Summary
This chapter reviews our current knowledge on the reactions leading to light-induced water splitting into molecular oxygen and hydrogen, bound in the form of plastoquinol. This process takes place in the multimeric protein complex of Photosystem II (PS II). After a brief description of the basic principles of biological solar energy exploitation through photosynthesis, general features of the kinetics, energetics and the structural array and nature of the cofactors of PS II are presented. The overall reaction pattern of the water:plastoquinone oxidoreductase comprises three types of sequences: (a) light-induced charge separation leading to the “stabilized” radical pair \( \text{P68}{0}^{+•}{\text{Q}}_{\text{A}}^{-•}\); (b) oxidative water splitting into molecular oxygen and four protons released into the lumen, driven by P680+• as oxidant and with tyrosine \( {\text{Y}}_{\text{Z}}\) involved as intermediate, and (c) two-step reduction of plastoquinone to quinol under uptake of two protons from the cytoplasm/stroma by the use of \( {\text{Q}}_{\text{A}}^{-•}\) as reductant. Evidence is presented that within the (Chl a)4 \( Phe{o}_{x}\) (x = 0, 1 or 2) unit, which constitutes the photoactive pigment P680, the lowest singlet exciton state predominantly located on Chl a D1 acts as electron donor for the primary charge separation and that the exceptionally high reduction potential of P680 mainly originates from a specific hydrophobic protein microenvironment. The key step of oxidative water splitting is the O-O bond formation which is shown to occur most likely at the level of a binuclearly complexed peroxide. It is emphasized that the protein matrix plays a key role in tuning the energetics and kinetics. Examples are presented for effects of protein relaxation and flexibility on the reaction pattern of PS II.
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Notes
- 1.
The term ‘‘proton gradient’’ is often used in an incorrect manner. A gradient of protons, by definition, is ∇[H+ (r→)] where ∇ is a vector operator (derivative to space coordinates) and[H+ (r→)] the scalar field of the local proton concentration at position r→. In fact the vector ∇[H+ (r→)] is the driving force for a vectorial process; however, the actual value of this space dependent quantity is very difficult to determine in biological membrane systems and requires detailed calculations. Therefore only the difference of the trans-membrane electrochemical potential of protons D m∼[H+] is known and this parameter should not be misleadingly referred to as “proton gradient."
- 2.
D1/D2/Cyt \( {b}_{559}\) preparations were first isolated by Nanba and Satoh (1987) and shown to enable light-induced P680+• Pheo-• formation. This reaction takes place with a high quantum yield (Kurreck et al., 1997a). However, these preparations which are completely deprived of \( {\text{Q}}_{\text{A}}\), \( {\text{Q}}_{\text{B}}\)and the NHFe (Kurreck et al., 1997b) cannot stabilize the primary charge separation. Therefore these samples are not an analogue of the isolated reaction centers from anoxygenic purple bacteria (PBRC) (for reviews, see Lancaster, 2008, Parson, 2008) and should not be termed “PS II reaction centers” although this terminology is widely used in the literature.
Abbreviations
- Ant:
-
antenna complexes of PS II;
- BChl, Bphe:
-
bacteriochlorophyll, bacteriopheophytin;
- Chl:
-
chlorophyll;
- \( {\text{Chl}}_{\text{D1}}\), \( {\text{Chl}}_{\text{D2}}\) :
-
– “monomeric chlorophylls” of the D1- and D2-branches, respectively of the RC;
- CP:
-
chlorophyll-binding proteins;
- CP43, CP47:
-
core antenna subunits of PS II;
- Cyt \( {b}_{559}\) :
-
– cytochrome \( {b}_{559}\)
- Cyt \( {b}_{6}f\) :
-
– cytochrome \( {b}_{6}f\) complex;
- DFT:
-
density functional theory;
- EET:
-
excitation energy transfer;
- \( {\text{E}}_{\text{m}}\) :
-
– midpoint redox potential;
- ENDOR:
-
electron nuclear double resonance;
- ERPE:
-
exciton-radical pair equilibrium;
- EPR:
-
electron paramagnetic resonance;
- ESSEM:
-
Electron spin echo envelope modulation;
- ETC:
-
electron transfer chain;
- EXAFS:
-
extended X-ray absorption fine structure;
- FTIR:
-
Fourier Transform Infrared;
- LHCII –:
-
light-harvesting complex II;
- \( {\text{M}}_{\text{j}}{\text{L}}_{\text{k}}{\text{W}}_{1}\) :
-
– detailed symbol for redox states \( {\text{S}}_{\text{i}}\)of the WOC;
- MLS:
-
EPR multiline signal for the \( {\text{S}}_{2}\) of the WOC;
- NET:
-
nonadiabatic electron transfer;
- NHFe:
-
non-heme iron center;
- PS II:
-
Photosystem II;
- PS II CC:
-
Photosystem II core complex;
- PBRC:
-
RC of purple bacteria;
- PG:
-
phosphatidylglycerol;
- \( \text{P68}{0}^{+•}\) :
-
– oxidized state of the RC pigments;
- \( {\text{P}}_{\text{D1}}\), \( {\text{P}}_{\text{D2}}\) :
-
– “special pair” chlorophylls of the D1- and D2-branches, respectively, of the RC;
- \( {\text{Pheo}}_{\text{D1}}\), \( {\text{Pheo}}_{\text{D2}}\) :
-
– pheophytins of the D1- and D2-branches, respectively, of the RC;
- PQ:
-
plastoquinone;
- QA, QB :
-
– plastoquinones of the RC;
- QENS:
-
quasielastic neutron scattering;
- RC:
-
reaction center;
- RC-PC:
-
reaction center pigment complex;
- \( {\text{(RC - PC)}}^{*}\), \( \text{P68}{0}^{*}\) :
-
– excited singlet state of RC pigments;
- WOC:
-
water oxidizing complex;
- \( {\text{Y}}_{\text{Z}}\), \( {\text{Y}}_{\text{D}}\) :
-
– redox active tyrosines of the D1- and D2-branches, respectively;
- XANES:
-
X-ray absorption near edge spectroscopy
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Acknowledgements
I am very grateful to J. Eaton-Rye for critical reading of the manuscript and competent help in editing. Likewise, many thanks to J. Kern for providing Fig. 17.2 and together with J. Messinger for fruitful discussions, J.-R. Shen for providing Fig. 17.6, P. Kühn for the electronic version of Figs. 17.4–17.6, 17.9, J. Pieper for Fig. 17.10 and S. Renger for Figs. 17.1, 17.3, 17.7 and 17.8. I also thank S. Nothing for competent typing of the manuscript. The financial support by Deutsche Forschungsgemeinschaft (Sfb 429, TP A1) is gratefully acknowledged.
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Renger, G. (2012). Photosynthetic Water Splitting: Apparatus and Mechanism. In: Eaton-Rye, J., Tripathy, B., Sharkey, T. (eds) Photosynthesis. Advances in Photosynthesis and Respiration, vol 34. Springer, Dordrecht. https://doi.org/10.1007/978-94-007-1579-0_17
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