Xanthophyll Cycle-Dependent Energy Dissipation and Flexible Photosystem II Efficiency in Plants Acclimated to Light Stress

Abstract
The effect of an acclimation to light stress during the growth of leaves on their response to high photon flux densities (PFDs) was characterised by quantifying changes in photosystem II (PSII) characteristics and carotenoid composition. During brief experimental exposures to high PFDs sun leaves exhibited: (a) much higher levels of antheraxanthin + zeaxanthin than shade leaves, (b) a greater extent of energy dissipation in the light-harvesting antennae, and (c) a greater decrease of intrinsic PSII efficiency that was rapidly reversible. During longer experimental exposures to high PFD, deep-shade leaves but not the sun leaves showed slowly developing secondary decreases in intrinsic PSII efficiency. Recovery of these secondary responses was also slow and inhibited by lincomycin, an inhibitor of chloroplast-encoded protein synthesis. In contrast, under field conditions all changes in intrinsic PSII efficiency in open sun-exposed habitats as well as understory sites with intense sunflecks appeared to be caused by xanthophyll cycle-dependent energy dissipation. Furthermore, comparison of leaves with different maximal rates of electron transport revealed that all leaves compensated fully for these differences by dissipating very different amounts of absorbed light via xanthophyll cycle-dependent energy dissipation, thereby all maintaining a similarly low PSII reduction state. It is our conclusion that an increased capacity for xanthophyll cycle-dependent energy dissipation is a key component of the acclimation of leaves to a variety of different forms of light stress, and that the response of leaves to excess light experienced in the growth environment is thus likely to be qualitatively different from that to sudden experimental exposures to PFDs exceeding the growth PFD.