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Relationships Between Antioxidant Metabolism and Carotenoids in the Regulation of Photosynthesis

  • Chapter
The Photochemistry of Carotenoids

Part of the book series: Advances in Photosynthesis and Respiration ((AIPH,volume 8))

Summary

The large driving forces produced in photosynthesis require precise control and regulation to prevent potentially destructive side reactions involving active oxygen species that damage pigments and proteins. Oxygen can be activated either by energy transfer, yielding singlet oxygen, or by reduction, yielding superoxide or hydrogen peroxide. At Photosystem I, the lifetime of the excited chlorophyll singlet state within the antenna pigment bed is short and little threat is posed by formation of highly reactive singlet oxygen. The situation is completely different in Photosystem II. Here, the lifetime of singlet chlorophyll is sufficiently long to allow significant formation of chlorophyll triplet states able to transfer energy to ground state triplet oxygen, generating singlet oxygen. Carotenoids intervene to control these processes in at least two important ways. First, pigments such as β-carotene are capable of directly quenching both triplet chlorophyll and singlet oxygen states (the so-called triplet valve mechanism). Second, the xanthophyll cycle is involved in lowering the yield of triplet chlorophyll formation by pre-emptive quenching of excited singlet state chlorophyll, a mechanism that can have a high quantum yield value for energy dissipation. The chief difference between these two mechanisms is that the xanthophyll cycle is inducible and subject to regulation, whereas the triplet valve pathway is constitutive and unregulated. Although formation of singlet oxygen must be avoided or controlled, chloroplasts have exploited the potential of oxygen chemistry to drive and regulate metabolism while minimizing the deleterious effects of uncontrolled interactions with oxygen. Hence, while the potential of the chloroplast as a source of oxidative stress is large, in reality this organelle offers minimal risk because of pre-emptive regulation and effective defense. The production of superoxide and hydrogen peroxide by the thylakoid membranes is limited by efficient control of electron transport. This regulation limits the potential for oxidative damage and prevents high rates of electron transport to oxygen. Effective antioxidant defense ensrures rapid elimination of active forms of oxygen further preventing oxidative damage.

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Abbreviations

A1:

secondary electron acceptor in rcI

AA:

ascorbic acid

AOS:

active oxygen species

APX:

ascorbate peroxidase

Ci:

intracellular CO2 concentrations

1Chl*:

chlorophyll singlet state

DHA:

dehydroascorbate

DHAR:

dehydroascorbate reductase

φ eλ :

yield of photochemistry by rcII withoxidized QA

F:

Faraday constant

Fd:

ferredoxin

Fm:

relative yield or intensity of chlorophyll fluorescence whenall QA is reduced in the dark-adapted state

F′m:

as Fm but under conditions of irradiation

FNR:

ferredoxin-NADP reductase

FO:

relative yield or intensity of fluorescence when all QA is oxidized

Fν:

relative change in fluorescence yield or intensity produced by completely reducing the QA pool in the dark-adapted state

Fν:

as Fν but under conditions of irradiation

FÇ:

third electron acceptor in rcI

FÇ/B:

fourth electron acceptor in rcI

GR:

glutathione reductase

GSH:

reduced glutathione

GSSG:

glutathione disulfide

JPSI:

index of PS I electron transport, the product of φPS I and irradiance

LHCII:

light-harvesting chlorophyll a/b-binding protein associated with PS II

MDHA:

monodehydroascorbate

MDHAR:

monodehydroascorbate reductase

NPQ:

non-photochemical quenching of variable chlorophyll a fluorescence

P680:

chlorophyll pair that forms the primary electron donor in the reaction center of PS II

P700:

chlorophyll pair that forms the primary electron donor in the reaction center of PS I

PQ:

plastoquinone

PS:

photosystem

φPS I:

quantum yield for PS I electron transport

φPS II:

quantum yield for PS II electron transport

QA:

primary stable electron acceptor toPS II

QB:

terminal electron acceptor of the PS II reaction center

qQ:

probability that an exciton in PS II will encounter a rcII with anoxidized QA

R:

universal gas constant (8.31 JK−1mol−1)

rcI:

reaction center of PS I

rcII:

reaction center of PS II

Rubisco:

ribulose-1,5-bisphosphate carboxylase/oxygenase

RuBP:

ribulose-1,5-bisphosphate

SOD:

superoxide dismutase

T:

absolute temperature

ΔμH+:

transthylakoid proton potential

VDE:

violaxanthin deepoxidase

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Authors and Affiliations

  1. Department of Biochemistry and Physiology, Institute of Arable Crops Research (IACR), Rothamsted, Harpenden, Herts, AL5 2JQ, UK

    Christine H. Foyer

  2. ATO/DLO, Bornsesteeg 59, Postbus 17, NL-6700 AA, Wageningen, The Netherlands

    Jeremy Harbinson

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  1. Christine H. Foyer
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  1. University of Connecticut, Storrs, CT, USA

    Harry A. Frank

  2. Liverpool John Moores University, Liverpool, UK

    Andrew J. Young

  3. University of Liverpool, Liverpool, UK

    George Britton

  4. University of Glasgow, Glasgow, UK

    Richard J. Cogdell

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Foyer, C.H., Harbinson, J. (1999). Relationships Between Antioxidant Metabolism and Carotenoids in the Regulation of Photosynthesis. In: Frank, H.A., Young, A.J., Britton, G., Cogdell, R.J. (eds) The Photochemistry of Carotenoids. Advances in Photosynthesis and Respiration, vol 8. Springer, Dordrecht. https://doi.org/10.1007/0-306-48209-6_17

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