
reactions of PSII 44
3.3.2.4 Formation of reducing power 45
3.3.3 Proton-transfer pathways and formation
of ATP (photophosphorylation) 46
References 51
Chapter 4: Basics of Photosynthesis: The Carbon Reactions 59
4.1 C3 Photosynthesis: The Calvin-Benson Cycle 59
4.2 C4 Photosynthesis: The Hatch−Slack Pathway 63
4.3 Crassulacean Acid Metabolism 69
4.4 Translocation and Biomass Production 71
4.5 Alternate Pathways and Evolution 74
References 75
Chapter 5: Regulation of Photosynthesis 81
5.1 Background 81
5.2 State Transitions 82
5.3 Safety Valve: Dissipation of Electronic Energy
as Heat (“Quenching”) 86
5.4 Xanthophyll Cycles 87
5.5 Reaction Centers as Sinks for Excess
Excitation Energy 90
5.6 Quenching of Chlorophyll Triplets 93
5.7 Regulation of RuBisCO 94
5.8 Redox Regulation of Certain Calvin-Benson
Cycle Enzymes by Thioredoxin 95
5.9 Other Regulatory Mechanisms Related
to Photosynthesis 96
vi
Contents
5.10 Leaf Movement 97
5.11 Concluding Remarks 97
References 98
Chapter 6: Photosynthesis and Our Planet 107
6.1 Oxygenation of the Earth’s Atmosphere 107
6.2 Protection: Ozone Was Formed From Oxygen
and It Protects Us From the UV Radiation 108
6.3 Earth Temperature Over Time and the Effect
of the Biosphere on It 110
6.3.1 Influence of the Sun 110
6.3.2 Influence of the Earth’s atmosphere 111
6.4 Conclusion 115
References 117
Chapter 7: Anoxygenic Photosynthesis 123
7.1 Introduction 123
7.2 Anoxygenic Photosynthetic Organisms:
Their Reaction Centers and Pathways
for Carbon Assimilation 125
7.3 Relation between the RCs of Anoxygenic
and Oxygenic Photosynthetic Organisms 127
7.4 Photosynthetic N Assimilation 128
2
7.5 Rhodopsin-Based Phototrophy 131
References 132
Chapter 8: The Past, Present and the Future 135
8.1 Spread of Photosynthesis by Successive Endosymbiosis 135
8.2 Adapting Photosynthesis that has Evolved Underwater
for Life on Land 140
8.2.1 Grana 141
8.2.2 Hormones 142
8.2.3 Stomata 143
8.2.4 Reaching for light 145
vii
Contents
8.2.4.1 Evolution of trees 145
8.2.4.2 Cheating: Lianas (Climbers) and Epiphytes 145
8.3 The Role of Fungi 145
8.4 Making More Photosynthesis… More Biomass,
More Bioenergy, New Chemicals and Hydrogen 146
8.4.1 General: Classical breeding and genetic engineering 146
8.4.2 Conversion from C3 to C4 metabolism by genetic
engineering 146
8.4.3 Cyanobacterial genes in higher plants 147
8.4.4 Improvement of RuBisCO 148
8.4.5 Photorespiratory bypass 149
8.4.6 Photosynthetic H production 149
2
8.4.7 Biofuels and other chemicals 152
8.4.8 Other improvements 152
References 153
Chapter 9: The Ultimate: Artificial Photosynthesis 165
9.1 Objectives and Approaches 165
9.2 Water Oxidation Coupled to Hydrogen Production:
in Principle Simple, in Practice Not So Easy 166
9.3 Reduction of Carbon Dioxide 172
9.4 Conclusions 176
References 177
Index 183
viii
Preface
Jules Verne (1828–1905) wrote in 1874: “Water will one day
be employed as fuel—the hydrogen and oxygen that constitute
it, used singly or together, will furnish an inexhaustible source of
heat and light”.
Why should we focus on Photosynthesis, which, by using sunlight, becomes
an inexhaustible source of energy for the planet? We hope that our book
“Photosynthesis: Solar energy for life” will answer that for you. Photo-
synthesis is the most central and the most important process on the Earth,
providing “order” from “disorder” using the all-abundant sunlight. Each
year, photosynthesis fixes about 100 gigatons of carbon, supporting all life
on our Planet, by providing us with oxygen to breathe and food to survive.
Our understanding of photosynthesis can now be said to have reached
encyclopedic dimensions; there have been, in the past, many good books
at various levels (Appendix A1). Our book is intended to fulfill the
needs of undergraduate and beginning graduate students in all branches
of biology as well as biochemistry, biophysics, and bioengineering.
Photosynthesis will be central to future advances in producing more
1 Appendix A
During 1945–1956, Eugene Rabinowitch (1898–1973), a post-doc student of the 1926 Nobel
laureate, James Franck (1882–1964), wrote a 2088 page book in three physical volumes
(Vol. I [1945], Vol. II [1951], Part 1, and Vol., Part 2 [1956]). It was indeed like a “Bible”
of this field (available free at http://www.life.illinois.edu/govindjee/g/Books.html) at that
time. If we were to write a similar book (almost impossible for most of us), it will take
at least 30,000 pages! For college and university students, books by the following authors
are worth mentioning: Robert Blankenship (Molecular Mechanisms of Photosynthesis, 2nd
ed., 2014), Paul G. Falkowski & John A. Raven (Aquatic Photosynthesis, 2nd ed., 2007),
Bacon Ke (Photosynthesis, 2003), David W. Lawlor (Photosynthesis, 2001), Eugene Rab-
inowitch & Govindjee (Photosynthesis, 1969; available free at http://www.life.illinois.edu/
govindjee/g/Books.html.), and David Hall & Krishna K. Rao (Photosynthesis, 1967–1999).
For advanced graduate and doctoral students, there are indeed many edited books by, e.g.,
Govindjee (Bioenergetics, 1975), Achim Trebst and Mordhay Avron (Encyclopedia of Plant
Physiology, 1976), several volumes in “Topics in Photosynthesis”, edited by James Barber
(1976–1992), and more than 40 volumes in “Advances in Photosynthesis and Respiration”
since 1994 (Govindjee and Thomas Sharkey, series editors; now Sharkey and Julian
Eaton-Rye; see: http://www.springer.com/series/5599?detailsPage=titles).
ix