Photosystem II Photochemistry Research Articles

Photosystem II Tolerance to Excess Zinc Exposure and High Light Stress in Salvia sclarea L.

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.

Combining PSII photochemistry and hydraulics improves predictions of photosynthesis and water use from mild to lethal drought.

Rising temperatures and increases in drought negatively impact the efficiency and sustainability of both agricultural and forest ecosystems. Although hydraulic limitations on photosynthesis have been extensively studied, a solid understanding of the links between whole plant hydraulics and photosynthetic processes at the cellular level under changing environmental conditions is still missing, hampering our predictive power for plant mortality. Here, we examined plant hydraulic traits and CO2 assimilation rate under progressive water limitation by implementing Photosystem II (PSII) dynamics with a whole plant process model (TREES). The photosynthetic responses to plant water status were parameterized based on measurements of chlorophyll a fluorescence, gas exchangeand water potential for Brassica rapa (R500) grown in a greenhouse under fully watered to lethal drought conditions. The updated model significantly improved predictions of photosynthesis, stomatal conductanceand leaf water potential. TREES with PSII knowledge predicted a larger hydraulic safety margin and a decrease in percent loss of conductivity. TREES predicted a slower decrease in leaf water potential, which agreed with measurements. Our results highlight the pressing need for incorporating PSII drought photochemistry into current process models to capture cross-scale plant water dynamics from cell to whole plant level.

Mechanistic Approach on Melatonin-Induced Hormesis of Photosystem II Function in the Medicinal Plant Mentha spicata

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.

Impact of Coated Zinc Oxide Nanoparticles on Photosystem II of Tomato Plants.

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.

Evaluating photosynthetic performance of transgenic soybean harboring CKX13 under salinity conditions

Cytokinins (CKs) are considered one of the critical phytohormones with an important role in plant response to abiotic stresses. One of the key enzymes in regulating CK levels is CK oxidase/dehydrogenase (CKX), which catalyzes the irreversible CK degradation. Although CKX gene members in soybean (Glycine max) have been identified, their importance in plant tolerance against salinity has not yet been fully elucidated. In this study, GmCKX13, a gene with upregulated expression in various tissues of soybean at various stages under water deficit condition as shown in previous investigation, was selected for exploring its role in relation to photosynthetic performance under salt stress conditions. The results showed that the transgenic soybean overexpressing GmCKX13 could maintain a better assimilation rate compared to their wild-type (WT) counterparts under the salinity condition. This could be explained by the higher chlorophyll content and the less damage caused by the stress in the photosynthetic machinery of the transgenic plants. According to the analyses, the transformers displayed higher values of maximal photosystem II (PSII) photochemical efficiency (Fv/Fm), maximal quantum yield of primary PSII photochemistry (φPo), quantum yield for electron transport (φEo) and efficiency for electron being transferred from QA- to plastoquinone (ψEo). The evaluation for the overall photosynthetic performance using performance index in absorption basis (PIABS) also supported the differential photosynthesis potential between the two genotypes. Collectively, these findings suggested that appropriate GmCKX13 modulation could enhance photosynthetic adaptation to salt-stressed conditions in soybeans.

Optically detected magnetic resonance and mutational analysis reveal significant differences in the photochemistry and structure of chlorophyll f synthase and photosystem II

In cyanobacteria that undergo far red light photoacclimation (FaRLiP), chlorophyll (Chl) f is produced by the ChlF synthase enzyme, probably by photo-oxidation of Chl a. The enzyme forms homodimeric complexes and the primary amino acid sequence of ChlF shows a high degree of homology with the D1 subunit of photosystem II (PSII). However, few details of the photochemistry of ChlF are known. The results of a mutational analysis and optically detected magnetic resonance (ODMR) data from ChlF are presented. Both sets of data show that there are significant differences in the photochemistry of ChlF and PSII. Mutation of residues that would disrupt the donor side primary electron transfer pathway in PSII do not inhibit the production of Chl f, while alteration of the putative ChlZ, P680 and QA binding sites rendered ChlF non-functional. Together with previously published transient EPR and flash photolysis data, the ODMR data show that in untreated ChlF samples, the triplet state of P680 formed by intersystem crossing is the primary species generated by light excitation. This is in contrast to PSII, in which 3P680 is only formed by charge recombination when the quinone acceptors are removed or chemically reduced. The triplet states of a carotenoid (3Car) and a small amount of 3Chl f are also observed by ODMR. The polarization pattern of 3Car is consistent with its formation by triplet energy transfer from ChlZ if the carotenoid molecule is rotated by 15° about its long axis compared to the orientation in PSII. It is proposed that the singlet oxygen formed by the interaction between molecular oxygen and 3P680 might be involved in the oxidation of Chl a to Chl f.

Exposed anthocyanic leaves of Prunus cerasifera are special shade leaves with high resistance to blue light but low resistance to red light against photoinhibition of photosynthesis.

The photoprotective role of foliar anthocyanins has long been ambiguous: exacerbating, being indifferent to or ameliorating the photoinhibition of photosynthesis. The photoinhibitory light spectrum and failure to separate photo-resistance from repair, as well as the different methods used to quantify the photo-susceptibility of the photosystems, could lead to such a discrepancy. We selected two congeneric deciduous shrubs, Prunus cerasifera with anthocyanic leaves and Prunus triloba with green leaves, grown under identical growth conditions in an open field. The photo-susceptibilities of photosystem II (PSII) and photosystem I (PSI) to red light and blue light, in the presence of lincomycin (to block the repair), of exposed leaves were quantified by a non-intrusive P700+ signal from PSI. Leaf absorption, pigments, gas exchange and Chl a fluorescence were also measured. The content of anthocyanins in red leaves (P. cerasifera) was >13 times greater than that in green leaves (P. triloba). With no difference in maximum quantum efficiency of PSII photochemistry (Fv/Fm) and apparent CO2 quantum yield (AQY) in red light, anthocyanic leaves (P. cerasifera) showed some shade-acclimated suites, including lower Chl a/b ratio, lower photosynthesis rate, lower stomatal conductance and lower PSII/PSI ratio (on an arbitrary scale), compared with green leaves (P. triloba). In the absence of repair of PSII, anthocyanic leaves (P. cerasifera) showed a rate coefficient of PSII photoinactivation (ki) that was 1.8 times higher than that of green leaves (P. triloba) under red light, but significantly lower (-18 %) under blue light. PSI of both types of leaves was not photoinactivated under blue or red light. In the absence of repair, anthocyanic leaves exhibited an exacerbation of PSII photoinactivation under red light and a mitigation under blue light, which can partially reconcile the existing controversy in terms of the photoprotection by anthocyanins. Overall, the results demonstrate that appropriate methodology applied to test the photoprotection hypothesis of anthocyanins is critical.

Hormesis Responses of Photosystem II in Arabidopsis thaliana under Water Deficit Stress.

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.

PCA-based detection of phosphorous deficiency in wheat plants using prompt fluorescence and 820 nm modulated reflection signals.

Phosphorus deficiency induces biochemical and morphological changes which affect crop yield and production. Prompt fluorescence signal characterizes the PSII activity and electron transport from PSII to PSI, while the modulated light reflection at 820 (MR 820) nm investigates the redox state of photosystem I (PSI) and plastocyanin (PC). Therefore, combining information from modulated reflection at 820 nm with chlorophyll a fluorescence can potentially provide a more complete understanding of the photosynthetic process and integrating other plant physiological measurements may help to increase the accuracy of detecting the phosphorus deficiency in wheat leaves. In our study, we combined the chlorophyll a fluorescence and MR 820 signals to study the response of wheat plants to phosphorus deficiency as indirect tools for phosphorus plant status characterization. In addition, we studied the changes in chlorophyll content index, stomatal conductance (gs), root morphology, and biomass of wheat plants. The results showed an alteration in the electron transport chain as a specific response to P deficiency in the I-P phase during the reduction of the acceptor side of PSI. Furthermore, P deficiency increased parameters related to the energy fluxes per reaction centers, namely ETo/RC, REo/RC, ABS/RC, and DIo/RC. P deficiency increased the values of MRmin and MRmax and decreased νred, which implies that the reduction of PSI and PC became slower as the phosphorus decreased. The principal component analysis of the modulated reflection and chlorophyll a fluorescence parameters, with the integration of the growth parameters as supplementary variables, accounted for over 71% of the total variance in our phosphorus data using two components and provided a reliable information on PSII and PSI photochemistry under P deficiency.

Cyanobacterial phycobilisomes as a platform for the stable production of heterologous enzymes and other proteins

Efforts to stably over-express recombinant proteins in cyanobacteria are hindered due to cellular proteasome activity that efficiently degrades foreign proteins. Recent work from this lab showed that a variety of exogenous genes from plants, humans, and bacteria can be successfully and stably over-expressed in cyanobacteria, as fusion constructs with the abundant β-subunit of phycocyanin (the cpcB gene product) in quantities up to 10–15% of the total cell protein. The CpcB*P fusion proteins did not simply accumulate in a soluble free-floating form in the cell but, rather, they assembled as functional (α,β*P)3CpcG1 heterohexameric light-harvesting phycocyanin antenna discs, where α is the CpcA α-subunit of phycocyanin, β*P is the CpcB*P fusion protein, the asterisk denoting fusion, and CpcG1 is the 28.9 kDa phycocyanin disc linker polypeptide (Hidalgo Martinez et al., 2022). The present work showed that the CpcA α-subunit of phycocyanin and the CpcG1 28.9 kDa phycocyanin disc linker polypeptide can also successfully serve as leading sequences in functional heterohexameric (α*P,β)3CpcG1 and (α,β)3CpcG1*P fusion constructs that permit stable recombinant protein over-expression and accumulation. These were shown to form a residual light-harvesting antenna and to contribute to photosystem-II photochemistry in the cyanobacterial cells. The work suggested that cyanobacterial cells need phycocyanin for light absorption, photosynthesis, and survival and, therefore, may tolerate the presence of heterologous recombinant proteins, when the latter are in a fusion construct configuration with essential cellular proteins, e.g., phycocyanin, thus allowing their substantial and stable accumulation.

Sandstorms damage the photosynthetic activities of Haloxylon ammodendron seedlings

Haloxylon ammodendron is a preferred shrub species for buffering against wind and fixing sand in arid sandy areas of northwest China. To determine whether sandstorms cause damage to H. ammodendron seedlings, we investigated the effects of wind-blown sands on the photosynthetic function of H. ammodendron by simulating sand-carrying wind in the wind tunnel. The results showed that photosystem II (PSII) non-photochemical quenching (NPQ) was sensitive to wind erosion, and sustained blowing of sand-carrying winds enhanced NPQ reduction. The rapidly relaxing quenching NPQf made up the majority component of NPQ; the NPQf/NPQ ratio was approximately 64.4% in the sand-free wind group and nearly 56.2% in the sand-carrying wind group. The distribution of the quantum efficiency of the excitation energy indicated that the relative proportions of the quantum yield of PSII photochemistry ΦPSII, the quantum yield of quenching due to light-induced ΦNPQ, and non-light-induced ΦNO were influenced by both wind erosion and light intensity; the sand-carrying wind resulted in a relative decrease in ΦNPQ and a corresponding increase in ΦNO. The maximum quantum efficiency of PSII photochemistry Fv/Fm was relatively stable in the sand-free wind group, with an average value of approximately 0.81. Compared to sand-free wind, sand-carrying winds caused remarkable decreases in Fv/Fm. Light intensity was the main factor affecting ΦPSII, ΦNPQ, and relative electron transport rate (rETR); there was no interaction effect between the duration of sand-carrying wind and light intensity. Taken together, wind-blown sands cause irreversible damage to the photosynthesis of H. ammodendron seedlings, which is the main factor restricting population regeneration in shelterbelts.

Effects of exogenous NO on the growth and photosynthetic fluorescence characteristics of ryegrass seedlings under B[a]P stress

Benzoapyrene (B[a]P) pollution poses a threat to the environment and the food chain and consequently to human health. However, the alleviation of the harmful effects of B[a]P pollution in perennial ryegrass (Lolium perenne L.) by the application of exogenous nitric oxide (NO) has been ignored. Thus, in this paper the effects of exogenous sodium nitroprusside (SNP, a NO donor) on the growth, photosynthetic fluorescence characteristics, and antioxidant enzyme activity of ryegrass exposed to B[a]P stress are investigated. B[a]P stress induced the reduction of the aboveground and belowground dry weights, chlorophyll (a, b), the total chlorophyll contents, the carotenoid content, the net photosynthetic rate (Pn), the intercellular carbon dioxide concentration (Ci), the water use efficiency (WUE), the photosystem II (PSII) potential activity (Fv/F0), the maximum quantum yield of PSII photochemistry (Fv/Fm), the steady-state fluorescence yield (Fs), and the non-photochemical quenching (qN), while enhancement was recorded in response to the foliar spray of SNP at 200 and 300 μmol·L-1 under B[a]P stress. Gray correlation and principal component analyses show that 200 μmol·L 1 of SNP more drastically alleviated the damage caused by B[a]P stress than 300 μmol·L 1 of SNP. The exogenous NO-mediated alleviation of B[a]P toxicity in ryegrass was associated with preserved photosynthetic characteristics and activation of antioxidant enzymes.

Mechanistic Insights on Salicylic Acid Mediated Enhancement of Photosystem II Function in Oregano Seedlings Subjected to Moderate Drought Stress.

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).

Anthropogenic Dusts Influence Leaf Anatomical and Eco-Physiological Traits of Black Locust (Robinia pseudoacacia L.) Growing on Vesuvius Volcano

In the Mediterranean region, some areas of the Vesuvius National Park (southern Italy) are subjected to a severe anthropogenic impact, especially during spring and summer seasons. The continuous trampling of tourists and buses leads to the formation of “dust-clouds”, exposing plants, especially along the paths, to a great deposition of powder particles on leaves. The aim of this study was to analyze if the dust deposition induces changes in leaf morpho-anatomical and eco-physiological traits of the alien, invasive, species Robinia pseudoacacia L., with particular attention to the photosystem II (PSII) efficiency. We selected plants located near the paths with a high deposition of dust (HD) and plants far away from the paths (low deposition, LD), and tested them over three dates along summer. We analyzed PSII photochemistry, photosynthetic pigments content, and leaf functional (e.g., relative water content and leaf dry matter content) and morpho-anatomical traits (e.g., parenchyma thickness, mesophyll density). HD plants presented a more efficient PSII activity, indicated by the higher quantum yield of PSII electron transport (FPSII) (9%) and electron transport rate (ETR) (38%) in the end of July. Dust deposition also reversibly altered photosynthetic pigments concentration and some lamina traits, adjustable in the short-term (e.g., intercellular spaces and phenolics distribution). We hypothesize that HD leaves were shielded by dusts which would protect their photosynthetic apparatus from the excess of light.

Early Drought Stress Warning in Plants: Color Pictures of Photosystem II Photochemistry

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.

Temporal heterogeneity in photosystem II photochemistry in Artemisia ordosica under a fluctuating desert environment.

Acclimation strategies in xerophytic plants to stressed environmental conditions vary with temporal scales. Our understanding of environmentally-induced variation in photosystem II (PSII) processes as a function of temporal scales is limited, as most studies have thus far been based on short-term, laboratory-controlled experiments. In a study of PSII processes, we acquired near-continuous, field-based measurements of PSII-energy partitioning in a dominant desert-shrub species, namely Artemisia ordosica, over a six-year period from 2012-2017. Continuous-wavelet transformation (CWT) and wavelet coherence analyses (WTC) were employed to examine the role of environmental variables in controlling the variation in the three main PSII-energy allocation pathways, i.e., photochemical efficiency and regulated and non-regulated thermal dissipation, i.e., Φ PSII, Φ NPQ, and Φ NO, respectively, across a time-frequency domain from hours to years. Convergent cross mapping (CCM) was subsequently used to isolate cause-and-effect interactions in PSII-energy partitioning response. The CWT method revealed that the three PSII-energy allocation pathways all had distinct daily periodicities, oscillating abruptly at intermediate timescales from days to weeks. On a diurnal scale, WTC revealed that all three pathways were influenced by photosynthetically active radiation (PAR), air temperature (T a), and vapor pressure deficit (VPD). By comparing associated time lags for the three forms of energy partitioning at diurnal scales, revealed that the sensitivity of response was more acutely influenced by PAR, declining thereafter with the other environmental variables, such that the order of influence was greatest for T a, followed by VPD, and then soil water content (SWC). PSII-energy partitioning on a seasonal scale, in contrast, displayed greater variability among the different environmental variables, e.g., Φ PSII and Φ NO being more predisposed to changes in T a, and Φ NPQ to changes in VPD. CCM confirmed the causal relationship between pairings of PSII-energy allocation pathways, according to shrub phenology. A. ordosica is shown to have an innate ability to (i) repair damaged PSII-photochemical apparatus (maximum quantum yield of PSII photochemistry, with F v/F m > 0.78), and (ii) acclimatize to excessive PAR, dry-air conditions, and prolonged drought. A. ordosica is relatively sensitive to extreme temperature and exhibits photoinhibition.

Photosynthetic regulation in response to strontium stress in moss Racomitrium japonicum L.

Strontium (Sr2+) pollution and its biological effects are of great concern including photosynthetic regulation, which is fundamental to environmental responses, especially for bryophytes during their terrestrial adaptation. Alternative electron flows mediated by flavodiiron proteins (FLVs) and cyclic electron flow (CEF) in photosystem I (PSI) are crucial to abiotic stresses moss responses; however, little is known about the moss photosynthesis regulation under nuclide treatment. We measured chlorophyll fluorescence parameters in PSI, photosystem II (PSII) and the P700 redox state, oxidative stress in the moss Racomitrium japonicum under low (5mg/L), moderate (50mg/L) and high (500mg/L) Sr2+ stress level. Moderate and high Sr2+ stress triggered H2O2 and malondialdehyde (MDA) generation, and catalase (CAT) activity increases, which are involved in reactive oxygen species regulation. The significant PSII photochemistry (Fv/Fm), Chla/chlb, Y(I)/Y(II), Y(NA), Y(ND) and ETRI-ETRII decreases at moderate and high Sr2+, and the Y(I), Y(II) decreases at high Sr2+ revealed the photo-inhibition and photo-damage in PSI and PSII by moderate and high Sr2+ stress. The nonphotochemical quenching (NPQ) increased significantly at moderate and high Sr2+ stress, reflecting a heat-dissipation-related photo-protective mechanism in antenna system and reaction centers. Moreover, rapid re-oxidation of P700 indicated that FLV-dependent flows significantly regulated PSI redox state under moderate and high Sr2+ stress. and CEF upregulation was found at low Sr2+. Finally, photosynthetic acclimation to Sr2+ stress in R. japonicum was linked to FLVs and CEF adjustments.

A Hormetic Spatiotemporal Photosystem II Response Mechanism of Salvia to Excess Zinc Exposure.

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.

Fast chlorophyll a fluorescence induction (OJIP) phenotyping of chlorophyll-deficient wheat suggests that an enlarged acceptor pool size of Photosystem I helps compensate for a deregulated photosynthetic electron flow.

The wheat lines affected by a decrease in the leaf chlorophyll content typically experience a biomass loss. A known major problem of the chlorophyll-deficient wheat mutants is their limited prevention of Photosystem I (PSI) over-reduction brought about by an insufficient cyclic electron flow, potentially exposing them to a higher sensitivity to light fluctuations. However, the resistance of some mutant lines against fluctuating light suggests the occurrence of regulatory processes compensating for the defect in cyclic electron flow. In this study, a phenotyping approach based on fast chlorophyll a fluorescence induction (OJIP transient), corroborated by P700 redox kinetics, was applied to a collection of chlorophyll-deficient wheat lines, grown under continuous or fluctuating light. Quantitative parameters calculated from the OJIP transient are considered informative about Photosystem II (PSII) functional antenna size and photochemistry, as well as the functioning of the entire photosynthetic electron transport chain. The mutants tended to recover a wild-type-like chlorophyll content, and mature plants could hardly be distinguished based on their effective PSII antenna size. Nevertheless, specific OJIP-derived parameters were strongly correlated with the phenotype severity, in particular the amplitude of the I-P phase and the I-P/J-P amplitude ratio, which are indicative of a more capacitive pool of PSI final electron acceptors (ferredoxin and ferredoxin-NADP+ oxidoreductase, FNR). We propose that the enlargement of such pool of electron carriers is a compensatory response operating at the acceptor side of PSI to alleviate potentially harmful over-reduced states of PSI. Our results also suggest that, in chlorophyll-deficient mutants, higher FV /FM cannot prove a superior PSII photochemistry and wider I-P phase is not indicative of a higher relative content of PSI.

Paclobutrazol Ameliorates Low-Light-Induced Damage by Improving Photosynthesis, Antioxidant Defense System, and Regulating Hormone Levels in Tall Fescue.

Paclobutrazol (PBZ) is a plant-growth regulator (PGR) in the triazole family that enhances plant tolerance to environmental stresses. Low-light (LL) intensity is a critical factor adversely affecting the growth of tall fescue (Festuca arundinacea Schreb.). Therefore, in this study, tall fescue seedlings were treated with PBZ under control and LL conditions to investigate the effects of PBZ on enhancing LL stress resistance by regulating the growth, photosynthesis, oxidative defense, and hormone levels. Our results reveal that LL stress reduced the total biomass, chlorophyll (Chl) content, photosynthetic capacity, and photochemical efficiency of photosystem II (PSII) but increased the membrane lipid peroxidation level and reactive oxygen species (ROS) generation. However, the application of PBZ increased the photosynthetic pigment contents, net photosynthetic rate (Pn), maximum quantum yield of PSII photochemistry (Fv/Fm), ribulose-1,5-bisphosphate carboxylase (RuBisCO) activity, and starch content. In addition, PBZ treatment activated the antioxidant enzyme activities, antioxidants contents, ascorbate acid-glutathione (AsA-GSH) cycle, and related gene expression, lessening the ROS burst (H2O2 and O2∙−). However, the gibberellic acid (GA) anabolism was remarkably decreased by PBZ treatment under LL stress, downregulating the transcript levels of kaurene oxidase (KO), kaurenoic acid oxidase (KAO), and GA 20-oxidases (GA20ox). At the same time, PBZ treatment up-regulated 9-cis-epoxycarotenoid dioxygenase (NCED) gene expression, significantly increasing the endogenous abscisic acid (ABA) concentration under LL stress. Thus, our study revealed that PBZ improves the antioxidation and photosynthetic capacity, meanwhile increasing the ABA concentration and decreasing GA concentration, which ultimately enhances the LL stress tolerance in tall fescue.