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Salvia sclarea L. is a Cd2+ tolerant medicinal herb with antifungal and antimicrobial properties cultivated for its pharmacological properties. However, accumulation of high Cd2+ content in its tissues increases the adverse health effects of Cd2+ in humans. Therefore, there is a serious demand to lower human Cd2+ intake. The purpose of our study was to evaluate the mitigative role of excess Zn2+ supply to Cd2+ uptake/translocation and toxicity in clary sage. Salvia plants were treated with excess Cd2+ (100 µM CdSO4 ) alone, and in combination with Zn2+ (900 µM ZnSO4 ), in modified Hoagland nutrient solution. The results demonstrate that S. sclarea plants exposed to Cd2+ toxicity accumulated a significant amount of Cd2+ in their tissues, with higher concentrations in roots than in leaves. Cadmium exposure enhanced total Zn2+ uptake but also decreased its translocation to leaves. The accumulated Cd2+ led to a substantial decrease in photosystem II (PSII) photochemistry and disrupted the chloroplast ultrastructure, which coincided with an increased lipid peroxidation. Zinc application decreased Cd2+ uptake and translocation to leaves, while it mitigated oxidative stress, restoring chloroplast ultrastructure. Excess Zn2+ ameliorated the adverse effects of Cd2+ on PSII photochemistry, increasing the fraction of energy used for photochemistry (ΦPSII ) and restoring PSII redox state and maximum PSII efficiency (Fv/Fm), while decreasing excess excitation energy at PSII (EXC). We conclude that excess Zn2+ application eliminated the adverse effects of Cd2+ toxicity, reducing Cd2+ uptake and translocation and restoring chloroplast ultrastructure and PSII photochemical efficiency. Thus, excess Zn2+ application can be used as an important method for low Cd2+-accumulating crops, limiting Cd2+ entry into the food chain. © 2022 by the authors. Licensee MDPI, Basel, Switzerland.

Potato, Solanum tuberosum L., one of the most commonly cultivated horticultural crops throughout the world, is susceptible to a variety of herbivory insects. In the present study, we evaluated the consequence of feeding by the sap-sucking insect Halyomorpha halys on potato leaf photosynthetic efficiency. By using chlorophyll fluorescence imaging methodology, we examined photosystem II (PSII) photochemistry in terms of feeding and at the whole leaf area. The role of reactive oxygen species (ROS) in potato’s defence response mechanism immediately after feeding was also assessed. Even 3 min after feeding, increased ROS generation was observed to diffuse through the leaf central vein, probably to act as a long-distance signalling molecule. The proportion of absorbed energy being used in photochemistry (ΦPSII ) at the whole leaf level, after 20 min of feeding, was reduced by 8% compared to before feeding due to the decreased number of open PSII reaction centres (qp). After 90 min of feeding, ΦPSII decreased by 46% at the whole leaf level. Meanwhile, at the feeding zones, which were located mainly in the proximity of the leaf midrib, ΦPSII was lower than 85%, with a concurrent increase in singlet-excited oxygen (1O2 ) generation, which is considered to be harmful. However, the photoprotective mechanism (ΦNPQ), which was highly induced 90 min after feeding, was efficient to compensate for the decrease in the quantum yield of PSII photochemistry (ΦPSII ). Therefore, the quantum yield of non-regulated energy loss in PSII (ΦNO), which represents1O2 generation, remained unaffected at the whole leaf level. We suggest that the potato PSII response to sap-sucking insect feeding underlies the ROS-dependent signalling that occurs immediately and initiates a photoprotective PSII defence response to reduce herbivory damage. A controlled ROS burst can be considered the primary plant defence response mechanism to herbivores. © 2022 by the authors. Licensee MDPI, Basel, Switzerland.

The herbal plant Salvia sclarea L. (clary sage) is classified to cadmium (Cd) accumulators and considered as a potential plant for phytoremediation of heavy metal polluted soil. However, the effect of Cd only treatment on the function of the photosynthetic apparatus of S. sclarea, as well as the mechanisms involved in Cd tolerance have not yet been studied in detail. This study was conducted to examine the integrative responses of S. sclarea plants exposed to a high Cd supply (100 µM) for 3 and 8 days by investigating element nutrient uptake, oxidative stress markers, pigment composition, photosynthetic performance and leaf structure. Measurements of the functional activities of photosystem I (PSI, by P700 photooxidation), photosystem II (PSII, by chlorophyll fluorescence parameters), the oxygen-evolving complex (oxygen evolution by Joliot- and Clark-type electrodes), as well as the leaf pigment and phenolic contents, were used to evaluate the protective mechanisms of the photosynthetic apparatus under Cd stress. Data suggested that the molecular mechanisms included in the photosynthetic tolerance to Cd toxicity involve strongly increased phenolic and anthocyanin contents, as well as an increased non-photochemical quenching and accelerated cyclic electron transport around PSI up to 61%, which protect the function of the photosynthetic apparatus under stress. Furthermore, the tolerance of S. sclarea to Cd stress is also associated with increased accumulation of Fe in leaves by 25%. All the above, clearly suggest that S. sclarea plants employ several different mechanisms to protect the function of the photosynthetic apparatus against Cd stress, which are discussed here. © 2021 The Authors

In recent years, due to the development of industrial and agricultural production, heavy metal contamination has attracted increasing attention. Aromatic and medicinal plant Salvia sclarea L. (clary sage) is classified to zinc (Zn) accumulators and considered as a potential plant for the phytore-mediation of heavy metal polluted soils. In this study, an adaptation of clary sage to 900 µM (excess) Zn exposure for eight days in a hydroponic culture was investigated. The tolerance mechanisms under excess Zn exposure were assessed by evaluating changes in the nutrient uptake, leaf pigment and phenolic content, photosynthetic activity and leaf structural characteristics. The uptake and the distribution of Zn, as well as some essential elements such as: Ca, Mg, Fe, Mn and Cu, were examined by inductively coupled plasma mass spectrometry. The results revealed that Salvia sclarea is a Zn-accumulator plant that tolerates significantly high toxic levels of Zn in the leaves by increasing the leaf contents of Fe, Ca and Mn ions to protect the photosynthetic function and to stimulate the photosystem I (PSI) and photosystem II (PSII) activities. The exposure of clary sage to excess Zn significantly increased the synthesis of total phenolics and anthocyanins in the leaves; these play an important role in Zn detoxification and protection against oxidative stress. The lipid peroxidation and electrolyte leakage in leaves, used as clear indicators for heavy metal damage, were slightly increased. All these data highlight that Salvia sclarea is an economically interesting plant for the phytoextraction and/or phytostabilization of Zn-contaminated soils. © 2021 by the authors. Licensee MDPI, Basel, Switzerland.

Five-day exposure of clary sage (Salvia sclarea L.) to 100 μM cadmium (Cd) in hydroponics was sufficient to increase Cd concentrations significantly in roots and aboveground parts and affect negatively whole plant levels of calcium (Ca) and magnesium (Mg), since Cd competes for Ca chan-nels, while reduced Mg concentrations are associated with increased Cd tolerance. Total zinc (Zn), copper (Cu), and iron (Fe) uptake increased but their translocation to the aboveground parts de-creased. Despite the substantial levels of Cd in leaves, without any observed defects on chloroplast ultrastructure, an enhanced photosystem II (PSII) efficiency was observed, with a higher fraction of absorbed light energy to be directed to photochemistry (ΦPSΙΙ). The concomitant increase in the pho-toprotective mechanism of non-photochemical quenching of photosynthesis (NPQ) resulted in an important decrease in the dissipated non-regulated energy (ΦNO), modifying the homeostasis of reactive oxygen species (ROS), through a decreased singlet oxygen (1O2) formation. A basal ROS level was detected in control plant leaves for optimal growth, while a low increased level of ROS under 5 days Cd exposure seemed to be beneficial for triggering defense responses, and a high level of ROS out of the boundaries (8 days Cd exposure), was harmful to plants. Thus, when clary sage was exposed to Cd for a short period, tolerance mechanisms were triggered. However, exposure to a combination of Cd and high light or to Cd alone (8 days) resulted in an inhibition of PSII function-ality, indicating Cd toxicity. Thus, the rapid activation of PSII functionality at short time exposure and the inhibition at longer duration suggests a hormetic response and describes these effects in terms of “adaptive response” and “toxicity”, respectively. © 2020 by the authors. Licensee MDPI, Basel, Switzerland.

The role of anthocyanin accumulation in poinsettia leaves exposed to photo-oxidative stress was evaluated by comparing green (anthocyanin less) and reddish (anthocyanin well equipped) leaves, co-occurring in poinsettia (Euphorbia pulcherrima Willd. ex Klotzsch). For the assessment we compared the photoprotective and antioxidant mechanisms in the two leaf types in response to photo-oxidative stress, generated in chloroplasts by the application of methyl viologen (MV). MV accepts electrons from photosystem I (PSI) with subsequent transfer to molecular oxygen to produce superoxide anions (O2 [rad] –) that are converted by the chloroplast superoxide dismutase (SOD) to hydrogen peroxide (H2O2) that is reduced by ascorbate peroxidase (APX) to water and oxygen. At 90ömin after MV application, the decrease in the activity of the H2O2-scavenging enzyme APX resulted in increased H2O2 levels in both leaf types, but to a significantly lower level in the reddish leaves, possibly due to the significantly higher anthocyanin and phenolic content that are considered as H2O2-scavengers. Reddish poinsettia leaves having higher antioxidant activity possessed more effective photoprotective and reactive oxygen species (ROS)-scavenging mechanisms, including higher enzymatic (SOD and APX), and non-enzymatic (anthocyanin and phenolic) content over the green ones. Anthocyanin well equipped poinsettia leaves were in advantage in response to photo-oxidative stress over the anthocyanin less ones that appeared not to be equally protected. This superiority in the avoidance of the photo-oxidative stress by the reddish leaves was also associated to a higher effective quantum yield of PSII photochemistry (ΦPSII), a lower excitation pressure (1- qp) and a lower H2O2 generation, compared to their green counterparts. © 2020 Elsevier B.V.

Young and mature leaves of Arabidopsis thaliana were exposed by foliar spray to 30 mg L-1 of CuZn nanoparticles (NPs). The NPs were synthesized by a microwave-assisted polyol process and characterized by dynamic light scattering (DLS), X-ray diffraction (XRD), and transmission electron microscopy (TEM). CuZn NPs effects in Arabidopsis leaves were evaluated by chlorophyll fluorescence imaging analysis that revealed spatiotemporal heterogeneity of the quantum efficiency of PSII photochemistry (ϕPSII) and the redox state of the plastoquinone (PQ) pool (qp), measured 30 min, 90 min, 180 min, and 240 min after spraying. Photosystem II (PSII) function in young leaves was observed to be negatively influenced, especially 30 min after spraying, at which point increased H2O2 generation was correlated to the lower oxidized state of the PQ pool. Recovery of young leaves photosynthetic efficiency appeared only after 240 min of NPs spray when also the level of ROS accumulation was similar to control leaves. On the contrary, a beneficial effect on PSII function in mature leaves after 30 min of the CuZn NPs spray was observed, with increased FPSII, an increased electron transport rate (ETR), decreased singlet oxygen (1O2) formation, and H2O2 production at the same level of control leaves.An explanation for this differential response is suggested. © 2019 by the authors.

In this study, for a first time (according to our knowledge), we couple the methodologies of chlorophyll fluorescence imaging analysis (CF-IA) and laser ablation inductively coupled plasma mass spectrometry (LA-ICP-MS), in order to investigate the effects of cadmium (Cd) accumulation on photosystem II (PSII) photochemistry. We used as plant material Salvia sclarea that grew hydroponically with or without (control) 100 μM Cd for five days. The spatial heterogeneity of a decreased effective quantum yield of electron transport (ΦPSII) that was observed after exposure to Cd was linked to the spatial pattern of high Cd accumulation. However, the high increase of non-photochemical quenching (NPQ), at the leaf part with the high Cd accumulation, resulted in the decrease of the quantum yield of non-regulated energy loss (ΦNO) even more than that of control leaves. Thus, S. sclarea leaves exposed to 100 μM Cd exhibited lower reactive oxygen species (ROS) production as singlet oxygen (1O2). In addition, the increased photoprotective heat dissipation (NPQ) in the whole leaf under Cd exposure was sufficient enough to retain the same fraction of open reaction centers (qp) with control leaves. Our results demonstrated that CF-IA and LA-ICP-MS could be successfully combined to monitor heavy metal effects and plant tolerance mechanisms. © 2019 by the authors.

Some elemental levels, morphological and photosynthetic characteristics and cellular metabolites of wheat plants (Triticum aestivum L. cv. Vergina) growing in the field on an ore body (Cu concentration in the soil 3050 mug g-1) were compared to control plants growing in the same environment where the Cu concentration in the soil was 140 mug g-1. The concentrations of Cu, K, Pb, Zn, Mg and Fe were higher in the ore plants but Ca was lower. Growth of the ore plants was inhibited. with decreased height (25%), weight (5%), leaf area (7%) and leaf dry weight (5%) compared to the control plants. Leaf protein concentration of the ore plants was 16.2 mg cm-2 leaf area, 63% of that of the control plants. The ore plants were chlorotic and chlorophyll concentration was 3.8 mug cm-2 leaf area, 6.4 times lower than that in the control plants. Ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco, EC 4.1.1.39) activity was 13.4 mumol CO2 fixed (mg dry weight)-1 min-1, 164% of the activity in control plants. Therefore, growth inhibition did not appear to result from inhibition of the dark reactions of photosynthesis. High Rubisco activity appears to be maintained to permit maximal CO2 fixation rates whenever energy is available. Parameters of chlorophyll fluorescence F(m), F(v), t1/2 and F(v)/F(m) were lower in the ore-grown plants than in control plants; F0 was higher. These values indicate that there is a decrease in the pool size of the electron acceptors on the reducing side of photosystem II (PSII), a destruction of PSII centres and interference in the photochemistry of PSII. The nitrogen content, soluble sugars, starch and lipid content were lower in ore-grown plants. Lower carbohydrate levels appear to result from low photosynthetic activity. The fatty acid composition of lipids was similar in both groups. A lower proportion of polyunsaturated fatty acids was found in the ore-grown plants. Stress caused by high Cu concentration in the soil appears to affect the light reactions of photosynthesis leading to growth inhibition.