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Photophysics of Photosynthetic Pigment-Protein Complexes

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Non-Photochemical Quenching and Energy Dissipation in Plants, Algae and Cyanobacteria

Part of the book series: Advances in Photosynthesis and Respiration ((AIPH,volume 40))

Summary

Photosynthesis converts solar energy into energy of chemical bonds. This process is initiated when a photon of sunlight is absorbed by a photosynthetic pigment molecule, followed by a highly efficient transfer of the excitation energy, excitation trapping, and charge separation at the reaction center. The excited state dynamics initiated by light absorption are central to the primary reactions of photosynthesis. Unless successfully transferred away from the excited chromophore within the excitation lifetime, the excitation energy relaxes back to the electronic ground state, either via emission of a photon (radiative decay) or through various nonradiative processes. The photosynthetic machinery can control the nonradiative relaxation rate: it can increase it under stress conditions (e.g., high light) by adjusting electronic properties of chromophores as well as their interaction, or decrease it under optimal conditions reaching >90 % efficiency of energy transfer. Some background in photophysics is, therefore, needed to understand the mechanistic aspects of the initial events following photoexcitation of photosynthetic complexes. The goal of this chapter is to describe the excited states involved in photoreactions and to outline the physical basis of photophysical processes involved in photosynthesis. We introduce the principles of light absorption and the nature of electronic excited states and light-initiated dynamics in photosynthetic complexes. De-excitation pathways, rate constants, quantum yields and lifetimes of fluorescence, excitation energy transfer and related photophysics are discussed. In the concluding section, we present an overview of the mechanisms of non-photochemical quenching (NPQ) of chlorophyll fluorescence in terms of photophysics of the excited states of photosynthetic pigments.

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Abbreviations

A g , B u , E u :

Excited state symmetry;

a u , b g , e g :

Molecular orbital symmetry;

B, Qx, Qy :

Chlorophyll singlet states;

C n , S n , σ h , I, e :

Molecular symmetry operators;

Chl:

Chlorophyll;

EET:

Excitation energy transfer;

EM:

Electromagnetic;

FRET:

Förster resonance energy transfer;

HOMO:

Highest occupied molecular orbital;

IC:

Internal conversion;

ISC:

Intersystem crossing;

LUMO:

Lowest unoccupied molecular orbital;

LH2:

Light-harvesting complex 2 of purple bacteria;

LHC:

Light-harvesting complex;

LHCII:

Light-harvesting complex II, a major antenna of PS II in plants and green algae;

NPQ:

Non-photochemical quenching of chlorophyll fluorescence;

PS I, PS II:

Photosystem I, photosystem II;

PSU:

Photosynthetic unit;

RC:

Reaction center;

S0, S1, and S2 :

Ground, first excited and second excited singlet states of carotenoids, not to be confused with similar names for the states of the oxygen evolving complex, the 4Mn-Ca (water) complex;

TDC:

Transition density cube;

Φfl, Φ ET , Φ tr , Φ PQ , Φ NPQ :

Quantum efficiencies of fluorescence (fl), energy transfer (ET), trapping (tr), photochemical (PQ) and non-photochemical quenching (NPQ) respectively; note that Φ PQ is equivalent to Φ p used by other authors

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Acknowledgments

The authors thank R. van Grondelle and H. van Amerongen for their suggestions and critical comments. Authors also acknowledge the Natural Sciences and Engineering Research Council of Canada (NSERC) and the Defense Advanced Research Projects Agency (DARPA, QuBE) for financial support. Govindjee acknowledges the support and friendship of the photosynthesis community at large.

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Authors and Affiliations

  1. Department of Chemistry, University of Toronto, 80 St. George Street, Toronto, ON, M5S 3H6, Canada

    Evgeny E. Ostroumov & Yaser R. Khan

  2. Department of Chemistry, Princeton University, Washington Rd, Princeton, NJ, 08544, USA

    Gregory D. Scholes

  3. Department of Plant Biology, Department of Biochemistry and Center of Biophysics and Quantitative Biology, University of Illinois at Urbana-Champaign, Urbana, IL, 61801, USA

    Govindjee

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  1. Evgeny E. Ostroumov
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  2. Yaser R. Khan
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  3. Gregory D. Scholes
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  4. Govindjee
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Corresponding author

Correspondence to Gregory D. Scholes .

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Editors and Affiliations

  1. Ecology and Evolutionary Biology, University of Colorado, Boulder, Boulder, Colorado, USA

    Barbara Demmig-Adams

  2. Laboratory of Photosynthetic Membranes Biological Research Center, Institute of Plant Biology Hungarian Academy of Sciences, Szeged, Hungary

    Gyozo Garab

  3. Ecology and Evolutionary Biology, University of Colorado, Boulder, Colorado, USA

    William Adams III

  4. Department of Plant Biology 265 Morrill Hall, MC-116, University of Illinois at Urbana-Champaign, Urbana, Illinois, USA

    Govindjee

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Dedication

This chapter is dedicated to the late Professor Robert M. Clegg (July 18, 1945 — October 15, 2012) of the University of Illinois at Urbana-Champaign (UIUC). He was a pioneer of the physics of living cells, and a world leader in fluorescence lifetime-resolved imaging microscopy (FLIM) of biological systems. He was an authority on fluorescence spectroscopy and rapid kinetic methods as applied to biological systems. Bob was a great human being, an intellectual, a “living library” to his students and his colleagues, a “connoisseur” of creative literature (not only in English, but in French and German as well), a historian of science, a great teacher, a poet, a storyteller, a great chef of diverse cuisine for friends and family, and a friend to all with whom he associated. One of us (Govindjee) was fortunate to have worked with him since he came to UIUC; they used FLIM in studying photoprotection against excess light in the green alga Chlamydomonas reinhardtii, and in avocado, a higher plant.

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Ostroumov, E.E., Khan, Y.R., Scholes, G.D., Govindjee (2014). Photophysics of Photosynthetic Pigment-Protein Complexes. In: Demmig-Adams, B., Garab, G., Adams III, W., Govindjee, . (eds) Non-Photochemical Quenching and Energy Dissipation in Plants, Algae and Cyanobacteria. Advances in Photosynthesis and Respiration, vol 40. Springer, Dordrecht. https://doi.org/10.1007/978-94-017-9032-1_4

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