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Melatonin (MT) is considered as an antistress molecule that plays a constructive role in the acclimation of plants to both biotic and abiotic stress conditions. In the present study, we assessed the impact of 10 and 100 μM MT foliar spray, on chlorophyll content, and photosystem II (PSII) function, under moderate drought stress, on oregano (Origanum vulgare L.) plants. Our aim was to elucidate the molecular mechanism of MT action on the photosynthetic electron transport process. Foliar spray with 100 μM MT was more effective in mitigating the negative impact of moderate drought stress on PSII function, compared to 10 μM MT. MT foliar spray significantly improved the reduced efficiency of the oxygen-evolving complex (OEC), and PSII photoinhibition (Fv/Fm), which were caused by drought stress. Under moderate drought stress, foliar spray with 100 μM MT, compared with the water sprayed (WA) leaves, increased the non-photochemical quenching (NPQ) by 31%, at the growth irradiance (GI, 205 μmol photons m−2 s−1), and by 13% at a high irradiance (HI, 1000 μmol photons m−2 s−1). However, the lower NPQ increase at HI was demonstrated to be more effective in decreasing the singlet-excited oxygen (1O2) production at HI (−38%), in drought-stressed oregano plants sprayed with 100 μM MT, than the corresponding decrease in 1O2 production at the GI (−20%), both compared with the respective WA-sprayed leaves under moderate drought. The reduced 1O2 production resulted in a significant increase in the quantum yield of PSII photochemistry (ΦPSII), and the electron transport rate (ETR), in moderate drought-stressed plants sprayed with 100 μM MT, compared with WA-sprayed plants, but only at the HI (+27%). Our results suggest that the enhancement of PSII functionality, with 100 μM MT under moderate drought stress, was initiated by the NPQ mechanism, which decreased the 1O2 production and increased the fraction of open PSII reaction centers (qp), resulting in an increased ETR. [ABSTRACT FROM AUTHOR]

In recent years, inorganic nanoparticles, including calcium hydroxide nanoparticles [Ca Ca(OH)2 NPs], have attracted significant interest for their ability to impact plant photosynthesis and boost agricultural productivity. In this study, the effects of 15 and 30 mg L−1 oleylamine-coated calcium hydroxide nanoparticles [Ca(OH)2@OAm NPs] on photosystem II (PSII) photochemistry were investigated on tomato plants at their growth irradiance (GI) (580 μmol photons m−2 s−1) and at high irradiance (HI) (1000 μmol photons m−2 s−1). Ca(OH)2@OAm NPs synthesized via a microwave-assisted method revealed a crystallite size of 25 nm with 34% w/w of oleylamine coater, a hydrodynamic size of 145 nm, and a ζ-potential of 4 mV. Compared with the control plants (sprayed with distilled water), PSII efficiency in tomato plants sprayed with Ca(OH)2@OAm NPs declined as soon as 90 min after the spray, accompanied by a higher excess excitation energy at PSII. Nevertheless, after 72 h, the effective quantum yield of PSII electron transport (ΦPSII) in tomato plants sprayed with Ca(OH)2@OAm NPs enhanced due to both an increase in the fraction of open PSII reaction centers (qp) and to the enhancement in the excitation capture efficiency (Fv'/Fm') of these centers. However, the decrease at the same time in non-photochemical quenching (NPQ) resulted in an increased generation of reactive oxygen species (ROS). It can be concluded that Ca(OH)2@OAm NPs, by effectively regulating the non-photochemical quenching (NPQ) mechanism, enhanced the electron transport rate (ETR) and decreased the excess excitation energy in tomato leaves. The delay in the enhancement of PSII photochemistry by the calcium hydroxide NPs was less at the GI than at the HI. The enhancement of PSII function by calcium hydroxide NPs is suggested to be triggered by the NPQ mechanism that intensifies ROS generation, which is considered to be beneficial. Calcium hydroxide nanoparticles, in less than 72 h, activated a ROS regulatory network of light energy partitioning signaling that enhanced PSII function. Therefore, synthesized Ca(OH)2@OAm NPs could potentially be used as photosynthetic biostimulants to enhance crop yields, pending further testing on other plant species. [ABSTRACT FROM AUTHOR]

Water deficit is the major stress factor magnified by climate change that causes the most reductions in plant productivity. Knowledge of photosystem II (PSII) response mechanisms underlying crop vulnerability to drought is critical to better understanding the consequences of climate change on crop plants. Salicylic acid (SA) application under drought stress may stimulate PSII function, although the exact mechanism remains essentially unclear. To reveal the PSII response mechanism of celery plants sprayed with water (WA) or SA, we employed chlorophyll fluorescence imaging analysis at 48 h, 96 h, and 192 h after watering. The results showed that up to 96 h after watering, the stroma lamellae of SA-sprayed leaves appeared dilated, and the efficiency of PSII declined, compared to WA-sprayed plants, which displayed a better PSII function. However, 192 h after watering, the stroma lamellae of SA-sprayed leaves was restored, while SA boosted chlorophyll synthesis, and by ameliorating the osmotic potential of celery plants, it resulted in higher relative leaf water content compared to WA-sprayed plants. SA, by acting as an antioxidant under drought stress, suppressed phototoxicity, thereby offering PSII photoprotection, together with enhanced effective quantum yield of PSII photochemistry (ΦPSII) and decreased quantity of singlet oxygen (1O2) generation compared to WA-sprayed plants. The PSII photoprotection mechanism induced by SA under drought stress was triggered by non-photochemical quenching (NPQ), which is a strategy to protect the chloroplast from photo-oxidative damage by dissipating the excess light energy as heat. This photoprotective mechanism, triggered by NPQ under drought stress, was adequate in keeping, especially in high-light conditions, an equal fraction of open PSII reaction centers (qp) as of non-stress conditions. Thus, under water deficit stress, SA activates a regulatory network of stress and light energy partitioning signaling that can mitigate, to an extent, the water deficit stress on PSII functioning. [ABSTRACT FROM AUTHOR]

Photosystem II (PSII) functions were investigated in basil (Ocimum basilicum L.) plants sprayed with 1 mM salicylic acid (SA) under non-stress (NS) or mild drought-stress (MiDS) conditions. Under MiDS, SA-sprayed leaves retained significantly higher (+36%) chlorophyll content compared to NS, SA-sprayed leaves. PSII efficiency in SA-sprayed leaves under NS conditions, evaluated at both low light (LL, 200 μmol photons m−2 s−1) and high light (HL, 900 μmol photons m−2 s−1), increased significantly with a parallel significant decrease in the excitation pressure at PSII (1-qL) and the excess excitation energy (EXC). This enhancement of PSII efficiency under NS conditions was induced by the mechanism of non-photochemical quenching (NPQ) that reduced singlet oxygen (1O2) production, as indicated by the reduced quantum yield of non-regulated energy loss in PSII (ΦNO). Under MiDS, the thylakoid structure of water-sprayed leaves appeared slightly dilated, and the efficiency of PSII declined, compared to NS conditions. In contrast, the thylakoid structure of SA-sprayed leaves did not change under MiDS, while PSII functionality was retained, similar to NS plants at HL. This was due to the photoprotective heat dissipation by NPQ, which was sufficient to retain the same percentage of open PSII reaction centers (qp), as in NS conditions and HL. We suggest that the redox status of the plastoquinone pool (qp) under MiDS and HL initiated the acclimation response to MiDS in SA-sprayed leaves, which retained the same electron transport rate (ETR) with control plants. Foliar spray of SA could be considered as a method to improve PSII efficiency in basil plants under NS conditions, at both LL and HL, while under MiDS and HL conditions, basil plants could retain PSII efficiency similar to control plants. [ABSTRACT FROM AUTHOR]

High light (HL) intensity has a substantial impact on light energy flow and partitioning within photosynthetic apparatus. To realize the impact of HL intensity on zinc (Zn) tolerance mechanisms in clary sage (Salvia sclarea L., Lamiaceae) plants, we examined the effect of the altered chlorophyll and nutrient uptake under excess Zn supply on the response mechanism of photosystem II (PSII) photochemistry. Eight-week-old clary sage plants were treated with 5 μM Zn (control) or 900 μM Zn in Hoagland nutrient solution. Leaf elemental analysis for Zn, Mn, Mg, and Fe was performed by inductively coupled plasma mass spectrometry (ICP-MS), whereas PSII functioning under HL was evaluated by chlorophyll fluorescence imaging analysis. Exposure of S. sclarea plants to 900 μM Zn increased leaf Zn accumulation and decreased leaf Mg and chlorophyll. The decreased non-photochemical quenching (NPQ) provided evidence of the photoprotection offered by the smaller light-harvesting antennae due to the reduced chlorophyll. The increased Mn after Zn exposure corresponded with higher efficiency of the oxygen-evolving complex (OEC) that was significantly correlated with the maximum efficiency of photosystem II (PSII) photochemistry (Fv/Fm). An increased electron transport rate (ETR) coincided with increased leaf Fe, which is known to play a vital role in the enzymes engaged in ETR. The decreased (32%) NPQ after an 8-day exposure to Zn caused an increased (10%) quantum yield of non-regulated energy loss in PSII (ΦNO), indicative of an increased singlet oxygen (1O2) production. It is suggested that the decreased NPQ induced acclimation responses of clary sage plants to HL and excess Zn by increasing 1O2 production. The reduced (18%) excess excitation energy (EXC) at PSII and the increased (24%) quantum yield of PSII photochemistry (ΦPSII) and ETR indicated improved photosynthetic efficiency under excess Zn and HL intensity. Therefore, the exposure of medicinal plants to excess Zn not only boosts their photosynthetic efficiency, enhancing crop yields, but can also improve Fe and Zn content, ameliorating the human health deficiency of these two essential micronutrients. [ABSTRACT FROM AUTHOR]

Melatonin (MT) is considered a new plant hormone having a universal distribution from prokaryotic bacteria to higher plants. It has been characterized as an antistress molecule playing a positive role in the acclimation of plants to stress conditions, but its impact on plants under non-stressed conditions is not well understood. In the current research, we evaluated the impact of MT application (10 and 100 μM) on photosystem II (PSII) function, reactive oxygen species (ROS) generation, and chlorophyll content on mint (Mentha spicata L.) plants in order to elucidate the molecular mechanism of MT action on the photosynthetic electron transport process that under non-stressed conditions is still unclear. Seventy-two hours after the foliar spray of mint plants with 100 μM MT, the improved chlorophyll content imported a higher amount of light energy capture, which caused a 6% increase in the quantum yield of PSII photochemistry (ΦPSII) and electron transport rate (ETR). Nevertheless, the spray with 100 μM MT reduced the efficiency of the oxygen-evolving complex (OEC), causing donor-side photoinhibition, with a simultaneous slight increase in ROS. Even so, the application of 100 μM MT decreased the excess excitation energy at PSII implying superior PSII efficiency. The decreased excitation pressure at PSII, after 100 μM MT foliar spray, suggests that MT induced stomatal closure through ROS production. The response of ΦPSII to MT spray corresponds to a J-shaped hormetic curve, with ΦPSII enhancement by 100 μM MT. It is suggested that the hormetic stimulation of PSII functionality was triggered by the non-photochemical quenching (NPQ) mechanism that stimulated ROS production, which enhanced the photosynthetic function. It is concluded that MT molecules can be used under both stress and non-stressed conditions as photosynthetic biostimulants for enhancing crop yields. [ABSTRACT FROM AUTHOR]

We recently proposed the use of engineered irregularly shaped zinc oxide nanoparticles (ZnO NPs) coated with oleylamine (OAm), as photosynthetic biostimulants, to enhance crop yield. In the current research, we tested newly engineered rod-shaped ZnO nanorods (NRs) coated with oleylamine (ZnO@OAm NRs) regarding their in vivo behavior related to photosynthetic function and reactive oxygen species (ROS) generation in tomato (Lycopersicon esculentum Mill.) plants. ZnO@OAm NRs were produced via solvothermal synthesis. Their physicochemical assessment revealed a crystallite size of 15 nm, an organic coating of 8.7% w/w, a hydrodynamic diameter of 122 nm, and a ζ-potential of −4.8 mV. The chlorophyll content of tomato leaflets after a foliar spray with 15 mg L−1 ZnO@OAm NRs presented a hormetic response, with an increased content 30 min after the spray, which dropped to control levels 90 min after the spray. Simultaneously, 90 min after the spray, the efficiency of the oxygen-evolving complex (OEC) decreased significantly (p < 0.05) compared to control values, with a concomitant increase in ROS generation, a decrease in the maximum efficiency of PSII photochemistry (Fv/Fm), a decrease in the electron transport rate (ETR), and a decrease in the effective quantum yield of PSII photochemistry (ΦPSII), indicating reduced PSII efficiency. The decreased ETR and ΦPSII were due to the reduced efficiency of PSII reaction centers (Fv'/Fm'). There were no alterations in the excess excitation energy at PSII or the fraction of open PSII reaction centers (qp). We discovered that rod-shaped ZnO@OAm NRs reduced PSII photochemistry, in contrast to irregularly shaped ZnO@OAm NPs, which enhanced PSII efficiency. Thus, the shape and organic coating of the nanoparticles play a critical role in the mechanism of their action and their impact on crop yield when they are used in agriculture. [ABSTRACT FROM AUTHOR]

Zinc oxide nanoparticles (ZnO NPs) have emerged as a prominent tool in agriculture. Since photosynthetic function is a significant measurement of phytotoxicity and an assessment tool prior to large-scale agricultural applications, the impact of engineered irregular-shaped ZnO NPs coated with oleylamine (ZnO@OAm NPs) were tested. The ZnO@OAm NPs (crystalline size 19 nm) were solvothermally prepared in the sole presence of oleylamine (OAm) and evaluated on tomato (Lycopersicon esculentum Mill.) photosystem II (PSII) photochemistry. Foliar-sprayed 15 mg L−1 ZnO@OAm NPs on tomato leaflets increased chlorophyll content that initiated a higher amount of light energy capture, which resulted in about a 20% increased electron transport rate (ETR) and a quantum yield of PSII photochemistry (ΦPSII) at the growth light (GL, 600 μmol photons m−2 s−1). However, the ZnO@OAm NPs caused a malfunction in the oxygen-evolving complex (OEC) of PSII, which resulted in photoinhibition and increased ROS accumulation. The ROS accumulation was due to the decreased photoprotective mechanism of non-photochemical quenching (NPQ) and to the donor-side photoinhibition. Despite ROS accumulation, ZnO@OAm NPs decreased the excess excitation energy of the PSII, indicating improved PSII efficiency. Therefore, synthesized ZnO@OAm NPs can potentially be used as photosynthetic biostimulants for enhancing crop yields after being tested on other plant species. [ABSTRACT FROM AUTHOR]

Most agricultural land, as a result of climate change, experiences severe stress that significantly reduces agricultural yields. Crop sensing by imaging techniques allows early-stage detection of biotic or abiotic stress to avoid damage and significant yield losses. Among the top certified imaging techniques for plant stress detection is chlorophyll a fluorescence imaging, which can evaluate spatiotemporal leaf changes, permitting the pre-symptomatic monitoring of plant physiological status long before any visible symptoms develop, allowing for high-throughput assessment. Here, we review different examples of how chlorophyll a fluorescence imaging analysis can be used to evaluate biotic and abiotic stress. Chlorophyll a is able to detect biotic stress as early as 15 min after Spodoptera exigua feeding, or 30 min after Botrytis cinerea application on tomato plants, or on the onset of water-deficit stress, and thus has potential for early stress detection. Chlorophyll fluorescence (ChlF) analysis is a rapid, non-invasive, easy to perform, low-cost, and highly sensitive method that can estimate photosynthetic performance and detect the influence of diverse stresses on plants. In terms of ChlF parameters, the fraction of open photosystem II (PSII) reaction centers (qp) can be used for early stress detection, since it has been found in many recent studies to be the most accurate and appropriate indicator for ChlF-based screening of the impact of environmental stress on plants. [ABSTRACT FROM AUTHOR]

Since drought stress is one of the key risks for the future of agriculture, exploring the molecular mechanisms of photosynthetic responses to water deficit stress is, therefore, fundamental. By using chlorophyll fluorescence imaging analysis, we evaluated the responses of photosystem II (PSII) photochemistry in young and mature leaves of Arabidopsis thaliana Col-0 (cv Columbia-0) at the onset of water deficit stress (OnWDS) and under mild water deficit stress (MiWDS) and moderate water deficit stress (MoWDS). Moreover, we tried to illuminate the underlying mechanisms in the differential response of PSII in young and mature leaves to water deficit stress in the model plant A. thaliana. Water deficit stress induced a hormetic dose response of PSII function in both leaf types. A U-shaped biphasic response curve of the effective quantum yield of PSII photochemistry (ΦPSII) in A. thaliana young and mature leaves was observed, with an inhibition at MiWDS that was followed by an increase in ΦPSII at MoWDS. Young leaves exhibited lower oxidative stress, evaluated by malondialdehyde (MDA), and higher levels of anthocyanin content compared to mature leaves under both MiWDS (+16%) and MoWDS (+20%). The higher ΦPSII of young leaves resulted in a decreased quantum yield of non-regulated energy loss in PSII (ΦNO), under both MiWDS (−13%) and MoWDS (−19%), compared to mature leaves. Since ΦNO represents singlet-excited oxygen (1O2) generation, this decrease resulted in lower excess excitation energy at PSII, in young leaves under both MiWDS (−10%) and MoWDS (−23%), compared to mature leaves. The hormetic response of PSII function in both young and mature leaves is suggested to be triggered, under MiWDS, by the intensified reactive oxygen species (ROS) generation, which is considered to be beneficial for activating stress defense responses. This stress defense response that was induced at MiWDS triggered an acclimation response in A. thaliana young leaves and provided tolerance to PSII when water deficit stress became more severe (MoWDS). We concluded that the hormesis responses of PSII in A. thaliana under water deficit stress are regulated by the leaf developmental stage that modulates anthocyanin accumulation in a stress-dependent dose. [ABSTRACT FROM AUTHOR]

Dramatic climate change has led to an increase in the intensity and frequency of drought episodes and, together with the high light conditions of the Mediterranean area, detrimentally influences crop production. Salicylic acid (SA) has been shown to supress phototoxicity, offering photosystem II (PSII) photoprotection. In the current study, we attempted to reveal the mechanism by which SA is improving PSII efficiency in oregano seedlings under moderate drought stress (MoDS). Foliar application of SA decreased chlorophyll content under normal growth conditions, but under MoDS increased chlorophyll content, compared to H2O-sprayed oregano seedlings. SA improved the PSII efficiency of oregano seedlings under normal growth conditions at high light (HL), and under MoDS, at both low light (LL) and HL. The mechanism by which, under normal growth conditions and HL, SA sprayed oregano seedlings compared to H2O-sprayed exhibited a more efficient PSII photochemistry, was the increased (17%) fraction of open PSII reaction centers (qp), and the increased (7%) efficiency of these open reaction centers (Fv′/Fm′), which resulted in an enhanced (24%) electron transport rate (ETR). SA application under MoDS, by modulating chlorophyll content, resulted in optimized antenna size and enhanced effective quantum yield of PSII photochemistry (ΦPSII) under both LL (7%) and HL (25%), compared to non-SA-sprayed oregano seedlings. This increased effective quantum yield of PSII photochemistry (ΦPSII) was due to the enhanced efficiency of the oxygen evolving complex (OEC), and the increased fraction of open PSII reaction centers (qp), which resulted in an increased electron transport rate (ETR) and a lower amount of singlet oxygen (1O2) production with less excess excitation energy (EXC). [ABSTRACT FROM AUTHOR]

Chromium [Cr(VI)] pollution is a major environmental risk, reducing crop yields. 5-Aminolevunic acid (5-ALA) considerably improves plant abiotic stress tolerance by inducing hydrogen peroxide (H2O2) and nitric oxide (NO) signalling. Our investigation aimed to uncover the mechanism of tomato tolerance to Cr(VI) toxicity through the foliar application of 5-ALA for three days, fifteen days before Cr treatment. Chromium alone decreased plant biomass and photosynthetic pigments, but increased oxidative stress markers, i.e., H2O2 and lipid peroxidation (as MDA equivalent). Electrolyte leakage (EL), NO, nitrate reductase (NR), phytochelatins (PCs), glutathione (GSH), and enzymatic and non-enzymatic antioxidants were also increased. Foliar application of 5-ALA before Cr treatment improved plant growth and photosynthetic pigments, diminished H2O2, MDA content, and EL, and resulted in additional enhancements of enzymatic and non-enzymatic antioxidants, NR activity, and NO synthesis. In Cr-treated tomato seedlings, 5-ALA enhanced GSH and PCs, which modulated Cr sequestration to make it nontoxic. 5-ALA-induced Cr tolerance was further enhanced by sodium nitroprusside (SNP), a NO donor. When sodium tungstate (ST), a NR inhibitor, was supplied together with 5-ALA to Cr-treated plants, it eliminated the beneficial effects of 5-ALA by decreasing NR activity and NO synthesis, while the addition of SNP inverted the adverse effects of ST. We conclude that the mechanism by which 5-ALA induced Cr tolerance in tomato seedlings is mediated by NR-generated NO. Thus, NR and NO are twin players, reducing Cr toxicity in tomato plants via antioxidant signalling cascades. [ABSTRACT FROM AUTHOR]

The consequences of climate change, and the increased accumulation of metalloids, like arsenic (As), in the environment, are significantly affecting crop performance and yield. Arsenic interferes with various plant biochemical and physiological processes, which result in diminished plant growth and development. Magnesium oxide nanoparticles (MgO-NPs) can improve plant growth and contribute to plant tolerance of heavy metal/metalloid toxicity. During current research, the efficacy of MgO-NPs was assessed for lessening arsenic (As) toxicity in soybean plants. In our experiment As uptake, plant growth, antioxidant enzyme activity, nutrient content, photochemical efficiency and photosynthetic performance were evaluated with/without exogenous application of 500 mg L−1 MgO-NPs in the presence/absence of 150 µM As in soybean plants. Foliar application of MgO-NPs, in the presence of As, enhanced plant height and dry weight by 17% and 15% respectively, and improved net photosynthetic rate by 12.9%, stomatal conductance by 13.4%, intercellular CO2 concentration by 15.3% and transpiration rate by 14.7%, as well as nutrient uptake and photosystem II (PSII) efficiency. In contrast, it decreased As uptake and oxidative stress as evaluated with hydrogen peroxide (H2O2) and lipid peroxidation (MDA). Hence, field tests may be implemented to formulate MgO-NPs use in agriculture, in order to obtain sustainable crop production in arsenic-contaminated soils. [ABSTRACT FROM AUTHOR]

Drought, the major limiting factor for plant growth and crop productivity, affecting several physiological and biochemical processes, is expected to increase in duration, intensity, and frequency as a consequence of climate change. Plants have developed several approaches to either avoid or tolerate water deficit. Plants as a response to drought stress (DS), close stomata, reducing carbon dioxide (CO2) entry in the leaf, thus decreasing photosynthesis which results in reduced synthesis of essential organic molecules that sustain the life on earth. The reduced CO2 fixation, decreases electron transport rate (ETR), while the absorbed light energy overdoes what can be used for photochemistry resulting in excess reactive oxygen species (ROS) and oxidative stress. Current imaging techniques allow non-destructive monitoring of changes in the physiological state of plants under DS. Thermographic visualization, near-infrared imaging, and chlorophyll a fluorescence imaging are the most common verified imaging techniques for detecting stress-related changes in the display of light emission from plant leaves. Chlorophyll a fluorescence analysis, by use of the pulse amplitude modulation (PAM) method, can principally calculate the amount of absorbed light energy that is directed for photochemistry in photosystem II (PSII) (ΦPSII), dissipated as heat (ΦNPQ), or dissipated by the non-radiative fluorescence processes (ΦNO). The method of chlorophyll a fluorescence imaging analysis by providing colour pictures of the whole leaf PSII photochemistry, can successfully identify the early drought stress warning signals. Its implementation allowed visualization of the leaf spatial photosynthetic heterogeneity and discrimination between mild drought stress (MiDS), moderate drought stress (MoDS), and severe drought stress (SDS). The fraction of open reaction centers of PSII (qp) is suggested as the most sensitive and suitable indicator of an early drought stress warning and also for selecting drought tolerant cultivars. [ABSTRACT FROM AUTHOR]

Exposure of Salvia sclarea plants to excess Zn for 8 days resulted in increased Ca, Fe, Mn, and Zn concentrations, but decreased Mg, in the aboveground tissues. The significant increase in the aboveground tissues of Mn, which is vital in the oxygen-evolving complex (OEC) of photosystem II (PSII), contributed to the higher efficiency of the OEC, and together with the increased Fe, which has a fundamental role as a component of the enzymes involved in the electron transport process, resulted in an increased electron transport rate (ETR). The decreased Mg content in the aboveground tissues contributed to decreased chlorophyll content that reduced excess absorption of sunlight and operated to improve PSII photochemistry (ΦPSII), decreasing excess energy at PSII and lowering the degree of photoinhibition, as judged from the increased maximum efficiency of PSII photochemistry (Fv/Fm). The molecular mechanism by which Zn-treated leaves displayed an improved PSII photochemistry was the increased fraction of open PSII reaction centers (qp) and, mainly, the increased efficiency of the reaction centers (Fv′/Fm′) that enhanced ETR. Elemental bioimaging of Zn and Ca by laser ablation–inductively coupled plasma–mass spectrometry (LA–ICP–MS) revealed their co-localization in the mid-leaf veins. The high Zn concentration was located in the mid-leaf-vein area, while mesophyll cells accumulated small amounts of Zn, thus resembling a spatiotemporal heterogenous response and suggesting an adaptive strategy. These findings contribute to our understanding of how exposure to excess Zn triggered a hormetic response of PSII photochemistry. Exposure of aromatic and medicinal plants to excess Zn in hydroponics can be regarded as an economical approach to ameliorate the deficiency of Fe and Zn, which are essential micronutrients for human health. [ABSTRACT FROM AUTHOR]

Clary sage (Salvia sclarea L.) is a medicinal plant that has the potential to be used for phytoextraction of zinc (Zn) and cadmium (Cd) from contaminated soils by accumulating these metals in its tissues. Additionally, it has been found to be more tolerant to excess Zn than to Cd stress alone; however, the interactive effects of the combined treatment with Zn and Cd on this medicinal herb, and the protective strategies of Zn to alleviate Cd toxicity have not yet been established in detail. In this study, clary sage plants grown hydroponically were simultaneously exposed to Zn (900 µM) and Cd (100 μM) for 8 days to obtain more detailed information about the plant responses and the role of excess Zn in mitigating Cd toxicity symptoms. The leaf anatomy, photosynthetic pigments, total phenolic and anthocyanin contents, antioxidant capacity (by DPPH and FRAP analyses), and the uptake and distribution of essential elements were investigated. The results showed that co-exposure to Zn and Cd leads to an increased leaf content of Fe and Mg compared to the control, and to increased leaf Ca, Mn, and Cu contents compared to plants treated with Cd only. This is most likely involved in the defense mechanisms of excess Zn against Cd toxicity to protect the chlorophyll content and the functions of both photosystems and the oxygen-evolving complex. The data also revealed that the leaves of clary sage plants subjected to the combined treatment have an increased antioxidant capacity attributed to the higher content of polyphenolic compounds. Furthermore, light microscopy indicated more alterations in the leaf morphology after Cd-only treatment than after the combined treatment. The present study shows that excess Zn could mitigate Cd toxicity in clary sage plants. [ABSTRACT FROM AUTHOR]

Salicylic acid (SA), an essential plant hormone, has received much attention due to its role in modulating the adverse effects of biotic and abiotic stresses, acting as an antioxidant and plant growth regulator. However, its role in photosynthesis under non stress conditions is controversial. By chlorophyll fluorescence imaging analysis, we evaluated the consequences of foliar applied 1 mM SA on photosystem II (PSII) efficiency of tomato (Solanum lycopersicum L.) plants and estimated the reactive oxygen species (ROS) generation. Tomato leaves sprayed with 1 mM SA displayed lower chlorophyll content, but the absorbed light energy was preferentially converted into photochemical energy rather than dissipated as thermal energy by non-photochemical quenching (NPQ), indicating photoprotective effects provided by the foliar applied SA. This decreased NPQ, after 72 h treatment by 1 mM SA, resulted in an increased electron transport rate (ETR). The molecular mechanism by which the absorbed light energy was more efficiently directed to photochemistry in the SA treated leaves was the increased fraction of the open PSII reaction centers (qp), and the increased efficiency of open reaction centers (Fv'/Fm'). SA induced a decrease in chlorophyll content, resulting in a decrease in non-regulated energy dissipated in PSII (ΦNO) under high light (HL) treatment, suggesting a lower amount of triplet excited state chlorophyll (3Chl*) molecules available to produce singlet oxygen (1O2). Yet, the increased efficiency, compared to the control, of the oxygen evolving complex (OEC) on the donor side of PSII, associated with lower formation of hydrogen peroxide (H2O2), also contributed to less creation of ROS. We conclude that under non stress conditions, foliar applied SA decreased chlorophyll content and suppressed phototoxicity, offering PSII photoprotection; thus, it can be regarded as a mechanism that reduces photoinhibition and photodamage, improving PSII efficiency in crop plants. [ABSTRACT FROM AUTHOR]

Simple Summary: Potato is one of the most universally cultivated horticultural crops and is vulnerable to a range of herbivorous insects. One of them is the brown marmorated stink bug, an invasive polyphagous sap-sucking agricultural insect pest that penetrates the phloem to sieve elements and removes sap via a specialized mouthpart, the stylet. By using the chlorophyll fluorescence imaging methodology, we examined potato photosystem II (PSII) photochemistry responses in the area of feeding on the whole leaf area. Highly increased reactive oxygen species (ROS) generation was observed as rapidly as 3 min after feeding to initiate defence responses and can be considered the primary plant defence response mechanism against herbivores. Our experimental results confirmed that chlorophyll fluorescence imaging methodology can detect spatial heterogeneity of PSII efficiency at the whole leaf surface and is a promising tool for investigating plant response mechanisms of sap-sucking insect herbivores. We suggest that PSII responses to insect feeding underlie ROS-dependent signalling. We conclude that the potato PSII response mechanism to sap-sucking insect herbivores is described by the induction of the defence response to reduce herbivory damage, instead of induction of tolerance, through a compensatory photosynthetic response mechanism that is observed after chewing insect feeding. 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 represents 1O2 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. [ABSTRACT FROM AUTHOR]

Bisphenol A (BPA), an intermediate chemical used for synthesizing polycarbonate plastics, has now become a wide spread organic pollutant. It percolates from a variety of sources, and plants are among the first organisms to encounter, absorb, and metabolize it, while its toxic effects are not yet fully known. Therefore, we experimentally studied the effects of aqueous BPA solutions (50 and 100 mg L−1, for 6, 12, and 24 h) on photosystem II (PSII) functionality and evaluated the role of reactive oxygen species (ROS) on detached leaves of the model plant Arabidopsis thaliana. Chlorophyll fluorescence imaging analysis revealed a spatiotemporal heterogeneity in the quantum yields of light energy partitioning at PSII in Arabidopsis leaves exposed to BPA. Under low light PSII function was negatively influenced only at the spot-affected BPA zone in a dose- and time-dependent manner, while at the whole leaf only the maximum photochemical efficiency (F v /F m) was negatively affected. However, under high light all PSII photosynthetic parameters measured were negatively affected by BPA application, in a time-dependent manner. The affected leaf areas by the spot-like mode of BPA action showed reduced chlorophyll autofluorescence and increased accumulation of hydrogen peroxide (H2O2). When H2O2 was scavenged via N-acetylcysteine under BPA exposure, PSII functionality was suspended, while H2O2 scavenging under non-stress had more detrimental effects on PSII function than BPA alone. It can be concluded that the necrotic death-like spots under BPA exposure could be due to ROS accumulation, but also H2O2 generation seems to play a role in the leaf response against BPA-related stress conditions. [ABSTRACT FROM AUTHOR]

By analyzing chlorophyll fluorescence data, Guo et al., revealed that gliotoxin (GT), a fungal secondary metabolite, affects both PSII electron transport at the acceptor side, and the reduction rate of PSI end electron acceptors' pool. Chlorophyll fluorescence imaging analysis for elucidating the mechanism of photosystem II acclimation to cadmium exposure in the hyperaccumulating plant Noccaea caerulescens. Bisphenol A (BPA), an intermediate chemical used for synthesizing plastic materials, has a spot-like mode of action on I Arabidopsis thaliana i leaves that was revealed by Adamakis et al., using chlorophyll fluorescence imaging analysis. Keywords: photochemical efficiency; photosynthetic heterogeneity; reactive oxygen species; environmental stress; energy partitioning; oxidative stress; photosystem II; non-photochemical quenching EN photochemical efficiency photosynthetic heterogeneity reactive oxygen species environmental stress energy partitioning oxidative stress photosystem II non-photochemical quenching N.PAG N.PAG 3 05/17/21 20210414 NES 210414 Chlorophyll I a i fluorescence that results from the absorbed light energy can be interpreted in terms of photosynthetic activity to obtain information about the state of photosynthetic apparatus and especially of photosystem II (PSII) (Krause and Weis, [11]; Murchie and Lawson, [17]). [Extracted from the article]

Photosynthetic activity, oxidative stress, and Cu bioaccumulation in the seagrass Cymodocea nodosa were assessed 4, 12, 24, 48, and 72 h after exposure to two copper oxide nanoparticle (CuO NP) concentrations (5 and 10 mg L). CuO NPs were characterized by scanning electron microscopy (SEM) and dynamic light scattering measurements (DLS). Chlorophyll fluorescence analysis was applied to detect photosystem II (PSII) functionality, while the Cu accumulation kinetics into the leaf blades was fitted to the Michaelis-Menten equation. The uptake kinetics was rapid during the first 4 h of exposure and reached an equilibrium state after 10 h exposure to 10 mg L and after 27 h to 5 mg L CuO NPs. As a result, 4-h treatment with 5 mg L CuO NPs, decreased the quantum yield of PS II photochemistry (Φ) with a parallel increase in the regulated non-photochemical energy loss in PSII (Φ). However, the photoprotective dissipation of excess absorbed light energy as heat, through the process of non-photochemical quenching (NPQ), did not maintain the same fraction of open reaction centers ( q ) as in control plants. This reduced number of open reaction centers resulted in a significant increase of HO production in the leaf veins serving possibly as an antioxidant defense signal. Twenty-four-hour treatment had no significant effect on Φ and q compared to controls. However, 24 h exposure to 5 mg L CuO NPs increased the quantum yield of non-regulated energy loss in PSII (Φ), and thus the formation of singlet oxygen (O) via the triplet state of chlorophyll, possible because the uptake kinetics had not yet reached the equilibrium state as did 10 mg L. Longer-duration treatment (48 and 72 h) had less effect on the allocation of absorbed light energy at PSII and the fraction of open reaction centers, compared to 4-h treatment, suggesting the function of a stress defense mechanism. The response of C. nodosa leaves to CuO NPs fits the 'Threshold for Tolerance Model' with a threshold time (more than 4 h) required for induction of a stress defense mechanism, through HO production. [ABSTRACT FROM AUTHOR]

A population of the metallophyte Noccaea ( Thlaspi) caerulescens originating from a Zn-enriched area at Røros Copper Mine (Norway) was studied. N. caerulescens tolerance to accumulate Cd and Zn was evaluated in hydroponic experiments by chlorophyll fluorescence imaging analysis. In the field-collected N. caerulescens mother plants, Zn shoot concentrations were above Zn hyperaccumulation threshold while, in hydroponic experiments under 40-μM Cd exposure, shoot Cd concentrations were clearly above Cd hyperaccumulation threshold. Cadmium ions and, to a less extent, Zn were mainly retained in the roots. Exposure to Cd enhanced Zn translocation to the shoot, while decreased significant total Ca uptake, suggesting that Cd uptake occurs through Ca transporters. Nevertheless, it increased Ca translocation to the leaf, possibly for photoprotection of photosystem II (PSII). Exposure to 800 μM Zn or 40 μM Cd resulted in increased Fe uptake suggesting that in N. caerulescens, Cd uptake does not take place through the pathway of Fe uptake and that conditions that lead to Cd and Zn accumulation in plants may also favor Fe accumulation. Despite the significant high toxicity levels of Zn and Cd in leaves, under Zn and Cd exposure, respectively, the allocation of absorbed light energy at PSII did not differ compared to controls. The results showed that N. caerulescens keep Cd and Zn concentrations in the mesophyll cells in non-toxic forms for PSII and that the increased Ca and Fe accumulation in leaves alleviates the toxicity effects. Chlorophyll fluorescence imaging revealed that PSII of N. caerulescens resisted better the phytotoxic effects of 20 times higher Zn than Cd exposure concentration. Overall, it is concluded that the use of chlorophyll fluorescence imaging constitutes a promising basis for investigating heavy metal tolerance of plants. [ABSTRACT FROM AUTHOR]

Exposure of Arabidopsis thaliana young and mature leaves to the herbicide paraquat (Pq) resulted in a localized increase of hydrogen peroxide (H2O2) in the leaf veins and the neighboring mesophyll cells, but this increase was not similar in the two leaf types. Increased H2O2 production was concomitant with closed reaction centers (qP). Thirty min after Pq exposure despite the induction of the photoprotective mechanism of non-photochemical quenching (NPQ) in mature leaves, H2O2 production was lower in young leaves mainly due to the higher increase activity of ascorbate peroxidase (APX). Later, 60 min after Pq exposure, the total antioxidant capacity of young leaves was not sufficient to scavenge the excess reactive oxygen species (ROS) that were formed, and thus, a higher H2O2 accumulation in young leaves occurred. The energy allocation of absorbed light in photosystem II (PSII) suggests the existence of a differential photoprotective regulatory mechanism in the two leaf types to the time-course Pq exposure accompanied by differential antioxidant protection mechanisms. It is concluded that tolerance to Pq-induced oxidative stress is related to the redox state of quinone A (QA). [ABSTRACT FROM AUTHOR]