Xanthophyll Cycle Components Research Articles

Kinetics of the xanthophyll cycle and its role in photoprotective memory and response

Efficiently balancing photochemistry and photoprotection is crucial for survival and productivity of photosynthetic organisms in the rapidly fluctuating light levels found in natural environments. The ability to respond quickly to sudden changes in light level is clearly advantageous. In the alga Nannochloropsis oceanica we observed an ability to respond rapidly to sudden increases in light level which occur soon after a previous high-light exposure. This ability implies a kind of memory. In this work, we explore the xanthophyll cycle in N. oceanica as a short-term photoprotective memory system. By combining snapshot fluorescence lifetime measurements with a biochemistry-based quantitative model, we show that short-term memory arises from the xanthophyll cycle. In addition, the model enables us to characterize the relative quenching abilities of the three xanthophyll cycle components. Given the ubiquity of the xanthophyll cycle in photosynthetic organisms the model described here will be of utility in improving our understanding of vascular plant and algal photoprotection with important implications for crop productivity.

Seasonal Differences in Leaf Photoprotective Potential between Adults and Juveniles of Two Mediterranean Perennials with Distinct Growth Forms: A Comparative Field Study.

The photosynthetic differences between adult and juvenile Mediterranean plants were previously studied under field conditions, yet the corresponding differentiation of their photoprotective efficiency has not been sufficiently investigated. The present study aims to examine possible differences in the photoprotective potential between adults and juveniles of two native Mediterranean plants with distinct growth forms. Thus, the seasonal variations in individual carotenoids, electron transport rate (ETR), and non-photochemical quenching (NPQ) were monitored in fully exposed mature leaves from adults and juveniles of the winter deciduous tree Cercis siliquastrum L. and the evergreen sclerophyllous shrub Nerium oleander L. All plants were grown under apparently similar field conditions. In both species, juveniles displayed substantially lower ETR and increased NPQ values than adults in spring, with the differences intensifying during summer drought and diminishing in autumn. Concomitantly, juveniles showed significantly higher chlorophyll-based total carotenoids in spring and summer mainly due to the higher investment in xanthophyll cycle components (VAZ), in combination with an increased mid-day de-epoxidation state (DEPS) and partial retention of zeaxanthin in the dark. In N. oleander, although ETR was lower in juveniles during winter, NPQ was extremely low in both ages. In conclusion, juveniles exhibit enhanced photoprotection potential, especially in the summer, due to their reduced photochemical capacity. The photosynthetic superiority of adults during the favorable spring period may be attributed to the needs of the co-existing reproductive effort.

Protective Responses at the Biochemical and Molecular Level Differ between a Coffea arabica L. Hybrid and Its Parental Genotypes to Supra-Optimal Temperatures and Elevated Air [CO2

Climate changes with global warming associated with rising atmospheric [CO2] can strongly impact crop performance, including coffee, which is one of the most world’s traded agricultural commodities. Therefore, it is of utmost importance to understand the mechanisms of heat tolerance and the potential role of elevated air CO2 (eCO2) in the coffee plant response, particularly regarding the antioxidant and other protective mechanisms, which are crucial for coffee plant acclimation. For that, plants of Coffea arabica cv. Geisha 3, cv. Marsellesa and their hybrid (Geisha 3 × Marsellesa) were grown for 2 years at 25/20 °C (day/night), under 400 (ambient CO2, aCO2) or 700 µL (elevated CO2, eCO2) CO2 L−1, and then gradually submitted to a temperature increase up to 42/30 °C, followed by recovery periods of 4 (Rec4) and 14 days (Rec14). Heat (37/28 °C and/or 42/30 °C) was the major driver of the response of the studied protective molecules and associated genes in all genotypes. That was the case for carotenoids (mostly neoxanthin and lutein), but the maximal (α + β) carotenes pool was found at 37/28 °C only in Marsellesa. All genes (except VDE) encoding for antioxidative enzymes (catalase, CAT; superoxide dismutases, CuSODs; ascorbate peroxidases, APX) or other protective proteins (HSP70, ELIP, Chape20, Chape60) were strongly up-regulated at 37/28 °C, and, especially, at 42/30 °C, in all genotypes, but with maximal transcription in Hybrid plants. Accordingly, heat greatly stimulated the activity of APX and CAT (all genotypes) and glutathione reductase (Geisha3, Hybrid) but not of SOD. Notably, CAT activity increased even at 42/30 °C, concomitantly with a strongly declined APX activity. Therefore, increased thermotolerance might arise through the reinforcement of some ROS-scavenging enzymes and other protective molecules (HSP70, ELIP, Chape20, Chape60). Plants showed low responsiveness to single eCO2 under unstressed conditions, while heat promoted changes in aCO2 plants. Only eCO2 Marsellesa plants showed greater contents of lutein, the pool of the xanthophyll cycle components (V + A + Z), and β-carotene, compared to aCO2 plants at 42/30 °C. This, together with a lower CAT activity, suggests a lower presence of H2O2, likely also associated with the higher photochemical use of energy under eCO2. An incomplete heat stress recovery seemed evident, especially in aCO2 plants, as judged by the maintenance of the greater expression of all genes in all genotypes and increased levels of zeaxanthin (Marsellesa and Hybrid) relative to their initial controls. Altogether, heat was the main response driver of the addressed protective molecules and genes, whereas eCO2 usually attenuated the heat response and promoted a better recovery. Hybrid plants showed stronger gene expression responses, especially at the highest temperature, when compared to their parental genotypes, but altogether, Marsellesa showed a greater acclimation potential. The reinforcement of antioxidative and other protective molecules are, therefore, useful biomarkers to be included in breeding and selection programs to obtain coffee genotypes to thrive under global warming conditions, thus contributing to improved crop sustainability.

Phenotypic Diversity and Plasticity of Photoresponse Across an Environmentally Contrasting Family of Phytoflagellates.

Organisms often employ ecophysiological strategies to exploit environmental conditions and ensure bio-energetic success. However, the many complexities involved in the differential expression and flexibility of these strategies are rarely fully understood. Therefore, for the first time, using a three-part cross-disciplinary laboratory experimental analysis, we investigated the diversity and plasticity of photoresponsive traits employed by one family of environmentally contrasting, ecologically important phytoflagellates. The results demonstrated an extensive inter-species phenotypic diversity of behavioural, physiological, and compositional photoresponse across the Chlamydomonadaceae, and a multifaceted intra-species phenotypic plasticity, involving a broad range of beneficial photoacclimation strategies, often attributable to environmental predisposition and phylogenetic differentiation. Deceptively diverse and sophisticated strong (population and individual cell) behavioural photoresponses were observed, with divergence from a general preference for low light (and flexibility) dictated by intra-familial differences in typical habitat (salinity and trophy) and phylogeny. Notably, contrasting lower, narrow, and flexible compared with higher, broad, and stable preferences were observed in freshwater vs. brackish and marine species. Complex diversity and plasticity in physiological and compositional photoresponses were also discovered. Metabolic characteristics (such as growth rates, respiratory costs and photosynthetic capacity, efficiency, compensation and saturation points) varied elaborately with species, typical habitat (often varying more in eutrophic species, such as Chlamydomonas reinhardtii), and culture irradiance (adjusting to optimise energy acquisition and suggesting some propensity for low light). Considerable variations in intracellular pigment and biochemical composition were also recorded. Photosynthetic and accessory pigments (such as chlorophyll a, xanthophyll-cycle components, chlorophyll a:b and chlorophyll a:carotenoid ratios, fatty acid content and saturation ratios) varied with phylogeny and typical habitat (to attune photosystem ratios in different trophic conditions and to optimise shade adaptation, photoprotection, and thylakoid architecture, particularly in freshwater environments), and changed with irradiance (as reaction and harvesting centres adjusted to modulate absorption and quantum yield). The complex, concomitant nature of the results also advocated an integrative approach in future investigations. Overall, these nuanced, diverse, and flexible photoresponsive traits will greatly contribute to the functional ecology of these organisms, addressing environmental heterogeneity and potentially shaping individual fitness, spatial and temporal distribution, prevalence, and ecosystem dynamics.

Particle film technology modulates xanthophyll cycle and photochemical dynamics of grapevines grown in the Douro Valley

Field-grown grapevines are often exposed to multiple environmental stresses, which challenges wine-growers to develop sustainable measures to sustain vine growth, yield, and quality. Under field conditions this task is demanding, due to differences in the magnitudes of stresses and associated plant responses. In this study we explored the hypothesis that kaolin-particle film application improves grapevine photoprotection through the regulation of xanthophyll cycle genes, limiting the thermal dissipation of excess energy under harsh environmental conditions. Hence, we selected two grapevine varieties, Touriga-Nacional (TN) and Touriga-Franca (TF), grown in the Douro Demarcated Region, and evaluated changes in light dissipation mechanisms, xanthophyll cycle components, and the expression of xanthophyll cycle genes during the 2017 summer season. The results showed that, from veraison to ripening, kaolin triggered the up-regulation of violaxanthin de-epoxidase (VvVDE1) and zeaxanthin epoxidase (VvZEP1) genes, indicating optimised regulation of the xanthophyll cycle. Kaolin treatment also decreased chlorophyll (Chla, Chlb, Chl(a+b)) and carotenoid (Car) accumulation under increasing summer stress conditions in both varieties and lowered the non-photochemical quenching (NPQ) of grapevines on ripening, suggesting a long-term response to summer stress. In addition, kaolin-treated grapevines showed increased Chla/Chlb and lower Chl(a+b)/Car ratios, displaying some features of high light adapted leaves. Overall, this study suggests that kaolin application enabled grapevines to benefit from fluctuating periods of summer stress by managing chlorophyll and carotenoid content and limiting down-regulation of both photochemistry and photoinhibition processes. Under Mediterranean field conditions, kaolin application can be considered an efficient method of minimising summer stress impact on grapevines.

Effect of drought and heat stresses on photosynthesis, pigments, and xanthophyll cycle in alfalfa (Medicago sativa L.)

Climate changes severely influence photosynthetic activity of plants. Studies on the combined effect of drought and heat stress on photosynthesis, pigments, and xanthophyll cycle particularly in alfalfa (Medicago sativa L.) are limited. Thus, this study investigated the combined effects of drought and heat stress in two alfalfa varieties ('Deqin' and 'Algonguin'). Results revealed that the values of the net photosynthetic rate demonstrated a significantly decreasing tendency under stresses. The effect of stresses on ribulose 1,5-bisphosphate carboxylase activity in both varieties showed a trend of first declining and then rising. Meanwhile, phosphoenolpyruvate carboxylase activity showed an increasing tendency. The intensification of stresses led to reduced photochemical efficiency and increased capability of thermal dissipation. A linear correlation was found between nonphotochemical quenching and xanthophyll cycle components. Thus, the change in carotenoids demonstrated an important role in photoprotection, the xanthophyll cycle was the main way of dissipation of excess energy under stress conditions in alfalfa.

Photochemical and antioxidative responses of the glume and flag leaf to seasonal senescence in wheat.

The non-leaf photosynthetic organs have recently attracted much attention for the breeding and screening of varieties of cereal crops to achieve a high grain yield. However, the glume photosynthetic characteristics and responses to high temperature at the late stages of grain filling are not well known in winter wheat (Triticum aestivum L.). In the present study, an experiment was conducted to investigate the anatomy, chloroplast temporal changes, chlorophyll fluorescence, xanthophyll cycle and antioxidative defense system in glumes of field-grown wheat during grain filling compared with flag leaves. Observations using a light microscope revealed that the glumes developed a solid structural base for performing photosynthesis. Compared with the flag leaves, the glumes preserved a more integral ultrastructure, as observed under transmission electron microscopy, and had higher values of Fv/Fm and ΦPSII at the maturity stage. Further analysis of the chlorophyll fluorescence demonstrated that the glumes experienced high non-photochemical quenching (NPQ) at the late stages. Determination of the pool size of the xanthophyll cycle suggested that the (A+Z)/(V+A+Z) ratio was consistently higher in glumes than in flag leaves and that the V+A+Z content was considerably higher in glumes at the maturity stage. In addition, the glumes exhibited a higher antioxidant enzyme activity and a lower accumulation of reactive oxygen species. These results suggest that the glumes are photosynthetically active and senesce later than the flag leaves; the advantages may have been achieved by coordinated contributions of the structural features, higher NPQ levels, greater de-epoxidation of the xanthophyll cycle components and antioxidative defense metabolism.

The non-foliar hypoxic photosynthetic syndrome: evidence for enhanced pools and functionality of xanthophyll cycle components and active cyclic electron flow in fruit chlorenchyma.

Green fruits display a high engagement in CEF and enhanced VAZ cycle activity as a response to the demands imposed by their internal aerial conditions, particularly low O 2 , due to gas exchange limitations. In the present study, we used HPLC analysis, post-illumination changes in fluorescence yield under varying O2 and CO2 partial pressures and absorbance changes at 820 nm induced by far-red light to assess the carotenoid composition, the functionality of the xanthophyll cycle (VAZ) and the possibility of an active cyclic e (-) flow (CEF) in the fully exposed green fruits from Nerium oleander and Rosa sp. Equally exposed, mature leaves served as controls. Compared to leaves, fruits display less total chlorophylls and carotenoids but higher Car/Chl ratio, mainly shaped by the increased pools of the VAZ cycle components, in both species. The enhanced VAZ pool size in fruits is combined with a higher mid-day de-epoxidation state (DEPS). Moreover, fruits exhibit considerably lower levels of oxidizable P700, a faster re-reduction of PSI and significantly higher relative magnitude of CEF, irrespective of the O2/CO2 levels applied. We conclude that the higher VAZ investment may serve the enhanced heat dissipation needs in fruits, in the presence of a suppressed linear e (-) flow. In addition, the elevated potential of CEF may replenish the ATP lost due to hypoxia and concurrently facilitate the development of adequate non-photochemical quenching (NPQ), through its contribution to ΔpH increase. Since other non-foliar green organs exhibit a similar photosynthetic pattern, we argue that this may reflect a common strategy for green tissues under similar micro-environmental conditions, particularly hypoxia.

Effects of Exogenous Putrescine on Chlorophyll Fluorescence Imaging and Heat Dissipation Capacity in Cucumber (Cucumis sativus L.) Under Salt Stress

The objective of this study was to identify the effects of exogenous putrescine on photosynthetic performance and heat dissipation capacity in cucumber seedlings under salt stress. The stress of 75 mM NaCl for 7 days caused a significant decrease in net photosynthetic rate (P N ). The experiment employed a chlorophyll fluorescence imaging technique and demonstrated that the maximal quantum yield of photosystem II photochemistry (Fv/Fm) and the actual photochemical efficiency of photosystem II (ΦPSII) were reduced by salt stress. Moreover, salt stress markedly reduced the photochemical quenching coefficient (qP) and non-photochemical quenching coefficient (qN), and significantly increased non-regulated heat dissipation (ΦNO). However, stressed plants supplied with exogenous putrescine exhibited higher P N and ΦPSII, which indicated that putrescine can alleviate the detrimental effects on photosynthesis induced by salt stress. Putrescine sprayed on stressed plants significantly enhanced the regulated energy dissipation (ΦNPQ) and decreased ΦNO. Application of exogenous putrescine also changed the levels of xanthophyll cycle components and further enhanced the de-epoxidation state of xanthophyll cycle pigments under salt stress. Under control conditions, putrescine exerted little influence on the photosynthetic parameters in cucumber leaves. In conclusion, the application of exogenous putrescine may improve the heat dissipation capacity by promoting the xanthophyll cycle to reduce the damage caused by excess excitation energy, thus enhancing the salt tolerance of cucumber seedlings.

Photoprotection in the green tidal alga Ulva prolifera: role of LHCSR and PsbS proteins in response to high light stress

Ulva prolifera, an intertidal macroalga, has to adapt to wide variations in light intensity, making this species particularly rewarding for studying the evolution of photoprotective mechanisms. Intense light induced increased non-photochemical quenching (NPQ) and stimulated de-epoxidation of xanthophyll cycle components, while DTT-treated samples had lower NPQ capacity, indicating that the xanthophyll cycle must participate in photoprotection. In this work, we found that the PsbS-related NPQ was maintained in U.prolifera. According to analysed gene expression, both LhcSR and psbS were up-regulated in high light, suggesting that these two genes are light-induced. LHCSR and PsbS proteins were present at different light intensities and accumulated under high light conditions, and PsbS concentrations were higher than LHCSR, showing that the NPQ mechanism of U.prolifera is more dependent on PsbS protein concentration. Moreover, the level of both LHCSR and PsbS proteins was high even in the darkness, and neither the transcript level nor protein content of LhcSR and psbS genes varied significantly following short-term exposure to intense light. These findings suggest that this alga can modulate NPQ levels through regulation of the xanthophyll cycle and concentrations of PsbS and/or LHCSR.

Molecular characterization and primary functional analysis of PeVDE, a violaxanthin de-epoxidase gene from bamboo (Phyllostachys edulis)

PeVDE was expressed primarily in bamboo leaves, which was up-regulated under high light. The protein encoded by PeVDE had enzyme activity of catalyzing violaxanthin (V) to zeaxanthin (Z) through antheraxanthin (A) as assay shown in vitro. Violaxanthin de-epoxidase (VDE), a key enzyme of xanthophyll cycle, catalyzes conversion from violaxanthin (V) to zeaxanthin (Z) through antheraxanthin (A) to protect photosynthesis apparatus. A cDNA, PeVDE, encoding a VDE was isolated from bamboo (Phyllostachys edulis) by RT-PCR and RACE methods. PeVDE is 1,723bp and contains an ORF encoding 451 amino acids, with a transit peptide of 103 amino acids. The mature protein is deduced to have 348 amino acids with a calculated molecular weight of 39.6kDa and a theoretic isoelectric point of 4.5. Semi-quantitative RT-PCR assay indicated that the highest expression level of PeVDE was in leaf, which agreed with the accumulation pattern of PeVDE protein. Real time PCR results showed that PeVDE was up-regulated and reached the highest level after the treatment (1,200μmo1m(-2)s(-1)) for 2h, then decreased and kept at the level similar to that of 0.5h after treatment for 8h. To investigate the function of PeVDE, mature protein was heterologously expressed in Escherichia coli and the enzymatic activity assay was carried out using V as substrate. The pigments that formed in the reaction mixture were extracted and analyzed by HPLC method. Besides V, A and Z were detected in the reaction mixture, which indicated that the recombinant protein exhibited enzymatic activity of catalyzing V into Z through A. This study indicates that PeVDE functions through regulating the components of xanthophyll cycle, which might be one of the critical factors that contribute to the growth of bamboo under naturally varying light conditions.

Photosynthesis and photoinhibition in two differently coloured varieties of Oxalis triangularis - the effect of anthocyanin content

The purpose of this study was to clarify effects of anthocyanins on photosynthesis and photoinhibition in green and red leaves of Oxalis triangularis. Gas analysis indicated that green plants had the highest apparent quantum yield for CO2 assimilation [0.051 vs. 0.031 μmol(CO2) μmol-1(photon)] and the highest maximum photosynthesis [10.07 vs. 7.24 μmol(CO2) m-2 s-1], while fluorescence measurements indicated that red plants had the highest PSII quantum yield [0.200 vs. 0.143 μmol(e-) μmol-1(photon)] and ETRmax [66.27 vs. 44.34 μmol(e-) m-2 s-1]. Red plants had high contents of anthocyanins [20.11 mg g-1(DM)], while green plants had low and undetectable levels of anthocyanin. Red plants also had statistically significantly (0.05>p>0.01) lower contents of xanthophyll cycle components [0.63 vs. 0.76 mg g-1(DM)] and higher activities of the reactive oxygen scavenging enzyme ascorbate peroxidase [41.2 vs. 10.0 nkat g-1(DM)]. Anthocyanins act as a sunscreen, protecting the chloroplasts from high light intensities. This shading effect causes a lower photosynthetic CO2 assimilation in red plants compared to green plants, but a higher quantum efficiency of photosystem II (PSII). Anthocyanins contribute to photoprotection, compensating for lower xanthophyll content in red plants, and red plants are less photoinhibited than green plants, as illustrated by the Fv/Fm ratio.

Roles of xanthophylls and exogenous ABA in protection against NaCl-induced photodamage in rice (Oryza sativa L) and cabbage (Brassica campestris)

Changes in actual efficiency of PS II photochemistry, non-photochemical quenching (NPQ), content of xanthophylls and kinetics of de-epoxidation were studied in ABA-fed and non-ABA-fed leaves of rice and cabbage under NaCl stress. Salt stress induced more progressive decrease in actual efficiency of PS II photochemistry (ФPS II), higher reduction state of PS II, and a small significant increase in NPQ in NaCl-sensitive rice plants as compared with NaCl-tolerant cabbage plants, whereas exogenously supplied ABA alleviated the decrease in actual efficiency of PS II photochemistry (ФPS II), induced a lower reduction state of PS II, and caused higher capacity of NPQ in ABA-fed plants than in non-ABA-fed plants. As a result, there were higher activities of photosynthetic electron transport, higher capacity of energy dissipation, and lower cumulation of excess light in cabbage than in rice plants, and in ABA-fed leaves than in non-ABA-fed leaves. The effect of ABA was more efficient in cabbage than in rice plants. Addition of exogenous ABA resulted in enhancement of the size of the xanthophyll cycle pool, promotion of de-epoxidation of the xanthophyll cycle components, and a rise in the level of NPQ by altering the kinetics of de-epoxidation of the xanthophyll cycle. Protection from photodamage appears to be achieved by coordinated contributions by exogenous ABA and xanthophyll cycle-mediated NPQ. This variety of photoprotective mechanisms may be essential for conferring photodamage tolerance under NaCl stress.

Salt-induced oxidative stress in rosemary plants: Damage or protection?

Mechanisms of photoprotection and antioxidant protection, including changes in chlorophylls, xanthophyll cycle components and levels of low-molecular-weight chloroplastic antioxidants (lutein, β-carotene and α-tocopherol) were studied together with levels of malondialdehyde, a product of lipid peroxidation, in the response of rosemary (Rosmarinus officinalis L.) plants to salt stress. Plants were exposed to increasing NaCl concentrations (50, 100 and 150mM) for 6 weeks, and two concentrations of the following chloride salts: KCl, CaCl2, MgCl2 and FeCl3, were used together with 100mM NaCl to explore the extent to which these salts can alter the mechanisms of photoprotection, antioxidant protection and malondialdehyde accumulation in leaves. Increasing concentrations of NaCl decreased leaf water contents and photosynthetic pigment levels, while the contents of α-tocopherol and malondialdehyde increased, but with completely different kinetics. α-Tocopherol levels increased in a dose-dependent manner as stress progressed, while malondialdehyde levels increased at the highest dose (150mM NaCl) but only during early phases of stress. Furthermore, although the addition of chloride salts to NaCl-treated plants apparently improved leaf physiological status, in terms of water and chlorophyll contents, plants showed an increased photoprotective demand and increased oxidative stress, particularly in FeCl3-treated plants. It is concluded that (i) rosemary plants can withstand moderate doses of NaCl in the medium (at least 150mM NaCl for 6 weeks), (ii) oxidative stress may be a mechanism for protecting plants from moderate doses of salt stress rather than causing damage to plants, and (iii) the addition of chloride salts to NaCl-treated plants may dramatically increase the photoprotective demand and oxidative stress of leaves, while plant growth is not negatively affected.

Dehydration rate and time of desiccation affect recovery of the lichenic algae Trebouxia erici: alternative and classical protective mechanisms

The mechanisms involved in desiccation tolerance of lichens and their photobionts are still poorly understood. To better understand these mechanisms we have studied dehydration rate and desiccation time in Trebouxia, the most abundant chlorophytic photobiont in lichen. Our findings indicate that the drying rate has a profound effect on the recovery of photosynthetic activity of algae after rehydration, greater than the effects of desiccation duration. The basal fluorescence (F'(o)) values in desiccated algae were significantly higher after rapid dehydration, than after slow dehydration, suggesting higher levels of light energy dissipation in slow-dried algae. Higher values of PSII electron transport were recovered after rehydration of slow-dried Trebouxia erici compared to rapid-dried algae. The main component of non-photochemical quenching after slow dehydration was energy dependent (q (E)), whereas after fast dehydration it was photoinhibition (q (I)). Although q (E) seems to play a role during desiccation recovery, no significant variations were detected in the xanthophyll cycle components. Desiccation did not affect PSI functionality. Classical antioxidant activities like superoxide dismutase or peroxidase decreased during desiccation and early recovery. Dehydrins were detected in the lichen-forming algae T. erici and were constitutively expressed. There is probably a minimal period required to develop strategies which will facilitate transition to the desiccated state in this algae. In this process, the xanthophyll cycle and classical antioxidant mechanisms play a very limited role, if any. However, our results indicate that there is an alternative mechanism of light energy dissipation during desiccation, where activation is dependent on a sufficiently slow dehydration rate.

Role of xanthophyll cycle-mediated photoprotection in Arbutus unedo plants exposed to water stress during the Mediterranean summer

We analyzed the response of potted strawberry tree (Arbutus unedo L.) seedlings exposed to water stress by withholding water for 10 d (WS). Leaf water potential, net CO2 assimilation, and stomatal conductance decreased with increasing water deficit. A 30 % reduction of chlorophyll (Chl) content in the antenna complexes was observed in WS-plants. Simultaneously, a decline of photochemical efficiency (Fv/Fm) occurred as a result of an excess of solar radiation energy when carbon assimilation was limited by stomata closure due to soil water deficit. The non-photochemical quenching of Chl fluorescence (ΦNPQ) significantly increased, as well as the leaf contents of zeaxanthin (Z) and antheraxanthin (A) at the expense of violaxanthin during the WS-period. Elevated predawn contents of de-epoxidized xanthophyll cycle components were associated with a sustained lowering of predawn photosystem 2 efficiency; this suggested an engagement of Z+A in a state primed for energy dissipation. Thus, the ability of strawberry trees to maintain the functionality of the xanthophyll cycle during the Mediterranean summer is an efficient mechanism to prevent irreversible damages to the photosynthetic machinery through thermal energy dissipation in the antenna and the reduction in photochemical efficiency.

Involvement of betacyanin in chilling-induced photoinhibition in leaves of Suaeda salsa

Seeds of Suaeda salsa were cultured in dark for 3 d and betacyanin accumulation in seedlings was promoted significantly. Then the seedlings with accumulated betacyanin (C+B) were transferred to 14/10 h light/dark and used for chilling treatment 15 d later. Photosystem 2 (PS2) photochemistry, D1 protein content, and xanthophyll cycle during the chilling-induced photoinhibition (exposed to 5 °C at a moderate photon flux density of 500 µmol m-2 s-1 for 3 h) and the subsequent restoration were compared between the C+B seedlings and the control (C) ones. The maximal efficiency of PS2 photochemistry (Fv/Fm), the efficiency of excitation energy capture by open PS2 centres (Fv'/Fm'), and the yield of PS2 electron transport (ΦPS2) of the C+B and C leaves both decreased during photoinhibition. However, smaller decreases in Fv/Fm, Fv'/Fm', and ΦPS2 were observed in the C+B leaves than in C ones. At the same time, the deepoxidation state of xanthophyll cycle, indicated by (A+Z)/(V+A+Z) ratio, increased rapidly but the D1 protein content decreased considerably during the photoinhibition. The increase in rate of (A+Z)/(V+A+Z) was higher but the D1 protein turnover was slower in C+B than C leaves. After photoinhibition treatment, the plants were transferred to a dim irradiation (10 µmol m-2 s-1) at 25 °C for restoration. During restoration, the chlorophyll (Chl) fluorescence parameters, D1 protein content, and xanthophyll cycle components relaxed gradually, but the rate and level of restoration in the C+B leaves was greater than those in the C leaves. The addition of betacyanins to the thylakoid solution in vitro resulted in similar changes of Fv/Fm, D1 protein content, and (A+Z)/(V+A+Z) ratio during the chilling process. Therefore, betacyanin accumulation in S. salsa seedlings may result in higher resistance to photoinhibition, larger slowing down of D1 protein turnover, and enhancement of non-radiative energy dissipation associated with xanthophyll cycle, as well as in greater restoration after photoinhibition than in the control when subjected to chilling at moderate irradiance.

Photoinhibition and xanthophyll cycle activity in bayberry (Myrica rubra) leaves induced by high irradiance

The effect of high irradiance (HI, photosynthetically active photon flux density of 1 300 µmol m−2 s−1) on net photosynthetic rate (P N), chlorophyll fluorescence parameters, and xanthophyll cycle components were studied in fruit tree bayberry leaves. HI induced the photoinhibition and inactivation of photosystem 2 (PS2) reaction centres (RCs), which was characterized by decreased P N, maximum yield of fluorescence after dark adaptation (Fm), photochemical efficiency of PS2 (Fv/Fm) and quantum yield of PS2 (ΦPS2), and increased reduction state of QA (1-qP) and non-photochemical quenching (NPQ). Initial fluorescence (F0) showed a decrease after the first 2 h, and subsequently increased from the third hour exposure to HI. Furthermore, a greater increase in the ratio (Fi-F0)/(Fp-F0) which is an expression of the proportion of the QB non-reducing PS2 centres, whereas a remarked decrease in the slope of Fi to Fp which represents the rate of QA reduction was observed in leaves after HI exposure. Additionally, HI caused an increase in the pool size of the xanthophyll cycle pigments and sustained elevated contents of zeaxanthin (Z), antheraxanthin (A), and de-epoxidation state (DES) at the end of the irradiation period. During HI, decreased Fm, Fv/Fm, ΦPS2, NPQ, slope of Fi to Fp, V+A+Z, and DES, and increased F0, 1-qP, ratio (Fi-F0)/(Fp-F0), and V were observed in dithiothreitol (DTT)-fed leaves compared to control ones under the same conditions. Hence photoinhibition caused by HI in bayberry was probably attributed to inactivation of PS2 RCs, and photoprotection from photodamage were mainly related to the xanthophyll cycle-dependent heat dissipation in excess photons.

The Photoprotective Role of Epidermal Anthocyanins and Surface Pubescence in Young Leaves of Grapevine (Vitis vinifera)

Depending on cultivar, surfaces of young leaves of Vitis vinifera may be glabrous-green ('Soultanina') or transiently have anthocyanins ('Siriki') or pubescence ('Athiri'). A test is made of the hypothesis that anthocyanins and pubescence act as light screens affording a photoprotective advantage to the corresponding leaves, and an assessment is made of the magnitude of their effect. Measurements were made on young leaves of the three cultivars in spring under field conditions. Photosynthetic gas-exchange and in vivo chlorophyll fluorescence were measured. Photosynthetic and photoprotective pigments were analysed by HPLC. Compared with glabrous-green leaves, both anthocyanic and pubescent leaves had greater dark-adapted PSII photochemical efficiency and net photosynthesis. In leaves possessing either anthocyanins or pubescence, the ratio of xanthophyll cycle components to total chlorophyll, and mid-day de-epoxidation state of the xanthophyll cycle were considerably smaller, than in glabrous-green leaves. These differences were more evident in pubescent leaves, probably indicating that trichomes were more effective in decreasing light stress than anthocyanins in the epidermis. Light screens, especially in the form of pubescence, decrease the risk of photoinhibition whilst allowing leaves to maintain a smaller content of xanthophyll cycle components and depend less on xanthophyll cycle energy dissipation. This combination of photoprotective features, i.e. decreased photon flux to the photosynthetic apparatus and lower xanthophyll cycle utilization rates may be particularly advantageous under stressful conditions.

Diurnal variation of gas exchange, chlorophyll fluorescence, and xanthophyll cycle components of maize hybrids released in different years

Diurnal variation of gas exchange, chlorophyll (Chl) fluorescence, and xanthophyll cycle components of three maize (Zea mays L.) hybrids released in different years, i.e. Baimaya (1950s), Zhongdan2 (1970s), and Nongda108 (1990s), were compared. On cloudless days, the newer hybrids always had higher net photosynthetic rate (P N), especially at noon, than the older ones. At noon, all the hybrids decreased their maximal yield of photosystem 2 (PS2) photochemistry (Fv/Fm) and actual quantum yield of PS2 (ΦPS2), the newer ones always showing higher values. Generally, the newer hybrids displayed higher photochemical quenching of Chl (qP) and lower non-photochemical quenching (NPQ). The interhybrid differences in P N may be owing to their differential photochemical efficiency. A midday depression in P N occurred in all hybrids, which might be caused by serious photoinhibition or by decreased stomatal conductance. However, midday depression in P N was more obvious in the older hybrids, especially when leaves were senescent. The higher de-epoxidation state of the xanthophylls was noted in older hybrids, which was confirmed by their larger NPQ. The newer maize hybrids did not need a strong de-epoxidation state since they had a better photosynthetic quantum conversion rate and a lower NPQ.