Molecular Mechanisms of Photosynthesis

Molecular Mechanisms of Photosynthesis
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MOLECULAR MECHANISMS OF PHOTOSYNTHESIS <p><b>Rediscover the foremost introduction to molecular photosynthesis on the market today</b> <p>In the comprehensively revised Third Edition of <i>Molecular Mechanisms of Photosynthesis</i>, distinguished researcher and professor Robert E. Blankenship delivers a brand-new update to the most authoritative textbook on the subject of photosynthesis. In addition to thorough coverage of foundational topics in photosynthesis, the book discusses cutting-edge advances in research in this area, including new structures and new information about the mechanism of oxygen production. <p>The author also describes advancements in the understanding of the regulation of photosynthesis and the critical process of photoprotection, as well as newly discovered pigments and organisms that extend oxygenic photosynthesis deeper into the near infrared spectral region. <p>Readers will also benefit from the inclusion of a fulsome appendix that incorporates a detailed introduction to the physical basis of photosynthesis, including thermodynamics, kinetics, and spectroscopy. A companion website offers downloadable figures as PowerPoint slides ideal for teaching. The book also includes: <ul><li>Thorough introductions to the basic principles of photosynthetic energy storage, photosynthetic organisms and organelles, and the history and early development of photosynthesis</li><li>An expansive discussion of photosynthetic pigments, including their structure and spectroscopy</li><li>Explorations of antenna complexes, energy transfer processes, reaction centers, and electron transport pathways in anoxygenic phototrophs and oxygenic photosynthetic organisms</li><li>Comprehensive treatments of chemiosmotic coupling, ATP synthesis, and carbon metabolism</li><li>Authoritative discussions of the evolution of photosynthesis and artificial photosynthesis</li></ul> <p>Perfect for advanced undergraduate and beginning graduate students in biochemistry and biophysics, <i>Molecular Mechanisms of Photosynthesis</i> will also earn a place in the libraries of students studying plant biology and seeking a one-stop resource in the field of molecular photosynthesis.

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Robert E. Blankenship. Molecular Mechanisms of Photosynthesis

Table of Contents

List of Tables

List of Illustrations

Guide

Pages

Molecular Mechanisms of Photosynthesis

Introduction to the third edition

Acknowledgements

About the companion website

Chapter 1 The basic principles of photosynthetic energy storage. 1.1 What is photosynthesis?

1.2 Photosynthesis is a solar energy storage process

1.3 Where photosynthesis takes place

1.4 The four phases of energy storage in photosynthesis

1.4.1 Antennas and energy transfer processes

1.4.2 Primary electron transfer in reaction centers

1.4.3 Stabilization by secondary reactions

1.4.4 Synthesis and export of stable products

References

Chapter 2 Photosynthetic organisms and organelles. 2.1 Introduction

2.2 Classification of life

2.2.1 Nomenclature

2.3 Prokaryotes and eukaryotes

2.4 Metabolic patterns among living things

2.5 Phototrophic prokaryotes

2.5.1 Purple bacteria

2.5.2 Green sulfur bacteria

2.5.3 Filamentous anoxygenic phototrophs

2.5.4 Heliobacteria

2.5.5 Chloracidobacteria

2.5.6 Gemmatimonadetes

2.5.7 Cyanobacteria

2.6 Photosynthetic eukaryotes

2.6.1 Algae

2.6.2 Plants

References

Chapter 3 History and early development of photosynthesis

3.1 Van Helmont and the willow tree

3.2 Carl Scheele, Joseph Priestley, and the discovery of oxygen

3.3 Ingenhousz and the role of light in photosynthesis

3.4 Senebier and the role of carbon dioxide

3.5 De Saussure and the participation of water

3.6 The equation of photosynthesis

3.6.1 The balanced equation for photosynthesis

3.7 Early mechanistic ideas of photosynthesis

3.7.1 Van Niel and the redox nature of photosynthesis

3.7.2 The Hill reaction: separation of oxidation and reduction reactions

3.8 The Emerson and Arnold experiments

3.9 The controversy over the quantum requirement of photosynthesis

3.10 The red drop and the Emerson enhancement effect

3.11 Antagonistic effects

3.12 Early formulations of the Z scheme for photosynthesis

3.13 ATP formation

3.14 Carbon fixation

References

Chapter 4 Photosynthetic pigments: structure and spectroscopy

4.1 Chemical structures and distribution of chlorophylls and bacteriochlorophylls

4.1.1 Chlorophyll a

4.1.2 Chlorophyll b

4.1.3 Chlorophyll c

4.1.4 Chlorophyll d

4.1.5 Chlorophyll e

4.1.6 Chlorophyll f

4.1.7 Bacteriochlorophyll a

4.1.8 Bacteriochlorophyll b

4.1.9 Bacteriochlorophylls c, d, e, and f

4.1.10 Bacteriochlorophyll g

4.2 Pheophytins and bacteriopheophytins

4.3 Chlorophyll biosynthesis

4.4 Spectroscopic properties of chlorophylls

4.5 Carotenoids

4.6 Bilins

References

Chapter 5 Antenna complexes and energy transfer processes. 5.1 General concepts of antennas and a bit of history

5.2 Why antennas?

5.3 Classes of antennas

5.4 Physical principles of antenna function

5.4.1 The funnel concept

5.4.2 Fluorescence analysis of antenna organization

5.4.3 Fluorescence excitation spectra – direct evidence for energy transfer

5.4.4 Förster theory of energy transfer

5.4.5 Exciton coupling

5.4.6 Relation of Förster transfer to exciton coupling

5.4.7 Quantum coherence effects in antenna systems

5.4.8 Trapping models – coupling of antennas to a reaction center

5.5 Structure and function of selected antenna complexes

5.5.1 Purple bacterial LH2 and LH1

5.5.2 Kinetics of energy transfer and trapping in purple bacteria

5.5.3 The LHCII complex from plants and algae

5.5.4 Phycobilisomes

5.5.5 Peridinin–chlorophyll protein

5.5.6 Chlorosome and FMO protein

5.6 Regulation of antennas

5.6.1 State transitions

5.6.2 Nonphotochemical quenching and the xanthophyll cycle

References

Chapter 6 Reaction centers and electron transport pathways in anoxygenic phototrophs

6.1 Basic principles of reaction center structure and function

6.2 Development of the reaction center concept

6.3 Purple bacterial reaction centers. 6.3.1 Structural features of purple bacterial membranes and reaction centers

6.3.2 Kinetics and pathways of electron transfer within purple bacterial reaction centers

6.4 Theoretical analysis of biological electron transfer reactions

6.5 Quinone reductions, the role of the Fe and pathways of proton uptake

6.6 Organization of electron transfer pathways

6.7 Completing the cycle – the cytochrome bc1 complex

6.7.1 The mechanism of electron and proton transfer in the cytochrome bc1 complex

6.8 Membrane organization in purple bacteria

6.8.1 Other electron transport pathways in purple bacteria

6.9 Electron transport in other anoxygenic phototrophic bacteria

References

Chapter 7 Reaction centers and electron transfer pathways in oxygenic photosynthetic organisms

7.1 Spatial distribution of electron transport components in thylakoids of oxygenic photosynthetic organisms

7.2 Noncyclic electron flow in oxygenic organisms

7.3 Photosystem II overall electron transfer pathway

7.4 Photosystem II forms a dimeric supercomplex in the thylakoid membrane

7.5 The oxygen‐evolving complex and the mechanism of water oxidation by Photosystem II

7.6 The structure and function of the cytochrome b6f complex

7.7 Plastocyanin donates electrons to Photosystem I

7.8 Photosystem I structure and electron transfer pathway

7.9 Ferredoxin and ferredoxin‐NADP reductase complete the noncyclic electron transport chain

7.9.1 Other roles of ferredoxin and cyclic electron transfer around Photosystem I

References

Chapter 8 Chemiosmotic coupling and ATP synthesis

8.1 Chemical aspects of ATP and the phosphoanhydride bonds

8.2 Historical perspective on ATP synthesis

8.3 Quantitative formulationof proton motive force

8.4 Nomenclature and cellular location of ATP synthase

8.5 Structure of ATP synthase

8.6 The mechanism of chemiosmotic coupling

References

Chapter 9 Carbon metabolism

9.1 The Calvin–Benson cycle is the primary photosynthetic carbon fixation pathway

9.1.1 The three phases of the Calvin–Benson cycle are carboxylation, reduction, and regeneration

9.1.2 Rubisco is the carboxylation enzyme

9.1.3 Details of the other Calvin–Benson cycle reactions

9.1.4 The stoichiometry and efficiency of the Calvin–Benson cycle reactions

9.1.5 The Calvin–Benson cycle is regulated by disulfide reduction of key enzymes and ion movements

9.2 Photorespiration is a wasteful competitive process to carboxylation

9.3 The C4 carbon cycle minimizes photorespiration

9.4 Crassulacean acid metabolism avoids water loss in plants

9.5 Algae and cyanobacteria actively concentrate CO2

9.6 Sucrose and starch synthesis

9.7 Other carbon fixation pathways in anoxygenic phototrophs

References

Chapter 10 Genetics, assembly, and regulation of photosynthetic systems

10.1 Gene organization in anoxygenic photosynthetic bacteria

10.2 Gene expression and regulation of purple photosynthetic bacteria

10.3 Gene organization in cyanobacteria

10.4 Chloroplast genomes

10.5 Pathways and mechanisms of protein import and targeting in chloroplasts

10.6 Gene regulation and the assembly of photosynthetic complexes in cyanobacteria and chloroplasts

10.7 The regulation of oligomeric protein stoichiometry

10.8 Assembly, photodamage, and repair of Photosystem II

References

Chapter 11 The use of chlorophyll fluorescence to probe photosynthesis

11.1 The time course of chlorophyll fluorescence

11.2 The use of fluorescence to determine the quantum yield of Photosystem II

11.3 Fluorescence detection of nonphotochemical quenching

11.4 The physical basis of variable fluorescence

References

Chapter 12 Origin and evolution of photosynthesis. 12.1 Introduction

12.2 Early history of the Earth

12.3 Origin and early evolution of life

12.4 Geological evidence for life and photosynthesis

12.5 The nature of the earliest photosynthetic systems

12.6 The origin and evolution of metabolic pathways with special reference to chlorophyll biosynthesis

12.7 Origin and evolution of photosynthetic pigments

12.8 Evolutionary relationships among reaction centers and other electron transport components

12.9 Do all photosynthetic reaction centers derive from a common ancestor?

12.10 The origin of linked photosystems and oxygen evolution

12.11 Origin of the oxygen‐evolving complex and the transition to oxygenic photosynthesis

12.12 Antenna systems have multiple evolutionary origins

12.13 Endosymbiosis and the origin of chloroplasts

12.14 Most types of algae are the result of secondary endosymbiosis

12.15 Following endosymbiosis, many genes were transferred to the nucleus, and proteins were reimported to the chloroplast

12.16 Evolution of carbon metabolism pathways

References

Chapter 13 Bioenergy applications and artificial photosynthesis. 13.1 Introduction

13.2 Solar energy conversion

13.3 What is the efficiency of natural photosynthesis?

13.3.1 Different types of efficiency of photosynthesis

13.4 Calculation of the energy storage efficiency of oxygenic photosynthesis

13.5 Why is the efficiency of photosynthesis so low?

13.6 How might the efficiency of photosynthesis be improved?

13.7 Artificial photosynthesis

13.7.1 Artificial photosynthetic systems that mimic reaction centers

13.7.2 Artificial antenna systems

13.7.3 Artificial water‐splitting systems

13.7.4 Semiconductor‐based artificial photosynthetic systems

13.7.5 Multiple junction artificial photosynthetic systems

13.7.6 Concluding thoughts on artificial photosynthesis

References

Appendix 1 Light, energy, and kinetics

A.1 Light

A.2 Thermodynamics

A.2.1 Thermodynamic functions: work, heat, internal energy, enthalpy, entropy, and free energy

A.2.2 The first law of thermodynamics

A.2.3 Entropy and the second law of thermodynamics

A.2.4 Gibbs free energy

A.2.5 Chemical potentials and equilibrium constants

A.3 Boltzmann distribution

A.4 Electrochemistry: reduction–oxidation reactions

A.4.1 Measurement of midpoint potentials

A.5 Chemical kinetics

A.5.1 First‐order kinetics

A.5.2 Parallel first‐order reactions

A.5.3 Sequential first‐order reactions

A.5.4 Temperature dependence of reaction rates

A.6 Quantum versus classical mechanics

A.7 Quantum mechanics – the basic ideas

A.7.1 Quantum mechanical formalism

A.7.2 The Heisenberg uncertainty principle

A.8 Molecular energy levels and spectroscopy. A.8.1 Absorption and fluorescence

A.8.2 Molecular potential energy curves

A.8.3 Harmonic oscillator

A.8.4 Molecular electronic energy levels: singlets and triplets

A.8.5 Electronic transitions

A.8.6 Absorption intensity, oscillator strength, and transition dipole moments

A.8.7 Einstein coefficients

A.8.8 Fluorescence quantum yield and lifetime

A.9 Practical aspects of spectroscopy. A.9.1 Beer–Lambert law of absorption

A.9.2 Absorption measurements

A.9.3 Difference spectroscopy and flash photolysis

A.9.4 Fluorescence measurements

A.9.5 Fluorescence lifetime measurements

A.9.6 Fluorescence polarization and anisotropy

A.10 Photochemistry

A.10.1 Excited state redox potentials

References

Further reading

Further advanced reading

Index

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Third Edition

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Figure 1.5 Schematic diagram of the noncyclic electron transfer pathway found in oxygenic photosynthetic organisms. The upper diagram (a) is an energetic picture of the electron transport pathway, incorporating the major reactions of photosynthesis into what is called the Z‐scheme of photosynthesis. The lower diagram (b) is a spatial picture, showing the major protein complexes whose energetics are shown in the Z‐scheme, and how they are arranged in the photosynthetic membrane. Neither view alone gives a complete picture, but together they summarize much information about photosynthetic energy storage.

Source: (a) Hohmann‐Marriott and Blankenship (2011) (p.532)/Annual Reviews. Reproduced with permission of Annual Reviews. (b) Courtesy of Dr. Jonathan Nield.

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