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Summary: The Photochemical Reflectance Index (PRI) provides an optical indicator of photosynthetic light‐use efficiency, photoprotection, and stress in plants. Although PRI can be applied in remote sensing, its interpretation depends on irradiance, which is hard to obtain from satellite or airborne imagery.To quantify forest photoprotective responses remotely, we developed a framework for modeling and interpreting PRI‐light responses of individual trees and species using airborne imaging spectrometry coupled with georeferenced forest inventory data from a temperate broad‐leaved forest. We derived an irradiance proxy, used hierarchical modeling to analyze PRI‐light responses, and developed a framework of physiological interpretations of model parameters as facultative and constitutive components of photoprotection.Photochemical Reflectance Index declined with illumination, and PRI‐light relationships varied with landscape position and among tree crowns and species. More sun‐exposed foliage had lower intercepts and slopes of the relationship, indicating greater constitutive, but less facultative, photoprotection.We show that tree photoprotective strategies can be quantified at multiple scales using airborne hyperspectral data in structurally complex forests. Our findings and approach have important implications for the remote sensing of forest stress by offering a new way to assess functional diversity through dynamic differences in photoprotection and photosynthetic downregulation and providing previsual indicators of forest stress. [ABSTRACT FROM AUTHOR]

Summary: Xanthophyll cycles are broadly important in photoprotection, and the reversible de‐epoxidation of xanthophylls typically occurs in excess light conditions. However, as presented in this review, compiling evidence in a wide range of photosynthetic eukaryotes shows that xanthophyll de‐epoxidation also occurs under diverse abiotic stress conditions in darkness. Light‐driven photochemistry usually leads to the pH changes that activate de‐epoxidases (e.g. violaxanthin de‐epoxidase), but in darkness alternative electron transport pathways and luminal domains enriched in monogalactosyl diacyl glycerol (which enhance de‐epoxidase activity) likely enable de‐epoxidation. Another 'dark side' to sustaining xanthophyll de‐epoxidation is inactivation and/or degradation of epoxidases (e.g. zeaxanthin epoxidase). There are obvious benefits of such activity regarding stress tolerance, and indeed this phenomenon has only been reported in stressful conditions. However, more research is required to unravel the mechanisms and understand the physiological roles of dark‐induced formation of zeaxanthin. Notably, the de‐epoxidation of violaxanthin to antheraxanthin and zeaxanthin in darkness is still a frequently ignored process, perhaps because it questions a previous paradigm. With that in mind, this review seeks to shed some light on the dark side of xanthophyll de‐epoxidation, and point out areas for future work. [ABSTRACT FROM AUTHOR]

Summary: The xanthophyll cycle is the metabolic process by which the carotenoid violaxanthin is de‐epoxidated to zeaxanthin, a xanthophyll with a crucial photoprotective role in higher plants and mosses. The role of zeaxanthin is still unclear in green algae, and a peculiar violaxanthin de‐epoxidating enzyme was found in the model organism Chlamydomonas reinhardtii. Here, we investigated the molecular details and functions of the xanthophyll cycle in the case of Chlorella vulgaris, one of the green algae most considered for industrial cultivation, where resistance to high light stress is a prerequisite for sustainable biomass production.Identification of the violaxanthin de‐epoxidase enzyme in C. vulgaris was performed by genome mining and in vitro analysis of the catalytic activity of the gene product identified. The photoprotective role of zeaxanthin was then investigated in vivo and in isolated pigment‐binding complexes.The results obtained demonstrate the functioning, even though with a different pH sensitivity, of a plant‐like violaxanthin de‐epoxidase enzyme in C. vulgaris. Differently from C. reinhardtii, zeaxanthin accumulation in C. vulgaris was found to be crucial for photoprotective quenching of excitation energy harvested by both photosystem I and II.These findings demonstrate an evolutionary divergence of photoprotective mechanisms among Chlorophyta. [ABSTRACT FROM AUTHOR]

Diatoms contain a highly flexible capacity to dissipate excessively absorbed light by nonphotochemical fluorescence quenching ( NPQ) based on the light-induced conversion of diadinoxanthin (Dd) into diatoxanthin (Dt) and the presence of Lhcx proteins. Their NPQ fine regulation on the molecular level upon a shift to dynamic light conditions is unknown., We investigated the regulation of Dd + Dt amount, Lhcx gene and protein synthesis and NPQ capacity in the diatom Phaeodactylum tricornutum after a change from continuous low light to 3 d of sine ( SL) or fluctuating ( FL) light conditions. Four P. tricornutum strains with different NPQ capacities due to different expression of Lhcx1 were included., All strains responded to dynamic light comparably, independently of initial NPQ capacity. During SL, NPQ capacity was strongly enhanced due to a gradual increase of Lhcx2 and Dd + Dt amount. During FL, cells enhanced their NPQ capacity on the first day due to increased Dd + Dt, Lhcx2 and Lhcx3; already by the second day light acclimation was accomplished. While quenching efficiency of Dt was strongly lowered during SL conditions, it remained high throughout the whole FL exposure., Our results highlight a more balanced and cost-effective photoacclimation strategy of P. tricornutum under FL than under SL conditions. [ABSTRACT FROM AUTHOR]

The photochemical reflectance index ( PRI) reflects diurnal xanthophyll cycle activity and is also influenced by seasonally changing carotenoid : Chl pigment ratios. Both changing pigment pools and xanthophyll cycle activity contribute to photoprotection in evergreen conifers exposed to boreal winters, but they operate over different timescales, and their relative contribution to the PRI signal has often been unclear., To clarify these responses and their contribution to the PRI signal, leaf PRI, pigment composition, temperature and irradiance were monitored over 2 yr for two evergreen conifers ( Pinus contorta and Pinus ponderosa) in a boreal climate., PRI was affected by three distinct processes operating over different timescales and exhibiting contrasting spectral responses. Over the 2 yr study period, the greatest change in PRI resulted from seasonally changing carotenoid : Chl pigment ratios, followed by a previously unreported shifting leaf albedo during periods of deep cold. Remarkably, the smallest change was attributable to the xanthophyll cycle., To properly distinguish these three effects, interpretation of PRI must consider temporal context, physiological responses to evolving environmental conditions, and spectral response. Consideration of the separate mechanisms affecting PRI over different timescales could greatly improve efforts to monitor changing photosynthetic activity using optical remote sensing. [ABSTRACT FROM AUTHOR]

Terrestrial laser scanning ( TLS) data allow spatially explicit ( x, y, z) laser return intensities to be recorded throughout a plant canopy, which could considerably improve our understanding of how physiological processes vary in three-dimensional space. However, the utility of TLS data for the quantification of plant physiological properties remains largely unexplored. Here, we test whether the laser return intensity of green (532-nm) TLS correlates with changes in the de-epoxidation state of the xanthophyll cycle and photoprotective non-photochemical quenching ( NPQ), and compare the ability of TLS to quantify these parameters with the passively measured photochemical reflectance index ( PRI)., We exposed leaves from five plant species to increasing light intensities to induce NPQ and de-epoxidation of violaxanthin (V) to antheraxanthin (A) and zeaxanthin (Z). At each light intensity, the green laser return intensity ( GLRI), narrowband spectral reflectance, chlorophyll fluorescence emission and xanthophyll cycle pigment composition were recorded., Strong relationships between both predictor variables ( GLRI, PRI) and both explanatory variables ( NPQ, xanthophyll cycle de-epoxidation) were observed., GLRI holds promise to provide detailed (mm) information about plant physiological status to improve our understanding of the patterns and mechanisms driving foliar photoprotection. We discuss the potential for scaling these laboratory data to three-dimensional canopy space. [ABSTRACT FROM AUTHOR]

• An earlier onset of photosynthesis in spring for boreal forest trees is predicted as the climate warms, yet the importance of soil vs air temperatures for spring recovery remains to be determined. Effects of various soil- and air-temperature conditions on spring recovery of photosynthesis in Scots pine ( Pinus sylvestris) seedlings were assessed under controlled environmental conditions. • Using winter-acclimated seedlings, photosynthetic responses were followed after transfer to different simulated spring conditions. • Recovery rates for photosynthetic electron transport and net CO2 uptake were slower in plants from cold or frozen soil compared with controls. In addition, a greater fraction of light absorbed was not used photochemically, but was dissipated thermally via xanthophyll cycle pigments. Intermittent frost events decreased photosynthetic capacity and increased thermal energy dissipation. Within a few days after frost events, photosynthetic capacity recovered to prefrost levels. After 18 d under spring conditions, no difference in the optimum quantum yield of photosynthesis was observed between seedlings that had been exposed to intermittent frost and control plants. • These results show that, if air temperatures remain favourable and spells of subfreezing air temperatures are only of short duration, intermittent frost events delay but do not severely inhibit photosynthetic recovery in evergreen conifers during spring. Cold and/or frozen soils exert much stronger inhibitory effects on the recovery process, but they do not totally inhibit it. [ABSTRACT FROM AUTHOR]

• The capability to withstand and to recover from severe summer droughts is becoming an important issue for tree species in central Europe, as dry periods are predicted to occur more frequently over the coming decades. • Changes in leaf gas exchange, chlorophyll a fluorescence and leaf compounds related to photoprotection were analysed in young Quercus pubescens trees under field conditions during two summers (2004 and 2005) of progressive drought and subsequent rewatering. • Photochemistry was reversibly down-regulated and dissipation of excess energy was enhanced during the stress phase, while contents of leaf pigments and antioxidants were almost unaltered. Plant water status was restored immediately after rewatering. Net photosynthesis ( Pn) measured at ambient CO2 recovered from inhibition by drought within 4 wk. Pn measured at elevated CO2– to overcome stomatal limitations – was restored after a few days. • A network of photoprotective mechanisms acted in preserving the potential functionality of the photosynthetic apparatus during severe drought, leading to a rapid recovery of photosynthetic activity after rewatering. Thus, Q. pubescens seems to be capable of withstanding and surviving extreme drought events. [ABSTRACT FROM AUTHOR]

• Here the mechanisms involved in excitation energy dissipation of Macrocystis pyrifera were characterized to explain the high nonphotochemical quenching of chlorophyll a (Chl a) fluorescence (NPQ) capacity of this alga. • We performed a comparative analysis of NPQ and xanthophyll cycle (XC) activity in blades collected at different depths. The responses of the blades to dithiothreitol (DTT) and to the uncoupler NH4Cl were also assayed. • The degree of NPQ induction was related to the amount of zeaxanthin synthesized in high light. The inhibition of zeaxanthin synthesis with DTT blocked NPQ induction. A slow NPQ relaxation upon the addition of NH4Cl, which disrupts the transthylakoid proton gradient, was detected. The slow NPQ relaxation took place only in the presence of de-epoxidated XC pigments and was related to the epoxidation of zeaxanthin. • These results indicate that in M. pyrifera, in contrast to higher plants, the transthylakoid proton gradient alone does not induce NPQ. The role of this gradient seems to be related only to the activation of the violaxanthin de-epoxidase enzyme. [ABSTRACT FROM AUTHOR]

• In the present study, we investigated the effects of CO2 availability on photosynthesis, photoinhibition and pigmentation in two species of amphibious plants, Lobelia cardinalis and Nesaea crassicaulis. • The plants were grown emergent under atmospheric conditions and submerged under low and high CO2 availability. • Compared with Lobelia, Nesaea had thin leaves and few stomata in all CO2 treatments. While Lobelia expressed no variation in anthocyanin content among treatments, Nesaea produced high concentrations of anthocyanin when submerged. Lobelia photosynthesis increased in response to increasing CO2 availability, and photoinhibition was negatively related to xanthophyll content. By contrast, Nesaea photosynthesis was highest under submerged conditions, and there was no relationship between photoinhibition and the xanthophyll content. • We conclude that the response of Lobelia to varying CO2 availability is similar to that of terrestrial plants and that this species relies on the xanthophyll cycle for nonphotochemical quenching (NPQ) and protection against photoinhibition. By contrast, the thin leaves, few stomata and low levels of chlorophylls and accessory pigments in Nesaea, relative to Lobelia, suggest adaptation to a submerged habitat. While Nesaea does not seem to rely on the xanthophyll cycle or other xanthophylls for NPQ, some role of anthocyanins in the protection against photoinhibition cannot be ruled out, owing to its effect as a sunscreen and as an efficient quencher of free radicals. [ABSTRACT FROM AUTHOR]

• Responses to simulated sunflecks were examined in upper canopy and coppice leaves ofNothofagus cunninghamiigrowing in an old-growth rainforest gully in Victoria, Australia.• Shaded leaves were exposed to a sudden increase in irradiance from 20 to 1500 µmol m−2 s−1. Gas exchange and chlorophyll fluorescence were measured during a 10 min simulated sunfleck and, in the ensuing dark treatment, we examined the recovery of PS II efficiency and the conversion state of xanthophyll cycle pigments.• Photosynthetic induction was rapid compared with tropical and northern hemisphere species. Stomatal conductance was relatively high in the shade and stomata did not directly control photosynthetic induction under these conditions. During simulated sunflecks, zeaxanthin was formed rapidly and photochemical efficiency was reduced. These processes were reversed within 30 min in coppice leaves, but this took longer in upper canopy leaves.• Poor drought tolerance and achieving a positive carbon balance in a shaded canopy may be functionally related to high stomatal conductance in the shade inN. cunninghamii. The more persistent reduction in photochemical efficiency of upper canopy leaves, which means less efficient light use in subsequent shade periods, but stronger protection from high light, may be related to the generally higher irradiance and longer duration of sunflecks in the upper canopy, but potentially reduces carbon gain during shade periods by 30%.New Phytologist(2004)doi: 10.1111/j.1469-8137.2004.01232.x© New Phytologist(2004) [ABSTRACT FROM AUTHOR]

Summary • Here, the effects of salinity (0–400 mM NaCl) on photosynthesis, photosystem II (PSII) efficiency and the xanthophyll cycle were investigated in the halophyte Atriplex centralasiatica grown under outdoor conditions. • Leaf sodium and chloride in leaves increased considerably whereas CO2 assimilation rate decreased. PSII efficiency (ΦPSII ) and the efficiency of excitation energy capture by open PSII reaction centres &formmu9; decreased whereas nonphotochemical quenching (NPQ) increased significantly. There was no change in photochemical quenching (qP ) in salt-adapted plants. Salinity induced no changes in the maximum efficiency of PSII photochemistry (F v /F m ) measured either at midday or at predawn. However, F v /F m values were c. 20% lower at midday than at predawn. • Contents of chlorophyll (a + b), neoxanthin, lutein and β-carotene were unchanged with increasing salt concentration, but zeaxanthan increased significantly, at the expense of violaxanthin. There was a linear relationship between the de-epoxidation state of the xanthophyll cycle and ΦPSII , &formmu0;, and NPQ. • Our results suggest that A. centralasiatica shows high tolerance to both high salinity and photoinhibition and that the xanthophyll cycle played an important role in protecting photosynthetic apparatus from photoinhibitory damage. Tolerance of PSII to salinity and photoinhibition can be viewed as an important strategy for A. centralasiatica to grow in very high saline soil during the summer season with high irradiance. [ABSTRACT FROM AUTHOR]

This study explored reflectance indices sampled with a `leaf reflectometer' as measures of pigment content for leaves of contrasting light history, developmental stage and functional type (herbaceous annual versus sclerophyllous evergreen). We employed three reflectance indices: a modified normalized difference vegetation index (NDVI), an index of chlorophyll content; the red/green reflectance ratio (RRED:RGREEN), an index of anthocyanin content: and the change in photochemical reflectance index upon dark light conversions (ΔPRI ), an index of xanthophyll cycle pigment activity. In Helianthus annuus (sunflower), xanthophyll cycle pigment amounts were linearly related to growth light environment; leaves in full sun contained approximately twice the amount of xanthophyll cycle pigments as leaves in deep shade, and at midday a larger proportion of these pigments were in the photoprotective, de-epoxidized forms relative to shade leaves. Reflectance indices also revealed contrasting patterns of pigment development in leaves of contrasting structural types (annual versus evergreen), in H. annuus sun leaves, there was a remarkably rapid increase in amounts of both chlorophyll and xanthophyll cycle pigments along a leaf developmental sequence. This pattern contrasted with that of Quercus agrifolia (coast live oak, a sclerophyllous evergreen), which exhibited a gradual development of both chlorophyll and xanthophyll cycle pigments along with a pronounced peak of anthocyanin pigment content in newly expanding leaves. These temporal patterns of pigment development in Q. agrifolia leaves suggest that anthocyanins arid xanthophyll cycle pigments serve complementary photoprotective roles during early leaf development. The results illustrate the use of reflectance indices for distinguishing divergent patterns of pigment activity in leaves of contrasting light history and functional type. [ABSTRACT FROM AUTHOR]

We have explored leaf-level plastic response to light and nutrients of Quercus ilex and Q. coccifera, two closely related Mediterranean evergreen sclerophylls, in a factorial experiment with seedlings. Leaf phenotypic plasticity, assessed by a relative index (PI = (maximum value - minimum)/maximum) in combination with the significance of the difference among means, was studied in 37 morphological and physiological variables. Light had significant effects on most variables relating to photosynthetic pigments, chlorophyll fluorescence and gas exchange, whereas nutrient treatment had a significant effect in only 10% of the variables. Chlorophyll content was higher in the shade whereas carotenoid content and nonphotochemical quenching increased with light. Nutrient limitations increased the xanthophyll-cycle pool but only at high light intensities, and the same interaction between light and nutrients was observed for lutein. Predawn photochemical efficiency of PSII was not affected by either light or nutrients, although midday photochemical efficiency of PSII was lower at high light intensities. Photosynthetic light compensation point and dark respiration on an area basis decreased with light, but photosynthetic capacity on a dry mass basis and photochemical quenching were higher in low light, which translated into a higher nitrogen use efficiency in the shade. We expected Q. ilex, the species of the widest ecological distribution, to be more plastic than Q. coccifera, but differences were minor: Q. ilex exhibited a significant response to light in 13% more of the variables than Q. coccifera, but mean PI was very similar in the two species. Both species tolerated full sunlight and moderate shade, but exhibited a reduced capacity to enhance photosynthetic utilization of high irradiance. When compared with evergreen shrubs from the tropical rainforest, leaf responsiveness of the two evergreen oaks was low. We suggest that the low leaf-level responsiveness found here is part of a conservative resource use strategy, which seems to be adaptive for evergreen woody plants in Mediterranean-type ecosystems.